Furcraea Foetida (L.) Haw

United States
Department
of Agriculture
Wildland Shrubs of the
Forest Service United States and Its
International Institute
of Tropical Forestry
San Juan, PR
Territories: Thamnic
Rocky Mountain
Research Station
Descriptions:
Fort Collins, CO
General Technical
Report IITF-GTR-26
Volume 1
July 2004
John K. Francis, Editor
Abstract ______________________________________
Francis, John K. ed. 2004. Wildland shrubs of the United States and its Territories:
thamnic descriptions: volume 1. Gen. Tech. Rep. IITF-GTR-26. San Juan, PR: U.S.
Department of Agriculture, Forest Service, International Institute of Tropical Forestry, and
Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain
Research Station. 830 p.
A discussion of the general characteristics of shrubs as a life form and their distribution
within the United States is followed by 311 short monographs containing general descrip-
tions, ranges, ecology, reproductive habits, growth and management, and benefits to
humans, animals, and the environment.
Keywords: Asteraceae, Rosaceae, Fabaceae, Cactaceae, woody plants, hardwoods, shrubs,

perennials
The U.S. Department of Agriculture, Forest Service, International Institute of Tropical Forestry
operates in cooperation with the University of Puerto Rico, Río Piedras, PR, 00936-4984
——————
The use of trade or firm names in this publication is for reader information and does not
imply endorsement by the U.S. Department of Agriculture of any product or service
Pesticide Precautionary Statement
This publication reports research involving pesticides. It does not

contain recommendations for their use, nor does it imply that the uses
discussed here have been registered. All uses of pesticides must be
registered by appropriate State and/or Federal agencies before they can
be recommended.
CAUTION: Pesticides can be injurious to humans, domestic animals,

desirable plants, and fish or other wildlife—if they are not handled or
applied properly. Use all pesticides selectively and carefully. Follow
recommended practices for the disposal of surplus pesticides and
pesticide containers.
CAUTION:
PESTICIDES
Wildland Shrubs of the United States and Its
Territories: Thamnic Descriptions:
Volume 1
John K. Francis, Editor

Foreword
Shrubs are an important life form that is difficult to circumscribe because of variations in
individual species that may be classified as shrubs in some circumstances but as trees or
herbs in others. Nevertheless the term shrub conjures up an image of a woody plant of
limited stature; usually with mature specimens of one-third to twice human heights and
often with multiple stems. Shrubs are important components in many kinds of habitats
including a role as landscape dominants or icons as well as integrated components of
other vegetation types. Whereas there are numerous national and regional handbooks on
trees and grasses and floras of continental, regional, state, and local scales, handbooks of
shrubs are limited to regional areas or to horticultural, agronomic, ornamental, or wildlife
biology applications. This volume begins an effort to provide important characteristics
for native and naturalized shrubs of the United States and its territories. Editor John
Francis and other contributors provide descriptive, distributional, ecological,
reproductive, growth, management, and use information on more than 300 shrub species
in this volume; in all there are more than 5,000 shrub species in the United States and its
territories. There is more work to do.
I believe that it is appropriate that this effort was undertaken by the USDA Forest Service
Institute of Tropical Forestry and the Rocky Mountain Station’s Shrub Sciences
Laboratory. These institutions represent foci of significant shrub diversity. Shrub
diversity is enhanced both by warm, mesic and tropical conditions and by aridity and
diverse topography. The two institutions—one in the tropics and one in the continental
Western United States desert--represent those contrasting situations.
E. Durant McArthur
Project Leader
Shrubland Biology and Restoration Work Unit
Shrub Sciences Laboratory
Rocky Mountain Research Station
i
Introduction
The evolution of the Wildland Shrub Manual begins a long time ago with the observation and
description of plants; however, its direct ancestor is the U.S. Silvics Manual: “Silvics of North America”
Volumes 1 and 2 (Burns and Honkala 1990a, 1990b) in which the editor participated as a species author.
After this, a regional silvics manual developed at the International Institute of Tropical Forestry was
published as single species’ separate publications and later translated into Spanish and compiled into a
single volume, “Bioecología de Árboles y Exóticos de Puerto Rico y las Indias Occidentales” (Francis and
others 2000). Besides the numerous botanical references cited in the following section and in the species
descriptions, other important references were important contributing sources for this effort. The “Woody-
Plant Seed Manual” (Schopmeyer 1974 and the update in progress, Bonner and Nisley 2003) is cited in
many of the species descriptions. The “Fire Effects Information System” database (Fire Sciences
Laboratory 2003) contributed a significant portion of the information used is a number of the mainland
U.S. species in this volume. The Wildland Shrub Manual differs from its predecessors by the extensive use
of Internet information sources that have not been available previously.
Latin family, genus, species, and authority of plant names were checked against the “Plants”
database (Natural Resources Conservation Service 2003). In a few cases, individual authors disagreed with
the name given in the database and have cited their reasons for using the name in the form they did.
Common names, for which there is really no central authority and which are likely to take another
generation to stabilize, were left to the discretion of the authors. Common names cited were generally
chosen from the many in existence for each species because they were more widely used or because they
were unique or especially descriptive.
Although somewhat simplified, the format of each shrub description generally follows that used in
the silvical descriptions. Sections: General Description, Range, Ecology, Reproduction, Growth and
Management, and Benefits, are followed by References cited. Each description carries a botanical drawing
or a photo to aid in recognition. Because the number of shrubs to be described is so great, it was decided to
make descriptions summaries rather than complete treatises of all information available. Also, to make the
information useful to as wide an audience as possible, authors were encouraged to use wording that is as
simple as possible without sacrificing technical accuracy. Although arranged in alphabetical order by genus
name, a table of contents is provided to make the process of turning to an individual species easier. A
glossary of technical terms and an index of common names are also provided.
These descriptions have been previously published on the Internet (Francis 2003) and the number
of shrub descriptions continues to be added to. The Internet allows revisions of existing descriptions so that
Internet versions of the descriptions may vary somewhat from descriptions contained in this volume.
Volume 1 of the Wildland Shrub Manual contains descriptions of 311 shrub species, a number determined
more by the time permitted for the project than anything else. It is anticipated that the process of compiling
information and describing shrubs will continue and other volumes will follow. The number of shrubs in
the United States and its Territories easily justifies the publication of five or more volumes of descriptions.
I sincerely hope that shrub scientists in the coming years will continue the effort of publishing thamnical
descriptions to facilitate the management of our wildland shrub natural resource.
References Cited
Bonner, F.T., tech. coord. And R.G. Nisley, manag. ed. 2003. Woody plant seed manual. U.S. Department
of Agriculture, Forest Service, Washington, DC. http://wpsm.net/ index.htm. [not paged].
Burns, R.M., and B.H. Honkala, eds. 1990a. Silvics of North America. Vol. 1, Conifers. U.S. Department of
Agriculture, Forest Service, Agriculture Handbook 654. Washington, DC. 675 p.
Burns, R.M., and B.H. Honkala, eds. 1990b. Silvics of North America. Vol. 2, Hardwoods. U.S. Department of
Agriculture, Forest Service, Agriculture Handbook 654. Washington, DC. 877 p.
ii
Fire Sciences Laboratory. 2003. Fire effects information system. U.S. Department of Agriculture, Forest
Service, Rocky Mountain Research Station, Ft. Collins, CO. http://www.fs.fed.us/database/feis/ [not
paged].
Francis, J.K., ed. 2003. Wildland shrubs of the United States and its territories. U.S. General Technical
Report IITF-WB-1. Department of Agriculture, Forest Service, International Institute of Tropical
Forestry, San Juan, PR and Rocky Mountain Experiment Station, Shrub Sciences Laboratory, Provo UT.
http://fs.fed.us/global/iitf/ wildland_shrubs.htm. [not paged].
Natural Resources Conservation Service. 2003. Plants database. U.S. Department of Agriculture,
Washington, DC. http://plants.usda.gov/ [not paged].
Schopmeyer, C.S., tech. coord. 1974. Seeds of woody plants in the United States. Agriculture Handbook
450. U.S. Department of Agriculture, Forest Service, Washington, DC. 883 p.
Acknowledgments
I especially want to thank John A. Parrotta, then Team Leader at the International Institute of
Tropical Forestry, USDA Forest Service, for his appreciation of the need to package species information
for popular benefit and his encouragement to embark on the Shrub Manual project. I must also thank Ariel
E. Lugo, IITF Director, for allowing me to dedicate almost 3 years nearly full time to shrub research and E.
Durant McArthur, Project Leader, Shrub Sciences Laboratory, Rocky Mountain Research Station, USDA
Forest Service, for graciously welcoming me to work at the Shrub Sciences Laboratory and furnishing
considerable financial support to the project. These and some 36 other authors wrote thamnical descriptions
to contribute to the book. More than 100 scientists participated in reviewing the manuscripts before they
were edited. Isaac Nordlund helped with graphics and programming. I owe as much as any other to Louise
Kingsbury, Director of Publishing Services for the Rocky Mountain Research Station, for personally and
promptly editing every one of the manuscripts.
John K. Francis
Research Forester, Retired
iii
Contents
Page Page
Forward and acknowledgements i Caesalpinia bonduc 129
Introduction ii Cajanus cajan 132
Contents iv Callicarpa americana 135
Wildland shrubs 1 Calotropis procera 137
Thamnical Descriptions Capparis indica 140
Abrus precatorius 12 Capraria biflora 142
Acacia farnesiana 14 Capsicum annuum 144
Acacia greggii 16 Capsicum frutescens 147
Acacia retusa 19 Casearia decandra 149
Acacia tortuosa 21 Casearia sylvestris 151
Acer glabrum 24 Cassine xylocarpa 154
Acer spicatum 27 Castela erecta 156
Acnistus arborescens 30 Ceanothus velutinus 158
Aegiphila martinicensis 32 Celastrus orbiculatus 161
Agave missionum 34 Celastrus scandens 164
Allamanda cathartica 36 Celtis reticulata 167
Amyris elemifera 38 Cephalanthus occidentalis 170
Aralia spinosa 40 Cercocarpus intricatus 173
Ardisia elliptica 43 Cercocarpus ledifolius 175
Argythamnia candicans 45 Cercocarpus montanus 178
Artemisia arbuscula 47 Cereus greggii 181
Artemisia bigelovii 50 Cestrum diurnum 184
Artemisia californica 52 Cestrum laurafolium 186
Artemisia cana 57 Chamaebiateria millefolium 188
Artemisia filifolia 60 Chenopodium oahuense 190
Artemisia frigida 63 Chiococca alba 192
Artemisia longiloba 66 Chromolaena geraniifolium 194
Artemisia lugoviciana 69 Chromolaena odoratum 196
Artemisia nova 72 Chrysobalanus icaco 199
Artemisia pygmaea 75 Chrysophyllum oliviforme 201
Artemisia ridgida 77 Chrysothamnus nauseosus 203
Artemisia rothrockii 79 Chrysothamnus parryi 206
Artemisia spinescens 83 Chrysothamnus viscidiflorus 209
Artemisia tridentata 85 Cissampelos pareira 212
Artemisia tripartita 88 Citrus aurantiifolia 214
Arundinaria gigantean 90 Clematis orientalis 217
Atriplex canescens 93 Clibadium erosum 219
Atriplex confertifolia 96 Clidemia cymosa 221
Atriplex corrugata 99 Clidemia hirta 223
Atriplex cuneata 101 Coccoloba krugii 225
Baccharis pilularis 103 Coccoloba microstachya 227
Barleria prionitis 105 Coccoloba uvifera 229
Batis maritima 107 Coffea arabica 232
Bidens menziesii 110 Colubrina arborescens 235
Bixa orellana 112 Comptonia peregrina 237
Bocconia frutescens 115 Conocarpus erectus 240
Bontia daphnoides 117 Corchorus hirsutus 243
Borrichia arborescens 119 Cordia lima 245
Bourreria virgata 121 Cordia polycephala 247
Brickellia californica 123 Cornus sericea 249
Bromelia penguin 125 Coursetia caribaea 252
Brunfelsia lactea 127 Crataegus douglasii 254
iv
Critonia portoricense 257 Jatropha gossypiifolia 399
Crossopetalum rhacoma 259 Juniperus communis 401
Crotalaria lotifolia 261 Juniperus osteosperma 404
Crotalaria spectabilis 263 Kalmia latifolia 407
Cuphea hyssopifolia 266 Kaonophyllon polyodon 411
Curatella americana 268 Krameria ixine 412
Cytisus scoparius 275 Lantana camara 414
Dalbergia ecastaphyllum 278 Lantana involucrata 417
Dasiphora floribunda 280 Larrea tridentata 419
Dasylirion wheeleri 283 Lasiacis divaricata 425
Datura metel 286 Leandra krugii 427
Dasmanthus virgatus 289 Ledum groenlandicum 429
Distictis lactiflora 291 Lepianthes peltata 432
Dodonaea viscosa 293 Lepidospartum burgessii 434
Dryas octopetala 295 Leucaena leucocephala 437
Duranta erecta 299 Lindera melissifolia 440
Encelia farinosa 302 Lonicera japonica 442
Ephedra nevadensis 307 Lotus scoparius 445
Ephedra viridis 310 Ludwigia octovalvis 449
Eriogonum fasciculatum 314 Lupinus arboreus 451
Erithalis fruticosa 319 Lycium pallidum 453
Ernodea littoralis 321 Macfadyena unguis-cati 456
Erythroxylum areolatum 323 Machaonia portoricensis 459
Erythroxylum brevipes 325 Mahonia aquifolium 461
Eugenia foetida 327 Mahonia repens 464
Eugenia pseudopsidium 329 Malvastrum americanum 467
Euonymus alatus 331 Mecranium latifolium 469
Euonymus americanus 333 Melochia nodiflora 471
Euonymus atropurpureus 335 Melochia tomentosa 473
Fallugia paradoxa 337 Miconia impetiolaris 475
Faramaea occidentalis 341 Miconia prasina 477
Forestiera acuminata 343 Miconia racemosa 480
Forestiera segregata 345 Mimosa aculeaticarpa 482
Fouquieria splendens 347 Mimosa arenosa 485
Furcraea foetida 350 Mimosa ceratonia 487
Gesneria pedunculosa 352 Mimosa pigra 489
Gonzalagunia hirsuta 354 Mimosa pudica 492
Gossypium hirsutum 356 Mimulus aurantiacus 495
Gouania lupuloides 359 Mitracarpus portoricensis 500
Grayia spinosa 361 Morella caroliniensis 502
Guettarda elliptica 363 Morinda citrifolia 505
Gundlachia corymbosa 365 Murraya exotica 508
Gutierreza sarothae 367 Myrica gale 511
Gynerium sagittatum 370 Neea buxifolia 514
Hamelia patens 372 Odontonema cuspidatum 516
Helicteres jamaicensis 374 Olyra latifolia 518
Hibiscus pernambucensis 376 Opuntia leptocaulis 520
Holodiscus discolor 379 Palicourea crocea 524
Holodiscus dumosus 382 Parathesis crenulata 527
Hyperbaena laurifolia 384 Parkinsonia aculeata 529
Indigofera suffruticosa 386 Parthenocissus quinquefolia 532
Isocoma tenuisecta 388 Passiflora edulis 535
Itea virginica 391 Paullinia pinnata 538
Ixora coccinea 393 Pavonia spinifex 540
Jacquinea arborea 395 Pennisetum purpureum 542
Jasminum fluminense 397 Penstemon ambiguous 545
v
Philadelphus lewisii 548 Securidaca virgata 689
Philadelphus microphyllus 551 Senecio flaccidus 691
Phoradendron quadrangulare 553 Senecio spartioides 694
Phragmites australis 555 Senna atomaria 696
Physocarpus malvaceus 558 Senna occidentalis 699
Picramnia pentandra 561 Senna polyphylla 702
Piper aduncum 564 Serenoa repens 704
Piper amalago 567 Serjania polyphylla 708
Piper swartzianum 569 Sesbania sericea 710
Piptocoma antillana 571 Sida fallax 712
Pisonia aculeata 573 Sida rhombifolia 714
Pithecellobium unguis-cati 575 Smilax domingensis 717
Pluchea carolinensis 577 Solanum drymophilum 719
Prosopis glandulosa 580 Solanum dulcamara 721
Proustia vanillosma 584 Solanum persicifolium 724
Prunus americana 586 Solanum torvum 726
Prunus emarginata 589 Sorbus scopulina 729
Prunus pumila 591 Spermacoce verticillata 732
Prunus virginiana 594 Strumpfia maritima 735
Psidium guajava 597 Styrax americanus 737
Psorothamnus scoparius 600 Suaeda suffrutescens 739
Psychotria brachiata 602 Suriana maritima 742
Psychotria deflexa 604 Symphoricarpos albus 744
Psychotria microdon 606 Tabebuia haemantha 747
Psychotria nervosa 608 Tamarix chinensis 749
Purshia tridentate 610 Tamonea boxiana 753
Quercus havardii 613 Taxus brevifolia 755
Randia aculeata 617 Taxus canadensis 758
Rauvolfia viridis 619 Tecoma stans 761
Rhododendron macrophyllum 621 Tetrapterys inaequalis 764
Rhododendron maximum 623 Thrinax morrisii 766
Rhus copallinum 625 Toxicodendron radicans 769
Rhus glabra 628 Toxicodendron rydbergii 772
Rhus microphylla 631 Trema lamarckianum 775
Rhus trilobata 634 Trichilia hirta 777
Ribes aureum 637 Trichostigma octandrum 779
Ribes cereum 641 Triumfetta semitriloba 781
Ricinus communis 643 Turbina corymbosa 783
Rondeletia inermis 645 Urena lobata 785
Rondeletia pilosa 647 Urera baccifera 788
Rosa woodsii 649 Vernonia albicaulis 790
Rubus discolor 652 Vernonia borinquensis 792
Rubus parviflorus 655 Vernonia proctori 794
Rubus rosifolius 658 Vernonia sericea 796
Salix arctica 660 Waltheria indica 798
Salix exigua 664 Wedelia reticulata 801
Salix scouleriana 668 Zisiphus obtusifolia 803
Salvia apiana 671 Zuckia brandegeei 806
Salvia mellifera 676 Conclusion
Sassafras albidum 681 Glossary of technical terms 808
Schaefferia frutescens 684 Common name index 812
Schinus terebinthifolius 686 Authors 824
vi
Wildland Shrubs
According to the dictionary (Merriam-Webster Inc 1984), the word “shrub” came
from the Old English scrybb wood, which is akin to the Norwegian skrubbebaer that means
a dwarf hardwood species. The implication seems to be that a shrub is a dwarf tree--not far
from the current usage of the word. The designation of this life form is driven by the need
for a category between trees and herbs.
If a tree is “a woody perennial plant, typically large and with a single well-defined
stem carrying a more or less definite crown” (Ford-Robertson 1971), then a shrub must
necessarily be smaller. Most definitions require that a shrub should have more than one main
stem caused by branching below or above the ground level (Allaby 1994, Viereck and Little
1972). Other frequent qualifications include the need for the plant to be perennial and to be
lignified (woody), at least in some of its parts (Allaby 1994, Ford-Robertson 1971, Orshan
1989). Shrubs are distinguished from herbs in that herbs do not develop persistent woody
tissue above ground (Ford-Robertson 1971). However, Lawrence (1955) admits that the
term shrub is “not subject to precise circumscription”.
1
There are several reasons why shrubs cannot be precisely defined. Many species are
trees in fertile habitat or favorable conditions and shrubs in difficult habitat. Certain species
may grow as shrubs in large portions of their range and become trees in limited areas. Most
large shrubs will produce an occasional individual with a clearly tree-like habit. It is not
possible to define exactly what “large” is (referring to stem size). A number of species
normally develop single stems but only reach stem sizes below that normally associated with
trees. The degree of lignification is another source of ambiguity. Many annual herbs become
quite lignified before dieing, and some perennial shrubs have little or no woody tissue.
Finally, the concept of perennial looses much of its meaning in the humid tropics where a
shrub may complete its life cycle in a year of continuous growth, and an herb may live
without dieback for many years.
There are many subtypes of shrubs including dwarf shrubs, half-shrubs (sub-shrubs),
cushion plants, woody vines (lianes and climbers), scrambling and prostrate forms, woody
parasitic plants, and various forms of succulents and semi-succulents. A description of these
forms can be found in Lawrence (1955), and a discussion of criterion for classification of
shrub forms may be found in Orshan (1989).
The concept of the shrub layer, which is populated by functional shrubs, sidesteps
the ambiguity of classifying species. In this system, the forest is divided into vertical layers
(canopies or horizons). In their simplest form, the layers are: the tree layer, consisting of
trees and tree-sized plants, the shrub layer, including all plants with any diameter at breast
height (d.b.h.) that is less than some arbitrary upper d.b.h. limit such as 5.0 cm, and the herb
layer, which does not reach breast height. The principal components of the shrub layer are
actually young trees. This concept is useful in studies of wildlife habitat, plant ecology, and
biomass and carbon distribution.
Because of their large size, influence on the environment, and considerable economic
value, trees have received most of the attention in forest research and ecological
descriptions. However, shrubs are more important than is generally realized, especially in
terms of biodiversity. Little (1979) lists 679 species of trees native and naturalized in the
continental United States. Another 60 native tree species with an additional number of
naturalized species grow in Hawaii (Little and Skolmen 1989). Puerto Rico and the U.S.
Virgin Islands support 547 native tree species (Little and others 1974) and at least 118
2
species of naturalized exotics (Francis and Liogier 1991). With allowances for species
represented in more than one area, the U.S. and its territories support about 1,300 tree
species. Shrub species in the same area are much more numerous.
Estimating the number of shrub species in the U.S. and its territories is challenging
because the life-form is subject to the perception of authors and compilers and not indicated
or consistently identified in many of the checklists and floras. However, it is possible to
count the shrub species indicated in the floras of several of the states or regions. Alaska has
about 110 species of shrubs (Viereck and Little 1972), California about 800 species
(McMinn 1951), Hawaii about 170 (Degener 1946, Degener 1957, Degener and Degener
1963), Idaho about 160 (Davis 1952), Pennsylvania about 264 (Rhoads and Klein 1993),
Puerto Rico about 540 (Liogier 1985, Liogeir 1988, Liogier 1994, Liogier 1995, Liogier
1997), and tropical Florida about 250 (Long and Lakela 1971). These totals (with
duplication eliminated) plus other miscellaneous sources (Abrams 1940, Abrams 1944,
Abrams 1951, Abrams and Ferris 1960, Benson 1969, Correll and Johnston 1970, Everitt
and Drawe 1993, Great Plains Flora Association 1986, Jones 1975, Morley 1969, Nelson
1996) exceed 2800 species and seems to account for most of the shrub species in the United
States and its territories. The Natural Resources Conservation Service (2003) “Plants”
database recently made it possible to search and list shrub species by region within the
United States and its territories (Puerto Rico and the U.S. Virgin Islands). Although sums for
growth habits (life-forms) are higher than if each species were exclusively assigned to a
single growth habit (eg. Salix exidua Nutt. appears as both tree and shrub), the figures are
instructive, especially for comparing areas, life-forms, and taxonomic divisions. Searching
all plant species that carry the growth habit designates, “shrub” or “sub-shrub,” yielded
5,281 species for the U.S. and territories. There was a great deal of variation between
political divisions (see figure below, not drawn to scale). California and Texas supported the
greatest numbers of species with 1,311 and 1,300 respectively. North Dakota and Alaska
had the lowest with 195 and 200 respectively. The U.S. Virgin Islands had the highest
average shrub biodiversity with 527 shrub species in only 349 km2 of territory.
3
Several factors influence the amount of shrub diversity found within a political unit.
Numbers of shrub species, as well as all plant species, increase as one travels from the harsh
boreal regions to the humid tropics. Diversity of habitat, such as mountain peaks, deserts,
riparian areas, and sea shore, is also very important. Southwestern deserts where shrubs are
the dominant vegetation also tend to have high shrub diversity.
Using searches of the “Plants” database mentioned above, the shrubs of the U.S. and
territories were found to be divided among 166 families. The most important families were:
Asteraceae, 618 species; Rosaceae, 510 species; Fabaceae, 342 species; Cactaceae, 193
species; Ericaceae, 189 species; Scrophulariaceae, 182 species; Rubiaceae, 165 species;
Malvaceae, 148 species; Euphorbiaceae, 128 species; Lamiaceae, 124 species;
Polygonaceae, 123 species; Companulaceae, 112 species; Boraginaceae, 106 species;
Ramnaceae, 103 species; and 152 additional families with 82 to 1 species each.
Reaching or maintaining a high biodiversity is important to ecosystem health.
Managers and the public are beginning to realize that subordinate species, as well as the tree
canopy, are important. It will not be enough, as formerly called for in management plans, to
establish a tree cover and wait for natural succession to fill in the subordinate species.
Shrubs, which are near-climax in ecosystems such as subtropical deserts, are, thus, the
principal species to be maintained or reestablished. In moister wildlands, wildlife managers
4
have long sought to achieve a certain portion of the landscape in the “brush” stage,
preferably in disbursed, irregular patches within more advanced forest.
The use of shrubs as well as all other types of native plants for reclamation and
restoration of damaged sites is becoming a very important topic (Hansen 1989). Shrubs are
planted as seedlings of various types and seeded using the same techniques employed with
trees (Alder and Ostler 1989), except that densities must often be higher. Because it costs
less, establishment by site manipulation and natural seeding and succession is preferred
whenever possible.
Shrubs yield many benefits to humankind directly and indirectly. Berries and similar
small fruits are the most important shrub-derived foods. There are many hundreds of kinds
of edible, wild berrylike fruits throughout the world. Seasonally harvested and preserved,
they were once very important to hunter-gatherer tribes and still are important in certain
rural areas. All our commercial berries descended from wild shrubs, and their wild ancestors
remain a source of genetic material for breeders. Wildland shrubs also furnish nuts, seeds,
herbs, greens, and medicinal materials to rural peoples.
Fuel is another major direct benefit from shrubs. Although wood from shrubs is not
present in quantities as great as tree wood in forests, its accessibility and ease of harvest
have made it a very important fuel source in underdeveloped areas and during recreational
camping, especially when collected by women and children. Shrubs are even harvested to
make charcoal in areas with few trees. It has also been suggested that shrub stands could be
harvested mechanically for industrial biomass fuel (Young and others 1989).
Humans benefit indirectly from shrubs through animals that eat them. Grazing
domestic goats, camels, and, to a lesser extent, sheep and cattle obtain a large part of their
forage from woody browse. Many wild ruminants preferentially browse shrubs even when
grass is available. Others supplement their diet of grasses and forbs with forage from shrubs.
A multitude of birds, mammals, amphibians, and reptiles obtain forage, mast (fruits and
seeds), and insects from shrub hosts. The cover provided by shrubs is critical for a vast
number of wild animals.
5
Finally, wild shrubs are an important esthetic component of our forests, prairies, and
deserts. As greenbelts and semi-wild urban parks become more widespread, wild shrubs will
be more appreciated and will be managed along with trees and herbaceous plants.
Shrubs also negatively impact humans. Undesirable shrubs often invade managed
and semi-managed pastures, excluding the more desirable grass, and become weeds in
croplands. They can temporarily suppress tree seedlings and slow reforestation. Shrubs,
especially the many thorny and a few poisonous species, are the bane of cross-country hikers
and woods workers. Forests with a heavy shrub layer are usually less visually pleasing than
open forests. Shrub understories often are a vehicle for explosive fire spread in seasonally
dry forest habitats.
In the early days of forestry, it seemed to be enough to protect and manage the forest
as a whole. Later, it was realized that we needed detailed information on each of the tree
species. To this end, a number of excellent references have been produced (Burns and
Honkala 1990a, Burns and Honkala 1990b, Burns and others 1998, Francis and Lowe 1999).
The time is coming when, if we are to manage the lesser species, we will need detailed
information on each or, at least, the most important shrubs. The objective of this work is to
6
provide an accessible reference of the biology, ecology, and management of individual
species of shrubs. The study of the biology, ecology, and management of trees is called
silvics, and the resulting descriptions are termed silvical descriptions. No such terms have
come to general use for the study and description of shrubs. Perhaps the terms should be
designated “thamnics” and “thamnical descriptions” from the Ancient Greek “thamnos”,
which means shrub.
References
Abrams, L. 1940. Illustrated flora of the Pacific states. Vol. 1. Stanford University Press,
Stanford, CA. 538 p.
Abrams, L., and R.S. Ferris. 1960. Illustrated flora of the Pacific states. Vol. 4. Stanford
University Press, Stanford, CA. 732 p.
Alder, G.M., and W.K. Ostler. 1989. Native shrub propagation and nursery stock
production. In: C.M. McKell, ed. The biology and utilization of shrubs. Academic Press,
Inc., San Diego, CA. p. 535-552.
Allaby, M. 1994. The concise Oxford Dictionary of ecology. Oxford University Press,
Oxford, UK. 415 p.
Benson, L. 1969. The cacti of Arizona. The University of Arizona Press, Tucson, AZ. 218 p.
7
Burns, R.M., and B.H. Honkala, eds. 1990a. Silvics of North America. Vol. 1, Conifers.
U.S. Department of Agriculture, Forest Service, Agriculture Handbook 654. Washington,
DC. 675 p.
Burns, R.M., and B.H. Honkala, eds. 1990b. Silvics of North America. Vol. 2, Hardwoods.
U.S. Department of Agriculture, Forest Service, Agriculture Handbook 654. Washington,
DC. 877 p.
Burns, R.M., M.S. Mosquera, and J.L. Whitmore, eds. 1998. Useful trees of the tropical
region of North America. Publication 3. North American Forestry Commission,
Washington, DC. 256 p.
Correll, D.S., and M.C. Johnston. 1970. Manual of the vascular plants of Texas. Texas
Research Foundation, Renner, TX. 1881 p.
Davis, R.J. 1952. Flora of Idaho. W.M.C. Brown Co., Dubuque, IA. 828 p.
Degener, O. 1946. Flora Hawaiiensis. Books 1-4. Otto Degener, Waialua, Oahu, HI. [Not
paged].
Degener, O. 1957. Flora Hawaiiensis. Book 5. Otto Degener, Waialua, Oahu, HI. [Not
paged].
Degener, O. and I. Degener. 1963. Flora Hawaiiensis. Book 6. Otto Degener, Waialua,
Oahu, HI. [Not paged].
Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs and cacti of South Texas. Texas Tech
University Press. 213 p.
Ford-Robertson, F.C., ed. 1971. Terminology of forest science, technology, practice, and
products. Society of American Foresters, Washington, DC. 349 p.
8
Francis, J.K. and H.A. Liogier. 1991. Naturalized exotic tree species in Puerto Rico. U.S.
Department of Agriculture, Forest Service, General Technical Report SO-82. Southern
Forest Experiment Station, New Orleans. 12 p.
Francis, J.K. and C.A. Lowe., eds. 1999. Bioecología de especies natives y exóticas de
Puerto Rico y las Indias Occidentales. U. S. Department of Agriculture, Forest Service,
General Technical Report IITF-15. International Institute of Tropical Forestry. Río
Piedras, PR. 583 p.
Great Plains Flora Association. 1986. Flora of the Great Plains. University Press of Kansas.
1391 p.
Hansen, D. 1989. Reclamation and erosion control using shrubs. In: McKell, C. M., ed. The
biology and utilization of shrubs. Academic Press, Inc, San Diego. 459-478.
Jones, F.B. 1975. Flora of the Texas coastal bend. Mission Press, Corpus Christi, TX. 262 p.
Lawrence, G.H.M. 1955. An introduction to plant taxonomy. The Macmillan Company,

New York. 179 p.
Liogier, H.A. 1985. Descriptive flora of Puerto Rico and adjacent islands. Vol. 1. Editorial
de la Universidad de Puerto Rico, San Juan, PR. 352 p.
9
Little, E.L., Jr. 1979. Checklist of United States trees (native and naturalized). U.S.
Department of Agriculture, Forest Service, Agriculture
Handbook 541. Washington, DC. 375 p.
Little, E.L., Jr., R.O. Woodbury, and F.H. Wadsworth. 1974. Trees of Puerto Rico and the
Virgin Islands. Vol. 2. U.S. Department of Agriculture, Forest Service, Agriculture
Handbook 449. Washington, DC. 1,024 p.
Little, E.L., Jr. and R. Skolmen. 1989. Common forest trees of Hawaii (native and
introduced). U.S. Department of Agriculture, Forest Service, Agriculture Handbook 679.
Long, R.W. and O. Lakela. 1971. A flora of tropical Florida. University of Miami Press,
Coral Gables, FL. 962 p.
Merriam-Webster Inc. 1984. Webster’s ninth new collegiate dictionary. Merriam-Webster

Inc., Publishers. Springfield, MA. 1,563 p.
McMinn, H. 1951. An illustrated manual of California shrubs. University of California

Press, Berkeley, CA. 663 p.
Morley, T. 1969. Spring flora of Minnesota. The University of Minnesota Press,

Minneapolis, MN. 283 p.
10
Natural Resource Conservation Service. 2003. Plants database. http://plants.usda.gov/. [not
paged].
Nelson, Gil. 1996. The shrubs and woody vines of Florida. Pineapple Press, Inc., Sarasota,
FL. 391 p.
Orshan, G. 1989. Shrubs as a growth form. In: McKell, Cyrus M., ed. The biology and
utilization of shrubs. Academic Press, Inc., San Diego, CA. p. 249-265.
Rhoads, A.F., and W.M. Klein, Jr. 1993. The vascular flora of Pennsylvania, annotated
checklist and atlas. American Philosophical Society, Philadelphia, PA. 636 p.
Viereck, L.A., and E.L. Little Jr. 1972. Alaska trees and shrubs. Agriculture Handbook 410.
U.S. Department of Agriculture, Forest Service, Washington, DC. 265 p.
Young, J.A., J.D. Budy, and R.A. Evans. 1989. Use of shrubs for fuel. In: C.M. McKell, ed.
The biology and utilization of shrubs. Academic Press, Inc, San Diego, CA. p. 479-492.
11
Abrus precatorius L. crab’s eye
FABACEAE
Synonyms: Abrus abrus (L.) W.F. Wright

Glycine abrus L.
John K. Francis
Ecology.—In Puerto Rico, crab’s eye grows in

subtropical moist (1000 to 2000 mm of
precipitation), subtropical dry forest (below 1000
mm of precipitation), and the lower extreme of the
subtropical wet forest type on all drainage classes
except very poorly drained and on soils of all
textures and parent materials. All types of
topography are colonized from near sea level to
1,000 m in elevation (in India) (Parrotta 2001).
Crab’s eye competes well with weeds and brush in
abandoned farmland, disturbed areas, and early
secondary forest. It requires disturbance to
maintain itself in dense, closed stands. Although
crab’s eye produces relatively few root nodules
compared to other legumes, nitrogenase activity is
comparatively high by nodule weight (Pokhriyal
and others 1997).
Reproduction.—A sample of crab’s eye seed

from Puerto Rico averaged 0.1088 + 0.0091
g/seed. These seeds, which were not treated in any
way, germinated at a final rate of 61 percent
between 11 and 182 days after sowing.
Germination is epigeous. Seeds are produced in
General Description.—Crab’s eye, also known as abundance. Seedlings are common in suitable
jumbie bean and licorice plant in English, peronías habitat, but few gain access to sufficient sunlight
and ojos de cangrejo in Spanish, and guen léglise to survive. Humans have been responsible for the
in French (Howard 1988, Liogier 1988), is a long-distance transport that has resulted in the
slender woody vine that climbs shrubs and low current pantropical distribution. Birds may move
trees. Its older stems are dark gray and the younger the seeds short distances either through curiosity
ones are green and very fine. The alternate, or by being momentarily deceived into thinking
pinnately compound leaves are 5 to 10 cm long that they are edible (personal communication with
and have five to 20 pairs of leaflets. The racemes Joseph Wonderly, IITF, Río Piedras, Puerto Rico).
have tight clusters of white to purple flowers Lateral extension of the vines also disperses the
(Howard 1988, Liogier 1988). The most notable seeds short distances. These plants can be
thing about this species is the 6-mm, spherical red controlled by heavy grazing, hand removal, and
and black seeds. Crab’s eye produces a tap and with herbicides.
lateral root system with abundant fine roots. The
larger roots are dark reddish brown. The lateral Growth and Management.—Seedlings grow at a
roots produce white nodules. moderate rate. It takes about 6 months after
germination for them to reach 30 cm in height.
Range.—Crab’s eye is possibly native of India Mature plants may grow 2 m or more per year in
(Acevedo-Rodríguez 1985) or of Guinea in Africa each leader. They eventually reach 5 to 10 m into
(Neal 1965), but today has naturalized throughout the canopy or a similar distance laterally
most of the tropics (Acevedo-Rodríguez 1985). (Acevedo-Rodríguez 1985). The longevity of
12
crab’s eye is not reported, but certainly it lives Southern Forest Experiment Station, New
longer than 3 years. New plants are easily grown Orleans, LA. 331 p.
in the nursery and probably can be established by
direct seeding into prepared seed spots. Because Bailey, L.H. 1941. The standard cyclopedia of
the species is exotic in all of the U.S. Territories horticulture. Vol. 1. The MacMillan Co., New
and because it becomes a weed in range and York. 1,200 p.
semimanaged pastures (Velez and von Overgeek
1950), it is difficult to see a justification for the Howard R.A. 1988. Flora of the Lesser Antilles,
establishment of plantations. Leeward and Windward Islands.
Dicotyledoneae. Vol. 4. Arnold Arboretum,
Benefits.—Crab’s eye seeds were used anciently Harvard University, Jamaica Plain, MA. 673 p.
as balance weights in the Eastern countries (Neal
1965) and as rosary beads by the Buddhists Liogier H.A. 1988. Descriptive flora of Puerto
(Bailey 1941) because they were so uniform in Rico and adjacent islands, Spermatophyta. Vol.
weight, as well as durable. Today, the seeds are 2. Editorial de la Universidad de Puerto Rico,
used to make necklaces and other jewelry and for Río Piedras, PR 481 p.
the sounding weights in maracas. The seeds
contain a protein-based poison called abrin Ndamba, J., N. Nyazema, N. Makaza, C.
(Parrotta 2001). One seed contains enough poison Anderson, and K.C. Kaondera. 1994. Traditional
to kill a human, but the seeds are very hard and herbal remedies used for the treatment of urinary
must be chewed or ground to release the poison schistosomiasis in Zimbabwe. Journal of
(Acevedo-Rodríguez 1985). However, cooking Ethnopharmacology 42(2): 125-132.
destroys the poison so that the seeds may be eaten
(Neal 1965). The leaves and roots contain Neal, M.C. 1965. In gardens of Hawaii. Spec. Pub.
glycyrrhizin, the principal component of licorice. 50. Bernice P. Bishop Museum Press, Honolulu,
These tissues prepared in various ways are used to HI. 924 p.
treat coughs and a number of other ailments
(Parrotta 2001). Crab’s eye holds out promise in Parrotta, J.A. 2001. Healing plants of Peninsular
the treatment of Schistosoma haematobium India. CAB International, Wallingford, UK.
infections. Extracts of the plant were shown to be 944 p.
lethal to adult schistosomes in hamsters (Ndamba
and others 1994). Crab’s eye vines are sometimes Pokhriyal, T.C., S.P. Chaukiyal, and H.B. Naithani.
grown for vegetative screens (Bailey 1941). 1997. Nodule biomass and nitrogen fixation
studies in some leguminous nitrogen fixing
References plants. Indian Forester 123(12): 1197-1198.
Acevedo-Rodríguez, P. 1985. Los bejucos de Vélez, I. and J. von Overgeek. 1950. Plantas
Puerto Rico. Vol. 1. Gen. Tech. Rep. SO-58. indeseables en los cultivos tropicales. Editorial
U.S. Department of Agriculture, Forest Service, Universitaria, Rìo Piedras, PR. 497 p.
13
Acacia farnesiana (L.) Willd. sweet acacia
FABACEAE
Synonyms: Acacia cavenia Bert.

Acacia leptophylla DC.
Vachellia farnesiana (L.) Wight & Arn.
John A. Parrotta
America, in South America as far south as Chile

and Argentina, and in many parts of the Old World
tropics and subtropics. Present on all continents
between 30° N. and 40° S. latitudes, it is the most
widely distributed species of Acacia (Siegler and
others 1986).
Ecology.—Sweet acacia is a drought-hardy, fire-

resistant species that does not tolerate frost and
grows well in areas receiving between 500 and
750 mm of rainfall with a dry season of 4 to 6
months (Webb and others 1980). Its best growth
occurs on well-drained soils. It tolerates heavy
clays to sands and a variety of soil conditions,
including saline soils, at elevations up to 2,000 m.
A light-demanding species, sweet acacia often
forms dense thickets on disturbed sites and is
associated with numerous other shrub and tree
species in secondary thorn woodlands, shrublands,
General Description.—Sweet acacia, also known and dry forests in its tropical and subtropical
as cassie, aroma, huisache, cambrón, espino American range (Rzendowski 1981). It is
blanco, and many other common names (Little and susceptible to attack by a number of insect
Wadsworth 1964), is a medium-sized shrub with species, leaf, stem, and root pathogens, though
many spreading branches and basal stems. The none appear to pose a serious threat to the species
leaves are alternate, bipinnately compound with (Parrotta 1992).
two to six pairs of pinnae, each with 10 to 25 pairs
of narrow leaflets 3 to 5 mm in length. The slightly Reproduction.—Sweet acacia produces small (to
zigzag twigs are dark brown with light-colored 5 mm in length) flowers that have functional male
dots (lenticels) and paired spines 3 to 20 mm in and female parts, borne in compact rounded heads
length at the nodes. The older bark is also dark 0.6 to 1.3 cm across. The flowers are very fragrant
brown and smooth. Its bright yellow or orange and are pollinated by bees and other insects. The
flowers, produced over a period of 2 to 4 months, thick, slightly flattened pods, 4 to 9 cm in length
depending on locality, are very fragrant and used and 0.5 to 1.3 cm broad, are produced in
in the perfume industry in France and elsewhere. abundance after about 3 years. They mature 4 to 6
months after flowering and contain a number of
Range.—Sweet acacia is believed to be native to hard-coated, brown seeds embedded in a pulpy
the American tropics, although precise information mesocarp. The species flowers and fruits between
is lacking about its range prior to the Spanish November and February in the Caribbean and
colonial era, during which it was introduced to between December and March in Central America
numerous countries throughout the tropics and (Hughes and Styles 1984, Little and Wadsworth
subtropics where it subsequently became 1964). Natural reproduction is abundant,
naturalized. Today sweet acacia is found in the particularly on disturbed sites and in active
southern United States from California to Florida, pastures where cattle readily consume the pods. In
throughout the West Indies, Mexico, Central nurseries, sweet acacia is usually propagated from
14
seed, although branch cuttings can also be rooted vegetation for food production in arid zones.
(Webb and others 1980). A collection of seeds General Technical Report RM-150. U.S.
from Puerto Rico contained 7,600 seeds/kg, began Department of Agriculture, Forest Service,
germinating in 6 days, and achieved 57 percent Rocky Mountain Forest and Range Experiment
germination after 13 days (Francis and Rodríguez Station, Fort Collins, CO. p. 90-98.
1993). Although pregermination treatment is not
necessary, cold- or hot-water soaking, chemical Francis, J.K. and A. Rodríguez. 1993. Seeds of
scarification, and, particularly, seed-coat Puerto Rican trees and shrubs: second
scarification by abrasion with sandpaper greatly installment. Research Note SO-374. U.S.
increase germination rates (Parrotta 1992). Department of Agriculture, Forest Service,
Southern Forest Experiment Station, New
Growth and Management.—Early growth is Orleans, LA. 5 p.
relatively rapid. About 1 m of height growth can
be obtained during the first year, although growth Hughes, C.E. and B.T. Styles. 1984. Exploration
rates of 30 to 50 cm during the first year under and seed collection of multi-purpose dry zone
semiarid field conditions are more typical trees in Central America. International Tree
(Foroughbakhch and others 1987). Depending on Crops Journal 3: 1-31.
the environment, maximum heights of plants
generally range from 3 to 5 m, with stem Liogier, H.A. 1990. Plantas medicinales de Puerto
diameters up to 5 cm. Because sweet acacia is Rico y del Caribe. Iberoamericana de Ediciones,
intolerant of shade and does not compete well with Inc., San Juan, PR. 566 p.
more aggressive woody vegetation such as
Prosopis L., management activities for enhancing Little, E.L., Jr. and F.H. Wadsworth. 1964.
growth and natural regeneration in natural and Common trees of Puerto Rico and the Virgin
plantation stands may include control of Islands. Agriculture Handbook 249. U.S.
competing vegetation and periodic soil Department of Agriculture, Forest Service,
disturbance. Washington, DC. 548 p.
Benefits.—Sweet acacia is planted in many parts Parrotta, J.A. 1992. Acacia farnesiana (L.) Willd.
of its natural and introduced tropical and –Aroma, huisache. Research Note SO-ITF-SM-
subtropical range for reforestation of degraded 49. U.S. Department of Agriculture, Forest
drylands, for fuelwood and small timber, livestock Service, Southern Forest Experiment Station,
fodder, and for its flowers used in the perfume New Orleans. 6 p.
industry. In some areas, it is considered a pest due
to its ability to colonize pastures and other Parrotta, J.A. 2001. Healing plants of peninsular
disturbed habitats. The tannin-rich bark is used for India. CAB International, New York. 944 p.
tanning leather, the gum obtained from the cut
bark is used as a substitute for gum arabic (from Rzendowski, J. 1981. Vegetacíon de México.
Acacia nilotica (L.) Delile), and a useful black dye Editorial Limusa, Mexico City. 432 p.
is obtained from the pods. Various parts of the
plant are used in traditional medicine (Liogier Siegler, D.S., S. Seilheimer, J. Keesy, and
1990, Parrotta 2001). In Mexico, for example, the H.F.Huang. 1986. Tannins from four common
flowers are used to treat headache and indigestion, Acacia species of Texas and northeastern
whereas a decoction of the green pods is used to Mexico. Economic Botany 40: 220-232.
treat dysentery and skin inflammations. In India,
the bark, heartwood, and leaves are all used Webb, D.B., P.J. Wood, and J. Smith. 1980. A
medicinally to treat a variety of ailments (Parrotta guide to species selection for tropical and
2001). subtropical plantations. Tropical Forestry Paper
15. Overseas Development Administration,
References Commonwealth Forestry Institute, University of
Oxford, London. 256 p.
Fourouhbakhch, R., R. Penaloza, and H. Stienen.
1987. Increasing productivity in the matorral of
northeastern Mexico: domestication of ten
native multipurpose tree species. In: Strategies
for classification and management of native
15
Acacia greggii Gray catclaw acacia
FABACEAE
Synonyms: Senegalia greggii (Gray) Britt. & Rose.
Juanita A. R. Ladyman
Range.—Catclaw acacia grows in New Mexico,

Arizona, Nevada, southern Utah, southern
Colorado, southern California, and Texas, and
southwards into the Mexican states of Coahuila,
Chihuahua, Sonora, and Baja California. Variety
greggii is the most widespread. Variety wrightii
grows in parts of western Texas and New Mexico
and in Sonora, Tamaupilas, and Nuevo León,
Mexico.
Ecology.—Catclaw acacia is a drought resistant,

deep-rooted plant that grows between 300 m and
Illustration source: USDA—Forest Service collection, 1,500 m in elevation in a variety of communities
Hunt Institute and soil types, from sandy or gravelly hills and
slopes, to canyon bottoms and along washes and
General Description.—A cacia greggii is streams (Dayton 1931, McAuliffe 1995,
commonly known as catclaw acacia, catclaw, Richardson 1995, Vines 1960). Although most
Texas mimosa, tearblanket, uña de gato and wait- commonly a 1- to 4-m tall shrub, it can grow to an
a-while (Epple 1995). Children sometimes call the 8- to 10-m tall tree, especially in deep, rich soils
plant “Frito” because the wide, flat fruits twist and (USDAFS 1988). It is a frequent member of desert
curl like corn chips. Many of the common names grasslands and is often associated with other
are similar to those applied to Mimosa acacia species and a variety of other shrubs
aculeaticarpa var. biuncifera which it superficially (Taylor and others 1997). It is not typically a
resembles. The genus name, Acacia, is derived dominant member of the community but may
from “akis” that means “thorn” in Greek (Gledhill reach co-dominance in washes (Schmutz and
1992). Catclaw acacia is a deciduous shrub, or others 1992, Uchytil 1990). Catclaw acacia is
small tree, with grayish-green bipinnately more cold tolerant than many acacia species
compound leaves that are up to 7.6 cm long. (Bowers 1993). However, in the northern part of
Catclaw acacia exhibits foliage dimorphism (Isely its range it generally grows no taller than 2 m
1973). The leaves clustered on spurs are small, whereas in its southern range it will grow to nearly
very short petioled, and usually have one to two 7 m. Catclaw acacia is a winter-deciduous shrub
pairs of pinnae whereas the foliage on new growth that grows only when leafy, remaining dormant in
is larger with petioles up to 1 cm long and with the winter and early spring (McClaran 1995).
one to three pairs of pinnae. The short, sharp, Observations suggest that it is a common host to
curved spines on the branches are approximately mistletoe. An ornithological study in Nevada noted
0.64 cm long and resemble cats’ claws (Epple that 67 percent of the catclaw acacia shrubs
1995). Two varieties exist, var. greggii and var. distributed along three 1-km transects were
wrightii (Kartesz 1994). Acacia greggii var. infested with large amounts of mistletoe,
greggii is synonymous with A. greggii var. Phorodendron californicum Nutt. (Blake 1984).
arizonica Isely; A. greggii var. wrightii (Benth.)
Isely, is synonymous with A. wrightii Benth. Reproduction.—Catclaw acacia flowers from
(Kartesz 1994). Variety wrightii can be April through October with the heaviest blooms in
distinguished from var. greggii by its wider April and May (Epple 1995, Everitt and Drawe
legume and larger leaflets. 1993). Variety wrightii flowers as early as March
(Vines 1960). Rain frequently prompts flowering.
The creamy-yellow, fragrant flowers are insect
16
pollinated (Bowers 1993). The fruit is a light Vines 1960). It is a food plant for both adult and
brown to reddish legume often becoming larval butterflies (Taylor and others 1997) and an
constricted between the seeds. The circular seeds important “bee plant” (Dayton 1931). The lac
are 5- to 7-mm in diameter (Isely 1973). The wax- insect, Tachardia lacca, feeds on the sap of
coated seeds delay germination for several years catclaw acacia and produces a resinous substance
and need to be scarified to germinate (Bowers that can be used in making varnish and shellac but
1993, Epple 1995). Regions where catclaw acacia probably not in commercial quantities (Powell
grows are prone to flash floods and the tumbling 1998). The wood has been used for fuel and tool
action of floodwaters in sandy, gravelly washes handles (Epple 1995). It is also in the ornamental
accomplish both seed scarification and seed and landscape trade and is a good hedge plant
dispersal (Bowers 1993). (Taylor and others 1997). It has been used in
projects to re-vegetate degraded land but is
Growth and Management.—Catclaw acacia is a sensitive to some pollutants such as those found in
thicket-forming shrub that has increased in copper mine wastes (Norem and others 1982).
abundance throughout the Southwestern United
States since the 1890s (Bahre 1995). This increase References
is likely influenced if not directly caused by
livestock overgrazing and fire suppression (Bahre Bahre, C.J. 1995. Human impacts on the
1995). Overgrazing has reduced native perennial Grasslands of southeastern Arizona. In: M.P.
grasses that, when healthy and dense, can reduce McClaran and T.R. Van Devender, eds. The
woody shrub seedling establishment and also Desert Grassland. The University of Arizona
provide sufficient fuel to carry a fire. Catclaw is Press, Tucson, AZ. p. 230-264.
susceptible to fire when young, but mature plants
re-sprout from the root crown following fire Blake, J.G. 1984. A seasonal analysis of bird
(Carmichael and others 1978, McPherson 1995). communities in southern Nevada. The
Catclaw is moderately resistant to phenoxy Southwestern Naturalist. 29(4): 463-474.
herbicides, and after only one application the
shrubs tend to re-foliate or re-sprout from the Bowers, J.E. 1993. Shrubs and trees of the
crown (Hibbert and others 1974). Successive Southwest Deserts. Southwest Parks and
herbicide applications kill the plants. Apparently Monuments Assoc., Tucson, AZ. 140 p.
the invasive resinbrush, Euryops multifidus
(Thunb.) DC. out-competes, or is in some other Carmichael, R.S., O.D. Knipe, C.P. Pase, and W.
way deleterious to, catclaw shrubs and may be a W. Brady. 1978. Arizona chaparral: Plant
threat in some parts of its range (McAuliffe 1995). associations and ecology. Research Paper RM-
202. U.S. Department of Agriculture, Forest
Benefits.—Catclaw acacia shrubs provide Service, Rocky Mountain Forest and Range
protective cover and shade for wildlife (Dayton Experiment Station, Fort Collins, CO. 17 p.
1931). Pods, twigs and foliage, especially new
growth, provides browse for a variety of wildlife, Dayton, W.A. 1931. Important western browse
for example jack rabbit, white-tailed and mule plants. Miscellaneous Publication 101. U.S.
deer, and the seeds are eaten by many species of Department of Agriculture, Washington, DC.
small mammals and birds, for example, ground 214 p.
squirrels, western white-winged dove, scaled and
bobwhite quail (Graham 1941, Everitt and Drawe Epple, A.O. 1995. A field guide to the plants of
1993, Taylor and others 1997). The leaves contain Arizona. LewAnn Publishing Company, Mesa,
16 to 20 percent crude protein (Everitt and Drawe AZ. 347 p.
1993). Cattle browse the shrubs especially early in
the spring before the new leaves appear, when the Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs
young green twigs provide some relatively and cacti of South Texas. Texas Tech University
succulent forage for a short period of time, and Press, Lubbock, TX. 231 p.
when other forage is limited (Graham 1941,
USDAFS 1988). The mature foliage and pods are Gledhill, D. 1992. The names of plants.
unpalatable to livestock (Graham 1941, Vines Cambridge University Press, Cambridge, UK.
1960). The fruits were ground into a flour, or meal, 202 p.
called pinole by Native Americans (Epple 1995,
17
Graham, E.H. 1941. Legumes for erosion control Norem, M.A., A.D. Day, and K.L. Ludeke. 1982.
and wildlife. Miscellaneous Publication 412. An evaluation of shrub and tree species used in
U.S. Department of Agriculture, Washington, revegetating copper mine wastes in the south-
DC. 153 p. western United States. Journal of Arid
Environments 5: 299-304.
Hibbert, A.R., E.A Davis, and D.G. Scholl. 1974.
Chaparral conversion potential in Arizona: Part Powell, A.M. 1998. Trees & shrubs of Trans-
I: water yield response and effects on other Pecos and adjacent areas. University of Texas
resources. Research Paper RM-126. U.S. University Press, Austin, TX (First ed. copyright
Department of Agriculture, Forest Service, Big Bend Natural History Assoc.) 498 p.
Rocky Mountain Forest and Range Experiment
Station, Fort Collins, CO. 36 p. Richardson, A. 1995. Plants of the Rio Grande
Delta. University of Texas Press, Austin, TX.
Isely, D. 1973. Leguminosae of the United States: 426 p.
I. Subfamily Mimosoideae. Memoirs of the New
York Botanical Garden 25(1) 1-152. Schmutz E.M., E.L Smith, P.R. Ogden, M.L. Cox,
J.O. Klemmedson, J.J. Norris, and L.C. Fierro.
Kartesz, J.T. 1994. A synonymized checklist of the 1992. Desert grasslands. In: Ecosystems of the
vascular flora of the United States, Canada and World. 8A. Natural Grasslands Introduction and
Greenland. Vol. Checklist. Second Ed. Timber Western Hemisphere. R.T. Coupland, ed.
Press Portland, OR. 622 p. Elsevier, London, UK. p. 337-362.
McAuliffe, J.R. 1995. Landscape evolution, soil Taylor, R.B., J. Zrutledge, J.G. Herrera. 1997. A
formation, and Arizona’s desert grasslands. In: field guide to common south Texas shrubs.
The Desert Grassland. M.P. McClaran and T.R. Texas Parks and Wildlife Press, Austin, TX.
Van Devender, eds. The University of Arizona 106 p.
Press, Tucson, AZ. p.100-129.
Uchytil, R.J. 1990. Acacia greggii. U.S.
McClaran, M.P. 1995. Desert grasslands and Department of Agriculture, Forest Service,
grasses. In: The Desert Grassland. M.P. Rocky Mountain Research Station, Fire Sciences
McClaran and T.R. Van Devender, eds. The Laboratory. Fire Effects Information System.
University of Arizona Press, Tucson, AZ. p. 1- http://www.fs.fed.us/database/feis/ [not paged].
30.
United States Department of Agriculture Forest
McPherson, G.R. 1995. The role of fire in desert Service. 1988. Range Plant Handbook. Dover
grasslands. In: The Desert Grassland. M.P. Publications, Inc. New York. 837 p.
McClaran and T.R. Van Devender, eds. The
University of Arizona Press, Tucson, AZ. p. Vines, R.A. 1960. Trees, shrubs, and woody vines
130-151. of the Southwest. Sixth printing. University of
Texas Press. Austin, TX. 1,104 p.
18
Acacia retusa (Jacq.) Howard zarza brava
FABACEAE
Synonyms: Acacia westiana DC

Acacia riparia authors, not Kunth
Senegalia westiana (DC.) Britt. & Rose
Mimosa retusa Jacq.
Mimosa paniculata West
Acacia guadalupensis DC.
Acacia sarmentosa Griseb.
John K. Francis
petioles and rachises are armed with recurved

spines that protect the plant and facilitate climbing.
The root system consists of many robust laterals
without an apparent taproot. Nodules were not seen
by the author. There may be single or multiple
stems from the ground level, but few branches
occur on the stem until it reaches increased light in
the canopy. The leaves are bipinnately compound
with four to 12 pinnae and 15 to 25 leaflet pairs per
pinnae (Liogier 1988). The inflorescence is a
panicle of small heads. The stamens are white. The
legumes are flat, brown, and 6 to 10 cm long and 1
to 2 cm wide. The seeds are oval, flat, and dark
brown (Howard 1988).
Range.—Zarza brava is native to Hispaniola,

Puerto Rico, the Virgin Islands, the Lesser Antilles,
Trinidad, and northern South America (Howard
1988). It is not known to have been planted or
naturalized elsewhere.
Ecology.—Zarza brava grows in habitat that

receives from as little as 700 mm to as much as
2200 mm of annual rainfall in Puerto Rico, and is
more common in the mid-range of this rainfall.
Soils from all parent materials, including limestone
and ultramafic rocks, are colonized. Soil textures
from sand to clays and pH’s from at least 5.0 to 8.0
General Description.—Zarza brava is also known are tolerated. Zarza brava can survive under a
as acacia zarza, amourette, and fleur du ben-aimé broken overstory. When mats are formed, only the
(Acevedo-Rodríguez 1985, Howard 1988, Liogier upper surface stems produce leaves. The species is
1988). The common name zarza brava is Spanish. more common in disturbed habitat than in remnant
The modifier “brava” means fierce, which aptly old growth forests. During the abandonment of
describes the way its thorns grab a person trying to agricultural land, it invades during the brushy
walk through it. Usually, zarza brava is a climber, pasture stage and becomes very common in early
but when open-grown, it scrambles or grows secondary forest. After the canopy closes, zarza
upright. It is also reported to occasionally become a brava declines in importance, but does not
slender tree (Howard 1988). The stems are gray, disappear.
squarish when young and angular or fluted when
older. The old and young stems and even the leaf Reproduction.—Zarza brava flowers in June and
19
December (Acevedo-Rodriguez 1985) and ripens scrambling patches in overgrazed cattle pasture. On
fruit in January and July. Fruit production can be the other hand, the species is a great nuisance to the
very heavy, and pods in one sample in Puerto Rico cattleman, woods workers, and hikers. It is
averaged 7.97 + 0.29 seeds/pod. The seeds frequently necessary to cut a path through zarza
averaged 0.041 + 0.001 g/seed or 24,000 seeds/kg. brava thickets with a machete, in order to move
Some of the seeds had been attacked by an through vegetation in the semiarid-zones within the
unknown species of beetle and a lepidopteran zarza brava range. When one contacts the vines,
larva. When planted in commercial potting mix, the the tips of the spines break off in the skin, causing
seeds began germinating in 4 days and by 11 days pain and sometimes resulting in infection. No
had reached their final 98 percent germination. specific control recommendations have been
Germination is epigeal. In the wild, seedlings are developed. In the past, control was attempted with
produced in abundance, but few survive beyond the mixed success by repeated cutting. Control could
first year. Zarza brava sprouts vigorously and probably be improved by treating the sprouts with
grows rapidly after fires or cutting. herbicides after cutting.
Growth and Management.— Seeds are dispersed References

by lateral extension of the vines, by pods tumbling
laterally through the air after becoming detached Acevedo-Rodríguez, P. 1985. Los bejucos de
from vines, and by accidental dispersal by grazing Puerto Rico. Vol. 1. General Technical Report
animals and humans. New seedlings make SO-58. U.S. Department of Agriculture, Forest
relatively rapid early growth. Older plants also add Service, Southern Forest Experiment Station,
length rapidly. The stems of zarza brava may New Orleans, LA. 331 p.
eventually reach as much as 10 cm in diameter and
as much as 12 m into tree canopies (Acevedo- Howard R.A. 1988. Flora of the Lesser Antilles,
Rodríguez 1985). No planting experience of zarza Leeward and Windward Islands.
brava is known, but if establishment of plants is Dicotyledoneae, Part 1. Vol. 4. Arnold
needed, doubtless the planting of potted nursery Arboretum, Harvard University, Jamaica Plain,
seedlings would succeed. The species is so MA. 673 p.
aggressive that direct seeding into prepared seed
spots in forest openings should also give good Liogier H.A. 1988. Descriptive flora of Puerto
results. Rico and adjacent islands, Spermatophyta. Vol.
2. Editorial de la Universidad de Puerto Rico,
Benefits.—Zarza brava is eaten by goats, but Río Piedras, PR. 481 p.
seldom touched by cattle. In fact, it survives as
20
Acacia tortuosa (L.) Willd. twisted acacia
FABACEAE
Synonyms: Mimosa tortuosa L.

Poponax tortuosa (L.) Raf.
Acacia parvifolia Duss
John K. Francis
do not open. The ovoid seeds are brown and hard
(Howard 1988, Liogier 1988, Little and others
1974). The chromosome number is 2n = 26 (Long
and Lakela 1971).
Range.—Twisted acacia is native to Jamaica,

Hispaniola, Puerto Rico, the Virgin Islands, the
Lesser Antilles, Trinidad, and in South America,
Venezuela, Colombia, and Ecuador including the
Galápagos Islands (Howard 1988, Instituto
Humboldt 2002, Liogier 1988, Little and others
1974). It is endangered in Florida where it may be
native or introduced (Little and others 1974,
Institute of Systematic Botany 2002). Although
several reputable sources list Texas, Mexico, and
Central America as being part of the A. tortuosa
range (e.g., Missouri Botanic Garden 2002, Nature
Conservancy 2002), Little and others (1974) and
Texas A & M Herbarium (2002) maintain that this
western population is in fact A. schaffneri (S.
Wats.) Herm.
Ecology.—Twisted acacia is intolerant of shade

and only moderately competitive. It finds
conditions for establishment and growth to
General Description.—Twisted acacia, also reproductive age in disturbed, dry areas.
known as acacia-bush, wild-poponax, sweet-briar, Disturbance since European colonization has been
cossie, aromo, cají torcido, and pompons jaunes, is by heavy grazing, fires, excessive timber harvest,
a deciduous shrub or small tree to 6 m in height and clearing for farming or pasture improvement.
and 15 cm in trunk diameter. The plant may have One of the most common plants of disturbed areas
single or multiple stems with gray-brown, finely in Curaçao is twisted acacia (Carambi Foundation
fissured bark and a light brown inner bark with 2002). It is found on shell mounds and along
darker streaks. The wood is light brown and hard. roadsides in Southern Florida (Nelson 1996). The
The twigs are slender with a zig-zag profile, species grows in coastal areas of Puerto Rico and
produce paired, straight spines at the nodes, and its offshore islands in areas that receive from 750
have whitish lenticels on the reddish-brown bark. to about 1100 mm of precipitation. The plant
Mature twisted acacia are flat topped and have a colonizes well-drained soils of the full range of
thin crown of feathery foliage. The bipinnately textures derived from both sedimentary and
compound leaves are 2.5 to 9 cm long with two to igneous rocks. Most of these soils are neutral to
eight pairs of pinnae and 10 to 20 pairs of leaflets mildly alkaline. Twisted acacia is not damaged by
per pinnae. Solitary or small clusters of heads of mild salt spray. It is usually ignored by cattle and
tiny, yellow-orange flowers are borne at the leaf tolerates browsing by goats.
axiles. Dark brown or black semiwoody, 8- to 14-
cm pods (legumes) are straight or slightly curved Reproduction.—Twisted acacia flowers
and somewhat constricted between the seeds. They intermittently throughout the year (Little and others
21
1974). Bees and other insects pollinate the flowers
(Treeguide 2002). A collection of pods from Carambi Foundation. 2002. Vegetation. http://
Puerto Rico contained an average of 12.4 www.curacao.com/carambi/veget.htm. 1 p.
seeds/pod and a range of eight to 15 seeds/pod. The
air-dried seeds separated from them averaged
0.0440 + 0.0004 g/seed or 22,700 seeds/kg. These Howard R.A. 1988. Flora of the Lesser Antilles,
were mechanically scarified (nicked) and incubated Leeward and Windward Islands.
on moist blotter paper. Ninety-nine percent Dicotyledoneae, Part 1. Vol. 4. Arnold
germinated between 5 and 21 days after sowing Arboretum, Harvard University, Jamaica Plain,
(author’s observation). Because of the hard, MA. 673 p.
impermeable seed coat, germination without
scarification is low and erratic. Any technique that Institute of Systematic Botany. 2002. Atlas of
perforates the seed coat will work. One source Florida Plants: Acacia tortuosa. University of
(Treeguide 2002) recommends placing the seeds in South Florida, Tampa, FL.
boiling water, immediately removing the heat http://plantatlas.usf.edu/main.asp?plantID=1545.
source and letting the seeds soak for 24 hours. 1 p.
Germination is epigeal. The seeds are dispersed by
herbivores, particularly ruminants that eat the pods. Instituto Humboldt. 2002. Plantas vasculares:
Seedlings often emerge in large numbers, but angiospermas y gimnospermas. http://www.
nearly all are consumed by grazing animals before humboldt.org.co/diqadeladiversidad/resultados/v
they have a chance to become woody and thorny. asculares.html. 6 p.
Growth and Management.—Although acacias Island Resource Foundation. 2002. Threatened and
can be propagated with cuttings, the use of seeds is endangered birds of the Insular Caribbean.
recommended (Bonsai Clubs International 2002). http://www.irf.org/bbarden.htm. 13 p.
No management experience has been published;
however, nursery production and field Liogier H.A. 1988. Descriptive flora of Puerto
establishment will probably be as easy as it is for Rico and adjacent islands, Spermatophyta. Vol.
other acacias. Control of twisted acacia along with 2. Editorial de la Universidad de Puerto Rico,
other secondary dry-forest species has been Río Piedras, PR. 481 p.
accomplished in several areas in Puerto Rico and
the U.S. Virgin Islands by bulldozing. The use of Little, E.L., Jr., R.O. Woodbury, and F.H.
heavy disks might be a better option where Wadsworth. 1974. Trees of Puerto Rico and the
removal of woody debris is not required. Virgin Islands. Vol. 2. Agriculture Handbook
449. U.S. Department of Agriculture,
Benefits.—Twisted acacia helps protect the soil in Washington, DC. 1,024 p.
its disturbed habitat. It is listed as a nitrogen-fixing
species (Winrock International 2002). The seeds Long, R.W. and O. Lakela. 1971. A flora of
are eaten by the threatened yellow-shouldered tropical Florida. University of Florida Press,
amazon parrot (Amazona barbadensis) in Bonaire Coral Gables, FL. 962 p.
and probably other parts of its range (Island
Resource Foundation 2002). The species serves as Missouri Botanical Garden. 2002. Current
cover for wildlife species. Cattle and other specimen list for Acacia tortuosa.
livestock eat the pods while goats and sheep http://mobot.mobot.org/cgi-bin/search_vast. [not
browse the foliage, especially when other forage is paged].
scarce. The wood is used to a limited extent for
fuel. Twisted acacia plants are used to create Nature Conservancy. 2002. The history of the
bonsai plants (Bonsai Clubs International 2002). Plains and its biota.
The species has been grown and pruned for hedges http://www.greatplains.org/resource/biodiver/bio
(Little and others 1974). stat/ecoregio.htm. 13 p.
References Nelson, G. 1996. The shrubs and woody vines of

Florida. Pineapple Press, Inc., Sarasota, FL.
Bonsai Clubs International. 2002. Acacia—Acacia 391 p.
sp. http://www.bonsai-bci.com/species/acacia.
html. 3 p.
22
Texas A&M Herbarium. 2002. Checklist of the Species.asp?SpeciesID=108&Region=NorthAme
vascular plants of Texas. http://csdl.tamu.edu/ rica. 2 p.
FLORA/cgi/msproj/shao/query.pl?query=Acacia
+tortuosa &rform=. 1 p. Winrock International. 2002. Nitrogen fixing trees
and shrubs. http://winrock.org/forestry/factpub/
Treeguide. 2002. Huisachillo, Acacia tortuosa, nftlist.htm. 29 p.
Willd. Fabaceae. http://treeguide.com/nn/
23
Acer glabrum Torr. Rocky Mountain maple
ACERACEAE
Synonyms: Acer tripartitum Nutt.
John K. Francis
Range.—Rocky Mountain maple occurs from

southern Alaska south into northern Mexico and
east from the Pacific Coast to Alberta, South
Dakota and Nebraska (Anderson 2001). There are
currently six recognized varieties: var. diffusum
from the Pacific Southwest, var. douglasii from
the Pacific Northwest, var. greenei from
California, var. glabrum from the inland West, var.
meomexicanum from the inland Southwest, and
var. torreyi from Oregon, California, and Nevada
(Natural Resources Conservation Service 2003).
They differ at least in leaf shape, size, and
thickness.
Ecology.—Rocky Mountain maple is moderately

shade tolerant. It can grow (slowly) in the
understory and midstory of closed coniferous
forest but grows in full sun as well. A pioneer in
disturbed areas, it starts from sprouts and seeds in
burned areas, clearcuts, avalanche fields, and road
sides, cuts and fills. It may grow along streams in
drier areas. Rocky Mountain maple is a member of
General Description.—Rocky Mountain maple, a huge number of plant associations, and is
also known as mountain maple and Douglas frequently dominant during seral stages of
maple, is an upright, deciduous shrub or small tree, succession (Anderson 2001, Johnson 1995). The
that ranges in height from 2 to 10 m, depending on species grows in soils derived from most rock
variety and habitat. It usually grows with multiple types, in all soil textures, and in skeletal soils.
stems, suckering from the base, and narrow These are moist, well-drained to somewhat poorly
crowns, and may be spindly in closed stands. The drained soils that are moderately acid to slightly
stem bark is thin, smooth, and gray, brown, or alkaline. It may be found at elevations from 350 m
reddish brown. Branches are opposite and sparse in British Colombia to 3,900 m in Colorado and
with reddish twigs. Roots are woody, extensive, Utah. Rocky Mountain maple is slightly more
and often deep. Leaves vary considerably, common in southern than northern aspects. Some
depending on variety. They are opposite, glabrous, of its locations have mean annual precipitation as
and have three to five lobes and rarely three low as 230 mm and as high as 1,500 mm
distinct leaflets, and range from 2 to 10 cm wide (Anderson 2001). It tolerates competition well,
and long. The margins are serrate or doubly usually outgrowing and overtopping young trees
serrate. They may be papery or leathery, and most shrub species. Dull red leaf spots are
depending on location and variety. Rocky caused by an unknown pathogen and bright red
Mountain maple occurs as monoecious and eriophyid mite colonies develop on the underside
dioecious trees that have small greenish flowers of leaves in some areas. The effects of these and
borne in loose, few-flowered terminal corymbose other diseases appear to be negligible to
cymes. Fruits are light brown (when mature) populations of the species.
double samaras that diverge at angles less than 45°
with wings about 2 cm long. (Abrams 1951, Reproduction.—Flowering occurs in early spring,
Anderson 2001, Wasser 1982, Welsh and others depending on the elevation and latitude (Wasser
1987). 1982). Fruits mature in late summer or early
24
autumn. Good seed crops occur every 1 to 3 years. shoots may be cooked and eaten as a vegetable
There are about 30,000 cleaned seeds/kg (Zasada (Pratt and others 2003). Native Americans made
and Strong 2003). Seeds disperse by whirling infusions and decoctions of the twigs and wood to
sideways as they descend, often carried by the treat swelling and nausea, respectively (Moerman
wind. Seeds sometimes lie dormant for one or two 1986). Rocky Mountain maple is widely used in
growing seasons before germinating. Germination environmental restoration projects within its native
is epigeal. Natural seedlings can be common to range and is an important ornamental species
abundant. valued for its petite size, tolerance to shade, and
bright red, yellow, and orange autumn colors
Growth and Management.—Rocky Mountain (Hansen 2003).
maple seedlings grow about 30 cm/year (Hansen
2003). On good sites sprouts may reach 1.3 m References
within 2 years and 3 m in 10 years. Maximum
heights are reached in 30 to 40 years (Anderson Abrams, L. 1951. Illustrated flora of the Pacific
2001). Fruits can be picked by hand from low States. Vol. 3. Stanford University Press,
shrubs or clipped or shaken from tall shrubs and Stanford, CA. 866 p.
small trees. Mechanical dewinging may be
advantageous to reduce bulk and improve handling Anderson, M.D. 2001. Acer glabrum. In: U.S.
before planting. The seeds can be stored for up to Department of Agriculture, Forest Service,
3 years (Hansen 2003). Stratification of 180 days Rocky Mountain Research Station, Fire Sciences
at 20 to 30 °C followed by 180 days at 3 to 5 °C is Laboratory, Fire Effects Information System.
recommended before sowing. Germination of http://www.fs.fed.us/database/feis/plants/shrub/
about 40 percent can be expected in about 30 days acegla/all.html. 62 p.
(Zasada and Strong 2003). Alternately, seeds are
sown after collection and beds are maintained for 2 Hansen, W.W. 2003. Native plants of the
years because a substantial amount of the Northwest: Acer glabrum (Douglas maple,
germination will occur after the second winter. Rocky Mountain maple, mountain maple).
Seeds are sown 0.6 to 2.5 cm deep and mulched in http://www.nwplants.com/plants/trees/aceraceae
shaded beds. Seedlings are produced as bare-root /acer_glabrum/ 5 p.
stock or containerized plants (Zasada and Strong
2003). The species can be propagated by layering Johnson, F.D. 1995. Wild trees of Idaho.
and rooted cuttings (Hansen 2003). University of Idaho Press, Moscow, ID. 212 p.
Benefits.—Rocky Mountain maple is an important Moerman, D.E. 1986. Medicinal plants of Native
part of the forest understory and pioneer America. Technical Reports 19. The Museum of
vegetation following fires and clearcuts. It helps Anthropology, University of Michigan, Ann
protect the soil, adds beauty to the forest, and Arbor, MI. 534 p.
furnishes food and cover for wildlife. It is an
important browse species for domestic livestock, Natural Resources Conservation Service. 2003.
especially sheep, and wild ungulates. Samples of Plants profile: Acer glabrum Torr., Rocky
summer growth contained about 8.7 percent crude Mountain maple. http://plants.usda.gov/
protein, 3.1 percent fat, 34.0 percent crude fiber, cgi_bin/plant_profile.chi?symbol=ACGL. 5 p.
51.0 percent N-free extract, and 3.2 percent ash.
Samples of winter browse contained 5.9 percent Pratt, M., J. Bowns, R. Banner, and A. Rasmussen.
crude protein, 2.4 percent fat, 33.3 percent crude 2003. Range plants of Utah: Rocky Mountain
fiber, and 54.2 percent N-free extract (Anderson maple. Utah State University Extension.
2001). The seeds and vegetative parts are http://extension.usu.edu/coop/natres/range/
consumed by ruffed and blue grouse, grosbeaks, Woody/rockymtnmaple.htm. 4 p.
and small mammals (Anderson 2001). The wood is
hard, close-grained, and flexible. It was used by Wasser, C.H. 1982. Ecology and culture of
Native Americans for arrow shafts, snowshoe selected species useful in revegetating disturbed
frames, ceremonial equipment, hut frames, and lands in the West. FWS/OBS-82/56. Fish and
drying and smoking racks. The inner bark Wildlife Service, U.S. Department of Interior,
furnished material for mats and ropes. Today, the Washington, DC. 347 p.
wood is mainly used for firewood (Hansen 2003)
and smoking meat (Johnson 1995). The young
25
Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. Zasada, J.C. and T.F. Strong. 2003. Acer L. In:
Higgins (eds.). 1987. A Utah flora. Great Basin F.T. Bonner, and R.G. Nisley, eds. Woody Plant
Naturalist Memoirs 9. Brigham Young Seed Manual. U.S. Department of Agriculture,
University, Provo, UT. 894 p. Forest Service, Washington, DC. 24 p.
26
Acer spicatum Lam. mountain maple
ACERACEAE
Synonyms: none
Paula M. Pijut
lesser extent. Staminate flowers drop soon after the

pollen is shed (De Jong 1976). Each flower is
borne on a slender stalk about 1.3 cm long and the
panicles are found at the branchlet tips. The bark
is thin, brown to reddish-brown, smooth,
becoming furrowed or warty with age (Dirr 1998).
The fruit is a 2.5 cm long, two-winged samara,
bright red when mature, with divergent wings at an
acute or right angle. The chromosome number of
mountain maple is 2n = 26 (Gleason and Cronquist
1991).
Range.—Mountain maple occurs from

Newfoundland to Saskatchewan, south to
Connecticut, Pennsylvania, Ohio, and northeastern
Iowa, and in the mountains to North Carolina and
Tennessee (Gleason and Cronquist 1991).
Illustration source: USDA—Forest Service collection,
Hunt Institute
Ecology.—Mountain maple prefers rich, moist,
General Description.—Mountain maple is a tall, acid, well-drained soils with low, diffuse light
deciduous shrub or clumped small tree that grows (Dirr 1998; Sullivan 1993). These sites include
3 to 10 m tall and can spread to form dense cool woods with humid climate and year-round
thickets (Dirr 1998; Gleason and Cronquist 1991; precipitation, rocky slopes and flats, along
van Gelderen and others 1994). In the northern streams, in ravines, and on moist hillsides.
part of its native range mountain maple reaches a Mountain maple also grows well on podzol soils,
maximum height of 6 m (Sullivan 1993). Leaves talus slopes, and in forested bogs (Sullivan 1993).
are opposite, simple, three-lobed or obscurely five- Mountain maple can form a canopy on cliff faces.
lobed, coarsely and irregularly serrate, dark Mountain maple is an understory or subcanopy
yellowish-green, glabrous with impressed veins shrub that is tolerant of deep shade, but also grows
(adaxial), lighter green and pubescent (abaxial), well in full sun, has medium moisture and fertility
truncate to cordate at the base, and 5 to 13 cm long requirements, and colonizes the understory as
and wide (Dirr 1998; Gleason and Cronquist 1991; pioneer tree species decline (Sullivan 1993;
van Gelderen and others 1994). The leaves turn USDA, NRCS 2002). Mountain maple is not salt
yellow, orange, or most frequently brilliant red in tolerant, and has low drought tolerance. Release
the fall. Stems are grayish pubescent when young, of mountain maple by canopy tree harvest allows
developing a purplish-red color on the upper the shrub to dominate the site within 5 to 10 years,
surface. Leaf scars are narrowly crescent shaped suppressing the growth of spruce and fir seedlings
(Dirr 1998). Buds are approximately 0.6 cm long, for at least 35 years (Sullivan 1993). In
stalked, pointed, red with minute, appressed, undisturbed, mature red pine-white pine sites in
grayish hairs, and with two to four valvate scales northeastern Minnesota, mountain maple forms a
(Dirr 1998; Gleason and Cronquist 1991). The dense, high shrub layer with beaked hazel and
flowers are small, greenish yellow, in fascicles of American hazel that inhibits reproduction of later
two to four, borne in erect 3 to 8 cm long panicles. successional species, such as balsam fir and spruce
Each flower is functionally either pistillate or (Sullivan 1993). Eradication of mountain maple is
staminate, although the non-functioning structures sometimes necessary for tree regeneration. The
still exist in the flower, reduced to a greater or weevil, Phyllobius oblongus L., is especially
27
damaging to mountain maple (Johnson and Lyon 1993). Propagation of mountain maple by cuttings
1991). The larvae feed on the roots, mature, and taken early in the summer produces strong plants
overwinter in the soil. Adults are brown weevils (van Gelderen and others 1994).
that defoliate mountain maple. They are found
throughout the northeast into Michigan and the Growth and Management.—Mountain maple is a
border provinces of Canada (Johnson and Lyon slow to medium growing shrub or small, short
1991). Kabatiella apocrypta (Ell. & Ev.) v. Arx (crooked) trunked tree of bushy appearance (Dirr
(anthracnose pathogen) causes minor damage to 1998). It is hardy in zones three to seven at high
mountain maple, although severe infections can elevations (USDA Plant Hardiness), and starts to
led to premature defoliation (Sinclair and others decline at 40 to 50 years of age (Sullivan 1993).
1987). Mountain maple is also susceptible to Mountain maple is girdled at the root collar by
attack by Verticillium dahilae Kleb. (verticillium low, surface fires, and can re-sprout from
wilt), Phyllactinia guttata (Wallr.:Fr.) Lev. underground stems (Sullivan 1993). A severe, hot
(powdery mildew), Nectria sp. (nectria canker), fire is necessary in order to kill the roots.
Cristulariella depraedens (Cooke) Hohn. (leaf Mountain maple can withstand repeated and heavy
spot/blight), Pseudomonas syringae pv. syringae browsing, producing new growth even when 80
van Hall (bacterial leaf spot and dieback), percent of the annual twig growth is removed each
Phyllosticta minima (Berk. and M.A.Curtis) year (Sullivan 1993). Mountain maple is an
Underw. and Earle (leaf spot), Rhytisma sp. (tar understory or subcanopy component in a number
spots), and Venturia acerina Plakidas ex Barr (leaf of northeastern forest associations (Sullivan 1993).
blotch) (Farr and others 1989, Jones and Benson Cutting mountain maple near the ground level,
2001, Sinclair and others 1987). clear-cutting, and spraying with 2,4-D in the
spring will increase the availability of new growth
Reproduction.—Mountain maple staminate for deer and moose browse (Sullivan 1993).
flowers and pistillate flowers occur on the same Burning can be used to suppress mountain maple
plant. Some individuals may flower completely in order to promote reproduction of other, more
with staminate flowers. Mountain maple is desirable tree species.
predominately duodichogamous (producing
flowers in three consecutive phases, male-female- Benefits.—Mountain maple is a nutritious browse
male, during anthesis), with a small part of the species for moose, white-tailed deer, cottontails,
population protogynous (stigma receptive before snowshoe hares, woodland caribou, beavers, and
the pollen is shed) (van Gelderen and others 1994). ruffed grouse (Sullivan 1993). Mountain maple is
Low temperatures appear to favor development of sometimes planted as an ornamental, but has little
female flowers and flowers with less reduction of landscape value excluding its brief fall color
the non-functioning parts. (van Gelderen and display and colorful samaras.
others 1994). The shrub blossoms in May to June
after the leaves are fully developed; fruit ripens in References
September to October, with seed dispersal in
October to December (Zasada and Strong 2000). De Jong, P.C. 1976. Flowering and sex expression
Mountain maple is insect pollinated (Sullivan in Acer L. A biosystematic study. Mededelingen
1983) and the seed is wind disseminated. Seeds of Landbouwhogeschool Wageningen 76 (2): 1-
mountain maple need to be stratified for 90 to 120 202.
days at 5 ºC for good germination (Dirr 1998).
Mountain maple does not build up a seed-bank and Dirr, M.A. 1998. Manual of woody landscape
seedling establishment is better on undisturbed plants, Their identification, ornamental
soils (Sullivan 1993). Seedling reproduction of characteristics, culture, propagation, and uses.
mountain maple does not result in dense, Stipes Publishing, Champaign, IL. 1,187 p.
competitive stands, as can be the case with its
vegetative reproduction. Mountain maples Farr, D.F., G.F. Bills, G.P. Chamuris, and A.Y.
frequently layer, giving them the potential to Rossman. 1989. Fungi on plants and plant
develop relatively complex clones of varying size products in the Untied States. The American
and morphology (Zasada and Strong 2000). Phytopathological Society, St. Paul, MN.
Mountain maple can reproduce by sprouting from 1,252 p.
underground lateral stems, rarely from root
suckers, and colonies usually develop following Gleason, H.A. and A. Cronquist. 1991. Manual of
disturbances by browsing or cutting (Sullivan vascular plants of northeastern United States and
28
adjacent Canada. The New York Botanical Sullivan, J. 1993. Acer spicatum. In: U.S.
Garden, Bronx, NY. 910 p. Department of Agriculture, Forest Service,
Rocky Mountain Research Station, Fire Sciences
Johnson, W.T. and H.H. Lyon. 1991. Insects that Laboratory. Fire Effects Information System.
feed on trees and shrubs. Cornell University http://www.fs.fed.us/
Press, Ithaca, NY. 560 p. database/feis/plants/shrub/acespi/index.html.
14 p.
Jones, R.K. and D.M. Benson. 2001. Diseases of
woody ornamentals and trees in nurseries. The USDA, NRCS. 2002. The PLANTS Database,
American Phytopathological Society, St. Paul, Version 3.5 National Plant Data Center, Baton
MN. 482 p. Rouge, LA. http://plants.usda.gov. [not paged].
Sinclair, W.A., H.H. Lyon, and W.T. Johnson. van Gelderen, D.M., P.C. de Jong, and H.J.
1987. Diseases of trees and shrubs. Cornell Oterdoom. 1994. Maples of the World. Timber
University Press, Ithaca, NY. 575 p. Press, Portland, OR. 458 p.
Sullivan, J.R. 1983. Comparative reproductive Zasada J.C. and T.F. Strong. 2000. Acer L. In:
biology of Acer pensylvanicum and A. spicatum Woody plant seed manual. F.T. Bonner, tech.
(Aceraceae). American Journal of Botany 70(6): coord., R.G. Nisley, ed. U.S. Department of
916-924. Agriculture, Forest Service, Washington, DC.
http://www.wpsm.net/ Acer.pdf. [not paged].
29
Acnistus arborescens (L.) Schlecht. gallinero
SOLANACEAE
Synonyms: Atropa arborescens L.

Cestrum cauliflorum Jacq.
Acnistus cauliflorus (Jacq.) Schott
Acnistus frutescens Bello
Cestrum macrostemon Sassé & Moç.
Dunalia arborescens (L.) Sleumer
John K. Francis
cluster, yellow or orange, globose, 5 to 11 mm in

diameter, and have a persistent calyx. The fruit
pulp is juicy, almost tasteless or slightly bitter, and
not edible to humans. Each fruit contains numerous
yellow, flattened seeds 1.5 to 2 mm wide with a
rolled embryo (Howard 1989, Liogier 1995, Little
and others 1974, Stevens and others 2001).
Range.—Gallinero is native to the Greater

Antilles, the Lesser Antilles, Trinidad and Tobago,
Mexico, Central America, and South America
through Brazil and Peru (Howard 1989, Little and
others 1974).
Ecology.—Gallinero is widespread and rare to

common on a wide range of soils over both igneous
and sedimentary rocks. In Puerto Rico, it grows in
areas that receive from about 1600 to 2500 mm of
mean annual precipitation. The species is most
common from 700 to 1,500 m in elevation in Costa
Rica (Haber 2002) and from 450 to 900 m in
Puerto Rico (Little and others 1974). The species is
General Description.—Gallinero is also known as frequently found on roadsides, landslides, old
hollow heart, wild tobacco, siyou, bastard sirio, pastures, and young secondary forests (Haber
galán arbóreo, mata gallina, tabaco de monte, 2002). Disturbance is necessary for establishment.
nigüito, marieneira, and tabak djab. It is an Gallinero is intolerant of shade; at least broken
evergreen shrub or small tree to 6 m in height and sunlight is necessary for long-term survival and
15 cm in stem diameter. Multiple stems are usual flowering. A number of insect species feed on the
for older plants. The trunk is supported by leaves and parasitize the fruits (Engriser 1995,
extensive lateral roots with “sinkers” and abundant Haber 2002, Simpson and others 2002).
fine roots. The roots are flexible, light tan, and
have furrowed bark. Stem bark is light brown or Reproduction.—Gallinero flowers in May and
gray and finely fissured. Stem wood is light brown June and fruits from May to September in Costa
and hard. Twigs of gallinero are stout, light brown Rica. Occasional individuals may be found in
or gray and finely hairy. The simple, alternate flower at any time during the year (Haber 2002). In
leaves are elliptical to lanciolate, 5 to 30 cm long Puerto Rico, the species flowers in spring and fruits
by 3 to 14 cm broad, entire, and pointed at both in late spring and summer (Little and others 1974).
ends. Gallinero bears tiny, fragrant, greenish-white The flowers are pollinated by bees, wasps, flies,
flowers in clusters of 30 or more in axillary butterflies, beetles, and occasionally hummingbirds
fascicles. The fruits (berries) are numerous in each (Haber 2002). A collection of fruits from Puerto
30
Rico averaged 0.2025 + 0.0084 g/fruit. Seeds Haber, W.A. 2002. Acnistus arborescens (L.)
separated from them weighed an average of Schltdl., Solanacear-Potatoe Family.
0.00074 + 0.00017 g/seed or 1,350,000 seeds/kg. http://www.cs.umb.edu/~whaber/Monte/Plant/
Sown on commercial potting mix, 88 percent Sola/Acn-arb. htm. 2 p.
germinated between 11 and 35 days after sowing.
Some of the seedlings were killed by “damping Howard, R.A. 1989. Flora of the Lesser Antilles,
off” fungi (species unknown). A fruiting tree Leeward and Windward Islands. Vol. 6. Arnold
produces an enormous number of seeds. Birds and Arboretum, Harvard University, Jamaica Plain,
bats eat the fruits and disperse the seeds (Engriser MA. 658 p.
1995). Layering (rooting) occurs whenever the
trunk or branches come in contact with the ground; Kupchan, S.M., W.K. Anderson, P. Bollinger,
sprouts from prostrate trunks and branches R.W. Doskotch, R.M. Smith, J.A. Saenz-
sometimes become independent plants. Renauld, H.K. Schnoes, A.L. Burlingaure, and
D.H. Smith. 1969. Tumor inhibitors. XXXIX.
Growth and Management.—Gallinero has a Active principles of Acnistus arborescens.
moderate growth rate and is relatively short-lived Isolation and structural and spectral studies of
(10 to 20 years). Nursery and management Withaferin A and Withacnistin. Journal of
experience has not been published. Organic Chemistry 34(12): 3,858-3,866.
Benefits.—Gallinero is used occasionally as an Liogier, H.A. 1995. Descriptive flora of Puerto

ornamental and for hedges and living fence posts. Rico and adjacent islands. Vol. 4. Editorial de
Chickens eat the fruits (Little and others 1974). la Universidad de Puerto Rico, San Juan, PR.
The wood is burned for fuel (Espinoza 1985). 617 p.
Practitioners of traditional herbal medicine use
extracts of leaves to treat cancerous growths. An Little, E.L., Jr., R.O. Woodbury, and F.H.
alcoholic extract was tested and found to inhibit Wadsworth. 1974. Trees of Puerto Rico and the
cancerous cells in vitro and in vivo. The active Virgin Islands. Vol. 2. Agriculture Handbook
ingredients were identified as Withaferin A and 449. U.S. Department of Agriculture,
Withacnistin (Kupchan and others 1969). Extracts Washington, DC. 1,024 p.
of gallinero were the most effective against
hamster lung fibroblasts of 31 Columbian plants Lopez de Cerain, A., R. Pinzón, J. Calle, A. Marín,
used in herbal medicine to treat cancer (Lopez de and A. Monge. 1996. Cytotoxic activities of
Cerain and others 1996). The species is used in Colombian plant extracts on Chinese hamster
Brazil as a diuretic, to treat liver infections, as well lung fibroblasts. Phytotherapy Research 10(5):
as an antitumor agent (Mingussi and Barata 2002). 431-432.
Gallinero helps reforest disturbed areas, protects
the soil, and furnishes food and cover for wildlife. Minguzzi, S. and L.E.S. Barata. 2002 Vitanolídeos
anti-câncer de Acnistus arborescens.
References http://www.sbq.org.br/ranteriores/
23/resumos/0398/ 2 p.
Engriser, E.M. 1995. The effect of insect larvae
infestation on fruit choice in phyllostomid fruit Simpson, S.E., H.N. Nigg, and J.L. Knapp. 2002.
bats: an experimental study. Biotropica 27(4): Host plants of diaprepes root weevil and their
523-525. implications to the regulatory process.
http://www.fcprac.ifas.ufl.edu/citrustopics/pest
Espinoza, P.L. 1985. Importance of the tree %20control/Diaprepes/Diaprepes%20Procee.
component in agroforestry-based coffee 18 p.
cultivation, with examples from Costa Rica.
Thesis. Institut fur Waldbau der Tropen und Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
Naturwaldforschung, University of Gottingen, Montiel, eds. 2001. Flora de Nicaragua.
Germany. 164 p. Monographs in Systematic Botany Vol. 85, No.
3. Missouri Botanical Garden, St. Louis, MO.
p. 1,911-2,666.
31
Aegiphila martinicensis Jacq. Caribbean spiritweed
VERBENACEAE
Synonyms: Aegiphila glandulifera Moldenke

Aegiphila panamensis Moldenke
John K. Francis
Howard 1989, Liogier 1995, Little and others

1974).
Range.—Caribbean spiritweed is native to Cuba,
Jamaica, Puerto Rico, the Virgin Islands, the
Lesser Antilles, Trinidad, Mexico to Panama,
Colombia, Venezuela, and French Guiana
1974, Missouri Botanic Garden 2003). It is not
known to have been planted or naturalized
elsewhere. Stevens and others (2001) treat the
Central American distribution as a separate species
(A. panamensis Moldenke) based on the shape of
the calyx.
Ecology.—Caribbean spiritweed is rare in Puerto

Rico (Little and others 1974) but is common in
Nicaragua (Stevens and others 2001). It occurs in
forests and brushy areas in foothills and lower
mountains of Puerto Rico from 60 to 244 m in
elevation in areas of limestone, other sedimentary,
and igneous rocks (Little and others 1974). In
Nicaragua, the species grows in forests and
pastures in wet to dry areas from near sea level to
900 m elevation (Stevens and others 2001).
Caribbean spiritweed is moderately intolerant of
General Description.—Caribbean spiritweed, shade and is able to flower and fruit in partial
also known as capaíllo, bois cabrit, bois de fer, shade.
bwa kabuit, sureau gros, and bois cabroit, is an
upright to climbing, evergreen shrub or small tree Reproduction.—Caribbean spiritweed flowers
up to 6 m in height and 7.5 cm in stem diameter. and fruits irregularly throughout the year (Little
The older stems are gray and furrowed, branches and others 1974). The fruits ripen a few at a time
are mostly smooth with lenticels, and twigs are in each infrutescence rather than all at once. A
four-angled. The ivory-colored wood is collection of fresh fruits from Puerto Rico weighed
moderately hard and brittle, and the stems have a an average of 0.496 + 0.026 g/fruit. Air-dried seed
2-mm pith. Roots are tan colored and flexible. The separated from them averaged 0.0331 + 0.0007
opposite, thin leaves are oblong-lanceolate to g/seed or 30,000 seeds/kg. Placed on moist blotter
oblong with entire edges, 8 to 16 cm long by 3 to 8 paper without pretreatment, 61 percent germinated
cm broad, rounded or pointed at the base and long- between 1 and 4.5 months after sowing (author’s
pointed at the tip. Inflorescences are axillary or observation). Germination was hypogeal
terminal, many-flowered cymes. Flowers are white (cotyledons remain below ground). Seeds are
or pale yellow, tubular, and 2 to 6 mm long. From probably dispersed by birds. Caribbean spiritweed
these develop globose to ovoid, yellow, orange, or sprouts when cut and may resprout to replace
red, 7- to 10-mm, fleshy fruits with persistent senescent stems.
calyxes, a grainy texture, and little flavor. Fruits
have one to four brown, elliptical seeds with Growth and Management.—Caribbean
longitudinal striations (author’s observations, spiritweed grows relatively rapidly, about 1
32
m/year. Stems may live 5 years or more. No Universidad de Puerto Rico, San Juan, PR.
planting or management experience has been 617 p.
published.
Little, E.L., Jr., R.O. Woodbury, and F.H.
Benefits.—Carabbean spiritweed helps protect the Wadsworth. 1974. Trees of Puerto Rico and the
soil and furnishes food and cover for wildlife. Virgin Islands. Vol. 2. Agriculture Handbook
Larva of the hawk moth, Munduca hannibal 449. U.S. Department of Agriculture,
hannibal, apparently feed exclusively on Washington, DC. 1,024 p.
Caribbean spiritweed (Oehlke 2003). Infusions of
the leaves and twigs have been used as a diuretic Missouri Botanic Garden. 2003. Flora of Panama
and as a pleasant tonic for treating asthma (Núñez- checklist: Aegiphila martinicensis Jacq. http://
Melédez 1982). In Costa Rica, the plant is mobot.mobot.org/cgi-bin/search_pick. 1 p.
considered an aphrodisiac (Duke 2003).
Núñez-Melédez, E. 1982. Plantas medicinales de
References Puerto Rico. Editorial de la Universidad de
Puerto Rico, Río Piedras, PR. 498 p.
Duke, J. 2003. Dr. Duke’s phytochemical and
ethnobotanical databases: Tico ethnobotanical Oehlke, B. 2003. Manduca hannibal hannibal.
dictionary. http://www.ars-grin.gov/duke/ http://www.silkmoths.bizland.com/mhannhan.
dictionary/tico/a.html. [not paged]. htm. 4p.
Howard, R.A. 1989. Flora of the Lesser Antilles. Stevens, R.D., C. Ulloa-U., A. Pool, and O.M.
Vol. 6. Arnold Arboretum, Harvard University, Montiel, eds. 2001. Flora de Nicaragua.
Jamaica Plain, MA. 658 p. Monographs in Systematic Botany Vol. 85, No.
3. Missouri Botanic Garden, St. Louis, MO.
Liogier, H.A. 1995. Descriptive flora of Puerto p. 911-2,666.
Rico and adjacent islands. Vol. 4. Editorial de la
33
Agave missionum Trel. corita
AGAVACEAE
Synonyms: Agave portoricensis Trel.
John K. Francis
elliptical or globose and 2.5 to 3.8 cm long and

contains numerous seeds, 6 to 8.5 mm long and 5
to 6 mm broad. If the flowers abort, numerous
bulbils are formed on the pedicels (author’s
observation, Britton and Wilson 1923).
Range.—Corita is native to Puerto Rico and the

offshore islands of Culebra and Vieques and St.
Thomas, St. Croix, St. John, Tortola, Virgin Gorda
and Anegada in the U.S. and British Virgin Islands
(Breckon and García 2001, Britton and Wilson
1923, University of the Virgin Islands
Conservation Data Center 2002). Based on work
by Trelease (1913), some sources (International
Plant Name Index 2002, Missouri Botanical
Garden 2002, Texas A & M University
Bioinformatics Working Group 2002) separate the
group into two species—A. missionum and A.
portoricensis Trel. However, Britton and Wilson
(1923) and Liogier and Martorell (2000) recognize
just A. missionum. Because ecological and
management differences between the two
populations are probably minor, they will be
treated here as a single species.
Ecology.—Corita grow naturally in well-drained to

excessively drained areas that receive between 750
mm and about 1700 mm of annual precipitation.
General Description.—Corita, also known as The species colonizes most soil types over both
maguey, is a succulent shrub with woody parts that sedimentary and igneous rock formations. Corita is
is 1.0 to 1.5 m in height in its vegetative stage and intolerant of shade and consequently does not grow
3 to 6 m when in flower. The plant is considered a well under a closed forest canopy and does not
shrub because it is of shrub size and has fibrous- compete well in tall grass swards. It is often found
woody tissue at the base of the leaf whorl, woody in disturbed areas, cliffs, and rocky terrain on the
thorns, and a woody flower stalk. The numerous coasts and inland. Corita plants are eaten by cattle
fibrous roots are about 5 mm in diameter and the when they are small but are generally avoided after
fibrous-woody base of the plant extends about 15 the thorns harden.
cm below the surface. Lanciolate to broadly
lanciolate, fleshy leaves are blue-green in color, Reproduction.—Flowering of corita appears to be
contain many fine, high-quality fibers running their triggered more by size and vigor than by age.
length, and thorns, 2 to 6 mm in length along their Probably, reserves of starch and other nutrients are
edges, and larger ones, 10 to 20 mm long, at the important. Flowering plants are often about 1 m or
tips. The leaves die when the plant is about more in height (tallest leaf), 2 m in diameter, and 5
midway through flowering. The flower stalks to 10 or more years old, depending on the quality
(panicles) are about 10 cm in basal diameter with of the habitat. Flowering proceeds from the bottom
soft wood and a pithy center. The yellow or of the inflorescence to the top. Although honeybees
yellow-green perianth is six-parted and funnel and other insects visit the flowers in great numbers,
shaped, 4.5 to 5.2 cm long. The capsules are
34
most plants never produce seeds. After the flowers former times.
fall off, bulbils form at the point where the flowers
were attached. Corita plants produce several References
hundred to several thousand bulbils each. A
collection of 100 bulbils from Puerto Rico weighed Breckon, G.J. and R.G. García. 2001. Vascular
(fresh) an average of 3.003 + 0.252 g/bulbil and plants of Susúa Forest. http://www.uprm.
ranged from 0.18 to 13.63 g. Placed with their edu/biology/profs.breckon/herbarium/florasusua.
bases slightly buried in moist potting mix, all of htm. 52 p.
them rooted within 1 week. They grew rapidly
under partial shade and were pricked into Britton, N.L. and P. Wilson. 1923. Botany of Porto
containers in about 3 weeks from sowing. After 2 Rico and the Virgin Islands, descriptive flora-
or 3 weeks they were moved to full sun and Spermatophyta. Scientific Survey of Porto Rico
reached about 15 cm in height in 2 additional and the Virgin Islands Vol. 5, Part 1. New York
months and were dispersed to the public (author’s Academy of Sciences, New York. 626 p.
observation). Plants originating from bulbils can be
numerous under fruiting plants. It is not known International Plant Name Index. 2002.
what specialized means of dispersal exist. International Plant Name Index Query.
However, new plants are frequently seen with no http://www.ipni.org/ipni/query_ipni.html. 1 p.
remains of flowering corita plants nearby.
Normally a plant dies after completing flowering, Liogier, H.A. and L.F. Martorell. 2000. Flora of
but many plants produce basal sprouts in the year Puerto Rico and adjacent islands, a systematic
of flowering, or before, that continue for another synopsis. 2nd ed. Editorial de La Universidad de
generation. Puerto Rico. 382 p.
Growth and Management.—Corita grows at a Missouri Botanical Garden. 2002. W3 tropicos.

steady, moderate rate from bulbils on well-watered http://mobot.mobot.org/cgi-bin/search_vast. 1 p.
soil and somewhat more slowly in poor, droughty
ground. Growth and development of seedlings has Texas A&M University Bioinformatics Working
not been reported. Transplanting is easy with little Group. 2002. Agavaceae (the century-plant
mortality. Because of low competition, disturbed, family). http://www.csdl.tamu.edu/FLORA/
dry, and rocky areas are probably the best sites for bonapfams/bonxxaga.htm. [not paged].
planting. Corita is recommended for planting on
shorelines and dunes in the Caribbean (United Trelease, W. 1913. Collections in Puerto Rico and
Nations Environment Program 1998). St. Thomas. Memoirs of the National Academy
of Sciences 11: 37.
Benefits.—Corita contributes to the biodiversity
and helps protect the soil in the sites where it United Nations Environment Program. 1998.
grows. Because of its hardiness in almost any well- Manual for sand dune management in the wider
drained soil, moderate size, and pleasing dark blue- Caribbean. http://www.cep.unep.org/issues/
green color, the species makes an excellent sanddunes.pdf. 73 p.
ornamental. Because exotic agaves were promoted,
it was used sparingly in the past but is beginning to University of the Virgin Islands Conservation Data
attract more attention. Corita appears to be an Center. 2002. U.S.V.I. vegetation classification
excellent honey source. The leaves contain strong system. http://cdc.uvi.edu/REAWEB/
fibers that probably were used for cordage in vegbody.html. [not paged].
35
Allamanda cathartica L. allamanda
APOCYNACEAE
Synonyms: Echites verticillata Sessé & Moç.

Orelia grandiflora Aublet
Allamanda grandiflora (Aublet) Poiret in Lam.
Allamanda hendersonii W. Bull ex Dombrain
John K. Francis
branches. The bright yellow flowers are 5 to 7.5 cm

across. Flowers of cultivated varieties are often
larger and may be colored white, cream, pink, or
orange. Capsules, which rarely occur in cultivated
varieties, are subglobose, 4 to 6 cm in diameter,
and densely prickled. They contain many tan,
flattened, winged seeds. There are 2n = 18
chromosomes (Howard 1989, Liogier 1995, Long
and Lakela 1976).
Range.—Allamanda is apparently native to

northern Brazil, Guyana, Surinam, and probably
French Guiana (Liogier 1995, Pacific Island
Ecosystems at Risk 2002, Tropilab Inc. 2002). The
species has been planted and has become
naturalized throughout the tropics (Howard 1989).
Ecology.—Wild and naturalized allamanda grows

on riverbanks in Suriname (Tropilab Inc. 2002), on
disturbed sites in Florida (Long and Lakela 1976),
along roads (Liogier 1995), and on abandoned
farms, house places, and around clandestine dumps
in Puerto Rico (author’s observation). The species
Illustration source: Missouri Botanical Garden is intolerant to intermediate in tolerance to shade. It
can grow below brush and a thin forest canopy,
General Description.—Allamanda, also known as usually without blooming, but blooms and makes
angel’s trumpet, golden trumpet, yellow bell, its greatest growth in full sun. Allamanda grows
canario, cautiva, dabel-da-dama, orelia, and liane à best in well drained, moist, sandy soils rich in
lait, is an evergreen, vine-like woody shrub. It may organic matter (Barcellos 2002). It does not
reach a free-standing height of 2 m and an tolerate salty soils, highly alkaline conditions, and
extension of 5 m or more. The species also climbs is killed by temperatures of -1 °C (Floridata 2002,
a few meters into the crowns of tall brush and low Tropilab Inc. 2002). In Nicaragua, the species
trees. Older plants often have multiple stems from occurs between 0 and 700 m in elevation (Stevens
the root crown and long stems with relatively few and others 2001). Allamanda has naturalized in
branches. Bark of lower stems is brown and Puerto Rico in areas that receive between about
furrowed. Twigs are green or yellow green. Stems 1000 and 2800 mm of mean annual precipitation.
and twigs exude a milky sap when cut. The
leathery, yellow-green to dark green leaves grow in Reproduction.—Allamanda blooms all year in
whorls of three or four, or are sometimes opposite. most habitats. Because capsules and seed are rarely
Leaves are 6 to 16 cm long, obovate to oblong- produced by cultivated varieties, naturalization is
lanceolate, pointed at both ends and have entire usually by vegetative means. In Puerto Rico, the
margins and short petioles. Inflorescences are few- species has been planted widely, persists
flowered, axillary cymes that grow near the ends of tenaciously, and spreads by layering as the vines
36
extend. In addition, trimmings dumped in vacant plantastoxicas.hpg.ig.com.br/toxicas/allcat.htm.
lots and wildlands root readily and start new 2 p.
colonies. Plants coppice vigorously when cut. The
species is invasive in Queensland, Australia Ecology and Evolutionary Biology Conservatory.
(Pacific Island Ecosystems at Risk 2002). 2002. Allamanda cathartica L. University of
Connecticut. http://florawww.eeb.uconn.edu/
Growth and Management.—Established acc_num/198500393.html. 3 p.
allamanda grows rapidly, adding 1 to 3 m to
extended length per year. Pruning is necessary to Floridata. 2002. Allamanda cathartica.
keep it under control in most landscaping http://www.floridata.com/ref/a/alla_cat.cfm. 3 p.
applications. There appears to be no reason to
establish it in wildlands, but the shrub could easily Howard, R.A. 1989. Flora of the Lesser Antilles,
be planted with cuttings placed directly in the soil. Leeward and Windward Islands. Dicotyledoneae.
Getting rid of spot infestations can be difficult. Part 3. Vol. 6. Arnold Arboretum, Harvard
Cutting is ineffective. For grubbing to work, University, Jamaica Plain, MA. 658 p.
removal of the plant stems and roots from the site
must be complete. Control with herbicides may Liogier, H.A. 1990. Plantas medicinales de Puerto
overcome some of these difficulties but is untested. Rico y del Caribe. Iberoamericana de Ediciones,
Inc. San Juan, PR. 566 p.
Benefits and Detriments.—In wild stands,
allamanda protects the soil and furnishes cover for Liogier, H.A. 1995. Descriptive flora of Puerto
wildlife. Year-long production of large, beautiful Rico and adjacent islands. Vol. 4. Editorial de la
flowers endears allamanda to gardeners and Universidad de Puerto Rico, San Juan, PR.
landscapers throughout the tropics and subtropics. 617 p.
The species is particularly useful because it will
grow in most areas with relatively little care. It is Long, R.W. and O. Lakela. 1976. A Flora of
used as ground cover, for hedges and screens, and Tropical Florida. Banyan Books, Miami, FL.
as upright shrubs. Because of its rapid growth, 962 p.
pruning is often necessary, which can expose
gardeners to the toxic sap that causes dermatitis Pacific Island Ecosystems at Risk. 2002.
symptoms of rash, blisters, and itch. Although Allamanda cathartica L., Apocynaceae.
incidence is much less common, plant parts are http://hear.org/pier_v3.3/alcat.htm. 2p.
also toxic if ingested. All parts contain the toxic
iridoid lactone, allamandin (Ecology and Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
Evolutionary Biology Conservatory 2002). In Montiel, eds. 2001. Flora de Nicaragua.
herbal medicine, teas prepared from leaves and Monographs in Systematic Botany Vol. 85, No.
roots are used as a strong purgative that must be 1. Missouri Botanic Garden Press, St. Louis,
used with caution (Liogier 1990). MO. 943 p.
References
Tropilab Inc. 2002. Allamanda cathartica L.
http://www.tropilab.com/allamanda.thml. 2 p.
Barcellos, D.C. 2002. Plantas ornamentais tóxicas:
Allamanda cathartica. http://www.
37
Amyris elemifera L. torchwood
RUTACEAE
Synonyms: Amyris maritima Jacq.

Amyris sylvatica Jacq.
Elemifera maritima (Jacq.) O. Kuntze
John K. Francis
are aromatic and contain one brown seed (Howard

1988, Liogier 1988, Little and Wadsworth 1964).
Range.—Torchwood is native to Florida, the

Bahamas, the Greater Antilles, the Lesser Antilles,
Trinidad, Guatemala, Belize, Honduras, and El
Salvador (Liogier 1988, Little and Wadsworth
1964).
Ecology.—Torchwood is moderately intolerant of

shade. Young plants compete well with shrubs and
small trees, but do not survive in dense grass
swards. In dry forests, understory plants linger for
decades until openings in the overstory allow them
to reach intermediate and codominant positions to
flower and fruit. The species may be found in both
remnant and secondary forests. In Puerto Rico,
torchwood grows in areas receiving from 750 to
over 2000 mm of mean annual precipitation. In the
upper end of the rainfall range, the species is
confined to excessively drained sites. Elevation
General Description.—Torchwood, also known as ranges from near sea level to about 600 m. It
candlewood, sea amyris, tea, cuabilla, bois tolerates a wide variety of soil textures, fertility
chandelle, bois flambeau, and bois pini, is an levels, and pH’s. Torchwood may be found in areas
evergreen shrub or small tree reaching maximum underlain by sedimentary rocks including
heights of 4 to 13 m, depending on the limestone, igneous, and metamorphic including
environment. It usually has a single stem emerging ultramafic rocks, and coastal sands. In the
from the ground, unless damaged in the past but Bahamas, torchwood grows in rocky and sandy
may begin branching near the ground when there is coastal thickets (Britton and Millspaugh 1962); in
plenty of sunlight. The trunk is smooth and gray, Florida, it grows in coastal areas and on rich
becoming rough with furrows and plates when old. hammocks farther inland (4-H Youth Development
The plants usually have a weak taproot, strong, 2001).
stiff laterals, and abundant fine roots. Torchwood
has a vertical branching habit. It may have a dense, Reproduction.—In Puerto Rico, torchwood
low crown in open-grown individuals. The twigs flowers irregularly through the year (Little and
are fine and yellowish-gray, becoming gray with Wadsworth 1964). Flowers are reported to appear
age. The foliage is pendulous and aromatic, pale- over several months in the fall in Florida (West and
green or pale-green with blue-green highlights. The Arnold 1952). Nursery plants in pots in Puerto
leaves are opposite or subopposite with three Rico bloomed at 2 years of age. Seeds cleaned
(sometimes five) leaflets on a 3-cm petiole. The from ripe fruits and sown without any pretreatment
blades are ovate, rhombic ovate, or lanciolate, with germinate normally. Birds are the principal seed
a pointed or long-pointed tip. The terminal or axial disbursers.
paniculate inflorescences contain many tiny
flowers. The fruits are black globose drupes that Growth and Management.—Nursery plants in
pots reached 1.8 m in 2 years. However, older trees
38
in the dry forests that torchwood inhabits grow Paper ITF-12. U.S. Department of Agriculture,
slowly and may live for 50 years or more (author’s Forest Service, Institute of Tropical Forestry,
observation). A release cut or thinning should be Río Piedras, PR. 28 p.
effective if it were necessary to improve the
position of torchwood in a stand. Hammer, R.L. 2001. Attracting birds to your
garden. Fairchield Tropical Garden.
Benefits.—The wood of torchwood is used for http://www.ftg.org/horticulture/habitat
fence posts today but was formerly used for fuel, plants.html. 4 p.
furniture, and torches--hence the name torchwood.
In a test of untreated service life of posts, 20 of 20 Howard R.A. 1988. Flora of the Lesser Antilles,
posts were still sound after 13 years in the ground Leeward and Windward Islands.
(Chudnoff and Goytia 1972). If the wood was more Dicotyledoneae, part 1. Vol. 4. Arnold
abundant and in larger pieces, it would be in Arboretum, Harvard University. Jamaica Plain,
demand for lumber because it is fragrant, fine- MA. 673 p.
grained, strong, durable, and repellent to dry-wood
termites (Little and Wadsworth 1964). Torchwood Liogier H.A. 1988. Descriptive flora of Puerto
is valued as a honey plant in the Dominican Rico and adjacent islands, Spermatophyta. Vol.
Republic (Marcano Fondeur 1973). Birds eat the 2. Editorial de la Universidad de Puerto Rico,
fruits, but there usually are many other more Río Piedras, PR. 481 p.
attractive foods present (West and Arnold 1952). In
Florida, torchwood is valued as a larval food for Liogier, H.A. 1990. Plantas medicinales de Puetro
the rare and endangered Schaus’s swallowtail Rico y del Caribe. Iberoamericana de Ediciones,
butterfly, Papilio aristodemus ponceanus (Hammer Inc., San Juan, PR. 566 p.
2001). Key deer (Odocollous virginanus clavium)
will not eat the foliage (Schaus and others 2001). Little, E.L., Jr. and F.L. Wadsworth. 1964.
However, feral goats eat at least the bark, which Common trees of Puerto Rico and the Virgin
has caused torchwood to decrease in abundance in Islands. Agriculture Handbook 249. U.S.
Mona Island, Puerto Rico (Little and Wadsworth Department of Agriculture, Washington, DC.
1964). Taxaline, an oxazole extracted from 548 p.
torchwood, had the most effective antibacterial
activity of several bioactive chemicals from West Marcano Fondeur, E. de J. 1973. La Flora Apícola
Indian plants against Mycobacterium tuberculosis de la República Dominicana. http.//marcano.
and two other Mycobacterium species (Rastogi and Freeservers.com/nature/estudios/apicola/dicotsp.
others 1998). Root and resin extracts from html. 11 p.
torchwood and others of the genus are used as an
expectorant (Liogier 1990). Rastogi, N, J. Abaul, G-K. Seng, A. Devallois, E.
Philogene, P. Bourgeois, and K.S Goh. 1998.
References Antimycobacterial activity of chemically
defined natural substances from the Caribbean
4-H Youth Development. 2001. Florida forest flora of Guadeloupe. FEMS-Immunology and
trees: torchwood (Amyris elemifera). Medical Microbiology 20(4): 267-273.
http://www.sfrc.ufl.edu/4h/Torchwood/
torchwoo.htm. 2 p. Schaus, C., S. Wade, and J. Dunan. 2001. Key
deer and plants they won’t eat.
Britton, N.L. and C.F. Millspaugh. 1962. The http://monroe.ifas.ufl.edu/key_deer_plants.htm.
Bahama flora. The New York Botanical Garden, 3 p.
New York. 695 p.
West, E. and L.E. Arnold. 1952. The native trees
Chudnoff, M. and E. Goytia. 1972. Preservative of Florida. University of Florida Press,
treatments and service life of fence posts in Gainesville, FL. 212 p.
Puerto Rico (1972 progress report). Research
39
Aralia spinosa L. devil’s walking stick
ARALIACEAE
Synonyms: Aralia leroana K. Koch.
Kristina Connor
1998). The dark green, glaucous leaflets are sessile,
5.1 to 10.2 cms long and 3.0 to 4.1 cm wide,
mostly paired, ovate, serrate, with pointed apex and
rounded base (Dirr 2002, Johnson and Hoagland
1999, Krüssmann 1986). There are prickles on the
veins and lower surfaces; they appear to be
outgrowths of epidermal and parenchyma cells
(Briand and Soros 2002). The wood is lightweight,
brown but streaked with yellow (Synor and Cowen
[no date]). It may be that the pithy nature of the
trunk of Devil’s walking stick allows for rapid
growth but ultimately restricts crown architecture
and maximum size attainable (Briand and others
1999). The species is deciduous and spreads by
underground runners.
Range.—Devil’s walking stick is found in the

Eastern United States, from Pennsylvania south to
Florida and west to Texas and southwestern Iowa.
It has escaped from cultivation and thus can be
found in New England, southern Ontario,
Michigan, Wisconsin, Oregon, Washington, and
western Europe (Sullivan 1992). It grows in
o
Illustration source: U.S. Department of Agriculture [not regions where temperatures fall to –29 C; such
dated] severe freezes can kill stems back to ground level
(Godfrey 1988, Frett and MacKenzie 1999,
General Description.—Devil’s walking stick, also Sullivan 1992, Scheper 2002).
known as angelica tree, American angelica-tree,
Hercules’ club, pigeon tree, pick tree, prickly ash, Ecology.—This highly adaptable species grows
prickly elder, toothache bush, toothache tree, and best in well drained soils of low and moist
shotbush, is a large, coarse-textured shrub or small woodlands but can be found growing in rocky, dry
tree, ranging from 6 to 10 m in height. The sturdy, or clay soils and under a range of pH conditions
ash gray to brown stems have dense, stout prickles, (Frett and MacKenzie 1999). It grows luxuriantly
and diameters to 15 cm are not uncommon (U.S. on good sites, but plants may live longer and be
Department of Agriculture [no date]). Young stems sturdier on poorer sites (Scheper 2002). It has a
are mostly unbranched with leaves clustered near fairly high heat tolerance. According to Russell
the top, producing a flat-topped, widely spreading (1997), it is shade tolerant, but Sullivan (1992)
crown. Stems tend to remain unbranched until the reports that it will decline if the overstory becomes
first terminal inflorescences are produced (Sullivan thick, a common characteristic of moderately
1992). Twigs are similar in color and armament to intolerant species. It has no serious pests, but
the stems and have a large pith (Synor and Cowen margined blister beetles may defoliate plants early
[no date]). Leaf scars are distinctive, lined with in the fall (Scheper 2002).
spines, and can extend half way around the twigs.
Leaves are alternate, bi- or tri-pinnately compound, Reproduction.—Devil’s walking stick spreads
and 0.9 to 1.8 m in length. The rachis is prickly and extensively by vegetative reproduction from
swollen at nodes (Bailey and Bailey 1976). The underground rhizomes. The first terminal
rachis and rachilla may be the functional equivalent inflorescences usually occur when plants are, on
of branches in this species (Briand and others the average, 3.5 years old (Sullivan 1992). The
40
white to cream colored flowers are small, perfect, However, it will send up shoots from its rhizomes
numerous, and occur in large clusters (umbels) in and can be somewhat invasive.
mid to late summer, depending on geographic area.
Panicles of flowers may be 30 to 46 cm long. Each References
flower, composed of five sepals, five reflexed
petals, and five stamens, is borne on a hairy stalk. Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third:
The ovary is inferior, and there are usually five A Concise Dictionary of Plants Cultivated in the
styles, united at the base (Brickell and Zuk 1996, United States and Canada. McMillan Publishing
U.S. Department of Agriculture [no date]). The Co., Inc. New York. 1,290 p.
purple-black fruits, maturing in late fall, are
produced in large quantities on pink-red stems. Briand, C.H., S.M. Campion, D.A. Dzambo, and
They are juicy, ovoid drupes up to 6.4 mm long K.A. Wilson. 1999. Biochemical properties of the
with three to five seed-like stones. The species can trunk of the Devil’s walking stick (Aralia
be propagated from seed sown outdoors in fall spinosa; Araliaceae) during the crown-building
(Russell 1997) or from suckers taken in late winter phase: Implications for tree architecture.
and root cuttings taken in late fall and overwintered American Journal of Botany 86(12): 1677-1682.
upside down in sand (Scheper 2002). Stored seeds
require 3 to 5 months stratification, followed by 1 Briand, C.H., A.D. Daniel, K.A. Wilson, and H.E.
to 4 months at 20 oC for germination. Woods. 1998. Allometry of axis length, diameter,
and taper in the Devil’s walking stick (Aralia
Growth and Management.Devil’s walking stick spinosa; Araliaceae). American Journal of Botany
can be maintained by mowing, cutting, and 85(9): 1201-1206.
burning, and will commonly form dense thickets
by resprouting after such disturbances. When the Briand, C.H. and C. Soros. 2002. Prickle
overstory becomes too dense, the species will distribution in Aralia spinosa.
decline. Defoliation by gypsy moth in http://henson1.salisbury.edu/~biology/
Pennsylvania and Maryland has increased the Briand/research/research.html. 5 p.
presence of Devil’s walking stick by thinning the
overstory (Sullivan 1992). The species can be Brickell, C. and J.D. Zuk. 1996. The American
controlled by injecting stems larger than 2 inches Horticultural Society A-Z Encyclopedia of
in diameter with herbicide (Loftis 1978). Garden Plants. DK Publishing, Inc., New York.
1,092 p.
Benefits.—The flowers provide a pollen and nectar
source to honeybees and a variety of other insects. Dirr, M.A. 2002. Aralia spinosa – Devil’s-
The berries are edible to wildlife, a favorite of walkingstick or Hercules-club. Noble Plants.
cedar waxwings and other birds, as well as other Plant information webpage,
frugivores and omnivores, including the black bear. http://www.nobleplants.com/Classnotes/
Deer browsing has been reported (Krüssmann fall/fallprofiles/aralia.htm
1986, Sullivan 1992). The bark, roots, and berries
have also been used for medicinal purposes by both Felter, H.W. and J.U. Lloyd. 1898. Aralia
Native Americans and early settlers. While various spinosa.-Prickly Elder. King’s American
plant parts, extracted in alcohol or water, have been Dispensatory. 18th ed., 3rd revision, 2 vol. Ohio
used to treat boils, fever, toothache, eye problems, Valley Co., Cincinnati, OH. 2,172 p. + index.
skin conditions, and snakebite, the raw berries can
be mildly toxic to humans if ingested, contact with Frett, J. and B. MacKenzie. 1999. Aralia spinosa.
the bark or roots can cause a brief skin irritation, The University of Delaware Botanical Garden.
and ingesting the fresh bark will cause vomiting Species descriptions. http://bluehen.ags.udel.edu.
(Felter and Lloyd 1898, Grieve 1971). The species 1 p.
is cultivated as an ornamental plant, the lacy
appearance of the foliage, along with the prolific Godfrey, R.K. 1988. Trees, shrubs, and woody
flowers and fruit making it an interesting vines of northern Florida and adjacent Georgia
landscape plant (Dirr 2002, Odenwald and others and Alabama. The University of Georgia Press,
1996). It is also an excellent barrier plant (Frett Athens, GA. 734 p.
and MacKenzie 1999, Michigan State University
Extension 1999) and is good for urban use. Grieve, M. 1971. A Modern Herbal. The
medicinal. culinary, cosmetic and economic
41
properties, collection and folk-lore of herbs, Russell, A.B. 1997. Trees of the Maritime Forest.
grasses, fungi, shrubs and trees with their modern Department of Horticultural Science, North
scientific uses. Dover Publications Inc., New Carolina State University,
York. 902 p. http://www.ces.ncsu.edu/depts/hort/
consumer/factsheets/maritime. 2 p.
Johnson, F.L. and B.W. Hoagland. 1999. Aralia
spinosa. Catalog of the Woody Plants of Scheper, J. 2002. Aralia spinosa. Floridata. http://
Oklahoma, Oklahoma Biological Survey www.floridata.com/ref/a/aral_spi.cfm. 4 p.
http://www. biosurvey.ou.edu. 1 p.
Sullivan, J. 1992. Aralia spinosa. In: U.S.
Krüssmann, G. 1986. Manual of Cultivated Broad- Department of Agriculture, Forest Service, Rocky
Leaved Trees and Shrubs. Volume I, A-D. Mountain Research Station Fire Sciences
Timber Press, Beaverton, OR. 448 p. Laboratory (2002 February). Fire Effects
Information System
Loftis, D.L. 1978. Preharvest herbicide control of http://www.fs.fed.us/database/feis. 10 p.
undesirable vegetation in southern Appalachian
hardwoods. Southern Journal of Applied Forestry Synor, T.D. and W.F. Cowen. [No date]. Ohio
2(2): 51-54. Trees. Aralia. Department of Natural Resources,
Ohio State University Extension Bulletin 700-00.
Michigan State University Extension. 1999. Aralia http://www.ag.ohio-
spinosa – Devil’s Walkingstick. Ornamental Plants state.edu/~ohioline/b700/index. html. 2 p.
plus Version 3.0.
http://www.msue.msu.edu/msue/imp/ modzz/ U.S. Department of Agriculture. [No date].
00001941.html. 1 p. Southern wetland flora: Field office guide to plant
species. USDA Soil Conservation Service, South
Odenwald, N.G., C.F. Fryling, Jr., and T.E. Pope. National Technical Center, Fort Worth, Texas.
1996. Plants for American Landscapes. Louisiana Jamestown, ND: Northern Prairie Wildlife
State University Press, Baton Rouge. 266 p. Research Center Home Page,
http://www.npwrc.usgs.gov/resource/1999
/soutflor/soutflor.htm. [not paged].
42
Ardisia elliptica Thunb. shoebutton ardisia
MYRSINACEAE
Synonyms: Ardisia ketoensis Hayata

Bladhia elliptica (Thunb.) Nakai
John K. Francis
and A. humilis Vahl under A. elliptica (Center for

Aquatic and Invasive Plants 2002, Neal 1965,
Pacific Island Ecosystems at Risk 2002). These
species are difficult to distinguish. Liogier (1995)
erroneously cites A. solanacea instead of A.
elliptica in Puerto Rico. Shoebutton ardisia is also
naturalized and escaped in Hawaii (Pacific Island
Ecosystems at Risk 2002), Florida (Center for
Aquatic and Invasive Plants 2002), and many other
areas.
Ecology.—Shoebutton ardisia principally grows

and is most aggressive in areas of wet soils. It
forms dense stands in low areas and intermittent
stream bottoms with clay soils in Puerto Rico. It is
General Description.—Shoebutton ardisia, also found in wet, lowland areas in Hawaii (Pacific
known as sea-shore ardisia, blackberry, Island Ecosystems at Risk 2002), in hammocks,
mameyuelo, mata pelandok, penah periah, ati disturbed wetlands, old fields, and tree islands in
popa’a, and atiu (Corner 1952, Pacific Island marshes in Florida (Center for Aquatic and
Ecosystems at Risk 2002), is an invasive shrub or Invasive Plants 2002), and in mangroves elsewhere
small tree to 6 m in height and 15 cm in basal (Tomlinson 1986). The annual precipitation of
diameter. The plants produce strong stems with these areas in Puerto Rico and probably elsewhere
gray bark. The stems are usually single, but is above 1500 mm/yr. The species is shade tolerant
additional sprouts may arise from the rootstalk, and usually grows under forest canopies. Seedlings
especially if the plant is injured. Shoebutton ardisia appear in well-established forest understories—
grows a strong taproot, much-branched laterals, disturbance is not required for establishment.
and fine roots with rhizomorphic tips. Leaves have Although significant insect herbivory does occur in
a rubbery or leathery texture and are pink when Florida, it does not appear to inhibit the advance of
young, turning dark green later. They are glabrous the species into new habitats (Dominguez and
and alternate with petioles about 1 cm long and others 2002).
blades 8 to 12 cm by about 3 cm. The
inflorescences are axillary, and umbellate, Reproduction.—Shoebutton ardisia blooms
corymbose, or racemose with numerous five- sporadically throughout the year (Miami-Dade
merous, pink flowers. The drupes are globose County 2002). The species is insect pollinated and
drupes 6 to 11 mm in diameter, dark purple to dull self-fertile (Pascarella 1997). A collection of fruits
black when ripe and contain one seed each from Puerto Rico weighed an average of 0.321 +
(author’s observation, Corner 1952, Howard 1989, 0.005 g/fruit. Seeds separated from them weighed
Pacific Island Ecosystems at Risk 2002). (air-dried) an average of 0.0463 + 0.001 g/seed.
Both the fruits and seeds are uniform in weight
Range.—The taxonomy and range of shoebutton (coefficients of variation 15 and 17, respectively).
ardisia are uncertain. Native areas for Ardisia Sown without any pretreatment in commercial
elliptica have been listed variously as Sri Lanka, potting mix, 75 percent germinated between 42 and
India, China, Taiwan, Malaya, South East Asia, 81 days after sowing. Germination is epigenous.
Indonesia, and the Philippines (Corner 1952, Although seedlings are subject to damping-off
Tomlinson 1986, Yuen-Po 1999). Some authors mortality right after germination, older seedlings
unite A. polycephala Wall., A. solanacea Roxb., transplant with little mortality or loss of vigor.
43
Birds are the principal seed dispersers, although 772 p.
raccoons and opossums in Florida also eat the
fruits and disperse seeds (Miami-Dade County Dominguez, J., A. Scott, T. Scott, G. Valdes, C.
2002). Glenn, and C. Moore. 2002. Herbivore damage
on the invasive exotic Ardisia elliptica and the
Growth and Management.—Although native A. escallonioides in Southeastern Florida.
shoebutton ardisia may grow as much as 1 m the NSF Research in Ecology: Invasive Species.
first year as a sprout, seedlings and established http://fig.cox.miami.edu/ys99/gladres.html. 3 p.
shrubs grow slowly, usually 0.25 to 0.5 m/year in
height. No figures are known for longevity, but 10 Francis, J.K. 2000. Estimating biomass and carbon
to 25 years or more are likely. It is often desirable content of saplings in Puerto Rican secondary
to control shoebutton ardisia in natural areas. Small forests. Caribbean Journal of Science 36(3-4):
numbers of seedlings and shrubs can be eliminated 346-350.
by pulling and grubbing. A broadcast spray of
glyphosate is effective in killing dense stands of Howard, R. A. 1989. Flora of the Lesser Antilles,
seedlings but will also kill native plants. Larger Leeward and Windward Islands. Vol. 6. Arnold
plants can be killed by applying a basal spray of Arboretum, Harvard University, Jamaica Plain,
triclopyhr mixed with an oil diluent (Pacific Island MA. 658 p.
Ecosystems at Risk 2002).
Liogier, H.A. 1995. Descriptive flora of Puerto
Benefits and Detriments.—Shoebutton ardisia Rico and adjacent islands. Vol. 4. Editorial de la
was once used heavily as an ornamental for accent Universidad de Puerto Rico, San Juan, PR.
and background plantings. Many plantings still 617 p.
remain, but the species is no longer cultivated
commercially (Miami-Dade County 2002). The Miami-Dade County. 2002. Shoebutton ardisia-
fruits are edible and taste slightly sour with a hint Ardisia elliptica, Ardisia humilis.
of starch but otherwise lack flavor. The species is http://www.co.miami-dade.fl.us/derm/
useful for fuel and vegetable stakes. The air-dry environment/badplants/plant%20…/shoebutton_
specific gravity of stem wood in Puerto Rico was ardisia.ht. 1 p.
measured at 0.48 + 0.01 g/cm3. Biomass (dry
weight) of plants from 137 cm in height to plants Neal, M.C. 1965. In gardens of Hawaii. Spec. Pub.
with a 5-cm D.B.H. may be calculated by the 50. Bernice P. Bishop Museum Press, Honolulu,
model: Total Dry Weight in grams = 22.020(stem HI. 924 p.
length in meters) (diameter at 30 cm)2 (Francis
2000). Shoebutton ardisia is extremely invasive. It Pacific Island Ecosystems at Risk. 2002. Invasive
forms dense stands under trees in wet sites in plant species: Ardisia elliptica Thunberg,
Florida, Hawaii, and Puerto Rico that suppress Myrsinaceae. http://www.hear.org/pier/arell.
native understory plants (Center for Aquatic and htm. 2 p.
Invasive Plants 2001).
Pascarella, J.B. 1997. Breeding systems of Ardisia
References Sw. (Myrsinaceae). Brittonia 49(1): 45-53.
Center for Aquatic and Invasive Plants. 2002. Tomlinson, P.B. 1986. The botany of mangroves.
Ardisia elliptica Thumb., Myrsinaceae/Myrsine Cambridge University Press, London. 413 p.
family. University of Florida, Gainesville, FL.
www.aquat1.itas.ufl.edu. 2 p. Yuen-Po, Y. 1999. An enumeration of
Myrsinaceae of Taiwan. Botanical Bulletin of
Corner, E.J.H. 1952. Wayside trees of Malaya. the Academy Sinensis 40: 39-47.
Government Printing Office, Singapore, Malaya.
44
Argythamnia candicans Sw. argythamnia
EUPHORBIACEAE
Synonyms: Tournesol candicans M. Gómez
John K. Francis
Range.—Howard (1989) lists the range of

agrythamnia as the Bahamas, the Greater Antilles,
Anguilla, St. Martin, St. Barts, Guadeloupe, and
Martinique. However, Britton and Millspaugh
(1962) state that the former listings for A.
candicans in the Bahamas should be A. lucayana
Millsp., an endemic.
Ecology.—Argythamnia inhabits areas of Puerto

Rico that receive from 750 to 1800 mm of annual
precipitation at elevations from near sea level to
about 450 m, usually within a few kilometers of the
sea. Soils of all textures derived from all local rock
types appear to be suitable, if well drained. Most
of the modern habitat is rocky, steep, or disturbed.
These sites include primary and secondary forests,
brushy natural pastures, and roadsides. The species
occurs as scattered individual plants or
occasionally in thickets. Argythamnia is
moderately intolerant of shade. It requires at least
good filtered sunlight to survive. Argythamnia
plants withstand light grazing well but tend to
disappear from areas that are overgrazed. It
resprouts after fires but is eliminated from large
General Description.—No common name for A. areas by repeated fires and competition from the
candicans is listed in the literature. The convention exotic grasses, Urochloa maximum (Jacq.) R.O.
among local natural resource workers, when a Webster and Bothriochloa pertusa (L.) A. Camus.
common name does not exist, is to use its genus or Frosts do not occur in the native range.
species name as a common name. Argythamnia is a
common upright, low or half shrub that reaches 1.5 Reproduction.—Argythamnia flowers and fruits
m in height and 2 cm of ground-line diameter. continuously. During the maturation and drying
Usual heights are from 0.5 to 1.0 m. It normally process, the capsules develop internal tension and
has a single stem, although branches may arise near eventually explode flinging seeds as much as 0.3 m
the base. It is supported by a tap and lateral root away from the parent plant. A collection of
system with fine tertiary roots. The branches and capsules from Puerto Rico ranged in fresh weight
twigs are thin and gray-green or whitish-green in from 0.0257 to 0.0381. Air-dry seeds collected
color. The leaves are green or gray-green above from them averaged 0.0023 + 0.0000 g/seed or
and reddish- or purplish-green and pubescent 429,000 seeds/kg. On moist filter paper, 97 percent
below. Petioles are 1.5 to 2.5 mm long. The blades of these seeds germinated between 10 and 82 days
are usually narrowly elliptic, 1.3 to 8.5 cm long after sowing.
and 0.9 to 3.6 cm wide. Argythamnia produces
tiny, inconspicuous staminate and pistilate flowers. Growth and Management.—Argythamnia is a
The latter develop into round, flattened capsules 5 relatively short-lived species. Seed-bearing plants
to 6 mm in diameter. The capsules have three examined by the author all had two or three growth
chambers, each with a globose, dark brown seed rings in the base of the stem. Establishment of
1.5 to 2 mm in diameter (Howard 1989, Liogier seedlings could be encouraged by soil disturbance
1988).
45
near seed-bearing plants. No planting experience Howard, R.A. 1989. Flora of the Lesser Antilles,
has been reported. Leeward and Windward Islands. Vol. 5. Arnold
Arboretum, Harvard University, Jamaica Plain,
Benefits.—Argythamnia is eaten by livestock and MA. 604 p.
contributes to ground cover and environmental
diversity. Liogier H.A. 1988. Descriptive flora of Puerto
Rico and adjacent islands, Spermatophyta. Vol.
References 2. Editorial de la Universidad de Puerto Rico,
Río Piedras, PR. 481 p.
Britton, N.L., and C.F. Millspaugh. 1962. The
Bahama Flora. The New York Botanical
Garden. New York. 695 p.
46
Artemisia arbuscula Nutt. low sagebrush
ASTERACEAE
Synonyms: Artemisia tridentata var. arbuscula (Nutt.) McMinn

Serphidium arbusculum (Nutt.) W.A. Weber
E. Durant McArthur and J.R. Taylor
disc flowers and is subtended by 10 to 15

canescent involucral bracts.
Taxonomy.—Three subspecies of Artemisia

arbuscula are currently recognized by the
International Plant Names Index (2003). These are
arbuscula, longicaulis Winward & McArthur, and
thermopola Beetle. Subspecies longicaulis, also
known as Lahonton low sagebrush, is endemic to
western Nevada (Winward and McArthur 1995). It
is probably a hybrid between Wyoming big
sagebrush (Artemisia tridentata ssp. wyomingensis
Beetle and Young) and low sagebrush (Winward
and McArthur 1995). Subspecies thermopola, also
known as hotsprings sagebrush, is a dwarf form
endemic to the Stanley Basin area of Idaho,
Jackson Hole area of Wyoming and east-central
Oregon. Beetle (1960) speculated that it is a hybrid
derived from low sagebrush and threetip sagebrush
(Artemisia tripartite Rydberg). Low sagebrush has
a base chromosome number of x = 9 and can be
diploid, tetraploid, or hexaploid depending on
population and subspecies (McArthur and
Sanderson 1999, Winward and McArthur 1995).
Range.—The range of low sagebrush extends

throughout Utah, Idaho, northern California,
Nevada, Oregon, Washington, southern Colorado,
and western Montana. It is usually found at
elevations ranging from 700 to 3,500 m. Low
sagebrush can grow well in mountains above 3,000
m, particularly in arid regions such as
southwestern Utah and Nevada.
General Description.—Low sagebrush is a short, Ecology.—Low sagebrush is adapted to dry,

spreading, irregularly-branched shrub up to 50 cm sterile, often rocky and alkaline clay soils. Mean
high. Twigs are slender and densely canescent annual precipitation throughout its range can vary
(silvery white pubescence) but may become nearly between 250 and 700 mm. Seedlings develop roots
glabrous, and thus darker green, in late summer. quickly to reduce the effects of soil surface
Leaves are broadly cuneate, 0.5 to 1.5 cm long, drought stress. Typical low sagebrush habitat is
and 0.3 cm wide. The leaf apex is usually three- generally drier and rockier than that of big
toothed or cleft, although leaves on the upper part sagebrush. It can, however, grow in mosaics with
of the flowering stalks may be entire. Flower big sagebrush where each species is confined to a
heads are arranged in elongate, narrow racemous particular soil type. Low sagebrush rarely grows in
panicles. Each head usually contains five to 11 mixed stands with black sagebrush (McArthur and
others 1979), and when it does, they don’t share
47
the same chromosome number (McArthur and References
Sanderson 1999). In areas where the distributions
of these two overlap, low sagebrush usually Beetle, A.A. 1960. A study of sagebrush, the
inhabits sites that have more moisture or are at section Tridentatae of Artemisia. Bulletin 368,
higher elevations than black sagebrush (A. nova University of Wyoming Agriculture Experiment
Nelson) (Tisdale and Hironaka 1981, Ward 1953, Station, Laramie, WY. 83 p.
Zamora and Tueller 1973). Low sagebrush is
usually associated with pinyon-juniper, mountain Brunner, J.R. 1972. Observations on Artemisia in
brush, and mountain big sagebrush communities. Nevada. Journal of Range Management 25: 205-
208.
Reproduction.—Flowering occurs from August to
September, depending upon strain and elevation. Dealy, J.E., DA. Leckenby, D.M. Concannon.
Seeds ripen in October and November (McArthur 1981. Wildlife habitats in managed
and others 1979). Seeds are usually wind dispersed rangelands—the Great Basin of southeastern
during late fall or winter, and seedlings emerge Oregon. General Technical Report PNW-120,
during the following spring. Seedlings emerge USDA Forest Service, Pacific Northwest Forest
rapidly and grow quickly, although root growth and Range Experiment Station, Portland, OR,
generally outpaces stem growth during early 66 p.
development. Plants can be established on adapted
sites by direct seeding, broadcasting, and drilling. International Plant Names Index. 2003.
Seeding should be conducted in the fall on a firm International plant names index.
seedbed and seeds should remain within 2 mm of http://www.ipni.org. [not paged].
the soil surface. There are approximately 1,800
cleaned seeds per gram (Meyer 2003) Kufeld, R.C., O.C. Wallmo, and C. Feddema.
1973. Foods of the Rocky Mountain mule deer.
Management.—Low sagebrush is a source of Research Paper RM-111, USDA Forest Service,
browse for livestock and big game during winter Rocky Mountain Forest and Range Experiment
months (Kufeld and others 1973, McArthur and Station, Fort Collins, CO. 31 p.
others 1979). Some forms are browsed more
heavily than others. A gray-green form in Nevada, McArthur, E.D., A.C. Blauer, A.P. Plummer, and
for example, may be heavily browsed, while the R. Stevens. 1979. Characteristics and
green form is only lightly browsed (Brunner hybridization of important Intermountain shrubs.
1972). Stand maintenance is often crucial because III. Sunflower family. Research Paper INT-220,
shrubs provide forage and cover under conditions USDA Forest Service, Intermountain Forest and
that are intolerable for most shrubs. Range Experiment Station, Ogden, UT. 82 p.
Attempts to seed low sagebrush out of its
ecological range have been unsuccessful. McArthur, E.D. and S.C. Sanderson. 1999.
Seedlings planted “offsite” usually establish but Cytogeography and chromosome evolution in
fail to reproduce. Stands recover quickly after subgenus Tridentatae of Artemisia (Asteraceae).
burning or other disturbance. Low sagebrush American Journal of Botany 86: 1,754-1,775.
spreads vigorously and can invade perennial
seeded grass stands. Perennial herb and grass Meyer, S.E. 2003. Artemisia. In: T.F. Bonner and
understory production usually decreases as stands R.G. Nisley, eds. Woody plant seed manual.
mature. Low sagebrush is not a good candidate for http://www.wpsm.net. 12 p.
mechanical control because habitat terrain is often
extreme. Tisdale, E.W. and M. Hironaka. 1981. The
sagebrush—grass region: a review of the
Benefits.—Low sagebrush is a native shrub that ecological literature. Bulletin 33. Forest,
adds structural and biological richness to arid Wildlife, and Range Experiment Station,
communities throughout the Intermountain West. University of Idaho, Moscow, ID. 31 p.
Low sagebrush communities provide important
habitat for a variety of domestic and wild animals Ward, G.H. 1953. Artemisia, section Seriphidium,
(Dealy and others 1981). Its ability to grow on in North America, a cytotaxonomic study.
exposed sites is important for wintering animals Contributions from the Dudley Herbarium 4:
and shrub cover helps reduce on-site soil erosion. 155-205.
48
Winward, A.H. and E.D. McArthur. 1995. Zamora, B. and P.T. Tueller. 1973. Artemisia
Lahontan sagebrush (Artemisia arbuscula ssp. arbuscula, A. longiloba, and A. nova habitat
longicaulis): a new taxon. Great Basin types in northern Nevada. Great Basin Naturalist
Naturalist 55(2): 151-157. 33: 225-242.
49
Artemisa bigelovii Gray Bigelow sagebrush
ASTERACEAE
Synonyms: Artemisia petrophila Wooton & Standley
E. Durant McArthur and Jeffrey R. Taylor
slender and erect and inflorescences form long

narrow panicles with short, recurved branches.
Flower heads normally bear one ray flower and
two disc flowers. Occasionally, however, heads
may include zero or two ray flowers and one or
three disc flowers. The turbinate involucre consists
of eight to 12 short, densely tomentose bracts 2 to
4 mm long and 1.5 to 2.4 mm wide.
Taxonomy.—This taxon is undivided, and

ecotype variation has not been documented.
Chromosome races from 2x to 8x, however, have
been documented (McArthur and others 1981;
McArthur and Sanderson 1999). Bigelow
sagebrush occupies a taxonomic position between
the true sagebrushes (subgenus Tridentatae) and
other Artemisia species (subgenus Artemisia). We
have chosen to treat Bigelow sagebrush as a
member of subgenus Tridentatae because of its
growth habit, wood anatomy, leaf form,
chromosomal karyotype, RAPD molecular genetic
markers, and ITS sequences in nuclear ribosomal
DNA (Kornkven and others 1998, McArthur and
others 1981, McArthur and others 1998). The
confusion exists because flower heads may include
one or two ray flowers in addition to the
characteristic disc flowers of the Tridentatae.
Range.—The distribution of Bigelow sagebrush is

more southern than other sagebrushes. It covers
approximately 88,000 km2 through western Texas,
southern Colorado, New Mexico, Arizona, Utah,
Nevada, and California between 900 and 2,400 m
elevation (Beetle 1960, Kearney and Peebles 1960,
Ward 1953).
General Description.—Bigelow sagebrush, also
known as flat sagebrush (Hall and Clements 1923), Ecology.—Bigelow sagebrush is one of the most
is a low shrub 20 to 40 cm high with numerous drought-tolerant sagebrushes. It typically grows in
spreading branches. Leaves are narrowly cuneate, canyons, gravelly draws, and dry flats. It also
1 to 2 cm long, 0.2 to 0.5 cm wide, and normally grows on rocky soils of the southern portion of the
tridentate, although tips may be variously shaped. pinyon-juniper woodlands (Hall and Clements
The leaves on vegetative branches are similar to 1923). It is often found in mixed stands with big
those of big sagebrush (Artemisia tridentata Nutt.), sagebrush, black sagebrush (Artemisia nova
although the lobes of Bigelow sagebrush are Nelson), leafless green rabbitbrush
shallower and more sharply dentate (McArthur and [Chrysothamnus nauseosus Pallas ex Pursh ssp.
others 1979). The odor of crushed leaves is mild junceus (Greene) Hall & Clements], shadscale
and pleasant. New growth is covered with a [Artriplex confertifolia (Torr. & Frem.) Wats.],
silvery-canescent pubescence. Flower stems are
50
and especially broom snakeweed [Gutierrezia Institution of Washington, Washington, DC.
sarothrae (Pursh) Britt. & Rusby]. Bigelow 355 p.
sagebrush is usually resistant to the common rust
diseases and insect galls common to other Kearney, T.H. and R.H. Peebles. 1960. Arizona
Tridentatae taxa (Beetle 1960). It is, however, flora, 2nd edition. University of California Press,
susceptible to wilt disease (Nelson and Krebill Berkeley, CA. 1,065 p.
1981). It grows in areas that usually receive 250 to
400 mm of annual precipitation. Kornkven, A.B., L.E. Watson, and J.R. Estes.
1998. Phylogenetic analysis of Artemisia section
Reproduction.—Flowering occurs from August to Tridentatae (Asteraceae) based on sequences
October and seeds mature during the following from the internal transcribed spacers (ITS) of
months. Seeds are wind dispersed in the fall and nuclear ribosomal DNA. American Journal of
winter, and seedlings emerge in spring. Seeds are Botany 85: 1787-1795.
among the smallest of sagebrush seeds, about 5.5
million cleaned seeds per kg (Meyer 2003). McArthur, E.D., A.C. Blauer, A.P. Plummer, and
Seedlings grow rapidly, and young shrubs are R. Stevens 1979. Characteristics and
robust and vigorous. Plants appear to have good hybridization of important intermountain shrubs.
drought tolerance, even as seedlings. III. Sunflower family. Research Paper INT-220.
Direct seeding has been limited to U.S. Department of Agriculture, Forest Service
experimental plantings. Seeded shrubs grow well Intermountain Forest and Range Experiment
and natural spread normally occurs around the Station, Ogden, UT. 82 p.
parental plant, suggesting that Bigelow sagebrush
can be successfully seeded. McArthur, E.D., C.L. Pope, and D.C. Freeman.
1981. Chromosomal studies of subgenus
Management.—Bigelow sagebrush is palatable to Tridentatae of Artemisia: evidence for
livestock and game throughout its distribution. Its autopolyploidy. American Journal of Botany 68:
twigs are less woody, aromatic, and bitter than 589-605.
most species within the big sagebrush complex
(Hall and Clements 1923). In addition, Bigelow McArthur, E.D. and S.C. Sanderson. 1999.
sagebrush is more grazing tolerant than most other Cytogeography and chromosome evolution of
sagebrushes. These shrubs, however, are generally subgenus Tridentatae of Artemisia (Asteraceae).
scattered, and extensive stands are found only near American Journal of Botany 86: 1754-1775.
the Four Corners area of Utah, Arizona, New
Mexico, and Colorado. McArthur, E.D., R. Van Buren, S.C. Sanderson,
and K.T. Harper. 1998. Taxonomy of
Benefits.—Bigelow sagebrush adds biological Sphaeromeria, Artemisia, and Tanacetum
diversity to shrub communities throughout the (Compositatae, Anthemideae) based on
Southwestern United States. This species improves randomly amplified polymorphic DNA (RAPD).
soil stability and reduces erosion. It also provides Great Basin Naturalist 58: 1-11.
palatable forage for livestock and wildlife and adds
structural diversity to the landscape. Meyer, S.E. 2003. Artemisia. In: T.F. Bonner and
References R.G. Nisley, eds. Woody plant seed manual.
http://www.wpsm.net 12 p.
Beetle, A.A. 1960. A study of sagebrush: the
section Tridentatae of Artemisia. Bulletin 368. Nelson, D.L. and R.G. Krebill. 1981. A sagebrush
Agricultural Experiment Station, University of wilt disease of unknown origin. Great Basin
Wyoming, Laramie, WY. 83 p. Naturalist 41: 184-191.
Hall, H.M. and F.E. Clements. 1923. The Ward, G.H. 1953. Artemisia, section Seriphidium,
phylogenetic method in taxonomy, the North in North America. A cytotaxonomic study.
American species of Artemisia, Chrysothamnus, Contributions from the Dudley Herbarium 4:
and Atriplex. Publication 326. Carnegie 155-205.
51
Artemisia californica Less. California sagebrush
ASTERACEAE
Synonyms: None
Arlee M. Montalvo and Catherine E. Koehler
sage scrub and coastal strand on dry slopes and

fans (Munz and Keck 1968, Hickman 1993). It is
an indicator species of sage scrub (Kirkpatrick and
Hutchinson 1977), occurs on virtually all soil types
except serpentine (Westman 1981a), and its range
is limited primarily by cold winter temperatures
(Malanson and O'Leary 1995). Leaves are
drought-deciduous and seasonally dimorphic
(Westman 1981b, Gray 1982). In California, shoot
and new leaf production begins with the winter
rains, usually in December, and continues
throughout winter and spring. Leaf drop and
production of smaller leaves occurs during the
summer drought (Westman 1981b). The shallow,
branched roots (Harrison and others 1971) allow
for rapid water absorption and growth response to
shallow rains (Gray 1982). The thin leaf cuticle
and numerous stomata allow a high photosynthetic
rate in response to water availability (Poole and
General Description.—Mature California Miller 1975, Gray 1983). Consequently, there is a
sagebrush are 0.6 to 2.5 m tall and abundantly higher transpiration rate and longer period of water
branched from the base (Munz and Keck 1968, stress compared to species with thick leaves and
Hickman 1993). The common name refers to its deep roots (Poole and Miller 1975, Gray 1982).
strong, sage-like aroma and being endemic to Roots form associations with arbuscular
California and Baja California. The stems are mycorrhizal (AM) fungi, but the beneficial nature
whitish with appressed hairs, long, slender, leafy, of the interaction may be facultative and is
and flexible. The somewhat hairy gray-green potentially affected adversely by nitrogen
leaves are soft, entire to divided into narrow linear deposition (Sigüenza 2000, Yoshida and Allen
segments, giving the entire shrub a wispy 2001).
appearance. The many small flower heads are less
than 5 mm wide, arranged in racemes, and have 15 Studies on chemistry of leaves, litter, and
to 30 disk florets per flower head. Plants are soil below shrubs documented the presence of both
diploid with n = 9 chromosomes (Hickman 1993). soluble compounds and volatile monoterpenes and
sesquiterpenes that inhibit germination or growth
Range.—California sagebrush occurs in California of some plants (Halligan 1973, 1975, 1976). Small
from Contra Costa Co. south into Baja California, mammalian herbivores and seed eaters often
including the Channel Islands, at elevations of less restrict foraging to beneath shrub canopies and to
than 800 m (Munz and Keck 1968, Hickman within a short distance of protective cover
1993). Historically, it was the dominant shrub on (Halligan 1973, 1974) and may contribute to
north-facing slopes in coastal sage scrub but has conspicuous bare zones on the edge of shrub
declined to about a third of its past abundance in stands and under canopies.
many areas of southern California, especially Reproduction.—Flowering of California
inland on north-facing slopes where it is being sagebrush tends to peak in late fall, but in Baja
replaced by exotic annual grasses (Minnich and California, summer rains may trigger flowering
Dezzani 1998). during summer (Minnich 1985). The reduced
flowers are typical of other wind-pollinated
Ecology.—California sagebrush is common in species of Artemisia. The single-seeded fruits
52
(achenes) ripen in December and January, with Fire Effects.—California sagebrush resprouts
some variation among years and habitats, and are after fire about 25 percent of the time (Keeley
primarily wind-dispersed (Eliason and Allen 1997, 1998). Resprouting appears to be lower in burns
DeSimone and Zedler 1999). Achenes are tiny through dense vegetation, where plants are older,
(about 60 micrograms) and in bulk amount to or if fire intensity is high (Malanson and O'Leary
about 14,300,000 seeds/kg. (personal 1982, Keeley 1998, Minnich and Dezzani 1998).
communication with S&S Seeds, Carpenteria, Seedling emergence after fire is variable and low
California). Seeds germinate in canopy openings (Zedler and others 1983, Keeley 1998). Seedlings
or in small grassland clearings generated by tend to appear the second year from seeds of
gophers (Thomomys spp.), but seedlings suffer resprouts or seeds blown in from adjacent areas.
high mortality from gopher activity (Eliason and Under high fire intensities or frequency, California
Allen 1997, DeSimone and Zedler 1999, 2001). sagebrush will likely be extirpated because of its
Seedlings emerge in the rainy season, and most poor resprouting ability and poor competitive
growth occurs by May. ability of seedlings (Malanson and O'Leary 1982,
Malanson and Westman 1991). Development,
Seed Germination.—Seeds will germinate when increased fire frequency, competition with exotic
fresh, but stored seeds may need cold stratification grasses, poor growth in grasslands, and air
to enhance germination (De Hart 1994). While pollution are each detrimental to this shrub’s
testing the combined effects of light and fire survival (Eliason and Allen 1997, Allen and others
components on germination, 73 percent of seeds 1998, Minnich and Dezzani 1998, Keeley 1998).
exposed to light germinated on soil, but seeds in
the dark did not germinate (Keeley 1987). These Horticulture.—California sagebrush can be
results are consistent with field data that show compact if kept pruned and subjected to occasional
germination may be limited by reduced light pinching. These aromatic plants remain attractive
(Elliason and Allen 1997, Montalvo and others in the dry season with occasional water (Schmidt
2002). Certain components of fire also influence 1980, Keator 1994). It prefers well-drained soils
germination. In light, treatment with leachate from and a low organic matter content (Wasowski and
charred wood (charrate) alone or with heat Wasowski 1995). Naturally occurring prostrate or
increases germination by 5 to 14 percent, but low-mounding varieties are in cultivation (Browse
germination is about a third lower when seeds are 1987, Perry 1992, Wasowski and Wasowski 1995)
subjected to high temperatures (70 to 120 oC). In and look good in dry borders or as a foreground to
contrast, in darkness both heat and charrate alone contrasting taller shrubs (Keator 1994).
stimulate germination, although heat and charrate
together generally decrease germination. Thus Benefits.—California sagebrush provides habitat
buried seeds may require some exposure to fire in for many plant and animal species and is an
order to germinate, but such exposure yields important component of critical habitat for the rare
inferior germination compared to light alone California gnatcatcher, Polioptila californica
(Keeley 1987). (Weaver 1998). The bird forages preferentially in
California sagebrush and the often co-dominant
Genetics and Geographic Variation.—Near the shrub Eriogonum fasciculatum Benth. (Beyers and
coast, plants vary in pubescence, color, chemistry, Wirtz 1997), in part because they house a high
and physiology (O'Brien 1980). Shrubs vary number and diversity of arthropods. Osborne
between a green form with sparse hairs on the (1998) found high diversity and abundance of
leaves to a form with dense hairs that give the arthropods in both species and concluded that A.
plants a gray hue. Populations differ in the californica, together with other sage scrub shrub
distinctness of forms and in their relative species, is important to the maintenance of high
frequency. Seedlings raised together generally arthropod species richness and abundance in
retain the grayness of their parents but can become coastal sage scrub. California sagebrush is also
grayer with age (O'Brien 1980). Differences in involved in species-specific interactions. For
pubescence also correspond to differences in water example, it is the host for Trirhabda
content, rate and timing of shoot elongation, leaf sericotrachyla, an herbivorous beetle whose
retention, and chemistry. The lower leaf-water developmental cycle is closely associated with the
content of gray plants negatively impacts larval seasonal growth of its only known host plant
growth and fecundity of the beetle, Trirhabda (O'Brien 1980).
sericotrachyla (O'Brien 1980).
53
Historical Uses.—Leaves of California sagebrush Endangered Species and Habitats. International
have been used by Native Americans for smoking, Assoc. of Wildland Fire, Fairfield, WA. p. 81-89.
in sweat-houses, and various other purposes (Bean
Bocek, B. 1984. Ethnobotany of Costanoan Indians,
and Saubel 1972). The Cahuilla used the plant to
California, based on collections by John P.
ensure proper maturation of girls into women. It is
Harrington. Economic Botany 38: 240-255.
said to stimulate the uterine mucosa, ensuring
rapid childbirth and, if regularly consumed as a Browse, P.M. 1987. Artemisia californica 'canyon
decoction prior to the onset of each menstruation, gray'. Pacific Horticulture 48: 56.
to prevent menstrual cramps and alleviate
menopausal trauma. Fresh or dried leaves were De Hart, J. 1994. Propagation Secrets for California
Native Plants. Jeanine De Hart, Encinitas, CA.
chewed to alleviate colds (Bean and Saubel 1972).
Costanoan Indians applied leaves to wounds or 28p.
aching teeth for pain reduction, and as a decoction DeSimone, S.A. and P.H. Zedler. 1999. Shrub
to bathe patients with colds, coughs, rheumatism, seedling recruitment in unburned Californian
or to be consumed and used as a poultice for coastal sage scrub and adjacent grassland.
treatment of asthma (Bocek 1984). Ecology 80: 2018-2032.
Growth and Management.—California DeSimone, S.A. and P.H. Zedler. 2001. Do shrub
sagebrush should be planted using shallow seeding colonizers of southern Californian grassland fit
methods such as hydroseeding or dry broadcasting generalities for other woody colonizers?
followed by seed imprinting. Shallow methods are Ecological Applications 11: 1,101-1,111.
superior to planting with a range drill because the Eliason, S.A. and E.B. Allen. 1997. Exotic grass
seeds need light for germination (Montalvo and competition in suppressing native shrubland re-
others 2002). Attempts to use annual legumes as establishment. Restoration Ecology 5: 245-255.
nurse plants can result in reduced seedling growth
and survival (Marquez and Allen 1996), but Gray, J.T. 1982. Community structure and
seedling survival may be unaffected when sown productivity in Ceanothus chaparral and coastal
with other native species at low density (Montalvo sage scrub of southern California. Ecological
and others 2002). Clearing weeds before planting Monographs 52: 415-435.
is critical. California sagebrush can return Gray, J.T. 1983. Nutrient use by evergreen and
vigorously to areas where all vegetation has first deciduous shrubs in southern California. Journal
been removed, even when sown with a variety of of Ecology 71: 21-41.
other species. Spring rains or artificial late fall and
spring irrigation may enhance survival of seedlings Halligan, J.P. 1973. Bare areas associated with
(Williams and Hobbs 1989, Elliason and Allen shrub stands in grassland: the case of Artemisia
1997, Padgett and others 2000). However, survival californica. BioScience 23: 429-432.
is not enhanced by late-season watering in wet Halligan, J.P. 1974. Relationship between animal
years (DeSimone and Zedler 2001). activity and bare areas associated with California
sagebrush in annual grassland. Journal of Range
References Management 27: 358-362.
Allen, E.B., P.E. Padgett, A. Bytnerowicz, and R. Halligan, J.P. 1975. Toxic terpenes from Artemisia
Minnich. 1998. Nitrogen deposition effects on californica. Ecology 56: 999-1003.
coastal sage vegetation of southern California. Halligan, J.P. 1976. Toxicity of Artemisia
General Technical Report 166, U.S. Department californica to four associated herb species. The
of Agriculture, Forest Service. p. 131-139. American Midland Naturalist 95: 406-421.
Bean, J.L. and K.S. Saubel. 1972. Temalpakh: Harrison, A.T., E. Small, and H.A. Mooney. 1971.
Cahuilla Indian knowledge and usage of plants. Drought relationships and distribution of two
Malki Museum Press, Morongo Indian Mediterranean-climate California plant
Reservation, CA. 225 p. communities. Ecology 52: 869-875.
Beyers, J.L. and W.O. Wirtz II. 1997. Vegetative Hickman, J.C, ed. 1993. The Jepson Manual:
characteristics of coastal sage scrub sites used by Higher Plants of California. University of
California gnatcatcher: implications for California Press, Ltd., Los Angeles, CA. 1,400 p.
management in a fire-prone ecosystem. In: J.
Greenlee, ed. Fire Effects on Rare and Keator, G. 1994. Complete Garden Guide to the
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Native Shrubs of California. Chronicle Books, Dissertation. University of California, Irvine. 125
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management: The October 1993 large fire episode Padgett, P.E., S.N. Kee, and E.B. Allen. 2000. The
in California. In: J. M. Moreno, ed. Large Forest Effects of irrigation of revegetation of semi-arid
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p. 69-90. Environmental Management 26: 427-435.
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scrub. Vegetatio 35: 21-33. conservation. Land Design Publishing, Claremont,
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Malanson, G.P. and J.F. O'Leary. 1982. Post-fire
regeneration strategies of Californian coastal sage Poole, D.K. and P.C. Miller. 1975. Water relations
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Malanson, G.P. and J.F. O'Leary. 1995. The coastal
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Malanson, G.P. and W.E. Westman. 1991. exotic grasses in southern California. Ph.D.
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pollution, and fire on California shrubland. 153 p.
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Wasowski, S. and A. Wasowski. 1995. Native
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invasive grass and native shrub in southern
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1983. Vegetation change in response to extreme
55
events: the effect of a short interval between fires
in California chaparral and coastal scrub. Ecology
64: 809-818.
56
Artemisia cana Pursh silver sagebrush
ASTERACEAE
Synonyms: Seriphidium canum (Pursh) W.A. Weber
contains from four to 20 disc flowers that are

subtended by ovate involucral bracts. Silver
sagebrush exhibits several levels of polyploidy
based on x = 9, varying by subspecies and
population. Diploid, tetraploid, and octaploid,
populations have been documented (McArthur and
others 1981, McArthur and Sanderson 1999, Ward
1953).
Taxonomy.—Silver sagebrush comprises three

subspecies: A. cana ssp. cana, A. cana ssp.
viscidula, and A. cana ssp. bolanderi. Ploidy levels
are octaploid, diploid or tetraploid, and diploid,
respectively (McArthur and others 1981,
McArthur and Sanderson 1999, Ward 1953).
Plains silver sagebrush (A. cana ssp. cana) and
mountain silver sagebrush (A. cana ssp. viscidula)
have a strong tendency to layer. Plains silver
sagebrush is an erect, round, canescent, freely
branched shrub up to 1.5 m tall. It has a more
eastern distribution than the other subspecies.
Putative natural hybrids have been found between
plains silver sagebrush and big sagebrushes
(Beetle 1960, Ward 1953). Plains silver sagebrush
has the ability to spread rapidly after burning by
resprouting and by rhizomes (Beetle 1960).
Mountain silver sagebrush (A. cana ssp.
viscidula) is an erect shrub usually not more than
1.0 m tall. It is distinguished from plains silver
General Description.—Silver sagebrush, also sagebrush by its smaller, darker leaves, and more
known as white sagebrush or hoary sagebrush, is western distribution. Leaves on vegetative stems
an erect, freely branched, rounded shrub up to 1.5 are 1 to 5 mm wide and up to 7 cm long. They are
m tall. Older stems have a dark brown, fibrous typically simple, entire, and grow in dark green
bark while younger stems are covered with a dense clusters. This subspecies varies in appearance but
white to yellowish-green tomentum. Branches is always darker than mountain big sagebrush,
often layer when in contact with the soil. Leaves with which it is often growing (Beetle 1960).
on vegetative branches are 1 to 10 mm wide and 2 Bolander silver sagebrush (A. cana ssp. bolanderi)
to 8 mm long. Leaf shape is linear to linear- has narrow leaves like mountain silver sagebrush,
oblanceolate, entire, or occasionally with one or but leaves are canescent like plains silver
two irregular lobes. Leaves are covered with a sagebrush.
silvery-white pubescence that becomes slightly
viscid with age. On flowering stalks, leaves are Range.—Aside from big sagebrush, silver
similar to those on vegetative branches but may be sagebrush is the most widely distributed member
smaller, especially toward the distal end. The of the genus. It occurs over approximately 140,000
foliage emits a mild to pungent odor when km2, from British Columbia and Saskatchewan in
crushed. Flower heads are arranged into dense, the north, to Arizona and New Mexico in the
narrow, leafy panicles, sometimes reduced to a south, and west to Oregon and California (Beetle
raceme or spike-like inflorescence. Each head
57
1960). Ranges for each subspecies, however, are spread by rhizomes. There are approximately
smaller. Plains silver sagebrush is found mostly 2,900 cleaned seeds per gram (Meyer 2003).
east of the Continental Divide, through Montana,
the Dakotas, Wyoming, western Nebraska, and Management.—Silver sagebrush can provide an
northern Colorado. Mountain silver sagebrush important source of browse and is used quite
inhabits mountainous regions (around 1,800 m or extensively by livestock and big game, especially
higher) of Montana, south along the Continental when other food sources are scarce (Kufeld and
Divide, and west to Arizona, Nevada, and Oregon. others 1973, Wasser 1982). In the western Great
The range of Bolander silver sagebrush is Plains area, silver sagebrush is an important
restricted, generally to enclosed basins, in central survival food for antelope. Domestic sheep and
Oregon and eastern California. mule deer found Bolander silver sagebrush among
the most preferable sagebrushes during winter and
Ecology.—Silver sagebrush grows on soils that fall feeding trials (Sheehy and Winward 1981).
are less mature, with less phosphorous, potassium, Sheep often browse silver sagebrush in the fall
nitrogen, organic matter, and lower cation after forbs and grasses are dry.
exchange capacity than soils supporting big Silver sagebrush can occupy areas at high
sagebrush (Hazlett and Hoffman 1975). It inhabits densities, and therefore is a candidate for plant
various environments, each subspecies adapting to control. It can be difficult to control because of its
different soil conditions. Plains silver sagebrush tendency to resprout from the crown and to spread
grows particularly well on well-watered, deep soils by rhizomes. It is not as susceptible to fire as other
of the northern Great Plains, especially along species of sagebrush (Wright and others 1979,
stream bottoms and drainageways (McArthur and White and Currie 1983). White and Currie (1983)
others 1979, Walton and others 1986). Mountain reported that silver sagebrush mortality after
silver sagebrush, on the other hand, grows in areas prescribed fall and spring burns was directly
with a heavy, lingering snow pack (Beetle 1960, related to fire intensity. In general, fall burns are
McArthur and others 1979, Tisdale and Hironaka more effective at reducing silver sagebrush than
1981, Winward 1980). Bolander silver sagebrush spring burns.
grows on poorly drained, alkaline soils (Beetle
1960, McArthur and others 1979, Tisdale and Benefits.—Silver sagebrush is an abundant native
Hironaka 1981, Winward 1980). Silver sagebrush shrub that provides additional diversity to
usually grows in areas with precipitation between shrubland ecosystems. It is an important winter
350 and 600 mm but it occurs in areas of water forage source for domestic sheep and game
accumulation, such as near streams or in enclosed animals and adds structural variety to the
basins. landscape. Native Americans traditionally used a
decoction from silver sagebrush to stop coughing
Reproduction.—Flowers bloom between August (Stubbendieck and others 1997). In addition, Euro-
and September and seed ripens in October and American settlers and Native Americans used it
November. Seeds are wind dispersed during the for fuel (McArthur and others 1979).
winter months. Establishment by direct seeding
has been successful (Kelsey 1986), especially References
when sown in the fall on the soil surface or at
shallow depths (Wasser 1982). Seedlings emerge Beetle, A.A. 1960. A study of sagebrush: the
in spring and grow rapidly. Seeding silver section Tridentatae of Artemisia. Bulletin 368,
sagebrush is best done in the late autumn or early Agricultural Experiment Station, University of
spring to maintain high viability and high Wyoming, Laramie, WY. 83 p.
germination of seeds (Hou and Romo 1998, Romo
and Grilz 2002, Romo and Young 2002). Harvey, S.J. 1981. Life history and reproductive
Seedlings are best established in lightly tilled soils strategies in Artemisia. M.S. thesis, Montana
so that stands of existing perennial grasses are State University, Bozeman, MT. 132 p.
maintained (Romo and Grilz 2002).
Plants respond dramatically to moist Hazlett, D.L. and G.R. Hoffman. 1975. Plant
conditions and may reach 35 to 50 cm in 1 year. species distributional patterns in Artemisia
Silver sagebrush can also spread asexually. tridentata—and Artemisia cana—dominatied
Harvey (1981) successfully propagated plants vegetation in western North Dakota. Botanical
through hardwood cuttings, and plants can also Gazette 136: 72-77.
58
Hou, J. and J.T. Romo. 1998. Cold-hardiness of Sheehy, D.P. and A.H. Winward. 1981. Relative
silver sagebrush seedlings. Journal of range palatability of seven Artemisia taxa to mule deer
Management 51: 704-708. and sheep. Journal of Range Management 34:
397-399.
Kelsey, R.G. 1986. Emergence, seedling growth,
and crude terpenoid concentration in a sagebrush Stubbendieck, J, Hatch, S.L., Butterfield, C.H.
garden. In: McArthur, E.D. and B.L Welch, 1997. North American Range Plants. University
comps. Proceedings—symposium on the biology of Nebraska Press, Lincoln, Nebraska. 229 p.
of Artemisia and Chrysothamnus; 1984 July 9-
13, Provo, UT. General Technical Report INT- Tisdale, E.W. and M. Hironaka. 1981. The
200. U.S. Department of Agriculture, Forest sagebrush—grass region: a review of the
Service, Intermountain Research Station, Ogden, ecological literature. Bulletin 33, Forest,
UT. p. 358-365. Wildlife, and Range Experiment Station,
University of Idaho, Moscow, ID. 31 p.
Kufeld, R.C., O.C. Wallmo, and C. Feddema.
1973. Foods of the Rocky Mountain mule deer. Walton, T.P., R.S. White, and C.L. Wambolt.
Research Paper RM-111. U.S. Department of 1986. Artemisia rproductive strategies: a review
Agriculture, Forest Service, Rocky Mountain with emphasis on silver sagebrush. In: E.D.
Forest and Range Experiment Station, Fort McArthur and B.L Welch, comps.
Collins, CO. 31 p. Proceedings—symposium on the biology of
Artemisia and Chrysothamnus; 1984 July 9-13,
McArthur, E.D., A.C. Blauer, A.P. Plummer, and Provo, UT. General Technical Report INT-200.
R. Stevens 1979. Characteristics and U.S. Department of Agriculture, Forest Service,
hybridization of important intermountain shrubs. Intermountain Research Station, Ogden, UT: 67-
III. Sunflower family. Research Paper INT-220. 74.
U.S. Department of Agriculture, Forest Service
USDA Forest Service, Intermountain Forest and Ward, G.H. 1953. Artemisia, section Seriphidium,
Range Experiment Station, Ogden, UT. 82 p. in North America. A cytotaxonomic study.
Contributions from the Dudly Herbarium 4: 155-
McArthur, E.D., C.L. Pope, and D.C. Freeman. 205.
1981. Chromosomal studies of subgenus
Tridentatae of Artemisia: evidence for Wasser, C.H. 1982. Ecology and culture of
autopolyploidy. American Journal of Botany 68: selected species useful in revegetating disturbed
589-605. lands in the west. FSW/OBS-82/56. U.S.
Department of the Interior Fish and Wildlife
McArthur, E.D. and S.C. Sanderson. 1999. Service. Washington, DC. 347 p.
Cytogeography and chromosome evolution of
subgenus Tridentatae of Artemisia (Asteraceae). Winward, A.H. 1980. Taxonomy and ecology of
American Journal of Botany 86: 1,754-1,775. sagebrush in Oregon. Station Bulletin 642.
Argicultural Experiment Station, Oregon State
Meyer, S.E. 2003. Artemisia. In: F.T. Bonner, and University, Corvallis, OR. 15 p.
R.G. Nisley, eds. Woody plant seed manual.
http://www.wpsm.net 12 p. White, R.S. and P.O. Currie. 1983. The effects of
prescribed burning on silver sagebrush. Journal
Romo, J.T. and R.W. Grilz. 2002. Establishment of Range Management 36: 611-613.
of silver sagebrush in the northern mixed prairie.
Journal of Range Management 55: 217-221. Wright, H.A., L.F. Neuenschwander, and C.M
Britton. 1979. The role and use of fire in
Romo, J. T. and J.A. Young. 2002. Temperature sagebrush-grass and pinyon-juniper plant
profiles and the effects of field environments on communites, a state-of-the-art review. General
germination of silver sagebrush. Native Plants Technical Report INT-158. U.S. Department of
Journal 3: 5-13. Agriculture, Forest Service, Intermountain Forest
and Range Experiment Station, Ogden, UT. 48 p.
59
Artemisia filifolia Torr. sandsage
ASTERACEAE
Synonyms: Oligosporus filifolius (Torr.) W.A.Weber
E.Durant McArthur and Jeffrey R. Taylor
into filiform divisions and are often fascicled.

Flower heads are arranged into leafy narrow
panicles. Each head usually contains two or three
fertile, pistilate, ray flowers and one to six perfect,
sterile, disc flowers. Five to nine canescent
involucral bracts subtend each head. The
receptacles and achenes are both glabrous.
Taxonomy.—Sandsage does not have subspecific

divisons. It has traditionally been assigned to the
subgenus Drancunlus based on floral
characteristics (McArthur 1979). Beetle (1979)
suggested that it might have an affinity with true
sagebrushes (subgenus Tridentatae). It does have
some cytological, chloroplast DNA, and chemical
similarities with Tridentatae (Kelsey and
Shafizadeh 1979, Korkven and others 1998, 1999,
McArthur and Pope 1979) although it differs from
Tridentatae in floral characteristics, wood
anatomy, and nuclear ribosomal DNA sequences
(Korkven and others 1998, Moss 1940). The
chromosome number for A. filifolia is 2n = 18; its
karyotype is similar to that of the true sagebrushes
(subgenus Tridentatae) (McArthur and Pope
1979).
Range.—Sandsage is common on sandy substrates

from the southern Black Hills in South Dakota
south to Texas and Chihuahua and west to Arizona
and Nevada (Hall and Clements 1923, McArthur
and others 1979, Rasmussen and Brotherson
1986). It is particularly important in the central
and southern Great Plains in bands along the sandy
mantles that parallel stream drainages (Berg 1994)
Ecology.—Sandsage grows almost exclusively on

sand. It is probably the most widespread shrub on
General Description.—Sandsage, also known as dunes and sandhills. In a study in southwestern
sand sagebrush and oldman sage or oldman Utah, Rasmussen and Brotherson (1986) found
sagebrush, is a freely branched, rounded shrub that that sandsage communities were less diverse than
grows up to 1.5 m tall. Young branches are adjacent communities, but plant densities were
covered with a canescent pubescence while the higher. They also reported that sandsage nutrient
older stems are covered by a dark gray-green or quality was not affected by the relative poor soil
blackish bark. Leaves are filiform, and covered nutrient profile characteristic of some sandsage
with a canescent pubescence. Leaf length can communities. Its accumulation of mineral nutrients
range from 3 to 8 cm, and leaf width is less than above that of the soil is apparently an adaptation to
0.5 cm. Leaves are entire or alternately divided sand habitats with inherently low soil fertility. It
60
grows in areas with an annual precipitation range Berg, W.A. 1994. Sand-sagebrush-mixed prairie,
of 220 to 600 mm. SRM 722. In: Shiflet, T. N., ed. Rangeland cover
types of the United States. Society for Range
Reproduction.—Blooming occurs during August Management, Denver, CO. p. 99.
and September, and seed ripens between October
and December (McArthur and others 1979). Seeds Hall, H.M. and F.E. Clements. 1923. The
are smaller than those of most sagebrushes, but phylogenetic method in taxonomy, the North
seedlings grow well once established. There are American species of Artemisia, Chrysothamnus,
approximately 3.2 million cleaned seeds per kg and Atriplex. Publication 326. Carnegie
(Meyer 2003). Seedlings emerge in the spring and Institution of Washington, Washington, DC.
grow and mature quickly. Survival is reduced, 355 p.
however, if it is planted in habitats that extend
beyond its natural range (Nelson and Krebill 1981). Kelsey, R.G. and F. Shafizadeh. 1979.
Sandsage has been seeded alone in most plantings Sesquiterpene lactones and systematics of the
but can be seeded with herbs adapted to arid sandy genus Artemisia. Phytochemistry 18: 1,591-
soils. It is easy to grow as bareroot stock and 1,611.
survives well when field planted. Wildings can be
transplanted in the spring or fall, and plants can be Korkven, A.B., L.E. Watson, and J.R. Estes. 1998.
easily reared from seed. Phylogenetic analysis of Artemisia section
Tridentatae (Asteraceae) based on sequences
Management.—The value of sandsage as a source from the internal transcribed spacers (ITS) of
of browse varies with community type. It is seldom nuclear ribosomal DNA. American Journal of
eaten in grasslands where other forage is adequate Botany 85: 1,787-1,795.
but is consumed by cattle, domestic sheep, and big
game in arid desert regions. It can be particularly Kornkven, A.B., L.E. Watson, and J.R. Estes.
important during dry years (Hall and Clements 1999. Molecular phylogeny of Artemisia section
1923, McArthur and others 1979). It is important in Tridentatae (Asteraceae) based on chloroplast
the vegetation cover of harsh sandy sites, although DNA restriction site variation. Systematic
limited water often reduces natural seed Botany 24: 69-84.
production. It is particularly useful in blackbrush
and pinyon-juniper communities where sandy McArthur, E.D. 1979. Sagebrush systematics and
outcrops occur. evolution. In: The sagebrush ecosystem: a
Sandsage resprouts vigorously after fire; symposium; 1978 April 27-28, Logan, UT.
fire may be useful in reducing canopy height and College of Natural Resources, Utah State
volume, thus allowing associated herbaceous University, Logan, UT. p. 14-22.
species to express more vigor (Vermeire and
others 2001. McArthur, E.D., A.C. Blauer, A.P. Plummer, and
R. Stevens 1979. Characteristics and
Benefits.—Sandsage can be an important, locally hybridization of important intermountain shrubs.
abundant, native shrub that provides habitat and III. Sunflower family. Research Paper INT-220.
forage for wildlife, including some small nongame U. S. Department of Agriculture, Forest Service
birds. This species helps reduce wind erosion by Intermountain Forest and Range Experiment
stabilizing light sandy soils. People in Mexico Station, Ogden, UT. 82 p.
have used a decoction of leaves to reduce the
effects of intestinal worms and other stomach McArthur, E.D. and C.L. Pope. 1979. Karyotypes
problems (Stubbendieck and others 1997). of four Artemisia species: A. carruthii, A.
filifolia, A. frigida, and A. spinescens. Great
References Basin Naturalist 39: 419-426.
Beetle, A.A. 1979. Autecology of selected Meyer, S.E. 2003. Artemisia. In: T.F. Bonner and
sagebrush species. In: The sagebrush ecosystem: R.G. Nisley, eds. Woody plant seed manual.
a symposium; 1978 April 27-28, Logan, UT. http://www.wpsu.net 12 p.
College of Natural Resources, Utah State
University, Logan, UT. p. 23-26. Moss, E.H. 1940. Interxylary cork in Artemisia
with a reference to its taxonomic significance.
American Journal of Botany 27: 762-768.
61
Stubbendieck, J., S.L. Hatch, and C.H. Butterfield
Nelson, D.L. and R.G. Krebill. 1981. A sagebrush 1997. North American Range Plants. Fifth
wilt disease of unknown origin. Great Basin Edition. University of Nebraska Press, Lincoln,
Naturalist 41: 184-191. NB. 231 p.
Rasmussen, L.L. and J.D. Brotherson. 1986. Vermeire, L.T., R.B. Mitchell, and S.C.
Habitat relationships of sandsage (Artemisia Fuhlendorf. 2001. Sand sagebrush response to
filifolia) in southern Utah. In: McArthur, E.D. fall and spring prescribed burns. In: McArthur,
and B.L. Welch, comps. Proceedings— E.D. and D.J. Fairbanks, comps. Shrubland
symposium on the biology of Artemisia and ecosystem genetics and biodiversity:
Chrysothamnus; 1984 July 9-13, Provo, UT. proceedings; 2000 June 13-15, Provo, UT. U.S.
U.S. Department of Agriculture, Forest Service, Department of Agriculture, Forest Service,
Intermountain Research Station, Ogden, UT. p. Rocky Mountain Research Station, Ogden, UT.
58-66. p. 233-235.
62
Artemisia frigida Willd. fringed sage
ASTERACEAE
Synonyms: Artemisia virgata Richardson
has a deep perennial taproot with numerous

extensive lateral roots as adaptations to withstand
drought.
Taxonomy.—Undoubtedly, fringed sage, with its

broad geographical and altitudinal distribution, has
considerable genetic variation. The taxon has three
varieties: the typical variety, Artemisia frigida
Willd. var. frigida, Artemisia frigida var.
gmelinianum Besser, and Artemisia frigida var.
williamsae S.L. Welsh (International Plant Names
Index 2003). Unlike several other widespread
Artemisia species, fringed sage is known only at
the diploid (2n = 2x = 18) chromosome level
(McArthur and Pope 1979). Fringed sage is a
member of the subgenus Artemisia (McArthur and
Pope 1979).
Range.—Fringed sage is probably the most widely

distributed and abundant species of Artemisia L.
Its range extends from Mexico in the south,
through most of the Western United States,
Western Canada, and Alaska in the north, and into
Siberia, Mongolia, and Kazakhstan (Harvey 1981;
USDA 1937). In the United States, it is most
abundant in the eastern and northern parts of its
range on the high plains along the eastern slope of
the Rocky Mountains. The altitudinal gradient of
General Description.—Fringed sage, known also fringed sage in the Intermountain area extends
as prairie sagewort, fringed sagewort and fringed from 900 to 3,500 m; in other parts of its range
sagebrush, is an aromatic, mat-forming, perennial (Great Plains and Asian Steppes) the lower
shrub 20 to 60 cm tall (McArthur and others 1979, elevational limits are somewhat less.
Stubbendieck and others 1997). The lower woody
stems are spreading and often branched, while Ecology.—Mean annual precipitation throughout
upper herbaceous stems are erect and leafy. the native range of fringed sage varies between
Adventitous roots may occur when stems come in 210 and 550 mm. It inhabits a wide variety of sites
contact with soil. The entire plant is densely silver- but typically grows in full sunlight on dry, coarse,
canescent. Leaves are abundant, 6 to 12 mm long, shallow soils. On winter livestock and game
and two or three times ternately (sub-pinnately) ranges in western Utah and eastern Nevada,
divided. Flowerheads are arranged into nodding fringed sagebrush may grow in dense stands along
racemes or open panicles. Small, densely hairy shallow depressions that collect moisture from
involucral bracts occur in series around each summer rains. In such areas, it is frequently
flowerhead. Each head contains 10 to 17 outer, associated with winterfat [Ceratoides lanata
seed-producing, pistillate, ray flowers and (Pursh) J. T. Howell], shadscale [Atriplex
numerous (25 to 50) tubular, funnelform, perfect confertifolia (Torr. & Frem.) Wats.], and
seed-producing disc flowers (McArthur and others rabbitbrushes (Chrysothamnus Nutt. species). On
1979, Stubbendieck and others 1997). This species plains, foothills, and mountain slopes, fringed sage
63
may grow in association with a variety of grasses Benefits.—Fringed sage is a good pioneer shrub
and forbs as well as with various shrubs, including for stabilizing disturbed sites. It is often used in
big sagebrush (Artemisia tridentata Nutt.), seeding Western strip mines, especially coal areas.
Bigelow sagebrush (Artemisia bigelovii Gray), Its strong taproot and numerous lateral roots help
sandsage (Artemisia filifolia Torr.), and especially stabilize gullies and reduce soil erosion. These
in overgrazed areas, with broom snakeweed rooting characteristics enable the shrub to resist
[Gutierrezia sarothrae (Pursh) Britt. & Rushby]. considerable grazing and trampling. It also
It is also a common understory shrub in ponderosa provides forage for livestock and forage and
pine communities in several Western States habitat for wildlife in both North America and
(McArthur and others 1979). Asia (Li and others 2002, USDA 1937). It is an
important food for sage grouse (Wallestad and
Reproduction.—Blooming occurs from June at others 1975). Fringed sage has some value as a
high elevations and latitudes, to November at medicinal plant (Hall and Clements 1923). Native
lower elevations and latitudes. Fringed sage is a Americans traditionally used fringed sagebrush as
prolific seed producer; each 2.5 cm length of toilet paper, to eliminate unpleasant odors from
inflorescence contains approximately 1,000 seeds dried meat, as a chewed application for wounds
(Harvey 1981). There are about 10 million cleaned prior to bandaging, and to make mats and fans
seed per kg (Plummer and others 1968). Seed (Stubbendieck and others 1997).
matures between September and December
(McArthur and others 1979) and are wind References
dispersed in winter. Seedlings emerge in spring.
Fringed sage can be established by seed, Alley, H. 1972. Range weed control. In:
transplanting young plants, or from rooted Proceedings 1970 Washington State Weed
segments of mature plants. Surface seeding on Control Conference, Yakima, WA. Washington
disturbed soils is recommended (Wasser 1982). State Weed Association. p. 39-45.
Management.—The forage value of fringed sage Cooperrider, A.Y. and J.A. Bailey. 1986. Fringed
varies considerably with location and season sagebrush (Artemisia frigida)—a neglected
(Dietz 1972). On Western ranges, it is most forage species of Western ranges. In: McArthur,
valuable to big game and livestock in late fall, E.D. and B.L Welch, comps. Proceedings—
winter, and early spring (Cooperrider and Bailey symposium on the biology of Artemisia and
1986, Kufeld 1973, USDA 1937, Wasser 1982). Chrysothamnus; 1984 July 9-13, Provo, UT.
Nutritive quality is highest in the spring but General Technical Report INT-200. U.S.
remains adequate throughout much of the year Department of Agriculture, Forest Service,
(Cooperrider and Bailey 1986, Rauzi 1982). Intermountain Research Station, Ogden, UT. p.
Individual plants are susceptible to overgrazing, 46-54.
but populations tend to increase under heavy
grazing pressure (Cooperrider and Bailey 1986). Dietz, D.R. 1972. Nutritive value of shrubs. In:
In the northwestern Great Plains, fringed sage may McKell, C.M., J.P. Blaisdell, and J.R. Goodin,
be less palatable than other available forages and tech. eds. Wildland shrubs—their biology and
occasionally invades deteriorated grasslands. management; 1971 July, Logan, UT. General
Pronghorn (Antilocapra Americana Ord) utilize Technical Report INT-1. U.S. Department of
fringed sage extensively during winter, and to a Agriculture, Forest Service, Intermountain Forest
lesser extent, year round (McArthur and others and Range Experiment Station, Ogden, UT. p.
1979). 289-302.
Fringed sage is well suited to grass and
broad-leafed herb communities. Seedlings are Hall, H.M. and F.E. Clements. 1923. The
competitive and can establish in areas with phylogenetic method in taxonomy, the North
herbaceous competition. It is adapted to mine American species of Artemisia, Chrysothamnus,
spoils, perhaps better than any other species of and Atriplex. Publication 326. Carnegie
Artemisia. It can be used as a biological control to Institution of Washington, Washington, DC.
reduce rapid weed expansion on large disturbed 355 p.
sites but can be controlled when it is too abundant
(Alley 1972). Harvey, S.J. 1981. Life history and reproductive
strategies in Artemisia. M.S. thesis, Montana
State University, Bozeman, MT. 132 p.
64
International Plant Plant Names Index. 2003. Utah. Publication 68-3. Utah Division of Fish
http//www.ipni.org.) [not paged]. and Game, Salt Lake City, UT. 183 p.
Kufeld, R.C. 1973. Foods eaten by the Rocky
Mountain elk. Journal of Range Management Rauzi, F. 1982. Seasonal variations in protein and
26: 106-113. mineral content of fringed sagewort (Artemisia
Li, F.R., A.f. Zhao, H.Y. Zhou, T.H. Zhang, and frigida). Journal of Range Management 35: 679-
X. Zhao. 2002. Effects of simulated grazing on 680.
growth and persistence of Artemisia frigida in a Stubbendieck, J., S.L. Hatch, and C.H.
semiarid sandy rangeland. Grass and Forage Butterfield, 1997. North American Range
Science 57: 239-246. Plants. Fifth Edition. University of Nebraska
Press, Lincoln, NB. 233 p.
McArthur, E.D., A.C. Blauer, A.P. Plummer, and
R. Stevens 1979. Characteristics and U. S. Department of Agriculture, Forest Service.
hybridization of important intermountain shrubs. 1937. Range plant handbook. United States
III. Sunflower family. Research Paper INT-220. Government Printing Office, Washington, DC.
U.S. Department of Agriculture, Forest Service, 816 p.
Intermountain Forest and Range Experiment
Station, Ogden, UT. 82 p. Wallestad, R., J.C. Peterson, and R.L. Eng. 1975.
Foods of adult sage grouse in central Montana.
McArthur, E.D. and C.L. Pope. 1979. Karyotypes Journal of Wildlife Management 39: 628-630.
of four Artemisia species: A. carruthii, A.
filifolia, A. frigida, and A. spinescens. Great Wasser, C.H. 1982. Ecology and culture of
Basin Naturalist 39: 419-426. selected species useful in revegetation disturbed
lands in the west. FSW/OBS-82/56. U. S.
Plummer, A.P., D.R. Christensen, and S.B. Department of the Interior, Fish and Wildlife
Monsen. 1968. Restoring big-game range in Service. Washington, DC. 347 p.
65
Artemisia longiloba (Osterhout) Beetle alkali sagebrush
ASTERACEAE
Synonyms: Artemisia spiciformis Osterhout var. longiloba Osterhout

Artemisia arbuscula Nutt. ssp. longiloba (Osterhout) L. Shultz
as low sagebrush (Artemisia arbuscula Nutt.) and

black sagebrush (Artemisia nova A. Nelson), are 3
mm wide or less. Beetle (1960) points out that this
species has been confused with silver sagebrush
(Artemisia cana Pursh) because of its large heads;
with big sagebrush (Artemisia tridentata Nutt.)
because of its broadly cuneate, three-lobed leaves;
and with low sagebrush because of its diminutive
size.
Taxonomy.—The taxon includes no recognized

subspecies or varieties. Although relegated to
subspecific status by some, e. g., the U. S.
Department of Agriculture, Natural Resource
Conservation Service plant data base 2003, we
prefer to maintain its specific status. Zamora and
Tueller (1974) differentiated it from typical low
sagebrush (Artemisia arbuscula Nutt.) in
ecological habitat preference. Its large, scattered,
distribution and its differential use by animals
suggest that it harbors considerable genetic
variation. Beetle (1960) suspected that it is a
parent to the narrowly endemic coaltown
sagebrush (Artemisia argillosa Beetle). Alkali
sagebrush has both diploid (2n = 18) and tetraploid
(2n = 36) populations (McArthur and others 1981).
Range.—Alkali sagebrush is found along foothills

and basins of the ranges that form the Continental
Divide in southwestern Montana, south through
Wyoming to northwestern Colorado, and scattered
westward through northern Utah and Idaho to
General Description.—Alkali sagebrush, also Nevada and Oregon (Beetle 1960, Winward 1980).
known as early sagebrush and low sagebrush, is a
small shrub up to 45 cm tall. It has lax, spreading Ecology.—Despite the common name of alkali
stems that frequently layer. The bark is dark sagebrush, the species also often grows well on
brown to black on older stems. The whole plant neutral soils (Passey and Hugie 1962, Tisdale and
has a dark gray-green appearance (Beetle 1960). Hironaka 1981). It is well adapted to xeric
Leaves on vegetative stems are broadly cuneate, conditions (Robertson and others 1966, Zamora
up to 2 cm long, and deeply three-lobed. Leaves and Tueller 1973). Mean annual precipitation
on flowering stems are similar in shape but throughout its range can vary between 220 and 500
smaller. Its large heads and early blooming mm. Alkali sagebrush typically grows in heavy,
readily distinguishes alkali sagebrush from other highly impermeable soils with dense “B” horizons
small sagebrushes (Beetle 1959). Heads contain (Passey and Hugie 1962, Tisdale and Hironaka
six to 11 disc flowers that are 3 to 5 mm wide. 1981). It does, however, grow on lighter soils as
Disc flowers of similarly sized sagebrushes, such well (McArthur and others 1979).
66
Reproduction.—Shrubs usually bloom in mid- Dealy, J.E., D.A. Leckenby, and D.M. Concannon.
June to early August and seed ripens in August 1981. Wildlife habitats in managed rangelands:
and September. Seeds are wind dispersed during The Great Basin of southeastern Oregon. Plant
late fall and winter, and seedlings emerge during communities and their importance to wildlife.
the following spring. Seeds are about the size of General Technical Report PNW-120. U.S.
low sagebrush seeds so we expect there would be Department of Agriculture, Forest Service,
about 2 million cleaned seeds per kg (Meyer Pacific Northwest Forest and Range Experiment
2003). Natural seeding occurs rapidly following Station, Portland, OR. 66 p.
fires and other disturbances, yet mechanical tillage
can limit establishment if seeds are sown too deep McArthur, E.D., A.C. Blauer, A.P. Plummer, and
or if the seedbed is disrupted (Monsen and Shaw R. Stevens 1979. Characteristics and
1986). hybridization of important intermountain shrubs.
III. Sunflower family. Research Paper INT-220.
Management.—Alkali sagebrush, depending on U.S. Department of Agriculture, Forest Service,
location and ecotype, can be a source of palatable Intermountain Forest and Range Experiment
browse for domestic sheep and provide habitat and Station, Ogden, UT. 82 p.
forage for other animals (Dealy and others 1981,
McArthur and others 1979). Stands of alkali McArthur, E.D., C.L. Pope, and D.C. Freeman.
sagebrush in good condition can support a diverse 1981. Chromosomal studies of subgenus
understory of perennial grasses and annual and Tridentatae of Artemisia: evidence for
perennial forbs. Such stands provide forage and autopolyploidy. American Journal of Botany 68:
cover for sage grouse, antelope, other wildlife, and 589-605.
domestic livestock (Dealy and others 1981,
Monsen and Shaw 1986). Some stands, however, Meyer, S.E. 2003. Artemisia. In: Bonner, F.T., and
are much less diverse and provide little forage or R. Nisley, eds. Woody plant seed manual.
habitat. Sites that lack a satisfactory understory http://www.wpsm.net. [not paged].
remain closed unless shrubs are reduced by fire or
other disturbance. Monsen and Shaw (1986) Monsen, S.B. and N.L. Shaw. 1986. Response of
reported that decadent stands can be rejuvenated an alkali sagebrush/fescue site to restoration
by mechanical means. treatments. In: McArthur, E.D. and B.L Welch,
This shrub has the potential to be a useful plant in comps. Proceedings—symposium on the biology
rehabilitation, especially on heavy, seasonally of Artemisia and Chrysothamnus; 1984 July 9-
dry soils. It has been observed invading 13, Provo, UT. General Technical Report INT-
roadcuts in Echo Canyon, Utah, and along 200. U.S. Department of Agriculture, Forest
roadsides near Kemmerer, Wyoming (McArthur Service, Intermountain Research Station, Ogden,
and others 1979). UT. p. 126-133.
Benefits.—Alkali sagebrush is a native shrub that Passey, H.B. and V.K. Hugie. 1962. Sagebrush on
provides habitat and forage for wildlife and relic ranges in the Snake River plains and
livestock. In addition, it has characteristics that northern Great Plains. Journal of Range
could make it useful in ecological restoration, Management. 15: 273-278.
rehabilitation, or soil conservation. It spreads well
through natural seeding and tolerates disturbance. Robertson, D.R., J.L. Nielsen, and N.H. Bare.
It also grows on argillic soils that may be too 1966. Vegetation and soils of alkali sagebrush
heavily textured for most other shrubs. and adjacent big sagebrush ranges in North Park,
Colorado. Journal of Range Management. 19:
References 17-20.
Beetle, A.A. 1959. New names within the section Tisdale, E.W. and M. Hironaka. 1981. The
Tridentatae of Artemisia. Rhodora 61: 82-85. sagebrush—grass region: a review of the
ecological literature. Bulletin No. 33, Forest,
Beetle, A.A. 1960. A study of sagebrush, the Wildlife, and Range Experiment Station,
section Tridentatae of Artemisia. Bulletin 368, University of Idaho, Moscow, ID. 31 p.
University of Wyoming Agriculture Experiment
Station, Laramie, WY. 83 p.
67
U. S. Department of Agriculture, Natural University, Oregon Agricultural Experiment
Resources Conservation Service— Station, Corvallis, OR. 15 p.
http//www.plants.usda.gov. Last visited August
25, 3003. Zamora, B. and P.T. Tueller. 1973. Artemisia
arbuscula, A. longiloba, and A. nova habitat
Winward, A.H. 1980. Taxonomy and ecology of types in northern Nevada. Great Basin Naturalist.
sagebrush in Oregon. Bull. 642. Oregon State 33: 225-242.
68
Artemisia ludoviciana Nutt. Lousiana sagewort
ASTERACEAE
Synonyms: Artemisia albula Wooton

Artemisia candicans Rydb.
Artemisia gnaphalodes Nutt.
Artemisia incompta Nutt.
Artemisia mexicana Willd. ex Spreng.
Artemisia redolens A.Gray
Artemisia sulcata Rydb.
Artemisia vulgaris var. americana Besser
Artemisia vulgaris ssp. ludoviciana H.M. Hall & Clem.
Artemisia vulgaris var. ludoviciana Kuntze
Jeffrey R. Taylor and E. Durant McArthur
green and are typically covered with a white to

gray tomentum. Leaves may be entire, tridentate,
variously-lobed oblanceolate, or linear. Lower
leaves are grayish green, oblanceolate, lobed, or
entire, 3 to 9 cm long and 0.5 to 2.5 cm long, while
upper leaves are mostly entire, gray to green
floccose to glabrous above, and white tomentose
beneath (Hall and Clements 1923). The leaf apex
is acute. Flower heads bear six to 12 pistilate ray
flowers and five to 12 perfect disc flowers,
arranged in elongate, leafy panicles 20 to 50 cm
long (Stubbendieck and others 1997). Louisana
sagewort has a strong tendency to spread through
rhizomes.
Taxonomy.—Artemisia ludoviciana is a highly

polymorphic species. Hall and Clements (1923)
chose to classify it within the Artemisia vulgaris
L. complex. This system of classification
consolidated 54 previously recognized species into
one species and 15 subspecies. Keck (1946)
suggested that the complex arose from one
cenospecies, and subsequently revised the
taxonomy. Keck’s revision classified A.
ludoviciana on the specific, rather than subspecific
level, with seven subspecies of its own. These
Illustration credit: USDA—Forest Service collection, subspecies include: typica, sulcata (Rydb.) Keck,
Hunt Institute albula (Woot.) Keck, candicans (Rydb.) Keck,
incompta (Nutt.) Keck, mexicana (Wild.) Keck,
General Description.—Louisana sagewort, also and redolens (Gray) Keck. Cytogenetic studies
known as cudweed sagewort, Louisiana sagebrush, show that the New World members within the
Louisiana wormwood, western mugwort, white vulgaris complex have a base chromosome
sage, sage-weed, estafiate, and iztafiate, is an number x = 9, as opposed to the Old World
aromatic, morphologically variable, herbaceous members, with a base chromosome number of x =
forb or sub-shrub 0.3 to 1 m tall. Stems are erect, 8. Lousiana sagewort and its close relatives (A.
single to numerous, and woody near the base. ludoviciana species complex) can be diploid,
Stems can be grayish-green, greenish-yellow, or tetraploid, or hexaploid (Estes 1969, Welsh and
69
others 1993). Estes (1969) suggested that Lousiana sagewort spreads asexually through
autopolyploidy has been the driving evolutionary rhizomes.
force within the complex. Molecular genetic
studies have also shown New World and Old Management.—Lousiana sagewort is used
World members of the complex to be different differentially by livestock and wildlife throughout
from one another (Vallès and McArthur 2001). its distribution. Populations in the southern
portions of its range, such as in southern Utah,
Range.—Lousiana sagewort is widely distributed New Mexico, and southern Colorado, are more
throughout Western North America. Its range palatable than those in the north (USDA Forest
spans from Ontario and Michigan in the east, to Service 1937, Stubbendieck and others 1997). In
British Columbia in the west, and south through these southern areas, it can provide valuable
Texas, Louisiana, California, and Mexico (USDA forage, especially on spring-fall and winter ranges.
Forest Service 1937). Lousiana sagewort grows at In more northern areas, it has a pungent taste and
various elevations from 250 to 3,050 m. is generally not eaten until after the first frost (Hall
and Clements 1923). Lousiana sagewort is
Ecology.—Lousiana sagewort grows on rocky, generally considered poor forage, although it is
sandy, or gravelly loams on open ridges, slopes, utilized by domestic sheep, deer, and elk to some
and mesas (USDA Forest Service 1937). It is also degree, and may be seasonally important (Shaw
common on plains and prairies and is usually a late and Monsen 1982, Hall and Clements 1923,
seral species that grows after grasses have declined USDA Forest Service 1937). Lousiana sagewort
(Hall and Clements 1923). Lousiana sagewort has also been used to revegetate road, logging, and
grows rapidly, is long lived, and is cold and other disturbances (Shaw and Monsen 1982). In a
drought tolerant (Shaw and Monsen 1982). It revegetation experiment at Mesa Verde National
grows well in sunny sites even though it is shade Park, CO, it survived well when transplanted in
tolerant. It is can be found in grasslands, roadcut sites (Paschke and others 1999).
sagebrush, ponderosa pine, and alpine
communities (Shaw and Monsen 1982). Lousiana Benefits.—Lousiana sagewort is an extensive and
sagewort often associates with asters (Aster L.), genetically diverse native plant that is widespread
goldenrods (Solidago L.), sunflowers (Helianthus throughout Western North America. It provides
L.), fringed sagebrush (Artemisia frigida Wild.), quick and excellent cover, reduces erosion, and
big sagebrush (Artemisia tridentata Nutt.), serves as a nurse plant for other species (Shaw and
tarragon (Artemisia dracunculus L.), needlegrass Monsen 1982). It also has pharmaceutical
(Stipa L.), blue grama [Bouteloua gracilis potential. Zavala-Sanchez and others (2002)
(H.B.K.) Lag. ex Steudel], muhly grasses found that an essential oil found in Lousiana
(Muhlenbergia Schreber), wheatgrasses sagewort, nonanal, inhibited diarrhea in mice. In
(Agropyron Gaertner), mountain dandelions addition, Lousiana sagewort extracts inhibited
[Agoseris glauca (Pursh) Raf.], and western reproduction in malarial parasites (Malagon and
yarrow (Achillea millefolium L.) (USDA Forest others 1997). The ssp. mexicana (estafiate) is used
Service 1937, Hall and Clements 1923). Mean for gastrointestinal pain, as a verimfuge, and as a
annual precipitation within its range varies bitter stimulant (Heinrich 2002). It has also been
between 400 and 1200 mm. used by Native Americans in ceremonial and
purification rituals, to deodorize feet, as pillows,
Reproduction.—Blooming begins in August and saddle pads, and to treat headaches, stomach
ends in September (Stubbendieck and others ailments, and coughs (Stubbendieck and others
1997). Seed matures in the fall and is dispersed by 1997).
wind during late fall or winter. Seeds are
relatively small, numbering about 8,360,000/kg References
(personal communication with S. Stranathan,
Upper Colorado Environmental Plant Center, Estes, J.R. 1969. Evidence for autoploid
Meeker, CO). Seed viability decreases after 2 or 3 evolution in the Artemisia ludoviciana complex
years of storage (personal communication with of the Pacific Northwest. Brittonia 21: 29-43.
K.R. Jorgensen, Great Basin Experimental Area
Ephraim, UT). Seeding aerially, by drill, or Hall, H.M. and F.E. Clements. 1923. The
broadcasting are all effective methods of planting phylogenetic method in taxonomy, the North
and should be conducted during fall or winter American species of Artemisia, Chrysothamnus,
(Shaw and Monsen 1982). In addition to seeds, and Atriplex. Publication no. 326, Carnegie
70
Institution of Washington, Washington, DC. Agriculture, Forest Service, Intermountain
355 p. Research Station, Ogden, UT. p. 123-131.
Heinrich, M. 2002. Ethnobotany, phytochemistry Stubbendieck, J., S.L. Hatch, and C.H Butterfield.
and biological/pharmacological activities of 1997. North American Range Plants. Fifth
Artemisia ludoviciana ssp. mexicana (estafiate). edition. University of Nebraska Press, Lincoln,
In: Wright, C.W., ed. Artemisia, medicinal and NB. 235 p.
aromatic plants Vol. 18. Taylor and Francis,
London, UK. p. 107-117. U.S. Department of Agriculture, Forest Service.
1937. Range plant handbook. United States
Keck, D.D. 1946. A revision of the Artemisia Government Printing Office, Washington, DC.
vulgaris complex in North America. 816 p.
Proceedings of the California Academy of
Sciences 25: 421-468. Vallès, J. and E.D. McArthur. 2001. Artemisia
systematics and phylogeny: cytogenetic and
Malagon, F., J. Vasquez, and A. Ruiz. 1997. molecular insights. In: McArthur, E.D. and D.J.
Antimalaric effect of an alchoholic extract of Fairbanks, comps. Shrubland ecosystem genetics
Artemisia ludoviciana mexicana in a rodent and biodiversity: proceedings. 2000 June 13-15,
malaria model. Parassitologia 39: 3-7. Provo, UT. Proceedings RMRS-P-21. U.S.
Department of Agrigulture, Forest Service,
Paschke, M.W., C. DeLeo, and E.F. Redente 1999. Rocky Mountain Research Station, Ogden, UT.
Revegetation of roadcut slopes in Mesa Verde p. 67-74.
National Park, U.S.A Restoration Ecology 8:
263-268. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
Higgins, eds. 1993. A Utah Flora. Second
Shaw, N. and S.B. Monsen. 1982. edition. Printing Services, Brigham Young
Nonleguminous forbs for rangeland sites. In: University, Provo, UT. 986 p.
Monsen, S.B. and N. Shaw, comps.
Proceedings: Managing Intermountain Zavala-Sanchez, M.A., S. Perez-Gutierrez, C.
Rangelands—Improvement of range and Perez-Conzalez, and D. Sanchez-Saldivar.
wildlife habitats; September 15-17, 1981; Twin 2002. Antidiarrhoeal activity of nonanal, an
Falls, ID, June 22-24, 1982; Elko, NV. Gen. aldehyde isolated from Artemisia ludoviciana.
Tech. Rep. INT-157. U.S. Department of Pharmaceutical Biology 40: 263-268.
71
Artemisia nova A. Nels. black sagebrush
ASTERACEAE
Synonyms: Artemisia tridentata ssp. nova (A. Nels.) H.&C.

Artemisia arbuscula ssp. nova (A. Nels.) McMinn
Artemisia arbuscula var. nova (A. Nels.) G. Ward
Serphidium novum (A. Nels.) W.A. Weber
involucral bracts. Leaf hairs, visible at 10x

magnification, can be useful diagnostic clues to
distinguish black sagebrush from other
sagebrushes.
Taxonomy.—Black sagebrush taxonomy is

relatively simple. It includes one species and two
varieties, A. nova var. duchesnicola and A. nova
var. nova (Welsh and Goodrich 1995). Beetle and
Johnson (1982) have found forms of black
sagebrush that are similar in appearance to
Wyoming big sagebrush. This, in addition to other
factors, has led some to suspect that black
sagebrush may have been involved in the
parentage of Wyoming big sagebrush (McArthur
1983, Winward 1976). Glandular trichomes are a
taxonomic feature that helps separate black
sagebrush from big sagebrush (A. tridentata Nutt.)
and low sagebrush (A. arbuscula Nutt.) (Kelsey
1984). Black sagebrush has a base chromosome
number of x = 9; both diploid (2n = 18) and
tetraploid populations are known. About two-
thirds of the cytologically known populations are
tetraploid (McArthur and Sanderson 1999).
Range.—Black sagebrush is one of the most

common shrubs in the Western United States.
General Description.—Black sagebrush is a small Populations can be found from California and
aromatic shrub generally 15 to 20 cm tall, although Oregon in the west, to Colorado in the east, and
it may occasionally exceed 75 cm. A dull grayish- from Montana in the north to northern Arizona and
tomentose layer causes most populations of black New Mexico in the south. Black sagebrush is most
sagebrush to appear darker than those of big common at elevations ranging between 1,500 to
sagebrush or low sagebrush. Branches are 2,400 m (McArthur and others 1979)
numerous and erect and arise from a spreading
base. Leaf base is typically cuneate, and the leaf Ecology.—Black sagebrush is well adapted to dry
surface is covered with a viscid glandular stony soils that are relatively shallow. Zamora and
pubescence. The leaf apex is three-toothed, Tueller (1973) found root restricting layers 28 to
although the uppermost leaves, particularly on 69 cm deep in half of their black sagebrush study
flowering stems, may be entire. Leaf dimensions sites. Gravelly or sandy loam strata are also
can vary between 0.5 to 2 cm long and 0.2 to 0.8 common subsurface soil profile characteristics in
cm wide. Flowering heads are grouped in narrow areas with black sagebrush. In addition, most soils
spike-like panicles that commonly extend above supporting black sagebrush are calcareous. Black
the foliage. Heads contain three to five disc sagebrush is not restricted to adverse
flowers subtended by eight to 12 greenish-yellow environments, however. If seeded, it can grow well
72
on sites normally occupied by big sagebrush. Benefits.—Black sagebrush is an important native
Mean annual precipitation throughout the natural shrub in many areas of the Western United States.
range of black sagebrush ordinarily is between 180 It provides structural diversity within native plant
and 320 mm. communities and provides an important browse
Most stands of black sagebrush do not source for native and introduced ungulates. It can
burn because populations are relatively sparse also reduce soil erosion on steep shallow slopes.
(Tisdale and Hironaka 1981). Exceptions to this ‘Gordon Creek’ is a tested germplasm from
rule, however, have been observed in central Utah, Carbon County, Utah, that has been demonstrated
where large stands of black sagebrush burned. In a to be high in nutritive quality and palatability to
comparative study, Nelson and Krebill (1981) wildlife (Welch and others 1994).
found that black sagebrush is less susceptible to a
wilt disease than other species of Artemisia grown References
under similar conditions.
Beetle, A.A. and K.L. Johnson. 1982. Sagebrush in
Reproduction.—Mature shrubs flower in late Wyoming. Bulletin 779. Agricultural
summer (usually August), and seed matures in Experiment Station, University of Wyoming,
September and October. Black sagebrush seed Laramie, WY. 68 p.
tends to be larger than those of other sagebrushes.
There are about 2,000 seeds/g (Deitschman 1974). Behan, B. and B.L. Welch. 1985. Black sagebrush:
Seeds are wind dispersed in late fall or early mule deer winter preference and monoterpenoid
winter. Seedlings emerge in the spring and often content. Journal of Range Management 38: 278-
grow rapidly. Once established, young plants can 280.
persist well even under adverse conditions. With
favorable moisture conditions, mature plants Clary, W.P. 1986. Black sagebrush response to
produce abundant seed, and natural spreading grazing in the east-central Great Basin. In: E.D.
occurs quickly. McArthur and B.L. Welch, comps. Proceedings:
symposium on the biology of Artemisia and
Management.—Black sagebrush is usually Chrysothamnus; 1984 July 9-13; Provo, UT.
considered a valuable browse species for wildlife General Technical Report INT-200. U.S.
and livestock, especially for deer, domestic sheep, Department of Agriculture, Forest Service,
and pronghorn (Clary 1986; McArthur and Intermountain Research Station, Ogden, UT:
Plummer 1978, USDA Forest Service 1937). 181-185 p.
Black sagebrush has good winter nutritive value,
but not as good as big sagebrush (Welch 1983). Deitschman, G.H. 1974. Artemisia. In: C.S.
Palatability, however, can vary significantly Schopmeyer, technical coordinator, Seeds of
between populations (Welch and others 1981). Woody Plants of the United States. Agriculture
Deer and elk preference studies for black and big Handbook 450. U.S. Department of Agriculture,
sagebrushes have yielded inconsistent results Forest Service, Washington, DC. 235-237 p.
(Scholl and others 1977, Nagy and Regelin 1977,
Smith 1950, Wambolt 1996). Welch and others Kelsey, R.G. 1984. Glandular trichomes: a helpful
have shown that some accessions are not eaten taxonomic character for Artemisia nova (black
while others are highly preferred (Behan and sagebrush). Journal of Range Management 37:
Welch 1985, Welch and others 1981), which may 370-372.
partially explain differences in preference studies.
Black sagebrush can also be a valuable McArthur, E.D. 1983. Taxonomy, origin, and
conservation species for dry, shallow, stony soils distribution of sagebrush (Artemisia tridentata)
because it produces abundant seed and seedlings and allies (subgenus Tridentatae). In: K.L.
establish readily. Due to its habitat, it is not Johnson, ed. Proceedings of the First Utah
usually a candidate for plant control. Black Shrub Ecology Workshop; 1981 September 9-
sagebrush can successfully be seeded in fall or 10; Ephraim, UT. College of Natural Resources,
early winter. Seeds should be covered with soil, Utah State University, Logan, UT. 3-13 p.
but remain within 6 mm of the surface. It grows
well with seeded herbs, but understory production McArthur, E.D., A.C. Blauer, A.P. Plummer, and
tends to decrease as the stand matures. R. Stevens 1979. Characteristics and
hybridization of important intermountain shrubs.
73
USDA Forest Service, Intermountain Forest and Welch, B.L. 1983. Improving the nutritive value of
Range Experiment Station, Ogden, UT. 82 p. winter range forage. In: S. B. Monsen and N.
Shaw, comps. Managing Intermountain
McArthur, E.D. and A.P. Plummer. 1978. rangelands—improvement of range and wildlife
Biogeography and management of native habitats: proceedings of a symposium; 1981
western North American shrubs: a case study, September 15-17; Twin Falls, ID; 1982 June 22-
section Tridentatae of Artemisia. Great Basin 24, Elko, NV. General Technical Report INT-
Naturalist Memoirs 2: 229-241. 157, U.S. Department of Agriculture, Forest
Service, Intermountain Forest and Range
McArthur, E.D. and S.C. Sanderson. 1999. Experiment Station, Ogden, UT: 158-164.
subgenus Tridentatae of Artemisia (Asteraceae). Welch, B.L., E.D. McArthur, and J.N. Davis.
American Journal of Botany 86: 1,754-1,775. 1981. Differential preference of wintering mule
deer for accessions of big sagebrush and black
Nagy, J.G. and W.L. Regelin. 1977. Influence of sagebrush. Journal of Range Management 34:
plant volatile oils on food selection by animals. 409-411.
Congress of Game Biologists 13: 225-230.
Welch, B.L., E.D. Nelson, S.A. Young, A.R.
Nelson, D.L. and R.G. Krebill. 1981. A sagebrush Sands, F.J. Wagstaff, and D.L. Nelson. 1994.
wilt disease of unknown origin. Great Basin ‘Gordon Creek’—a superior, tested germplasm
Naturalist 41: 184-191. of Wyoming big sagebrush. Research Paper INT-
461, U.S. Department of Agriculture, Forest
Smith, A.D. 1950. Sagebrush as a winter feed for Service, Intermountain Research Station. Ogden,
deer. Journal of Wildlife Management 14: 285- UT. 7 p.
289.
Welsh, S.L. and S. Goodrich. 1995. Plant novelties
Scholl, J.P., R.G. Kelsey, and F. Shafizadeh. 1977. in Lepidium (Cruciferae) and Artemisia
Involvement of volatile compounds of Artemisia (Compositae) from the Uinta Basin, Utah. Great
in browse preference by mule deer. Biochemical Basin Naturalist 55: 359-362.
Systematics and Ecology 5: 291-295.
Winward, A.H. 1976. Evolutionary development
Tisdale, E.W. and M. Hironaka. 1981. The of the Artemisia tridentata taxa. In: H. C. Stutz,
sagebrush—grass region: a review of the ed. Wildland shrubs: proceedings, symposium,
ecological literature. Bulletin 33, Forest, and workshop; 1975 November 4-6, Provo, UT.
Wildlife, and Range Experiment Station, College of Biology and Agriculture, Brigham
University of Idaho, Moscow, ID. 31 p. Young University, Provo, UT: 163.
U.S. Department of Agriculture, Forest Service. Zamora, B. and P.T. Tueller. 1973. Artemisia
1937. Range plant handbook. United States arbuscula, A. longiloba, and A. nova habitat
Government Printing Office, Washington, DC. types in northern Nevada. Great Basin Naturalist
816 p. 33: 225-242.
Wambolt, C.L. 1996. Mule deer and elk foraging
preference for 4 sagebrush taxa. Journal of
Range Management 49: 499-503.
74
Artemisia pygmaea Gray pygmy sagebrush
ASTERACEAE
Synonyms: Seriphidium pygmaeum (Gray) W.A.Weber
flowering branches are usually reduced and may

be entire. Flower heads contain three to five disc
flowers and are arranged into spike-like
inflorescences. Ray flowers are lacking. Twelve
to 18 greenish-yellow involucral bracts subtend
each head (Cronquist 1994, McArthur and others
1979).
Taxonomy.—No subspecific entities have been

described for pygmy sagebrush. Pygmy sagebrush
is known only at the diploid, 2n = 18, chromosome
level (McArthur and Sanderson 1999).
Range.—The distribution of pygmy sagebrush is

limited to approximately 54 km2 in eastern Utah,
western Nevada, and northern Arizona (Beetle
1960; McArthur and Plummer 1978; Ward 1953).
Ecology.—Pygmy sagebrush grows on calcareous

desert soils. Mean annual precipitation within its
range is approximately 200 to 300 mm. In
Nevada, this species is often associated with
halophytic threadleaf rubber rabbitbrush
[Chrysothamnus nauseosus (Pallas ex Pursh)
Britton ssp. consimilis (Greene) Hall & Clements].
In Utah, some relatively large stands are mixed
with black sagebrush (Artemisia nova A. Nelson).
Reproduction.—Blooming occurs between

August and September. Seed matures in October.
Seeds are relatively large compared to those of
other subgenus Artemisia subgenus Tridentatae
species (McArthur and others 1979). Seeds are
wind dispersed during late fall or winter.
Seedlings emerge during spring. Shrubs in
research plantings have spread naturally. It also
establishes well by transplanting divided plants.
General Description.—Pygmy sagebrush is a Although it spreads through naturally dispersed
dwarf, depressed, evergreen, cushion-like shrub seed, artificial seedings have been unsuccessful.
less than 20 cm tall. Bark on older stems becomes
dark brown and fibrous with age. Bark on younger Management.—Pygmy sagebrush provides little
branches is nearly white to straw-colored and browse because of its scarcity and size. It is
somewhat puberulent. Leaves on vegetative stems readily eaten, however, when available. It is a
are green, nearly glabrous, viscidulous, and are good candidate to plant in clay soils and may be of
pinnatified with three to 11 lobes, or sometimes considerable value for revegetating mine spoils
may be toothed. Leaf dimensions range between 2 and roadway disturbances.
to 4 mm wide and 2 to 8 mm long. Leaves on
75
Benefits.—Although it is not abundant, pygmy III. Sunflower family. Research Paper INT-220.
sagebrush does provide important soil cover in U.S. Department of Agriculture, Forest Service
dry, alkaline habitats where little else grows. It USDA Forest Service, Intermountain Forest and
also adds to the overall biological diversity to the Range Experiment Station, Ogden, UT. 82 p.
shrub communities where it is found.
McArthur, E.D. and A.P. Plummer. 1978.
References Biogeography and management of native
western shrubs: a case study, section Tridentatae
Beetle, A.A. 1960. A study of sagebrush, the of Artemisia. Great Basin Naturalist Memoirs 2:
section Tridentatae of Artemisia. Bulletin 368, 229-243.
University of Wyoming Agriculture Experiment
Station, Laramie, WY. 83 p. McArthur, E.D. and S.C. Sanderson. 1999.
Cronquist, A. 1994. Asterales, Intermountain flora, subgenus Tridentatae of Artemisia (Asteraceae).
volume 5. The New York Botanical Garden, American Journal of Botany 86: 1,754-1,775.
Bronx, NY. 496 p.
Ward, G.H. 1953. Artemisia, section Seriphicum,
McArthur, E.D., A.C. Blauer, A.P. Plummer, and in North America. A cytotaxonomic study.
R. Stevens. 1979. Characteristics and Contributions from the Dudley Herbarium 4:
hybridization of important intermountain shrubs. 155-205.
76
Artemisia rigida (Nutt.) Gray stiff sagebrush
ASTERACEAE
Synonyms: Artemisia trifida var. rigida Nutt.

Seriphidium rigidum W.A. Weber
silvery-canescent, spatulate, and usually deeply

divided into three to five narrowly linear lobes.
Occasionally, leaves are linear and entire
(Cronquist 1994, McArthur and others 1979).
Each head is composed of five to 16 perfect disc
flowers. The campanulate involucre is 4 to 5 mm
long with numerous canescent bracts. The
inflorescence is a leafy spike with heads sessile or
in small clusters in the axils of their subtending
leaves, which are generally longer than the heads.
This species somewhat resembles threetip
sagebrush (Artemisia tripartita Rydb.) in size,
silvery pubescence, and the deeply, narrowly lobed
leaves, but is distinguishable by the spike-like
inflorescence, large leafy bracts that subtend the
heads, and the deciduous leaves (Cronquist 1994,
McArthur and others 1979).
Taxonomy.—Stiff sagebrush does not include

subspecific taxa. However, both diploid, 2n = 18,
and tetraploid, 2n = 36 populations are known
(McArthur and others 1981, McArthur and
Sanderson 1999, Ward 1953).
Range.—Stiff sagebrush is found in the Columbia

and Snake River basins and the northern end of the
Great Basin between 910 to 1,500 m elevation. Its
range includes portions of Idaho, central and
eastern Oregon, and eastern Washington. The
report that its range extends into Montana is
apparently an error (Morris and others 1976;
Ecology.—This shrub grows primarily, if not

exclusively, on sites with rocky, basaltic parent
material (Daubenmire 1982, Tisdale and Hironaka
1981). Otherwise, its ecological niche is similar to
that of low sagebrush (Artemisia arbuscula Nutt.).
Stiff sagebrush grows in sites with annual
precipitation between 250 and 350 mm. Stiff
sagebrush is able to survive wildfires even though
General Description.—Stiff sagebrush, also it is not known to resprout or layer, probably
known as scabland sagebrush, is a low, pungently because it grows in scattered patches that carry
aromatic shrub with thick, rigid, somewhat brittle fires poorly. Plants grow slowly and require 2 to 4
branches. The bush grows up to 40 cm tall. years to attain a mature size (> 15 cm in height). It
Leaves, mostly 1 to 4 cm long, are deciduous, appears to have broader ecological amplitude than
77
its present range suggests (McArthur and others Hall, H.M. and F.E. Clements. 1923. The
1979). phylogenetic method in taxonomy; the North
American species of Artemisia, Chrysothamnus,
Reproduction.—Blooming begins in late August and Atriplex. Publication 326. Carnegie
and ends in early October, and seeds ripen in Institution of Washington, Washington, DC.
October and November. Seed is wind dispersed in 355 p.
late fall or winter, and seedlings emerge during the
ensuing spring. Seed production and seed quality McArthur, E.D., A.C. Blauer, A.P. Plummer, and
are low in comparison to other sagebrush species R. Stevens 1979. Characteristics and
(McArthur and others 1979). Attempts to seed and hybridization of important intermountain shrubs.
transplant this species in arid sagebrush ranges III. Sunflower family. Research Paper INT-220.
near Boise, Idaho, have been unsuccessful. In U.S. Department of Agriculture, Forest Service
such instances, plantings were made where USDA Forest Service, Intermountain Forest and
cheatgrass (Bromus tectorum L.) and other annuals Range Experiment Station, Ogden, UT. 82 p.
have invaded, and stiff sagebrush seedlings were
unable to compete. Plantings have successfully McArthur, E.D., C.L. Pope, and D.C. Freeman.
established on prepared seedbeds, however. 1981. Chromosomal studies of subgenus
Tridentatae of Artemisia: evidence for
Management.—Due to scant foliage and stiff autopolyploidy. American Journal of Botany 68:
branches, few animals except domestic sheep 589-605
browse stiff sagebrush (Hall and Clements 1923).
It provides forage on dry sites for domestic sheep McArthur, E.D. and S.C. Sanderson. 1999.
during midsummer when herbaceous plants are Cytogeography and chromosome evolution of
dry. subgenus Tridentatae of Artemisia (Asteraceae).
Benefits.—Stiff sagebrush is a native shrub that
adds biological diversity to Western ecosystems. Morris, M.S., R.G. Kelsey, and D. Griggs. 1976.
It is an important plant that provides cover and The geographic and ecological distribution of big
reduces erosion on rocky sites. Its ecology sagebrush and other woody Artemisias in
suggests that it could be used in reclaiming harsh, Montana. Proceedings Montana Academy of
disturbed sites. Sciences 36: 56-79.
References Tisdale, E.W. and M. Hironaka. 1981. The

sagebrush-grass region: a review of the
Cronquist, A. 1994. Asterales, Intermountain flora, ecological literature. Bulletin no. 33. University
volume 5. The New York Botanical Garden, of Idaho, College of Forestry, Wildlife and
Bronx, NY. 496 p. Range Sciences, Forest, Wildlife and Range
Experiment Station, Moscow, ID. 31 p.
Daubenmire, R. 1982. The distribution of
Artemisia rigidia in Washington: a challenge to Ward, G.H. 1953. Artemisia, section Seriphicum,
ecology and geology. Northwest Science 56: in North America. A cytotaxonomic study.
162-164. Contributions from the Dudley Herbarium 4:
155-205.
78
Artemisia rothrockii Gray. rothrock sagebrush
ASTERACEAE
Synonyms: Artemisia tridentata ssp. rothrockii Hall & Clements

Artemisia tridentata var. rothrockii McMinn
Sally A. Reynolds and Eric L. Berlow
surrounding slopes of lodgepole or red fir forest

but has expanded into mesic meadows since the
onset of livestock grazing in the 1850’s (Ratliff
1985, Odion and others 1988, Dull 1995). Grazing
and trampling along stream banks are proposed to
have caused channel incision and an associated
lowering of the water table in the adjacent meadow
(Kattelmann and Embury 1996, Kirchner and
others 1998, Knapp and Mathews 1996, USDA
Forest Service 1998, 2000). While more often
found on drier soils, rothrock sagebrush appears to
be tolerant of very high soil moisture as
individuals survive periods of standing water
following spring snowmelt (Berlow and others
2002). Additionally, in a pot experiment, rothrock
General Description.—Rothrock sagebrush, also
sagebrush seedlings showed 100 percent survival
known as timberline sagebrush, is a low, aromatic,
after 2 weeks and over 60 percent survival after 4
evergreen shrub, spreading to ascending from a
weeks of complete immersion in flowing water
narrow trunk, 20 to 60 cm tall. Bark is gray to dark
(Swartz, Berlow and D’Antonio unpublished data).
grayish brown, becoming fibrous with age. Young
The highest potential germination and seedling
twigs are densely tomentose. Leaves are dark
growth rates were observed in mesic, rather than
green, narrowly cuneate, ranging from 0.5 to 2 cm
xeric, meadow habitat. However, in these areas
long and 0.2 to 1.5 cm wide. Vegetative leaves are
micro-site availability and seed dispersal limit
three-toothed but, leaves on flowering stalks are
establishment, with seedlings preferentially
generally entire. Young leaves are canescent,
associated with small soil disturbances within 0.5
becoming glabrous with age (Shultz 1993).
m of a seed source (Berlow and others 2002).
Range.—Rothrock sagebrush is native and
Reproduction.—Flowers develop on narrow
endemic to California. It occurs at high elevations
inflorescences. Heads are sessile to short-
in the Sierra Nevada and the White and San
peduncled, 3 to 5 mm in diameter. Phyllaries
Bernardino Mountains. This species is commonly
(involucre bracts) occur in two series. Both series
found in the southern Sierra Nevada on the Kern
are shiny with wide margins. The outer phyllaries
Plateau, at Monache Meadows, Big Meadows,
are canescent with acute tips, whereas the inner
Ramshaw Meadows, Templeton Meadows, and
phyllaries are elliptic and only sparsely tomentose.
Mulkey Meadow. It is also found at Mono Pass,
Discoid flowers (ray florets are absent) have
Tuolumne Pass and Mount Dana in Tuolumne
between eight and 20 fertile florets. Seeds are
County, Angora Peak, and near Fallen Leaf Lake
small and resinous, ranging from 0.8 to 2 mm,
in El Dorado County (McMinn 1951). Its
with pappus present on outer seeds (Shultz 1993).
northernmost extent is Placer County (Sawyer and
Inflorescences begin development in late spring.
Keeler-Wolf 1995).
Flowering takes place at the end of the growing
season in early to late fall, and seed dispersal
Ecology. –Rothrock sagebrush generally vegetates
begins soon after. Seeds are primarily wind
well-drained, gravelly soils on the margins of
dispersed and do not appear to travel far, as
montane meadows from 2,000 to 3,500 m. In the
seedlings tend to be aggregated within 0.5 m of
southern Sierra Nevada, evidence suggests
adult shrubs (Berlow and others 2002).
rothrock sagebrush was historically limited to dry
Germination occurs immediately following spring
soils in the areas between the meadows and
79
snow-melt. Rothrock sagebrush does not form a Berlow, E.L., C.M. D’Antonio, and S.A.
large soil seedbank (Reynolds unpublished data). Reynolds. 2002. Shrub expansion in montane
An average sized adult plant can produce over meadows: the interaction of local-scale
5000 seeds (Reynolds unpublished data). disturbance and site aridity. Ecological
However, field germination rates are low (< 1 Applications 12: 1103-1118.
percent), and seed viability appears to decline
rapidly after 1 year (Berlow and others 2002, and Berlow, E.L., C.M. D’Antonio, and H. Swartz.
unpublished data). 2003. Response of herbs to shrub removal across
natural and experimental variation in soil
Growth and Management.—The growing season moisture: Implications for restoration. Ecological
for rothrock sagebrush in montane habitats is Applications. (in press).
relatively short, beginning in early spring and
continuing until the soil moisture is depleted in Dull, R.A. 1995. Palynological evidence for 19th
late summer. Under mesic conditions, rothrock century grazing-induced vegetation change in
sagebrush can begin flowering within 5 years of the southern Sierra Nevada, California. Journal
establishment (Berlow and others 2002). While of Biogeography 26: 899-912.
age to senescence is unknown, individuals up to 40
years old have been documented by counting Kattelmann, R. and M. Embury. 1996. Riparian
growth rings (Bauer and others 2002). Rates of areas and wetlands. In Sierra Nevada
increase in stem diameter have been observed to Ecosystem Project. University of California,
vary between 0.10 to 0.14 mm/year. Faster growth Centers for Water and Wildland Resources,
rates were found in microhabitats characterized by Davis, CA. p. 34-68.
moist surface soil and a shallow water table,
suggesting that this woody species can use similar Kirchner, J.W., L. Micheli, and J.D. Farrington.
water sources as herbaceous species (Bauer and 1998. Effects of herbaceous riparian vegetation
others 2002, Berlow and others 2003). Rothrock on streambank stability. Technical Completion
sagebrush seedlings are not palatable to livestock Report, Project #W-872, University of
or native grazers, which may contribute to their California Berkeley Water Resources Center.
encroachment into intensively grazed meadows. 43 p.
While large areas of sagebrush expansion are
associated with stream incision, rothrock Knapp, R.A. and K.R. Matthews. 1996.
sagebrush also has the demonstrated potential to Livestock grazing, Golden Trout, and streams
invade un-incised, mesic meadow habitat given the in the Golden Trout Wilderness, California:
presence of exposed soil and a nearby seed supply impacts and management implications. North
(Berlow and others 2002). Thus, on a large scale, American Journal of Fisheries Management 16:
preventing further stream incision is critical to 805-820.
slowing sagebrush invasion. Within mesic
meadow habitat, removal of scattered individual McMinn, H.E. 1951. An Illustrated Manual of
shrubs that serve as local seed sources may be the California Shrubs. University of California
most effective way to prevent rapid expansion into Press, Berkeley, CA. p. 608-609.
un-incised areas.
Odion, D.C., T.L. Dudley, and C.M. D'Antonio.
Benefits.—Rothrock sagebrush contributes to the 1988. Cattle grazing in southeastern Sierran
diversity of California’s montane plant meadows: Ecosystem change and prospects for
communities. It stabilizes sandy soil on the slopes recovery. In: C.A. Hall and V. Doyle-Jones,
of meadow fringes and provides habitat for a editors. Plant Biology of Eastern California,
variety of birds and small mammals. Mary Decker Symposium. White Mountain
Research Station, Los Angeles, CA. p. 277-292.
References
Ratliff, R.D. 1985. Meadows in the Sierra Nevada
Bauer, K., E.L. Berlow, and C.M. D’Antonio. of California: state of knowledge. General
2002. Shrub expansion in montane meadows: Technical Report PSW-84. U.S. Department of
The relationship between climate and Rothrock Agriculture, Forest Service, Pacific Southwest
sagebrush colonization patterns. Journal of Forest and Range Experiment Station.
Range Management. 55: 620-625. Berkeley, CA. 52 p.
80
Sawyer, J.O. and T. Keeler-Wolf. 1995. A Manual USDA Forest Service. 1998. Kern Plateau
of California Vegetation. California Native Plant Ecosystem Analysis. U.S. Department of
Society, Sacramento, CA. p. 188. Agriculture, Forest Service, Inyo National
Forest, Bishop, CA. 215 p.
Shultz, L.M. 1993. Artemisia. In: J.C. Hickman,
ed. The Jepson Manual: Higher Plants of USDA Forest Service. 2000. Templeton and
California. University of California Press, Whitney Grazing Allotments Environmental
Berkeley, CA. p. 202-205. Assessment. U.S. Department of Agriculture,
Forest Service, Inyo National Forest, Bishop,
CA. 111 p.
81
Artemisia spinescens Eaton budsage
ASTERACEAE
Syononyms: Picrothamnus desertorum Nutt.
(Cronquist 1994) states that “despite this proposed

segregation, its affinity with Artemisia is not in
question”. The inclusion or exclusion of segregate
genera from Artemisia including Picrothamnus has
not been confirmed by molecular genetic studies
as of yet (Vallès and McArthur 2001). Polyploidy
within the budsage has been documented; both
diploid, 2n = 18, and tetraploid, 2n = 36,
populations occur (McArthur and Pope 1979).
Range.—The range of budsage extends from

Wyoming to western Montana, southern Idaho,
and easten Oregon south to northwestern New
Mexico, northern Arizona, and southeastern
California (Cronquist 1994, McArthur and others
1979).
Ecology.—Budsage is well adapted to xeric

conditions. It is quite common in semiarid valley
bottoms, benches, and foothills throughout the
Interior Western United States (Hall and Clements
1923, Wood and Brotherson 1986). Mean annual
precipitation throughout its habitat varies between
150 and 250 mm. It has an extensive root system
General Description.—Budsage, also known as that grows primarily within the top 15 to 56 cm of
spring sage and bud sagebrush, is a low, the soil. Interxylary cork is formed over the
spinescent, pungently aromatic, rounded shrub 10 previous year’s wood that restricts the upward
to 50 cm high and profusely branched at the base. movement of water to the narrow zone of wood
Young branches and leaves are covered with a formed by the current year’s growth. The corky
white-tomentose pubescence that becomes stiff tissue develops during the early summer and
and gray with age. Leaves are small and normally reduces water loss during dormancy (Wood 1966).
2 cm or less in length. Leaves are three- to five- Summer dormancy is an adaptation that
palmately parted, again divided into three linear- reduces the effects of drought stress. Early in the
spatulate lobes. Leaves are crowded on short spring, when budsage first shows signs of breaking
stems, with those near the apex being smaller and dormancy and prior to bud elongation, the
more entire. Unlike most members of the genus, previous season’s bark growth can be removed
budsage is deciduous, losing most leaves by easily. This condition is known as “slipping” and
midsummer. Flowerheads are small and arranged is used as a sign to tell when the plant is palatable
in glomerate racemes of one to three heads in leaf to sheep (Wood and Brotherson 1986). Shrubs
axils on flower branches. Each head contains two subsequently produce new, bright-green leaves as
to six fertile, pistilate ray flowers and five to 13 soon as February or March. Terminal and lateral
perfect, but sterile, disc flowers. Four to eight buds generally expand and begin to elongate in
rounded involucral bracts subtend each flowerhead late March and early April. Although budsage
(Cronquist 1994, McArthur and others 1979). ordinarily begins to grow in early spring and
becomes dormant by midsummer, it occasionally
Taxonomy.—Budsage is distinct from closely may break summer dormancy after late summer
related Artemisia species and is sometimes storms.
considered a separate monotypic genus although
82
Reproduction.—Blooming normally begins Cronquist, A. 1994. Asterales, Intermountain flora,
during the last week in April and continues through Vol. 5. The New York Botanical Garden, Bronx,
the last week in May. Exceptions, however, have NY. 496 p.
been documented where blooming began as early
as late March or has extended as late as mid-June Hall, H.M., and F.E. Clements. 1923. The
(Wood 1966). Seeds normally mature in early phylogenetic method in taxonomy; the North
June. There are about 3 million cleaned seeds per American species of Artemisia, Chrysothamnus,
kg (Meyer 2003). Freezing temperatures in early and Atriplex. Publication 326. Carnegie
spring frequently kill developing embryos. Intact Institution of Washington, Washington, DC.
heads regularly fall from the plant without breaking 355 p.
apart to release seeds, although some seeds are
usually dispersed independently. Abundant natural Holmgren, R.C. and S.S. Hutchings. 1972. Salt
reproduction occurs in years when seed production desert shrub response to grazing use. In: McKell,
is plentiful and moisture conditions are favorable. C.M., J.P. Blaisdell, and J.R. Goodin, tech. eds.
Proceedings-symposium on wildland
Management.—Budsage is a palatable, nutritious shrubs--their biology and utilization, 1971 July;
winter forage for upland birds, small game, big Logan, UT. Gen. Tech. Rep. INT-1. U.S.
game, and domestic sheep. It is particularly Department of Agriculture, Forest Service,
preferred just after dormancy. Budsage is high in Intermountain Forest and Range Experiment
calcium, magnesium, phosphorous, and protein Station, Ogden, UT. p. 153-164.
(Wood and Brotherson 1986). It is usually more
palatable in late winter than early winter McArthur, E.D., A.C. Blauer, A.P. Plummer, and
(Holmgren and Hutchings 1972). Palatability R. Stevens 1979. Characteristics and
again decreases once twigs have elongated and hybridization of important intermountain shrubs.
volatile oil content increases (Cook and others III. Sunflower family. Research Paper INT-220.
1954). Budsage is especially important to grazing U.S Department of Agriculture, Forest Service,
animals during late winter in areas where there is Intermountain Forest and Range Experiment
an abundance of dry grass. Unfortunately, it does Station, Ogden, UT. 82 p.
not tolerate grazing well, and continual heavy
grazing may eliminate it from certain areas McArthur, E.D. and C.L. Pope. 1979. Karyotypes
(Holmgren and Hutchings 1972). In order to of four Artemisia species: A. carruthii, A.
maintain the species in plant communities, filifolia, A. frigida, and A. spinescens. Great
periodic rest from grazing is needed. This allows Basin Naturalist 39: 419-426.
seed and seedlings to develop.
Meyer, S. E. 2003. Artemisia. In: Woody plant
Benefits.—Budsage is an important native shrub seed manual. http://www.wpsu.net [not paged].
because of its forage value to various species of
wildlife and domestic livestock. Wildlife species Vallès, J. and E.D. McArthur. 2001. Artemisia
that utilize budsage include mule deer (Odocoileus systematics and phylogeny: cytogenetic and
heminonus Rafinesque), pronghorn (Antilocapra molecular insights. In: E.D. McArthur and D.J.
Americana Ord), mountain sheep (Ovis canadensis Fairbanks, comps. Shrubland ecosystem genetics
Shaw), black-tailed jackrabbits (Lepus californicus and biodiversity: proceedings, 2000 June 13-15;
Gray), and chucker partridge (Alectoris graeca Provo, UT. Proceedings RMRS-P-21. U.S
Meisner) (Wood and Brotherson 1986). In areas Department of Agriculture, Forest Service,
of local abundance, budsage can be an important Rocky Mountain Research Station, Ogden, UT.
source of protein during late winter months. p. 67-74.
References Wood, B. W. 1966. An ecological life history of

budsage in western Utah. M.S. thesis. Brigham
Cook, C.W., L.A, Stoddart, and L.E. Harris. 1954. Young University, Provo, UT. 85 p.
The nutritive value of winter range plants in the
Great Basin as determined with digestion trials Wood, B.W. and J.D. Brotherson. 1986.
with sheep. Bulletin 372. Utah State University, Ecological adaption and grazing response of
Agricultural Experiment Station, Logan, UT. budsage (Artemisia spinescens) in southwestern
56 p. Utah. In: McArthur, E.D. and B.L. Welch,
83
comps. Proceedings--symposium on the biology
of Artemisia and Chrysothamnus, 1984 July
9-13; Provo, UT. General Technical Report
INT-200. U.S. Department of Agriculture, Forest
Experiment Station, Ogden, UT. p. 75-92.
84
Artemisia tridentata Nutt. big sagebrush
COMPOSITAE
Synonyms: Artemisia tridentata var. angustifolia Gray

Artemisia angusta Rydb.
Artemisia tridentata ssp. typica Hall & Clements
Bruce L. Welch
6.5 cm long and 2 mm to 2 cm wide, and usually

are three-toothed at the apex (McArthur and others
1979). Leaf arrangement is spiral with a 2/5
divergence (Diettert 1938). Internodes are often so
short that the leaves form dense, rosette-like
clusters at the shoot tips (Diettert 1938). The
haploid chromosome number of big sagebrush is
nine (Diettert 1938, McArthur and Sanderson
1999). Two levels of ploidy have been reported
among the three subspecies of big sagebrush–
diploid (2n = 2x = 18) and tetraploid (2n = 4x =
36) (Diettert 1938, McArthur and Sanderson
1999). All three subspecies of big sagebrush
contain polyploid populations (McArthur and
Sanderson 1999).
Taxonomy.—There are three widely recognized

subspecies of big sagebrush (Beetle 1960, Beetle
and Young 1965). These are Wyoming big
sagebrush (A. t. ssp. wyomingensis), basin big
sagebrush (A. t. ssp. tridentata), and mountain big
sagebrush (A. t. ssp. vaseyana). Subspecies can be
separated on morphological, chemical, and
ecological characteristics (McArthur and others
1979, Welch 2002). Wyoming big sagebrush
occupies the drier sites that range from 20 to 32
cm of precipitation, 31 to 149 cm for mountain big
sagebrush with basin big sagebrush somewhat
between the other two subspecies (Welch 2002).
General Description.—Big sagebrush is an erect, Some authorities have divided mountain big
aromatic, evergreen species, that lacks winter sagebrush into three forms, varieties, or subspecies
buds. It has numerous ecotypes that range in size depending on the authority being quoted. One is
from 0.4 to 4.5 m tall (Beetle 1960, Beetle and called subalpine big sagebrush, A. t. ssp.
Young 1965, Diettert 1938, McArthur and others spiciformis, (McArthur and Goodrich 1986) or A.
1979). Growth forms are of two types; t. ssp. vaseyana f. spiciformis (Beetle and Johnson
multistemmed and single trunks and are correlated 1982). The second is referred to as “X” big
with subspecific taxonomy. This species produces sagebrush, or xeric big sagebrush or A. t. ssp.
two types of branches, vegetative and flowering xericensis (Welch 2002). Lastly, a variety of
(Diettert 1938). The bark on older vegetative mountain big sagebrush is called A. t. ssp.
branches is stringy and black or dark brown in vaseyana var. pauciflora (Welch 2002).
color. Bark on younger vegetative and
inflorescences is heavily covered with trichomes Range.—Beetle (1960) describes the range or
that give the branches a silvery green to gray color. distribution of big sagebrush in these terms:
The leaves are also silvery green or gray. Principal “occupying a great variety of sites at scattered
leaves are narrowly cuneate or oblanceolate, 1.0 to localities in southern Montana and central to
85
western Wyoming; southern-central British seed dispersal is highly variable among stands and
Columbia, south through central Washington, to somewhat dependent on elevation; some starting in
the Columbia River; from the south side of the mid-fall other not until early winter. Most seeds
Blue Mountains in northeastern Oregon, germinate immediately after snow melt or after
throughout central and southeastern Oregon, late-winter or early spring storms (Welch 2002).
southwestern and central Idaho to northern Utah, The seeds possess no special adaptations for wind
Nevada, and northwestern California; from the dispersal, but are wind dispersed to a maximum
northern great basin areas of California, Nevada, distance of 30 m from the mother plant. Hence
and Utah, extending southward at elevations from seedlings placement is heavily depended on wind
5,000 to 7,000 feet and extending into western direction (Welch 2002). Big sagebrush forms a
Colorado, northwestern New Mexico, northern weak soil seed bank (Welch 2002). A single
Arizona; at much lower elevations in southern mature big sagebrush plant can produced upward
California and northern Lower California, of 500,000 seeds, although, excessive browsing
Mexico.” It also occupies areas of the northern can reduce inflorescences production by a factor of
Great Plains in the States of North and South 20 (Welch 2002). The seeds must lie on the soil
Dakota, Montana, Wyoming, and Nebraska surface or be buried not more than 5 mm for
(Welch 2002). successful emergence (Welch 2002).
Ecology.—Big sagebrush grows in a variety of Growth and Management.—Vegetative growth

soils throughout its range from arid plains, valleys, starts in early spring and declines when soil
foothills, to mountain slopes, from 500 to 3,400 m moisture becomes deficient. In general, Wyoming
elevation (McArthur and others 1979, Welch big sagebrush, the smallest of the subspecies, has
2002). It has been found growing on five soil the slowest vegetative growth rate and produces
order: Alfisols, Aridisols, Entisols, Inceptisols, and the least amount of biomass. Basin big sagebrush
Mollisols. It can be found growing on all 12 soil has the fastest growth rate and is the largest of the
textural classes, but it is most often found on subspecies (McArthur and others 1979, Welch
loams or sandy loams (Welch 2002). Chemical 2002). In the past 50 years or more, management
properties of big sagebrush soils are highly has erroneously perceived big sagebrush to be a
variable (pH 5.9 to 10.0 and organic carbon 0.62 to highly competitive, dominating, and suppressive
4.14 percent) but two characteristics are common: species of grasses and forbs (Welch 2002). Thus
they are well-drained and contained low a variety of techniques have been developed to kill
concentrations of salts (Welch 2002). Depending big sagebrush (Welch 2002). As more studies
on subspecies, as many as 40 plant species show the importance of big sagebrush in
(grasses and forbs) can grow with big sagebrush maintaining it’s ecosystem, techniques have been
(Welch 2002). Big sagebrush is not fire tolerant and continue to be developed to restore big
and can be killed by a number of pathogenic fungi, sagebrush to its native sites (Welch 2002).
insects, and environmental conditions such as
winter kill or winter induced drought (Welch Benefits.—Big sagebrush is the keystone species
2002). of a large and highly variable ecosystem of
Western United States (Welch 2002). It is literally
Reproduction.—The growth of inflorescence a “nursing mother” to a host of organisms ranging
starts in late spring and is complete by late from microscopic to large mammals and is what
summer to early fall (Welch 2002). Flowering supports life in Bailey’s (1896) description: “One
occurs during early to late fall with fruit never recovers from his surprise that there should
development commencing almost simultaneously be so much life where apparently there is so little
with flowering. Thus reproductive growth occurs to support it.” Species that depend directly or
when water supplies and temperatures are not indirectly on big sagebrush are many: 31 species
particularly favorable; in fact, during this period of fungi, 52 species of aphids, 10 species of insects
big sagebrush plants have started shedding that feed on aphids, 42 species of midges and fruit
ephemeral leaves and neighboring plants species flies that induce galls, 20 species of insects that
are largely dormant (Welch 2002). However, parasitize the gall inducers, 6 species of insects
inflorescences do generate positive net that hibernate in big sagebrush galls, 18 species of
photosynthesis at a time when whole plant is in a beetles, 13 species of grasshoppers, 13 species of
water conservation mode (Welch 2002). Big shield-back katydids, 16 species of thrips, 74
sagebrush achenes or seeds are small, weighing species of spiders, 24 species of lichens, 16 species
from 0.00018 to 0.00025 g (Welch 2002). Time of of paintbrushes, 7 species of owl-clovers, 5 species
86
of bird’s beaks, 3 species of broom rapes, and a
host of large and small mammals, birds, and Krannitz, P.G. and M.M. Caldwell. 1995. Root
reptiles (Welch 2002). Soil fertility is higher growth responses of three Great Basin
under big sagebrush canopy cover than outside the perennials to intra-and inter specific contact with
canopy (Welch 2002). Roots of other plants can other roots. Flora. 190:161-167.
tap into these islands of fertility (Krannitz and
Caldwell 1995). Also, by a process known as McArthur, E.D., A.C. Blauer, A.P. Plummer, and
“hydraulic lift or hydraulic redistribution” R. Stevens. 1979. Characteristics and
associated plants receive part of their needs from hybridization of important Intermountain shrubs.
water pumped by big sagebrush to the soil surface III. Sunflower family. Research Paper INT-220.
from greater soil depths (Welch 2002). Lastly, big U.S. Department of Agriculture, Forest Service,
sagebrush provides a safe haven for grasses and Intermountain Forest and Range Experiment
forbs growing under its canopy from grazing Station, Ogden, UT. 82 p.
domestic livestock (Welch 2002).
McArthur, E.D. and S.K. Goodrich. 1986.
References Artemisia tridentate ssp. spiciformis:
distribution and taxonomic placement. In: E.D.
Bailey, W.W. 1896. The sage brush. American McArthur and B.L. Welch, comps. Proceedings:
Naturalist 3:356-360. Symposium on the biology of Artemisia and
Chrysothamnus; July 9-13, 1984; Provo, UT.
Beetle, A.A. 1960. A study of sagebrush–The Gen. Tech. Rep. INT-200. U.S. Department of
section Tridentatae of Artemisia. Bulletin 368. Agriculture, Forest Service, Intermountain
University of Wyoming, Agricultural Research Station, Ogden, UT. 55-57.
Experiment Station. Laramie. 83 p.
McArthur, E.D. and S.C. Sanderson. 1999.
Beetle, A.A. and K.L. Johnson. 1982. Cytogeography and chromosome evolution of
Sagebrush in Wyoming. B-779, University of subgenus Tridentatae of Artemisia (Asteraceae).
Wyoming, Agricultural Experiment Station, American Journal of Botany. 86:1,754-1,775.
Laramie. 68p.
Welch, B.L. [in press]. Big sagebrush: A sea
Beetle, A.A. and A. Young. 1965. A third fragmented into lakes, puddles, and ponds.
subspecies in the Artemisia tridentata complex. General Technical Report. U.S. Department of
Rhodora 67:405-406. Agriculture, Forest Service, Rocky Mountain
Research Station, Fort Collins, CO.
Diettert, R.A. 1938. The morphology of Artemisia
tridentata Nutt. Lloydia 1:3-74.
87
Artemisia tripartita Rydb. threetip sagebrush
ASTERACEAE
Synonyms: Artemisia trifida Nutt.

Seriphidium tripartitum (Rydb.) W.A.Weber
stems arising from the base. The bark on young

branches is canescent but becomes shredded and
grayish, light brown to dark brown or black, on
older stems. This species can layer and resprout
after fire or herbicide treatment (Beetle 1960,
Pechanec and others 1965, Schlatterer 1973,
Winward 1980). Leaves on vegetative branches are
canescent, 0.5 to 4 cm long, and typically deeply
divided into three linear or narrowly linear-
lanceolate lobes, which in turn may be three-cleft.
Upper leaves are often entire. Crushed foliage
emits a pungent odor. Flowerheads contain three to
11 disc flowers typically arranged into panicles.
Ray flowers are lacking. Eight to 12 canescent
involucral bracts subtend each head. Achenes are
resinous-granuliferous (Cronquist 1994, McArthur
and others 1979).
Taxonomy.—The taxon includes two subspecies:

Artemisia tripartita ssp. tripartita and Artemisia
tripartita ssp. rubicola Beetle. This species, as is
quite common for other species in the subgenus
Tridentatae, includes populations that are diploid,
2n = 18 and tetraploid, 2n = 36 (McArthur and
Sanderson 1999).
Subspecies tripartita, also known as tall
threetip sagebrush, is a freely-branching shrub up
to 1.8 m high. It can layer when conditions are
conducive, but it is seldom found layering in the
field. It can stump-sprout after fire (Beetle 1960).
Leaves on vegetative branches are 1.5 to 4 cm long
and deeply divided into three linear lobes, each
less than 1 mm wide, that may be further divided
(Beetle 1960). Flower heads bear four to eight disc
flowers arranged into panicles that may be reduced
to a spicate form.
Subspecies rubicola, also known as
Wyoming threetip sagebrush, is a dwarf shrub
usually less than 15 cm tall with decumbent
branches. It layers frequently and may have a
crown diameter of 30 to 50 cm (Beetle 1960).
Leaves on vegetative branches are often 3 cm long
and deeply divided into linear lobes, each at least 1
mm wide (Beetle 1959, 1960). Flower heads bear
General Description.—Threetip sagebrush is a
three to 11 disc flowers arranged in leafy,
round, evergreen shrub up to 1.8 m tall. It may
narrowly racemose panicles.
have a simple, trunk-like stem or several, lesser
88
Range.—Threetip sagebrush covers approximately sagebrush generally recovers well after
34,000 km2 in the Northern Rocky Mountains and disturbance, but hot summer fires can kill large
Great Basin States. It usually grows at elevations stands, inhibit resprouting, and significantly delay
between 910 to 2,700 m. It extends from British natural seedling recovery.
Columbia south through Montana and Wyoming to
Colorado, and west to Washington, Oregon, Benefits.—Three-tip sagebrush adds structural and
northern Nevada, and northern Utah (Beetle 1960). biological diversity to the landscape and reduces
Subspecies rubicola, however, is limited to areas the effects of wind and soil erosion. Although
of central and southeastern Wyoming and southern Beetle (1960) reported it was not palatable to
Oregon (Brunner 1972). livestock or game, Kufeld (1973) reported that elk
used it, and Kufeld and others (1973) also reported
Ecology.—Threetip sagebrush grows well on that deer use it. A form near Salmon, Idaho, is
moderate to deep, well-drained, loamy and sandy palatable to deer and may have introgressed into a
loam soils (Winward 1980). In some places, nearby population of Wyoming big sagebrush
particularly in Idaho, this species grows between (McArthur and others 1979).
the lower, hot, dry sites dominated by Wyoming
big sagebrush (Artemisia tridentata Nutt. ssp. References
wyomingensis Beetle & Young) and the higher,
cooler sites dominated by mountain big sagebrush Beetle, A.A. 1959. New names within the section
[Artemisia tridentata ssp. vaseyana (Rydb) Beetle] Tridentatae of Artemisia. Rhodora 61: 82-85.
(Schlatterer 1973). It often grows intermixed, but
as separate stands, with mountain big sagebrush, Beetle, A.A. 1960. A study of sagebrush. The
Wyoming big sagebrush, or alkali sagebrush section Tridentatae of Artemisia. Bulletin 368.
[Artemisia longiloba ([Osterh.) Beetle] in areas University of Wyoming, Agricultural
with 300 to 600 mm of annual precipitation. Experiment Station, Laramie, WY. 83 p.
Wyoming threetip sagebrush grows on rocky
knolls at high elevations (2,100 to 2,700 m) Brunner, J.R. 1972. Observations on Artemisia in
adjacent to mountain big sagebrush sites. Nevada. Journal of Range Management 25: 205-
208.
Reproduction.—Flowers bloom between July and
September and seed matures in October. Seeds are Cronquist, A. 1994. Asterales, Intermountain flora,
wind dispersed and seedlings emerge during the Vol. 5. The New York Botanical Garden, Bronx,
ensuing spring. Plants grow quickly and attain NY. 496 p.
maturity in 3 to 5 years. The seeds are smaller than
many Artemisia species; there are approximately 5 Kufeld, R.C. 1973. Foods eaten by the Rocky
million cleaned seeds/kg (Meyer 2003). Mountain elk. Journal of Range Management.
26: 106-113.
Management.—This species is a vigorous seeder.
Some forms are more palatable than others. Kufeld, R.C., O.C. Wallmo, and C. Feddema.
Threetip sagebrush establishes well by direct 1973. Foods of the Rocky Mountain mule deer.
seeding, rearing, or transplanting. Seedlings are Research Paper RM-11. U.S. Department of
able to persist with both a dense perennial and Agriculture, Forest Service, Rocky Mountain
annual understory, although it is no more effective Forest and Range Experiment Station, Fort
at excluding invasive annuals than other Collins, CO. 31 p.
sagebrushes. It frequently grows with a number of
other shrubs and herbs. It is compatible with many McArthur, E.D., A.C. Blauer, A.P. Plummer, and
introduced and native grasses including mountain R. Stevens 1979. Characteristics and
brome (Bromus carinatus H. & A.), slender hybridization of important intermountain shrubs.
wheatgrass [Elymus trachycaulus (Link) Gould ex III. Sunflower family. Research Paper INT-220.
Shinn], intermediate wheatgrass [Elymus hispidus U.S. Department of Agriculture, Forest Service,
(Opiz) Meld], and orchard grass (Dactylis Intermountain Forest and Range Experiment
glomerata L.). Station, Ogden, UT. 82 p.
Attempts to establish threetip sagebrush
on arid sites dominated by Wyoming big McArthur, E.D. and S.C. Sanderson. 1999.
sagebrush have been unsuccessful. Some ecotypes, Cytogeography and chromosome evolution of
however, can extend into drier sites. Threetip
89
subgenus Tridentatae of Artemisia (Asteraceae). Schlatterer, E.F. 1973. Sagebrush species and
American Journal of Botany 86: 1,754-1,775. subspecies. U.S. Department of Agriculture,
Forest Service, R-4 Range Improvement Notes
Meyer, S.E. 2003. Artemisia. In: F.T. Bonner, and 18(2): 1-11.
R.G. Nisley, eds. Woody plant seed manual.
http://www.wpsu.net [not paged]. Winward, A.H. 1980. Taxonomy and ecology of
sagebrush in Oregon. Bullitin 642. Oregon State
Pechanec, J. F., A.P. Plummer, J.H. Robertson, University, Oregon Agricultural Experiment
and A.C. Hull, Jr. 1965. Sagebrush control on Station, Corvallis, OR. 15 p.
rangelands. Agriculture. Handbook 277. U.S.
Department of Agriculture, Washington, DC.
40 p.
90
Arundinaria gigantea (Walt.) Muhl. giant cane
POACEAE
Synonyms: Arundinaria macrosperma Michx.

Arundinaria tecta (Walter) Muhl.
Arundinaria tecta var. decidua Beadle
Arundo gigantea Walter
Arundo tecta Walter
Kristina Connor
General Description.—Giant cane, also known as

cane or switchcane, is a perennial monocot, a
woody grass, and one of only two native bamboos.
With its stem-like rhizomes and hard, ‘woody’
stems, giant cane can grow to a height of 8 to 9 m
but is typically less. Lower sheaths are about half
as long as the culm internodes, while the upper six
to 10 sheaths culminate in 10 to 12 bristles that are
5 to 9 mm long (Hitchcock 1971). Leaves range
from 10 to 30 cm in length and from 2 to 3 cm
wide. They have rounded base, are lance shaped,
and have finely toothed margins (Bailey and
Bailey 1976). Leaves are typically pubescent
underneath and are either pubescent or glabrous
above. Leaves on stems 2 years and older grow on
side branches. Branches are short, less than 30 cm,
also with loose papery sheaths.
Range.—Giant cane is found at elevations ranging

from sea level in southern floodplains to 610 m
elevation in the Appalachian Mountains. It can
grow in dense thickets in the Mississippi Delta and
in other southern swamplands. Its range extends
from southern Maryland west into Ohio, Indiana,
and Illinois, south to the Gulf Coast and west to
Texas, Oklahoma, and Arkansas (Bailey and
Bailey 1976, Brickell and Zuk 1996, Walkup
1991). A recognized variety is A. gigantea subsp.
tecta (Walter) McClure.
Ecology.—Giant cane formerly occupied large

areas (canebrakes) in floodplains of southern
rivers; now these thickets are usually found only in
the Mississippi Delta where they form in low-
lying, shady moist areas. Elsewhere, giant cane is
usually intermixed with shrubs. It is fire dependent
and resprouts from rhizomes (Walkup 1991). It has
a broad tolerance for weather and can withstand
temperatures ranging from -23 to 41 oC. It grows
in a variety of soil types (muck lands to mountain
Illustration source: Grelen and Duvall 1966 slopes and rich alluvial soils) and is rugged, cold-
hardy and adaptable.
91
Reproduction.—Giant cane primarily spreads by References
rapid vegetative reproduction from large rhizomes.
It flowers infrequently, and at irregular intervals, Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third:
in early spring, forming simple panicles on the A Concise Dictionary of Plants Cultivated in the
branches. The panicles consist of a few racemes of United States and Canada. McMillan Publishing
five to 15 large, stalked spikelets, which may be Co., Inc. New York. 1,290 p.
purple (Bailey and Bailey 1976, Brickell and Zuk
1996). The inflorescence may be a single axis or Brickell, C. and J.D.. Zuk. 1996. The American
have branches. Seed production from these flowers Horticultural Society A-Z Encyclopedia of
is sparse and unpredictable. Garden Plants. DK Publishing, Inc., New York.
1,092 p.
Growth and Management.—Giant cane is a
short-lived, evergreen monocot. Individual stems Grelen, H.E. and V.L. Duvall. 1966. Common
survive less than 10 years, but the species sprouts plants of longleaf pine-bluestem range. Res. Pap.
prolifically from its rhizomes. In the presence of S0-23, U.S. Department of Agriculture Forest
fire and on the best-suited sites, it forms dense Service, Southern Forest Experiment Station,
thickets called canebrakes. While it can withstand New Orleans, LA. 96 p.
flooding, drought, and intense surface fires, giant
cane is sensitive to overgrazing (Grelen and Hitchcock, A.S. 1971. Manual of Grasses of the
Duvall 1966, Hitchcock 1971). One source reports United States. Vol. 1, 2nd Ed. Revised by A.
rapid growth (Brickell and Zuk 1996), especially Chase. Dover Publications Inc., New York. 569
after a fire. Fuel accumulates rapidly in a p.
canebrake, and, to eliminate the hazard, a short
burning cycle is advised (Hughes 1966, Walkup Hughes, R.H. 1966. Fire ecology of canebrakes.
1991). In: Proceedings 5th Annual Tall Timbers Fire
Ecology Conference, Tallahassee, FL. Tall
Benefits.—The dense thickets of giant cane Timbers Research Station. p. 148-158.
provide cover for nesting birds and small
mammals. In addition, the young shoots are edible, University of Florida. 2001. Website, Center for
sometimes used as a potherb (Hitchcock 1971), Aquatic and Invasive Plants. http://aquat1.ifas.
and of good nutritional quality. The leaves are a ufl.edu/arugig.html. 1 p.
preferred food for southern pearly eye butterfly
caterpillars (University of Florida 2001). The USDA, NRCS. 2001. The PLANTS database,
leaves extend well above the ground, plants can be National Plant Data Center, Baton Rouge, LA.
completely defoliated by cattle and are also http://plants.usda.gov. [not paged].
uprooted by swine (Hitchcock 1971, Walkup
1991). Thus, while grazing capacity of cane is Walkup, C. 1991. Arundinaria gigantea. In: U.S.
high, careful management is required to prevent Department of Agriculture, Forest Service, Rocky
deterioration of the plants (Grelen and Duvall Mountain Research Station Fire Sciences
1966). Laboratory. Fire Effects Information System.
http://www.fs.fed. us/database/feis. 9 p.
92
Atriplex canescens (Pursh) Nutt. fourwing saltbush
CHENOPODIACEAE
Synonyms: Calligonum canescens Pursh
E. Durant McArthur, Stewart C. Sanderson, and Jeffrey R. Taylor

borne in glomerules 2 to 3 mm wide (Blauer and
others 1976). Sexual expression varies between
populations and with ploidy level. Diploids are
usually monoecious, with a few exceptions. At
higher ploidy levels (tetraploids and hexaploids),
fourwing saltbush is trioecious (three sexual
states). In the trioecious condition, some plants are
consistently female and others are consistently
male and some are able to switch during stressful
conditions from the less physiologically taxing
female state to the male state (McArthur 1977,
McArthur and others 1992). Gender differences
appear to be adaptive and allow for greater species
radiation (Freeman and others 1993).
Taxonomy.—Fourwing saltbush is composed of a

number of polyploid races with a base chromosome
number of x = 9 (Sanderson and Stutz 1994, 2001).
Many have been named as varieties. It appears that
many of these are autopolyploids (Stutz 1978). The
common form of fourwing saltbush in the
Intermountain region (race Occidentalis) is
tetraploid (Sanderson and Stutz 2001). Of the
several distinct diploids the most widespread are
race Angustifolia of the Chihuahuan Desert and
race Linearis of the Sonoran Desert. Hexaploids
are also generally southern in distribution (Nevada,
Illustration source: USDA—Forest Service collection, New Mexico and Mexico) (Sanderson and Stutz
Hunt Institute
2001). Polyploids higher than 6x (up to 20x) are
usually restricted endemics (Sanderson and Stutz
General Description.—Fourwing saltbush, also
2001).
known as costilla de vaca and chamizo, is an
The North American perennial saltbushes
upright shrub between 10 and 200 cm tall. Stature
form a group of related species. Fourwing saltbush
varies with site conditions and genotype. Branches
apparently hybridizes, at least to some extent, with
arise freely from the base and are brittle and hard at
all of these species (Blauer and others 1976, Stutz
maturity. Young branches and leaves are covered
1978).
with inflated or collapsed bladder hairs that create
a scurfy, grey-green appearance. Leaves are
Range.—Fourwing saltbush is a widespread shrub
evergreen, linear to oblanceolate or spatutulate,
throughout much of Western North America. It
usually 1 to 4 cm long and 0.1 to 1.0 cm wide. The
grows from the Great Plains to the Pacific Coast
species is usually dioecious, although some are
and from Canada to Mexico at elevations ranging
monoecious or trioecious. Pistillate (female)
from below sea level to 2,400 m. Fourwing
flowers are small and inconspicuous. A pair of
saltbush has been planted in various parts of
small bracts united along their edges encloses each
Eurasia and Australia but naturalization has not
pistil. Each of these bracts has two ridges down the
been documented.
face that expand into “wings,” giving the utricle
(fruit) four “wings” at maturity. Staminate (male)
Ecology.—Fourwing saltbush is adapted to desert
flowers are small, yellow to red to brown, and
climates. Mean annual precipitation within its
93
distribution usually ranges between 150 and 400 and grows rapidly (Cibils 1998, McArthur and
mm. It grows on deep, usually halophytic, well- others 1983, Peterson and others 1987). Leaves,
drained sandy soils, gravelly washes, mesas, stems, and fruits provide browse throughout the
ridges, and slopes. Soil salts enhance growth of year. Winter protein content has been shown to
fourwing saltbush by increasing days to wilting, vary between accessions (Welch and Monsen
organic matter production, water use efficiency, 1981). Relative saponin concentrations may affect
and ability to extract water by means of osmotic palatability. In mixed species communities,
adjustments (Glenn and Brown 1998). It often herbage production of fourwing saltbush and forbs
grows intermixed with a wide variety of shrubs, remains quite high. Fourwing saltbush is a
grasses and forbs. facultative selenium absorber, making it mildly
poisonous to browsing animals in selenium-rich
Reproduction.—Flowering generally occurs soils (Davis 1972).
between May and September (Blauer and others When propagating fourwing saltbush, it is
1976). This period can vary, however, with important to select ecotypes that are adapted to the
genotype and location. Flowers are wind site. If possible, seed should be from a source with
pollinated, and seed matures in October. Most a similar climate unless the strain being planted is
wildland stands of fourwing saltbush produce known to have a broad ecological range. Climatic
abundant seed 3 out of every 5 years. and biological factors may affect plant persistence.
Seed fill in high quality lots may only Plants established from southern seed sources have
reach 50 percent, although rates of 40 percent or not persisted in colder northern areas. However,
lower are generally considered substandard (Meyer seed collected from colder areas have done well at
2003). Like many species of saltbush, Atriplex warmer sites (McArthur and others 1983, Plummer
canescens exhibits high levels of dormancy and others 1966).
controlled by multiple and complex mechanisms
(Meyer 2003). Woody bracts inhibit germination Benefits.—Fourwing saltbush is a valuable plant
mechanically and also contain water-soluble that provides livestock and wildlife habitat and
substances, salt and perhaps saponin, which inhibit food. It has been widely and successfully used in
germination (Clor and others 1989). Seed normally revegetation and restoration projects including soil
requires 20 to 30 days of stratification to assure stabilization (Blauer and others 1976, Plummer and
uniform germination. There are about 120,000 others 1966). Its ability to adapt and to hybridize
cleaned, dewinged seeds (utricles)/kg (Blauer and with other woody Atriplex can facilitate in situ
others 1976). selection during revegetation (Stutz 1982).
Artificial seeding should take place during
the fall or winter unless spring frosts threaten References
seedling survival. Fall and winter seeding promotes
seed stratification and overall germinability. When Barrow, J.R. 1997. Natural asexual reproduction
sown, seeds should remain within the top 1.25 cm in saltbush Atriplex canescens (Pursh) Nutt.
of soil. Broadcast seeding followed by chaining Journal of Arid Environments 36: 267-270.
has produced excellent stands. Shrubs can also be
established using drills alone or in separate rows Blauer, A.C., A.P. Plummer, E.D. McArthur, R.
among other species. Care should be taken to keep Stevens, and B.C. Giunta. 1976. Characteristics
herbs from displacing fourwing saltbush seedlings. and hybridization of important Intermountain
Seedlings grow vigorously and quickly produce a shrubs. II. Chenopod family. Research Paper
dense canopy although interspecific competition INT-177. U.S. Department of Agriculture, Forest
can reduce stand densities. Service, Intermountain Forest and Range
Shrubs can also be grown from nursery Experiment Station, Ogden, UT. 42 p.
stock, container stock, or by stem cuttings (Shaw
and Monsen 1984). Rooted stem cuttings have Cibils, A.F., D.M. Swift, and E.D. McArthur.
been used successfully to establish shrub seed 1998. Plant-herbivore interactions in Atriplex:
orchards with desirable traits (McArthur and others current state of knowledge. General Technical
1978, 1992). Some populations reproduce Report RMRS-GTR-14. U.S. Department of
asexually through root sprouting (Barrow 1997). Agriculture, Forest Service, Rocky Mountain
Research Station, Ogden, UT. 31 p.
Management.—Fourwing saltbush is a valuable
forage shrub because it is abundant, palatable, Clor, M.A., F.M.R. Al-Charachafchi, and N.
provides large quantities of forage, is nutritious, Mahmood. 1989. Seed dormancy and
94
germination inhibition of Atriplex canescens. Plummer, A.P., S.B. Monsen, and D.R.
Journal of Agriculture and Water Resources Christensen. 1966. Fourwing saltbush, a shrub
Research Plant Production 8: 55-64. for future game ranges. Publication 66-4. Salt
Lake City, UT: Utah State Department of Fish
Davis, A.M. 1972. Selenium accumulation in a and Game. 12 p.
collection of Atriplex species. Agronomy
Journal. 64: 823-824. Sanderson, S.C. and H.C. Stutz. 1994.
Chromosome numbers in Mojave and Sonoran
Freeman, D.C., E.D. McArthur, S.C. Sanderson, Desert Atriplex canescens (Chenopodiaceae).
and A.R. Tiedemann. 1993. The influence of American Journal of Botany 81: 1045-1053.
topography on male and female fitness
components of Atriplex canescens. Oecologia Sanderson, S.C. and H.C. Stutz. 2001.
93: 538-547. Chromosome races of fourwing saltbush
(Atriplex canescens), Chenopodiaceae. In: E.D.
Glenn, E.P. and J.J. Brown. 1998. Effects of salt McArthur and D.J. Fairbanks, comps.
levels on the growth and water use efficiency of Proceedings—shrubland ecosystem genetics and
Atriplex canescens (Chenopodicaceae) varieties biodiversity; 2000 June 13-15; Provo, UT. Proc.
in drying soil. American Journal of Botany 85: RMRS-P-21. U.S. Department of Agriculture,
10-16. Forest Service, Rocky Mountain Research
Station, Ogden, UT. p. 75-88.
McArthur, E.D. 1977. Environmentally induced
changes of sex expression in Atriplex canescens. Shaw, N. and S.B. Monsen. 1984. Nursery
Heredity 38: 97-193. propagation and outplanting of bareroot
chenopod seedlings. In: A.R. Tiedemann, E.D.
McArthur, E.D., D.C. Freeman, L.S. Luckinbill, McArthur, H.C. Stutz, R Stevens, and K.L.
S.C. Sanderson, and G.L. Noller. 1992. Are Johnson, K., comps. Proceedings-symposium on
triocy and sexual morphs of Atriplex canescens the biology of Atriplex and related chenopods,
genetically based: evidence from clonal studies. 1983 May 2-6; Provo, UT. Gen. Tech. Rep.
Evolution 46: 1708-1721. INT-172. U.S. Department of Agriculture, Forest
McArthur, E.D., A.P. Plummer, G.A. Van Epps, Experiment Station, Ogden, UT. p. 251-260.
D.C. Freeman, and K.R. Jorgensen. 1978.
Producing fourwing saltbush seed in seed Stutz, H.C. 1982. Broad gene pools required for
orchards. In: Hyder, D. N., ed. Proceedings of disturbed lands. In: E.F. Aldon and W. Oaks W.
the First International Rangeland Congress; eds. Reclamation of mined lands in the
1978 Aug. 14-18; Denver, CO. Denver, CO: southwest: Proceedings of the symposium; 1982
Society for Range Management: 406-410. Oct. 20-22; Albuquerque, NM. Soil
Conservation Society of America, New Mexico
McArthur, E.D., R. Stevens, and A.C. Blauer.
Chapter, Albuquerque, NM. p.113-118.
1983. Growth performance comparisons among
18 accessions of fourwing saltbush (Atriplex
Stutz, H.C. 1978. Explosive evolution of perennial
canescens) at two sites in central Utah. Journal
Atriplex in western America. Great Basin
of Range Management. 36: 78-81.
Naturalist Memoirs. 2: 161-168.
Meyer, S.E. 2003. Atriplex L. In: Woody Plant Welch, B.L. and S.B. Monsen. 1981. Winter crude
Seed Manual, http://www.wpsm.net [not paged]. protein among accessions of fourwing saltbush
grown in a uniform garden. Great Basin
Peterson, J.L., D.N. Ueckert, R.L. Potter, and J.E. Naturalist. 41: 343-346.
Huston. 1987. Ecotype variation in western
Texas fourwing saltbush. Journal of Range
Management. 40: 361-366.
95
Atriplex confertifolia (Torr. & Frém.) S. Wats. shadscale
CHENOPODIACEAE
Synonyms: Obione confertifolia Torr. & Fremont

Atriplex collina Woot. & Standl.
Stewart C. Sanderson, E. Durant McArthur, and Jeffrey R. Taylor

Taxonomy.—Shadscale includes several
chromosome races ranging from diploid (2n = 2x
= 18) to decaploid (2n = 10x = 90). Plant
similarities and the paucity of close relatives
suggest that these races are largely autoploid
(Stutz and Sanderson 1983). Of all the races,
tetraploids are most common (Sanderson and
others 1990). Generally, polyploid populations
occur in uniform stands at lower elevations and on
valley floors while diploid forms occur on uplands
in more diverse communities and are usually
larger in stature (Sanderson and others 1990, Stutz
and Sanderson 1983).
Range.—Shadscale is present throughout much of

Western North America, from Montana in the
north to Texas and Chihuahua in the south, and
from North Dakota in the east to California and
Oregon in the west (Stutz and Sanderson 1983). It
usually grows at elevations from 460 to 2,100 m.
Among the woody chenopods, it ranks next to
fourwing saltbush [Atriplex canescens (Pursh)
Nutt.] and winterfat [Ceratoides lantata (Pursh) J.
T. Howell] in frequency and distribution.
Ecology.—Shadscale usually grows on fine-

Illustration source: USDA—Forest Service collection, textured alkaline soils, although it can be found on
Hunt Institute coarser, sandier soils (Blaisdell and Holmgren
1984, McArthur and others 1978). It can occur in
General Description.—Shadscale is a compact nearly pure stands, but also grows intermixed with
spinescent shrub that is typically 15 to 120 cm tall many other shrubs, especially of the chenopod
and 30 to 170 cm wide. Branches are scruffy when and sunflower families, and with junipers
young but become rigid, brittle, and spiny with (Juniperus L.) and grasses. It tolerates alkali soils
age. Leaves are nearly circular to elliptic, oval, or better than most of these associates (McArthur and
oblong, 9 to 25 mm long and 4 to 20 mm wide others 1978). Relationships between microhabitat,
(Blauer and others 1976). Flowers are imperfect. synecology, and physiology have explained
Staminate (male) flowers are arranged in differences in distribution patterns within
glomerate inflorescences borne in leaf axils. intermixed stands of shadscale and winterfat
Pistilate (female) flowers are enclosed by foliose (Caldwell and others 1977, West and Gasto 1978).
bracts, 5 to 12 mm long, broadly ovate to almost Shadscale grows slowly, has a relatively short life
round, united at the base, and have entire, free, span and can be killed by drought or high water
somewhat spreading margins. Staminate flowers (Blaisdell and Holmgren 1984, McArthur and
are small and inconspicuous. Populations may others 1978). High water can kill plants directly,
have biased sex ratios and can include monocieous lead to anaerobiosis, induce abiotic disease, or
as well as dioecious individuals (Freeman and predispose plants to biotic pathogens, all of which
McArthur 1984). may have contributed to die-offs over the past
decades (Wallace and Nelson 1990). Shadscale
96
typically grows in arid climates where mean fruits have fallen in autumn. Grazing pressure can
annual precipitation can range between 180 and reduce more palatable shrub species, such as
360 mm. budsage and blacksage, resulting in increased
shadscale populations (Blaisdell and Holmgren
Reproduction.—Shadscale usually blooms during 1984). Shadscale may be more effective in
April and May. In isolated locations populations restoring and stabilizing salty areas than fourwing
may bloom as early as January and as late as June. saltbush or other saltbushes because it is more salt
Utricles mature, in general about mid-October. tolerant (McArthur and others 1978). In large
Shadscale seeds are largely dormant after stands, shadscale should be managed rather than
maturation but respond to cold treatment (chilling) mechanically controlled. Managing for shadscale
through dry after-ripening (Meyer and others is easier than reducing shrub densities and seeding
1998). Chilling and leaching soluble inhibitors in new species because limited soil moisture and
from seeds has been shown to increase precipitation make it difficult to manipulate these
germination (Garvin and others 1996). plant communities (Bleak and others 1965).
Interestingly, seeds from warm desert populations Aggressive annuals, such as cheatgrass, usually
responded to chilling more and after-ripened invade such areas if they are disturbed.
quicker than those from cold deserts (Meyer and
others 1998). Ecotypic germination variation is Benefits.—Shadscale is a genetically and
common in shadscale. morphologically diverse native shrub that occupies
Shadscale spreads naturally through seed a considerable range throughout the West. It
but is difficult to establish by artificial seeding provides soil cover, habitat for birds, and forage
(McArthur and others 1978, Monsen and for domestic livestock and wildlife. Some small
Richardson 1984). Low seed fill often compounds passerine birds, such as horned lark (Eremophila
seed dormancy and germination problems. Fall alpestris), Brewer’s sparrow (Spizella breweri),
seeding is recommended because it reduces the and sage thrashers (Oreoscoptes montanus) nest in
effects of dormancy, and utricles are more likely to shadscale stands (Medin 1990). Healthy stands
break down before spring. Seeds should remain also reduce the proliferation of exotic, undesirable
between 6 and 13 mm of the soil surface to annuals. In addition, seeds were historically
improve the likelihood of germination and ground into flour by Native-Americans for bread
establishment. Soil type and seedbed preparations (Stubbendieck and others 1997).
should be considered prior to seeding. Seed should
be planted at rates of 2.2 to 4.5 kg of pure live References
seed/ha and in separate rows from other seeded
species. Seedlings grow slowly and compete Blaisdell, J.P. and RC. Holmgren. 1984. Managing
poorly with herbs and aggressive grasses, Intermountain rangelands-salt-desert shrub
particularly within the first 2 years. Diploid races ranges. General Technical Report INT-163. U.S.
from pinyon-juniper woodlands and big sagebrush Department of Agriculture, Forest Service,
communities establish more readily in biologically Intermountain Forest and Range Experiment
diverse communities than other races. Station, Ogden, UT. 52 p.
Shadscale has also been successfully
transplanted when moisture conditions are right Blauer, A.C., A.P. Plummer, E.D. McArthur, R.
during spring or fall (Luke and Monsen 1984). Stevens, and B.C. Giunta. 1976. Characteristics
Transplants can be from container or bare-root and hybridization of important Intermountain
stock. Survival is usually poor unless care is taken shrubs. II. Chenopod family. Research Paper
during handling and planting. Young shrubs are INT-177. U.S. Deptment of Agriculture, Forest
particularly susceptible to overwatering. Service, Intermountain Forest and Range
Transplanting is most successful if plants are Experiment Station, Ogden, UT. 42 p.
dormant at the time of planting.
Bleak, A.T., N.C. Frischknecht, A.P. Plummer,
Management.—Despite its spiny nature, and R.E. Eckert, Jr. 1965. Problems in artificial
shadscale is grazed by livestock and game. Forage and natural revegetation of the arid shadscale
value is directly related to leaf persistence, which vegetation zone of Utah and Nevada. Journal of
is a characteristic that varies by habitat. Many Range Management. 18: 59-65.
species prefer the nutritious seeds (USDA 1937).
Livestock and wildlife tend to seek seed and leaf Caldwell, M.M., R.S. White, R.T. Moore, and L.B.
accumulations under parent shrubs after leaves and Camp. 1977. Carbon balance, productivity, and
97
water use by cold desert shrub communities Monsen, S.B. and B.Z. Richardson. 1984, Seeding
dominated by C3 and C4 species. Oecologia. 29: shrubs with herbs on a semiarid mine site with
275-300. and without topsoil. In: A.R. Tiedemann, E.D.
McArthur, H.C. Stutz, R. Stevens, and K.L.
Freeman, D. C. and E.D. McArthur. 1984. The Johnson, comps. Proceedings: Symposium on
relative influences of mortality nonflowering, the biology of Atriplex and related chenopods,
and sex change on the sex ratios of six Atriplex 1983 May 2-6; Provo, UT. Gen. Tech. Rep.
species. Botanical Gazette 145: 385-394. INT-172. U.S. Department of Agriculture,
Forest Service, Intermountain Forest and Range
Garvin, S.C., S. E. Meyer, and S. L. Carlson. 1996. Experiment Station, Ogden, UT. p. 298-305.
Seed germination studies in Atriplex confertifolia
(Torr. & Frem.) Wats. In: J.R. Barrow, E.D. Sanderson, S.C., H.C. Stutz, and E.D. McArthur
McArthur, R.E. Sosebee, and R.J. Tausch, 1990. Geographic differentiation in Atriplex
comps. Proceedings: shrubland ecosystem confertifolia. American Journal of Botany 77:
dynamics in a changing environment. 1995 May 490-498.
23-25; Las Cruces, NM. General Technical
Report INT-GTR-338. U.S. Department of Stubbendieck, J., S.L Hatch, and C.H. Butterfield
Agriculture, Forest Service, Intermountain 1997. North American Range Plants. Fifth
Research Station, Ogden, UT. p. 165-169. Edition. University of Nebraska Press, Lincoln,
NB. 305 p.
Luke, F. and S.B. Monsen. 1984. Methods and
costs for establishing shrubs on mined lands in Stutz, H.C. and S.C. Sanderson. 1983.
southwestern Wyoming. In: A.R. Tiedemann, Evolutionary studies of Atriplex: chromosome
E.D. McArthur, H.C. Stutz, R. Stevens, and K.L. races of A. confertifolia (shadscale). American
Johnson, comps. Proceedings-symposium on the Journal of Botany 70: 1536-1547.
biology of Atriplex and related chenopods, 1983
May 2-6; Provo, UT. Gen. Tech. Rep. INT-172. U.S. Department of Agriculture, Forest Service.
U.S. Department of Agriculture, Forest Service, 1937. Range Plant Handbook. U.S. Government
Intermountain Forest and Range Experiment Printing Office, Washington, DC. 816 p.
Station, Ogden, UT. p. 286-292.
Wallace, A. and D.L. Nelson. 1990. Wildland
McArthur, E.D., A.P. Plummer, and J.N. Davis. shrub dieoffs following excessively wet periods:
1978. Rehabilitation of game range in the salt a synthesis. In: E.D. McArthur, E.M. Romney,
desert. In: K.L Johnson, ed. Wyoming S.D. Smith, and P.T. Tueller, comps.
shrublands: proceedings 7th Wyoming shrub Proceedings: Symposium on cheatgrass
ecology workshop; 1978 May 31-June 1; Rock invasion, shrub die-off, and other aspects of
Springs, WY. University of Wyoming, Division shrub biology and management; 1989 April 5-7;
of Range Management, Laramie, WY. p. 23-50. Las Vegas NV. General Techical Report INT-
276. Ogden, UT. U.S. Department of
Medin, D.E. 1990. Birds of a shadscale (Atriplex Agriculture, Forest Service, Intermountain
confertifolia) habitat in east central Nevada. Research Station. p. 84-90.
Great Basin Naturalist 50: 295-298.
West, N.E. and J. Gasto. 1978. Phenology of the
Meyer, S.E., S.L. Carlson, and S.C. Garvin. 1998. aerial portions of shadscale and winterfat in
Seed germination regulation and field seed bank Curlew Valley, Utah. Journal of Range
carryover in shadscale (Atriplex confertifolia: Management. 31: 43-45.
Chenopodiaceae). Journal of Arid Environments
38: 255-267.
98
Atriplex corrugata S. Wats. mat saltbush
CHENOPODIACEAE
Synonyms: Atriplex nuttallii var. corrugata A. Nelson

Obione corrugata Ulbr.
Taxonomy.—This species will hybridize with

other saltbushes (Blauer and others 1976, Stutz
1978, 1984). Mat saltbush forms intermediates
with Atriplex confertifolia Torr. & Fremont and
Atriplex cuneata A. Nelson (Welsh and others
1993). Plants can be either diploid (2n = 18) or
tetraploid (2n = 36) (Sanderson and Stutz,
unpublished data).
Range.—Mat saltbush is an endemic of the

Colorado Plateau and Uintah Basin, found mainly
on Mancos shale formations in eastern Utah,
western Colorado, and northwestern New Mexico
between 1,200 and 2,100 m (Hall and Clements
1923; Hanson 1962).
Ecology.—Mat saltbrush grows on fine textured,

often saline, soils. It tolerates up to 13,000 ppm
soluble salts and is often the only perennial plant
present in such high-saline environments (Hanson
1962). Although it is probably the most halophytic
member of the genus, in less saline habitats,
patches of mat saltbush may alternate with patches
of winterfat [Ceratoides lanata (Pursh)
General Description.—Mat saltbush, also known
J.T.Howell], cuneate saltbush (Atriplex cuneata A.
as matscale, is a low shrub that forms dense,
Nelson), shadscale (Atriplex confertifolia Torr. &
prostrate (15 cm tall) and nearly white mats five to
Fremont), black greasewood (Sarcobatus
20 times wider than they are tall. Prostrate
vermiculatus Torr.), budsage (Artemisia
branches often form adventitious roots when in
spinescens Eaton), and greenmolly summercypress
contact with the soil. The bark is soft, spongy, and
(Kochia americana S. Wats.). It grows in areas
white. Leaves are sessile, evergreen, oblong,
with 150 to 280 mm of annual precipitation.
linear-oblancelate, or spatulate, rounded at the
apex, opposite on the lower portions of stems, and
Reproduction.—Blooming occurs between March
alternate above. Leaf blades are densely scurfy and
and May and fruits (utricles) ripen 6 to 10 weeks
7 to 18 mm wide (Blauer and others 1976,
later. Seed production is usually modest, although
McArthur and others 1978, Welsh and others
some seed is usually produced even during
1993). Individual shrubs are commonly dioecious
drought.
but may be monoecious. Staminate (male) flowers
Because germination is near zero without
are yellow to light-brown, 3 to 6 mm wide, and
cold treatment, fall is normally the recommended
borne in glomerulate spikes. Pistilate (female)
season for artificial seeding. Where possible,
flowers are enclosed by sessile or subsessile
midwinter planting is also successful. Preparatory
fruiting bracts. Fruiting bracts are 3 to 5 mm long,
measures are usually unnecessary if the site
4 to 6 mm wide, and united along two-thirds of
previously supported mat saltbush and is not
their length and densely tuberculate on the lower
infested with weeds. Recommended seeding rates
one-third (Blauer and others 1976, McArthur and
range between 2.2 and 4.5 kg of pure live seed/ha.
others 1978, Welsh and others 1993).
Seed should be sown in separate rows and covered
99
with a drag or harrow. Seeds should be planted References
near the soil surface and no more than 1.3 cm
deep. Seedlings emerge rapidly and grow Blauer, A.C., A.P. Plummer, E.D. McArthur, R.
vigorously during the first year. Growth continues Stevens, and B.C. Giunta. 1976. Characteristics
until soil moisture becomes limiting (usually by and hybridization of important Intermountain
mid-summer). There are about 220,000 cleaned shrubs. II. Chenopod family. Research Paper
seed per kg (Blauer and others 1976). INT-177. U.S. Department of Agriculture, Forest
Transplants from bare root stock, Service, Intermountain Forest and Range
wildings, or containers have all established Experiment Station, Ogden, UT. 42 p.
successfully. Wildings and bare root stock tolerate
considerable distress. Transplants survive Hall, H.M. and F.E. Clements. 1923. The
surprisingly well, even when planted on seemingly phylogenetic method in taxonomy, the North
infertile soils. Young plants have been American species of Artemisia, Chrysothamnus,
successfully transplanted into former black and Atriplex. Publication 326. Carnegie
greasewood communities in the Great Basin Institution of Washington, Washington, DC. 355
(Blauer and others 1976). p.
Management.—Mat saltbush is valuable winter Hanson, C.A. 1962. Perennial Atriplex of Utah and
forage for wildlife and domestic livestock. It often the northern deserts. M.S. thesis. Brigham
grows interspersed among other plant communities Young University, Provo, UT. 133 p.
on shale outcrops at medium elevations. Shrubs
often grow on windblown slopes and provide McArthur, E.D., A.P. Plummer, and J.N. Davis.
important, accessible winter forage. Big game 1978. Rehabilitation of game range in the salt
have been known to congregate on these areas desert. In: K.L. Johnson, ed. Wyoming
during periods of deep snow accumulation. Shrubs shrublands: proceedings of the 7th Wyoming
tolerate grazing and trampling and maintain vigor shrub ecology workshop; 1978 May 31-June1;
even after serious abuse. Rock Springs, WY. University of Wyoming,
Mat saltbush has been seeded with Division of Range Management, Laramie, WY.
selected grasses including Russian wildrye p. 23-50.
(Elymus junceus Fisch.), tall wheatgrass
[Agropyron elongatum (Host) P.Beauv.], and Sanderson, S.C., and H.C. Stutz. 2003.
crested wheatgrass [Agropyron cristatum (L.) Unpublished data. U.S. Department of
Gaertn.]. Although these species may increase Agriculture, Forest Service, Rocky Mountain
forage production, they do not usually persist in Research Station, Provo, UT
dense stands. Grasses and forbs will suppress
shrub establishment, but grasses interseeded into Stutz, H.C. 1978. Explosive evolution of perennial
established shrub stands do not reduce shrub Atriplex in western America. Great Basin
density or vigor. Naturalist Memoirs. 2: 161-168.
This species is useful in revegetation and
restoration projects and can help reclaim disturbed Stutz, H.C. 1984. Atriplex hybridization in western
areas with heavy textured soils such as road North America. In: A.R. Tiedemann, E.D.
construction sites or mine disturbances. McArthur, H.C. Stutz, R. Stevens, and K.L.
Johnson, comps. Proceedings: Symposium on
Benefits.—Mat saltbush is a native shrub that the biology of Atriplex and related chenopods,
occupies a specialized niche in Western 1983 May 2-6; Provo, UT. General. Technical
ecosystems. Its ability to grow on fine textured, Report INT-172. U.S. Department of
often saline soils helps reduce soil erosion on sites Agriculture, Forest Service, Intermountain Forest
that are inhospitable to many shrubs. It also and Range Experiment Station, Ogden, UT.
provides important forage for wildlife during p. 25-27.
winter when other sources of forage may be
inaccessible. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
Higgins, eds. 1993. A Utah flora. 2nd Ed.,
revised. Brigham Young University Print
Services, Provo, UT. 986 p.
100
Atriplex cuneata A. Nels. Castle Valley clover saltbush
CHENOPODIACEAE
Synonyms: Atriplex gardneri var. cuneata (A. Nels.) S.L.Welsh

Atriplex acanthocarpa var. cuneata Jones
Atriplex nuttalli subsp. cuneata H.M. Hall & Clem.
taxing female state, depending on climatic

conditions.
Taxonomy.—Castle Valley clover saltbush is one

of several related saltbush species. It hybridizes
with a number of other saltbushes, particularly
with fourwing saltbush (Atriplex canescens
(Pursh) Nutt.). Atriplex cuneata is tetraploid (2n =
36). Some taxonomists consider it to be a variety
or subspecies of other woody saltbush species. Its
exact taxonomic distinction remains somewhat
unclear. Hanson (1962) described one subspecies
for Atriplex cuneata, ssp. introgressa, found to be
diploid (2n = 18) (Sanderson and Stutz,
unpublished data). It is now considered to be a
variant of A. welshii Hanson, also diploid. Atriplex
cuneata hybridizes with other perennial Atriplex,
especially with A. confertifolia Torr. & Freemont,
a hybrid that has been named as A. X
neomexicana. Populations of this hybrid and its
derivatives may expand enormously during
favorable years but are eradicated during drought
(Sanderson and Stutz, unpublished data).
Range.—The distribution of Castle Valley clover

General Description.—Castle Valley clover saltbush includes portions of eastern Utah,
saltbush, also known as Castle Valley saltbush and southwestern Colorado, and northern New
cuneate saltbush, is a low shrub 10 to 45 cm tall Mexico. It usually grows at elevations between
with a more or less prostrate, woody, much- 1,220 and 2,170 m (Welsh and others 1993).
branched base with erect branches. Leaves are
evergreen, light-grey green, spatulate to broadly Ecology.—This species often grows on fine
elliptic, 2 to 6 cm long and 0.5 to 2.5 cm wide textured soils, typically clay or soils derived from
(Hall and Clements 1923). Staminate (male) shale, that are moderately to highly alkaline. It
flowers are dark, almost black, and are borne in grows on variably saline soils and may be
glomerules arranged in panicles. Pistilate (female) associated with many other shrubby chenopod
flowers are borne in axillary clusters and consist of species and several of the sagebrush (Artemisia L.)
pistils enclosed by wingless bracts. At maturity, species. It is often the dominant or codominant
bracts are 5 to 9 mm wide, irregularly toothed shrub with shadscale (Atriplex confertifolia Torr.
along their margins, and have numerous, crest-like & Frem.) or mat saltbush (Atriplex corrugata S.
tubercles on their side. Sex expression in Castle Wats.). It grows in areas with 160 to 310 mm of
Valley clover saltbush is trioecious (Freeman and precipitation.
McArthur 1984). In this system, some individuals
are consistently male, some are consistently Reproduction.—Castle Valley clover saltbush
female, and others can switch between the less blooms in April and May, or sometimes later
energistically expensive male state, and the more depending upon rainfall. Fruits (utricles) ripen
101
about 7 weeks after blooming (Hanson 1962). nutritious forage for wildlife and livestock on
Annual seed production is often erratic and only Western shrublands.
about 50 percent of utricles contain viable seed.
Germination rates are also erratic and vary among References
ecotypes. Fruits are small and generally number
about 180,000/kg (Blauer and others 1976). Blauer, A.C., A.P. Plummer, E.D. McArthur, R.
This species should not be seeded in areas Stevens, and B.C. Giunta. 1976. Characteristics
beyond its natural distribution. It is poorly adapted and hybridization of important Intermountain
to big sagebrush (Artemisia tridentata Nutt.) and shrubs. II. Chenopod family. Research Paper
other upland habitats but can be seeded into most INT-177. U.S. Department of Agriculture, Forest
salt desert shrublands. Young plants grow Service, Intermountain Forest and Range
vigorously and are competitive but can be Experiment Station, Ogden, UT. 42 p.
suppressed by perennial grasses and annual weeds.
It should not be seeded directly with herbaceous Freeman, D.C. and E.D. McArthur. 1984. The
species but competes well when seeded with relative influences of mortality, non-flowering,
fourwing saltbush [Atriplex canescens (Pursh) and sex change on the sex ratios of six Atriplex
Nutt.], green ephedra (Ephedra viridis Coville), species. Botanical Gazette 145: 385-394.
and spiny hopsage [Grayia spinosa (Hook.) Moq.].
Hall, H.M. and F.E. Clements. 1923. The
Management.—Castle Valley clover saltbush is phylogenetic method in taxonomy, the North
preferred by livestock throughout the year. It American species of Artemisia, Chrysothamnus,
remains green and succulent, even through winter. and Atriplex. Publication 326. Carnegie
It is an important source of protein in winter Institution of Washington, Washington, DC.
months for both livestock and game when most 355 p.
other forages are dry (McArthur and others 1978).
A stabilized hybrid between fourwing saltbush and Hanson, C.A. 1962. Perennial Atriplex of Utah and
Castle Valley clover saltbush has great potential to the northern deserts. M.S. thesis. Brigham
improve the forage quality on winter ranges. Such Young University, Provo, UT. 133 p.
a hybrid would be taller than Castle Valley clover
saltbush and retain more green foliage than McArthur, E.D., A.P. Plummer, and J.N. Davis.
fourwing saltbush. 1978. Rehabilitation of game range in the salt
This species shows particular promise in desert. In: K.L. Johnson, ed. Wyoming
winter game range restoration and on disturbed shrublands: proceedings of the 7th Wyoming
alkaline soils. Like other woody saltbushes, it shrub ecology workshop; 1978 May 31-June1;
grows on freshly disturbed soils, such as exposed Rock Springs, WY. University of Wyoming,
substrata created by mining or road construction Division of Range Management, Laramie, WY.
activities (Blauer and others 1976, McArthur and p. 23-50.
others 1978).
Sanderson, S.C. and H.C. Stutz. 2003.
Benefits.—Castle valley clover saltbush is a native Unpublished data. U.S. Department of
shrub that is adapted to fine textured soils that are Agriculture, Forest Service, Rocky Mountain
inhospitable to many other shrubs. It adds Research Station, Provo, UT
biological and structural diversity to the plant
communities on these otherwise difficult sites. It Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
helps stabilize the soil, can be useful in wildland Higgins, eds. 1993. A Utah Flora. Second
restoration projects, and provides valuable and edition, revised. Brigham Young University
Print Services, Provo, UT. 986 p.
102
Baccharis pilularis DC. coyote brush
ASTERACEAE
Synonyms: None
Christopher Ross
Oakland, California, apparently as a result of

decreased grazing and fire, and appears to favor
areas of bare soil (McBride and Heady 1968). It
may be a seral stage leading to oak and bay
woodlands in the San Francisco Bay area (McBride
1974). The annual grass Bromus mollis L. has been
found to interfere competitively with Baccharis
seedlings, especially in dry conditions (Da Silva
and Bartolome 1984). In coastal regions of central
California, a cecidomyiid midge, Rhopalomyia
californica Felt forms terminal galls on coyote
brush, and the midges are in turn preyed upon by
various parasites (Latto and Briggs 1995).
Reproduction.—Flowers are headed in leafy

panicles, with staminate heads 3.2 to 5 mm long,
and pistallate heads 3.5 to 5 mm long. Phyllaries
(involucre bracts) are in five to six series, linear-
lanceolate, hairy, and glandular, with convex
receptacle. Staminate flowers are 19 to 26, with 4
to 7 mm corollas and 3.5 to 4.5 mm pappus.
Pistillate flowers number 19 to 43, with 2.5 to 3.5
mm corollas. Seeds are glabrous, 1 to 2 mm long
with 8 to 10 ribs and a 5.5 to 9 mm pappus
(Sundberg 1993). Seed dispersal begins in
November, with germination in winter and early
General Description—Coyote brush, also known spring (McBride and Heady 1968). Coyote brush is
as chaparral broom, is a shrub to 3 m tall, with sometimes propagated horticulturally from tip
glabrous, sticky, oblanceolate to obovate leaves 8 cuttings.
to 55 mm long, with toothed or entire margins. It
may either be spreading or ascending in form Fire Effects.—Coyote brush has been observed to
(Sundberg 1993). burn readily in coastal California, despite the fact
that it is sometimes marketed as a fire-resistant
Range.—Coyote brush occurs from Oregon to species. Coyote brush seedlings and young plants
northern Mexico. In California it ranges from the are susceptible to fire damage, but older plants
northwestern coast south, including the peninsular quickly resprout from their base after fire (McBride
and transverse ranges and the Channel Islands, and and Heady 1968).
eastward to the Sierra Nevada foothills (Sundberg
1993). Growth and Management.—Coyote brush
seedlings are adversely affected by grazing and
Ecology.—Coyote brush is found on coastal bluffs trampling by livestock, although grazing animals
and oak woodlands from sea level to 1500 m. It may also foster its spread by decreasing grass
sometimes occurs on serpentine soils (Sundberg cover (McBride and Heady 1968). However,
1993). Coyote brush may invade stabilized dunes Williams and others (1987) found no proximal link
in northern California, especially after yellow bush between decreased grazing and fire and coyote
lupine has established (Pickart and Sawyer 1998). brush invasion. Mowing of specimens used as
It has been very invasive in grassland areas near
103
ground cover is effective in controlling height Latto, J. and C.J. Briggs. 1995. Factors affecting
without long-term detrimental effects (Hodel and distribution of the gall forming midge
Pittenger 1994). Coyote brush may be removed by Rhopalomyia californica (Diptera:
burning and then pulling out the roots. Cecidomyiidae). Environmental Entomology
24(3): 679-686.
Benefits.—Although several cultivated varieties
are often sold as a landscaping shrub or ground McBride, J. 1974. Plant Succession in the
cover, some growers caution that it is neither an Berkeley Hills, California. Madrono 22: 317-
especially attractive plant, nor as fire-proof as 380.
sometimes stated. It is of low value as livestock
forage (McBride and Heady 1968). Coyote brush McBride, J. and H.F. Heady. 1968. Invasion of
was used by Native Americans as a tea for use on Grassland by Baccharis pilularis DC. Journal
poison oak rash (Timbrook 1987). It is considered of Range Management 21: 106-108.
problematic in recreation areas due to its
impenetrability and frequent association with
Pickart, A.J. and J. O. Sawyer. 1998. Ecology and
poison oak (Rhus diversiloba T. & G.) (McBride
Restoration of Northern California Coastal Dunes.
and Heady 1968).
California Native Plant Society. Sacramento, CA.
152 p.
References
Sundberg, S. 1993. Baccharis. In J.C. Hickman,
Da Silva, P.G. and J. W. Bartolome. 1984.
ed. The Jepson Manual: higher plants of
Interaction between a shrub, Baccharis pilularis
California. University of California Press,
subsp. consanguinea (Asteraceae) and an annual
grass, Bromus mollis (Poaceae), in coastal Berkeley and Los Angeles. 1,400 p.
California. Madrono 31: 93-101.
Timbrook, J. 1987. Virtuous herbs: plants in
Hodel, D.R. and D.R. Pittenger. 1994. Responses Chumash medicine. Journal of Ethnobiology
of eight groundcover species to renovation by Winter 1987: 171-180.
mowing. Journal of Environmental Horticulture
12: 4-7. Williams, K., R.J. Hobbs, and S.P. Hamburg.
1987. Invasion of an annual grassland in
Northern California by Baccharis pilularis ssp.
consanguinea. Oecologia 72:461-465.
104
Barleria prionitis L. porcupine flower
ACANTHACEAE
Synonyms: none
John K. Francis
cultivated as an ornamental and has escaped in

many tropical countries including Hawaii (Neal
1965) and Puerto Rico (Liogier 1997).
Ecology.—Porcupine flower grows in a wide

variety of well-drained soils derived from igneous,
metamorphic (including ultramafics) and
sedimentary (including limestone) rocks. In the
Puerto Rican naturalized range, it grows in areas
receiving from about 750 to 900 mm of mean
annual precipitation (author’s observation). Within
the native range, it probably tolerates both higher
and lower precipitation, but no data are available.
Porcupine flower is found throughout the hotter
parts of India and defoliates annually during the
dry season (Parrotta 2001). The species is
moderately intolerant of shade, growing in both
full sunlight and under light forest canopies.
Porcupine flower may grow as single plants or in
large, tangled thickets. Dense stands eliminate
most other ground vegetation. However, the
species does not compete well in dense stands of
tall grass. It is most often found on roadsides,
bluffs and bars above streams, overgrazed range,
disturbed areas, and farmsteads.
General Description.—Porcupine flower, also Reproduction.—In the deciduous tropical forest

called espinosa amarilla, and picanier jaune, is a zone of central India, these shrubs flower from
flowering, spiny invader that reaches 1.5 m in September to December and fruit from January to
height and 2.5 cm in basal diameter. It has a tap April (Parrotta 2001). The flowering and fruiting
and lateral root system. Plants commonly have a seasons are about the same in Puerto Rico. Sixty
single stem but may have multiple stems or fruits collected in Puerto Rico averaged 0.0998 +
branches from near the ground. The stems and 0.0044 g/fruit. Air-dried seeds separated from the
branches are stiff, round, and light tan or light gray. same collection averaged 0.0305 + 0.0005 g/seed
The shrub is armed with 5- to 20-mm long spines or 33,000 seeds/kg. Forty-four percent of the seeds
in the leaf axils. Leaves are elliptic to oblong, 3 to sown on commercial potting mix germinated
10 cm long and 1.5 to 4 cm broad. The yellow, between 13 and 77 days following sowing. Natural
tubular flowers, 3 to 4 cm long and broad, are regeneration generally occurs within 1 or 2 m of
sessile in the leaf axils or in terminal branched the parent plant in Puerto Rico, perhaps because
spikes. The two-celled capsule is ovate-lanceolate animal vectors for seed dispersal are lacking.
and 1.3 to 2 cm long, with a sharp-pointed beak Seedlings can be abundant near fruiting plants.
about 6 mm long. The seeds are 8 by 5 mm,
flattened and covered with matted hairs (Howard Growth and Management.—Seedlings of
1989, Liogier 1997). porcupine flower grow slowly at first. However,
established plants add up to 0.5 m per year to their
Range.—Porcupine flower is a native of Tropical height and nursery seedlings reach 0.6 m in 6
East Africa and Asia (Burkill 1985). It is widely months. In Puerto Rico, these shrubs live about 4
105
years. Because it is spiny and generally ignored by Burkill, H.M. 1985. The useful plants of West
cattle, the species is considered a weed in much of Tropical Africa. Vol. 1. Royal Botanic Garden,
its range. Mowing, followed by spraying of the Kew, UK. 960 p.
sprouts with broadleaf herbicide, is a general
control measure for low shrubs and would probably Howard, R. A. 1989. Flora of the Lesser Antilles,
be effective against porcupine flower. Leeward and Windward Islands. Vol. 6. Arnold
Benefits.—Porcupine flower furnishes cover for MA. 658 p.
wildlife and protects the soil against erosion. It is
widely planted as an ornamental and cultivated in Liogier, H. A. 1997. Descriptive flora of Puerto
Asia as a hedge plant (Burkill 1985). Whole-plant Rico and adjacent islands. Vol. 5. Editorial de la
extracts of porcupine flower contain iridoid Universidad de Puerto Rico, San Juan, PR.
glycosides, barlerin, and verbascoside, which have 436 p.
shown potent activity against respiratory syncytial
virus in vitro and may account for the plant’s use in Neal, M.C. 1965. In gardens of Hawaii. Special
treating fever and several respiratory diseases in Publication 50. Bernice P. Bishop Museum
herbal medicine (Balick and others 1998). Extracts Press, Honolulu, HI. 924 p.
of the plant have also been shown to effectively
suppress the fungi Trichophyton mentagrophytes in Panwar, H.S., M.M. Nauriyal, and H.C. Joshi.
vitro (Panwar and others 1979). A mouthwash 1979. In vitro screening of certain indigenous
made from root tissue is used to relieve toothache plants for their antimycotic activity. Veterinary
and treat bleeding gums (Burkill 1985). The whole Research Bulletin 2(2): 164-167.
plant, leaves, and roots are used for a variety of
purposes in traditional Indian medicine. For Parrotta, J.A. 2001. Healing plants of Peninsular
example, the leaves are used to promote healing of India. CABI Publishing. Wellington, UK & New
wounds and to relieve joint pains and toothache York. 917 p.
(Parrotta 2001). Because of its antiseptic
properties, extracts of the plant are incorporated Prakruti. 2002. Suddh Bhangra (maka) oil. http://
into herbal cosmetics and hair products to promote www.prakrutiherbals.com/hairoil.htm 2 p.
skin and scalp health (Prakruti 2002, Probiotics
New Zealand 2002, Vaipani 2002). Probiotics New Zealand. 2002. Probiotics for
life—improving your health and quality of life
References through good bacteria. http://www.probiotics.
co.nz/prdcts_efml.asp?ProductID=5. 3 p.
Balick, M., P. Blanc, M. Morgan, J.L. Chen, R.
Cooper, M.R. Kernan, W. Nanakorn, N. Vaipani. 2002. Vaipani Herbal Ayurvedic
Parkinson, E.J. Rozhon, C.A. Stoddard, and Z.J. Partisthan—A herbal ayurvedic cosmetics
Yee. 1998. New iridoids from the medicinal company. http://www.vaipani.com/skincare/
plant Barleria prionitis with potent activity skin%20Care-Page3.html. 2 p.
against respiratory syncytial virus. Journal of
Natural Products 61(10): 1,295-1,297.
106
Batis maritima L. saltwort
BATACEAE
Synonyms: none
John K. Francis
Wiggins and Porter 1971).

Range.—Saltwort is native to coastal areas of
southern United States including California,
Mexico, Central America, South America to Brazil
and Peru, the West Indies, and the Galapagos
Islands (Howard 1988, Wiggins and Porter 1971).
It has been planted in a number of areas in the
tropics and has naturalized in at least Hawaii (Neal
1965).
Ecology.—Saltwort is uncommon to abundant in

low-laying areas near seashores. It grows in salt
marshes, at the upper edge of tidal flats, at the
edge of mangrove stands, and between scattered
mangroves. It is recognized as a major colonizer
after mangroves are destroyed by hurricanes.
Although it is not a water plant, it can endure brief
flooding and long periods of waterlogged soils
(Nelson 1996). Saltwort grows slowly in soils
with high salt concentrations and areas with
seawater overwash where it suffers little
competition from other plants. The species
manages salts by sequestering them in cell
vacuoles and eventually shedding the leaves
(Barbuda Turf Company 2002). It also grows in
soils without salt but is vulnerable to competition
from nonhalophytes. The soils are usually sandy,
marly, or gravely. Deposits of wrack (dead plant
material) by high tides have been shown to be
beneficial to the species (Pennings and Richards
General Description.—Saltwort, also known as 1998). Saltwort is intolerant of shade. The species
turtleweed, pickleweed, barilla, planta de sal, is not seriously affected by insects, disease, or
camphire, herbe-à-crâbes, and akulikuli-kai, is a grazing.
low, yellow-green shrub with succulent leaves. It
is prostrate or upright and occasionally reaches 1 Reproduction.—In Florida, saltwort flowers in
m in height, 2 m in lateral extent, and 5 cm in the spring and fruits in the summer (Long and
basal diameter. Stems are usually multiple as Lakela 1976). Flowering and fruiting occurs year-
sprouts from the root crown, and as they become round in Puerto Rico. Little is known about seed
tall and heavy, lie down and root along the stems production or germination. Most effective
forming loose mats. Weak roots with light-tan, reproduction of the species appears to be
corky bark form tap and lateral root systems. The vegetative. Sprouting from the root crown occurs
bark is grayish white; stem wood is weak and with and without disturbance. Layering is a
brittle. The glabrous leaves are succulent, linear or constant process of prostrate stems. New plants
narrowly oblanceolate and round or three- to four- can be started by cuttings and probably broken
angled in cross section. Tiny, white male and pieces of plants are carried to new habitat by water
female flowers occur on different plants. Fruits are and machinery. It is speculated that whole plants
axillary drupaceous, yellow-green syncarps 1 to 2 washed to sea during torrential rains in South
cm long (Liogier 1985, Stevens and others 2001,
107
America were carried by currents to establish the Indies. http://www.geocities.com/barbudaturf/
species in the Galapagos Islands (Trillmich 2002). hallophites%20in%20deep.htm. 3 p.
Growth and Management.—Saltwart plants Big Island Invasive Species Committee. 2002. Pest
established in experiments from cuttings and prevention/suppression: eradication of incipient
irrigated with fresh and brackish water grew 65 cm forest pest plants by the Big Island Species
and reached plant dry weights of 120 g in 13 Committee. http://www.hear.org/
weeks. Plants receiving saltier water grew much operationmicronia/2002biiscannual.pdf. 9 p.
less but still added height and weight with water
almost twice as salty as sea water. Tissue ash Florida Cooperative Extension Service. 2002.
concentrations did not differ significantly between Butterfly gardening in Florida. University of
treatments (Miyamoto and others 1996). Wild Florida, Gainesville, FL. http://edis.ifas.ufl.edu/
plants in stands in Puerto Rico appear to extend BODY_UW057. 24 p.
their stems about 1 m/year. Leaf succulence
increases as much as 1.5 times during the dry Howard R.A. 1988. Flora of the Lesser Antilles,
season to maintain a near static solute (chiefly Na+, Leeward and Windward Islands.
Cl-, and S04--) concentration (Lüttge and others Dicotyledoneae, Part 1. Vol. 4. Jamaica Plain,
1989). Probably all planting for landscaping and MA: Arnold Arboretum, Harvard University.
environmental restoration is by rooted cuttings or 673 p.
uprooted wild plants. Except in Hawaii, where it is
being suppressed as an exotic plant (Big Island Liogier H.A. 1985. Descriptive flora of Puerto
Invasive Species Committee 2002), there is little Rico and adjacent islands, Spermatophyta. Vol.
reason to control stands of saltwort. It grows in 1. Editorial de la Universidad de Puerto Rico,
disturbed areas where few other plants can survive. Río Piedras, PR. 352 p.
Benefits.—The principal benefit of saltwort is that Liogier, H.A. 1990. Plantas medicinales de Puerto
it grows in, covers, and protects salty low-laying Rico y del Caribe. Iberoamericana de Ediciones,
areas where few other species will grow. It is used Inc., San Juan, PR. 566 p.
as an ornamental and ground cover on similar
sites. Salt tolerance of the species allows it to be Long, R.W. and O. Lakela. 1976. A flora of
irrigated with brackish and sea water, as well as Tropical Florida. Banyon Books, Miami, FL.
fresh water (Miyamoto and others 1996). Ashes of 962 p.
the plant were once used in the manufacture of
glass and soap. The leaves are sometimes eaten as Lüttge, U., M. Popp, E. Medina, W.J. Cram, M.
a salad (Neal 1965). However, it is toxic in large Diaz, H. Griffiths, H.S.J. Lee, C. Schäfer, J.A.
quantities (Austin 1998). Saltwort serves as the Smith, and K.H. Stimmel. 1989. Ecophysiology
larval and adult hosts for the great Southern white of xerophytic and halophytic vegetation of a
and Eastern pigmy blue butterflies (Florida coastal alluvial plain in Northern Venezuela.
Cooperative Extension Service 2002) and is an New Phytologist 111(2): 283-291.
important food source for the Galapagos marine
iguana (Amblyrhynchus cristatus) (Wrege 2002). Miyamoto, S., E.P. Glenn, and M.W. Olsen. 1996.
The species has applications in herbal medicine to Growth, water use and salt uptake of four
treat eczema, psoriasis, and other skin conditions, halophytes irrigated with highly saline water.
rheumatism, gout, blood and vein disorders Journal of Arid Environments 32(2): 141-159.
(Liogier 1990).
Neal, M.C. 1965. In gardens of Hawaii. Spec. Pub.
References 50. Bernice P. Bishop Museum Press, Honolulu,
HI. 924 p.
Austin, D.F. 1998. Poisonous plants of southern
Florida. http://www.fau.edu/divdept/science/ Nelson, G. 1996. The shrubs and woody vines of
envsci/poison-pl.html. 9 p. Florida. Pineapple Press, Inc., Sarasota, FL.
391 p.
Barbuda Turf Company. 2002. Halophytes in
deep. Barbuda Turf Company, Codrington Pennings, S.C. and C.L. Richards. 1998. Effects of
Village, Barbuda, Antigua-Barbuda, West wrack burial in salt-stressed habitats: Batis
108
maritima in a southwest Atlantic salt marsh. scope.org/downloadpubs/scope45/ch01-1.3.1.
Ecography 21(6): 630-638. html. [not paged].
Stevens, W.D., C. Ulloa-U., A. Pool, and O.H. Wiggins, I.L. and D.M. Porter. 1971. Flora of the
Montiel, eds. 2001. Flora de Nicaragua. Galápagos Islands. Stanford University Press,
Monographs of Systematic Botany Vol. 85, No. Stanford, CA. 998 p.
1. Missouri Botanical Garden Press. 943 p.
Wrege, P.H. 2002. Bee-eater research: Galapagos!
Trillmich, F. 2002. El Niño in the Galapagos Department of Ecology and Evolutionary
Islands: a natural experiment. International Biology, Cornell University, Ithaca, New York.
Council of Scientific Unions. http://www.icsu- http://www.eeb.cornell.edu/wrege/pages/iggires.
html. 4 p.
109
Bidens menziesii (Gray) Sherff ko'oko'olau
ASTERACEAE
Synonymns: Coreopsis menziesii A. Gray

Bidens menziesii subsp. filiformis (Sherff) Ganders & Nagata
Bidens m. var. filiformis Sherff
Bidens lepida Degener & Sherff
Sarah A. Taylor and Randy S. Senock
the island of Hawaii, the species is a minor

component of the Ohi'a forest. In montane dry
shrublands of leeward Mauna Kea and Mauna Loa,
ko'oko'olau becomes structurally codominant with
a'alii. Due to its tall growth habitat, it appears
dominant over other species when it is in flower.
Ko'oko'olau appears intollerant of wildfire and is
conspicuously absent from burned areas on
leeward Hawaii. Ko'oko'olau does not resprout
following fire (Sherry and others. 1999). The plant
is worth preserving because, like the
honeycreepers, Loxops spp. and Hemignathus spp.,
ko'oko'olau serves as an example of the adaptive
radiation that occurred in Hawaii, with all 19
species of Bidens having a single ancestor.
General Description.―Bidens menziesii, known
Reproduction.―Seed production is fairly prolific
as ko'oko'olau, is a sparingly branched shrub with
as the inflorescence is a large terminal panicle
erect stems reaching 1 to 4 m in height. Leaves are
composed of 8 to 100 disc florets. Fruits are
4 to 26 cm long, including petiole, bipinnately
achenes. Chromosome number has been identified
divided into long linear divisions and may be
as 2n = 72 (Wagner 1990). Interestingly, the
glaborous or pubescent. Margins are entire.
species hybridizes readily and for this reason
Densely branched compound cymes terminate the
individuals unique to a particular local should not
main stem and lateral branches. Flowers measure 5
be introduced to a different zone, in order to
to 7 cm in diameter. Ray florets are 4 to 5 per
protect the integrity of endemic species.
head, disk florets 6 to 8 per head. Corollas are
yellow. Fruits are indehiscent, dry, grayish black,
Growth and Management.―Reasonably fast
straight, wingless or very rarely winged, barbless
growing, ko'oko'olau may not be long-lived.
and 9 to 12 mm long (Wagner 1990).
Ko'oko'olau can be grown easily from seeds or
from terminal cuttings using a root hormone. It
Range.―Ko'oko'olau can be found on Molokai
prefers light to moderate watering and full sun
and West Maui scattered on arid, leeward slopes
(Krauss 1998).
and cliffs in shrubland vegetation, generally
between the elevations of 200 to 750 m. On
Benefits.―Ko'oko‘olau was widely used by
leeward sides of Mauna Loa and Mauna Kea of
Hawaiians prior to European arrival and is still
Hawaii, this species is common on slopes of cinder
sold as tea, although the introduced pantropical B.
cones in montane a'ali'i shrublands, and in
pilosa is usually the species incorrectly labeled
subalpine forest, from elavations of 750 to 2,200
and sold as the traditional Hawaiian tea.
m. The species is one of 19 Bidens unique to the
Hawaiian islands (Wagner 1990).
References
Ecology.―Ko'oko‘olau is an important dryland
Carlquist, S. 1970. Hawaii--a natural history. The
shrub and forest plant of ecosystems seriously
Natural History Press, Garden City, NY. 484 p.
under threat by agriculture and development. On
110
Krauss, B. 1998. How to plant a native Hawaiian area, Hawaii. 1999 Hawaiian Conservation
garden. http://www.hawaii.gov/health/oeqc/ Conference, Honolulu, HI.
garden/index.html. [not paged].
Wagner, W.L., D.R. Herbst, and S.H. Sohmer.
Sherry, K., J.M Castillo, and R.B Shaw. 1999. 1990. Manual of the Flowering Plants of
Effects of wildfire on vegetation and rare plants Hawaii. University of Hawaii Press, Honolulu.
in arid montane shrublands, Pohakaloa training 1,854 p.
111
Bixa orellana L. annatto
BIXACEAE
Synonym: Bixa katagensis Delpierre
John K. Francis
neglected or abandoned farmland, some of which

has grown up to early secondary forest. Rainfall
ranges from 1000 mm to 3000 mm per year in
areas where the species grows naturally or under
cultivation. Soils with textures from sands to clays
are colonized. The species tolerates relatively low
base saturation and moderate compaction. Anatto
is vulnerable to overtopping and smothering by
trees, shrubs, vines, and grass. Plants that have
become overtopped and shaded cease to flower
and bear fruit. The species is frost sensitive (von
Carlowitz 1991).
Reproduction.—Annatto plants flower and fruit

heavily and almost continuously in favorable
General Description.—Annatto, which grows as
habitat. Cuttings taken from flowering plants will
a shrub or a small tree, is known by many other
produce flowers and fruits and a smaller shrub than
common names including lipstick plant, roucou,
plants of seed origin (Bailey 1941). Belfort and
achiote, bija, urucú, and shambu (Little and others
others (1992) report that seeds dried to moisture
1974). Its single or multiple stems are light brown.
contents of 10 to 15 percent germinated at from 8
The bark is more or less smooth with many warty
to 58 percent compared to fresh seeds (65 percent
lenticels, but may become fissured in old
moisture) that gave 96 percent germination.
individuals. If given ample space, annatto
Scarification was reported to improve germination
generally branches several times near the ground
of fresh seeds in another study. Mechanical
and develops a dense, spreading crown. The plant
scarification proved superior to acid or hot water
roots firmly with a thick taproot and finer laterals.
treatments (Amaral and others 1995). Air-dry fruits
The alternate leaves have long petioles, thin ovate
collected in Puerto Rico weighed an average of
blades with long-pointed tips. Panicles at the
1.701 + 0.078 g. Seeds from that collection
branch tips have few to many pink or white
averaged 0.029 + 0.008 g/seed or 35,000 seeds/kg.
flowers. The fruits are spiny capsules that dry and
Of these seeds that received no pregermination
split open in two parts to expose red seeds on their
treatment, 60 percent germinated between 11 and
inner surfaces (Howard 1989, Liogier 1995).
110 days after sowing. The resulting plants were
ready to prick out (10 cm in height) about 3 months
Range.—Although annatto is native to continental
after germination. Success in air layering of shoots
tropical America, it is unclear where the original
ranged from 93 to 100 percent except when
native stands of the species were located because
sawdust was used as substrate (7 percent) (Barbosa
the shrub has been cultivated since ancient times.
e Silva and others 1993). Stem cuttings rooted (up
It now grows in cultivation and naturalized from
to 60 percent) when treated with IAA or IBA.
Mexico to Argentina and throughout the Caribbean
Untreated cuttings did not root (Thirunavoukkarasu
Islands. The plant is also widely cultivated and
and Saxena 1997). Annatto can be grafted by
naturalized in tropical and subtropical regions
several techniques. The best method is budding (70
throughout the rest of the world.
percent success in tests) (Bruckner and others
1991).
Ecology.—Annatto is shade-intolerant and must
have disturbance or a broken forest canopy to
Growth and Management.—Annatto shrubs will
become established. All the naturally growing
bear fruit when 2 years old in Hawaii (Neal 1965).
annatto shrubs in Puerto Rico are found on
112
Under good management, plants will fruit within 1 Barbosa e Silva, K.M., F.C.G. Almeida, F.A.G.
year of planting (Nepstad and others 1991). An Almeida, P.S. Lima e Silva, J.J.L. de
Indian plantation yielded 529 kg/ha of seed at 2 Albuquerque, and J.J.L. de Albuquerque. 1993.
years old and 2,483 kg/ha of seed at 3 years old Effect of substrate on rooting of air-layered
(Kanjilal and Singh 1995). Annatto seldom shoots of annatto. Pesquisa Agropecuaria
reaches more than 5 m in height and 10 cm in stem Brasileira 28(1): 101-106.
diameter (Little and others 1974). Pruning of
ornamentals is recommended to shape and thicken Belfort, A.J.L., O.R. Kato, and M. do S.A. Kato.
the crowns (Warren 1997). 1992. Practical method of drying annatto seeds
for seedling production. Circular Tecnica 67.
Benefits.—Annatto, obtained from the oily arils of Centro de Pesquisa Agropecuaria do Tropico
the seeds is the world’s second most important Umido, EMBRAPA-CPATU, Belem, Brazil.
(after caramel) natural colorant (Mercadante and 14 p.
Pfander 1998), yielding yellow to red colors. The
colors are produced by several apocarotenoides Bruckner, C.H., S.S. Khouri, and A.V. Melgaco.
and may reach up to 7 percent of the seed’s dry 1991. Propagation of annatto (Bixa orellana L.)
mass (Katzer 1999). World production of annatto by five grafting methods. Revista Ceres 38: 218,
seed, both for commercial and home use, was 340-344.
estimated in 1990 at 10,000 tons per year (Arkcoll
1990). Brazil is the world’s largest exporter Howard, R.A. 1989. Flora of the Lesser Antilles.
(Katzer 1999). Not only was the dye used Vol. 6. Arnold Arboretum, Harvard University,
anciently to color food, but also to dye cloth and Jamaica Plain, MA. 658 p.
paint the skin (which is still done today). The
species is also planted as an ornamental, Kanjilal, P.B. and R.S. Singh. 1995. Agronomic
particularly the varieties with bright pink pods. evaluation of annatto (Bixa orellana L.). Journal
Branches with the dry pods are used in dry floral of Herbs, Spices, and Medicinal Plants 3(3): 13-
arrangements (Warren 1997). Bees collect nectar 17.
from its flowers to make honey. The wood is light-
weight (specific gravity 0.4), weak, and not Katzer, G. 1999. Annatto (Bixa orellana L.).
durable. It was used in former times to start fires http://www-ang.kfunigraz.ac.at/~katzer/engl/
by friction. Ropes and twine were made from the Bixa_ore.html. 3 p.
fibrous bark (Little and others 1974). The pulp
surrounding the seeds is widely used in herbal Liogier, H.A. 1990. Plantas medicinales de Puerto
medicine to treat burns, bleeding, dysentery, Rico y del Caribe. Iberoamericana de Ediciones,
gonorrhea, constipation, and fever (Parrotta 2001). Inc., San Juan, PR. 563 p.
Extracts of leaves, bark, and roots are reported to
be antidotes for poisoning from Manihot esculenta Liogier, H.A. 1995. Descriptive flora of Puerto
Crantz, Jatropha curcas L., and Hura crepitans L. Rico and adjacent islands. Vol. 4. Editorial
(Liogier 1990). de la Universidad de Puerto Rico, San Juan, PR.
617 p.
References
Amaral, L.I.V., M. de F.A. Pereira, and A.L. Wadsworth. 1974. Trees of Puerto Rico and the
Cortelazzo. 1995. Dormancy breaking in seeds Virgin Islands. Vol. 2. Agriculture Handbook
of Bixa orellana. Revista Brasiliera de 449. U.S. Department of Agriculture,
Fisiologia Vegetal 7(2): 151-157. Washington, DC. 1,024 p.
Arkcoll, D. 1990. New crops from Brazil. In: J.

Janick and J.E. Simon, eds. Advances in new Mercadante, A.Z. and H. Pfander. 1998.
crops. Timber press, Portland, OR. p. 367-371. Carotenoids from annatto: a review. Recent
Research Developments in Agriculture and Food
Bailey, L.H. 1941. The standard cyclopedia of Chemistry 2(1): 79-91.
horticulture. Vol. 1. The MacMillan Co. New
York, NY. 1,200 p. Neal, M.C. 1965. In gardens of Hawaii. Special
Publication 50. Bernice P. Bishop Museum
Press, Honolulu, HI. 924 p.
113
Nepstad, D.C., C. Uhl, and E.A.S. Serrao. 1991. on rooting of Bixa orellana L. stem cuttings.
Recuperation of a degraded Amazonian Orissa Journal of Horticulture 25(1): 84-86.
landscape: forest recovery and agricultural
restoration. Ambio 20(6): 248-255. von Carlowitz, P.G. 1991. Multipurpose trees and
shrubs, sources of seeds and inoculants.
Parrotta, J.A. 2001. Healing plants of Peninsular International Council for Research in
India. CAB International, Wallingford, UK. Agroforestry, Nairobi, Kenya. 328 p.
944 p.
Warren, W. 1997. Tropical plants for home and
Thirunavoukkarasu, M. and H.O. Saxena. 1997. A garden. Thames and Hudson, New York. 240 p.
short note on the effect of auxins (IAA, IBA)
114
Bocconia frutescens L. pan cimarrón
PAPAVERACEAE
Synonyms: Bocconia frutescens var. cernua Moc. & Sessé ex DC.

Bocconia glauca Salisb.
Bocconia pearcei Hutch.
Bocconia quercifolia Moench
Bocconia sinuatifolia Stokes
Bocconia subtomentosa L’Hér. ex Stahl
John K. Francis
3 cm long, and blades are 14 to 50 cm long.

Panicles up to 50 cm long have lax branches. The
numerous small flowers are yellow-cream in color.
The fruits are ellipsoidal capsules each containing a
black seed with a red, fleshy aril running along one
side (Howard 1988, Liogier 1985, Little and others
1974).
Range.—Pan cimarrón is reported to be native

from the middle of Mexico through Central
America to Argentina and Bolivia in South
America, and throughout the Caribbean (Howard
1988, Liogier 1985, Stevens and others 2001, New
York Botanical Garden 2002). It has naturalized in
forests in the islands of Maui and Hawaii in the
State of Hawaii (University of Hawaii Botany
2002).
Ecology.—Pan cimarrón grows in a wide variety

of soil types at middle elevations up to 2,150 m in
Mexico (Secretaría de Madio Ambiente y Recursos
Naturales 2002). In Puerto Rico, it grows in
subtropical dry forest along streams (750 to 1000
General Description.—Pan cimarrón is also mm/year precipitation), in subtropical moist forest
known as parrot weed, celadonia, plume-poppy, (1000 to 2000 mm/year precipitation), and in
palo de toro, panilla, llorasangre, curarador, palo subtropical wet forests (2000 to 3000 mm/year
amarillo, saúco, yagrumito, chocolate blanco, precipitation). In Nicaragua, the species grows
gordolobo, guacamayo, trompeto, tabaquillo, even in the cloud forests (Stevens and others 2001).
grande chelidoine, and bois codine. It is a shrub or In Puerto Rico, pan cimarrón is most common
small tree, usually 2 to 3 m in height and 2 to 3 cm along streams, road cuts, and landslides. The
in stem diameter. Branching near the ground species also grows widely scattered in brush lands,
normally results in multiple stems. The plant is which develop from abandoned fields and pastures,
supported by a tap and lateral system of orange-red and in secondary forest. Pan cimarrón is intolerant
roots that are fleshy outside and fibrous within. of shade. It competes vigorously with herbs and
There are relatively few branches. The outer bark is brush after establishment. The shrub does not form
light brown, smooth to lightly fissured; the inner continuous stands in Puerto Rico, but occurs as
bark is orange-red and bitter. An orange or yellow scattered individuals or small patches. However, as
sap exudes from wounds. The deeply lobed and an invader in Hawaii, it does form large, dense
toothed leaves are alternate but usually clustered at stands in dry and mesic habitat (University of
the ends of older branches. Petioles are stout, 1.5 to Hawaii Botany 2002).
115
Reproduction.—Pan cimarrón flowers and fruits de Publicaciones. San Salvador, El Salvador.
throughout the year (Little and others 1974, 703 p.
Stevens and others 2001). The infrutescences may
contain tens to hundreds of fruits. Fruits in the Howard R.A. 1988. Flora of the Lesser Antilles,
infrutescences mature a few at a time beginning at Leeward and Windward Islands.
the top and proceeding downward. A collection of Dicotyledoneae, Part 1. Vol. 4. Arnold
seeds from Puerto Rico weighed (air-dried) an Arboretum, Harvard University, Jamaica Plain,
average of 0.0146 + 0.0002 g/seed (author’s MA. 673 p.
observation). Germination begins in 25 to 55 days
with 40 percent germinating. Germination is Jardín Botánico de Bogotá José Mutis. 2002.
epigeal (Ricardi and others 1977). The seeds are Conservación ex situ. http://www.jbb.gov.co/
dispersed by birds (Environmental Protection conservacion/ex_situ/semillas/body_semillas.
Agency 2002). Damaged plants usually resprout. html. 4 p.
Growth and Management.—Once established, Liogier H.A. 1985. Descriptive flora of Puerto
pan cimarrón has relatively rapid growth (Jardín Rico and adjacent islands, Spermatophyta. Vol.
Botánico de Bogotá José Mutis 2002). Only in 1. Editorial de la Universidad de Puerto Rico,
Hawaii is the species mentioned as a nuisance Río Piedras, PR. 352 p.
where it displaces native vegetation and threatens
the endangered Blackburn’s sphinx moth Little, E.L., Jr., R.O. Woodbury, and F.H.
(Environmental Protection Agency 2002). In the Wadsworth. 1974. Trees of Puerto Rico and the
absence of tests of control measures, such standard Virgin Islands. Vol. 2. Agriculture Handbook
practices as grubbing and spraying with broadleaf 449. U.S. Department of Agriculture,
herbicides are recommended. Washingtion, DC. 1,024 p.
Benefits.—Pan cimarrón contributes to New York Botanical Garden. 2002. Specimens

biodiversity, helps protect the soil, and furnishes search results: Bocconia frutescens.
food and cover for wildlife. The sap has been used http://scisun.nybg.org.8890/searchdb/owa/www
as a dye (Little and others 1974). The orange wood catalog.search_list. [not paged].
is soft and brittle with a thick pith, and is of little
use. Extracts of various tissues of pan cimarrón are Pérez-Arbelaez, E. 1978. Plantas útiles de
employed in herbal medicine to control mange, Colombia. Litografía Arco, Bogota, Colombia.
lice, and intestinal worms, to treat ulcers of the 831 p.
eyes, to treat wounds, and to treat edema and
jaundice (Pérez-Arbelaez 1978). A dose of 15 Ricardi, M., F. Torres, C. Hernandez, and R.
drops of the sap causes a strong purgative effect Quintero. 1977. Morphology of seedlings of
(Guzmán 1975). The sap will cure warts and if Venezuelan trees. Revista Forestal Venezuelana
injected under the skin acts as a local anesthetic 27: 15-56.
although the injection itself causes considerable
pain. The bark contains the alkaloids, Secretaría de Madio Ambiente y Recursos
bocconietrine, cocconichlorine, bocconiyoidine, Naturales. 2002. Especies con usos no
and bocconixatine (Secretaría de Madio Ambiente maderables en bosques tropicales y
y Recursos Naturales 2002). Pan cimarrón is being subtropicales: Bocconia frutescens L.
evaluated as a species for restoration of disturbed http://www.semarnat.gob.mx/pfnm2/fichas/bocc
sites (Jardín Botánico de Bogotá José Mutis 2002). onia_frutescens.htm. 2 p.
References Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.

Montiel, eds. 2001. Flora de Nicaragua.
Environmental Protection Agency. 2002. Proposed Monographs in Systematic Botany, Vol. 85, No.
endangered status for Blankburn’s sphinx moth. 3. Missouri Botanical Garden, St. Louis, MO.
Federal Register Vol. 62, No. 63. http:// p. 1,911-2,666.
www.epa.gov/fedrgstr/EPA-SPECIES/1997/
April/Day-02/e8550/htm. 12 p. University of Hawaii Botany. 2002. Bocconia
frutescens, bocconia, Papaveraceae.
Guzmán, D.J. 1975. Especies útiles de la flora http://www.botany. hawaii.edu/faculty/
salvadoreña. Ministerio de Educación, Dirección cw_smith/boc_fru.htm. 1p.
116
Bontia daphnoides L. white alling
MYOPORACEAE
Synonyms: Bontia daphnifolia Salisb.

Bontia daphnoides var. menor (Gaertn. f.) A. DC.
Bontia menor Gaertn. f.
John K. Francis
shelled stone (Howard 1989, Liogier 1997, Little
and others 1974).
Range.—White alling is native to the Bahamas, the

Greater and Lesser Antilles, Trinidad, Venezuela,
and Guyana (Howard 1989, Little and others
1974). It is cultivated as an ornamental in Hawaii
(Hawaiian Ecosystems at Risk 2001) and Spain
(Sánchez 2001) and has naturalized in Florida
(Nelson 1996).
Ecology.—White alling is most common in coastal

thickets between the tidal mangroves and the
upland forests. These areas are flooded during
storm surges and receive a moderate amount of salt
spray. The soils vary from sandy to clayey, are
usually high in organic matter, and range in pH
from 7.0 to 8.5. The water table is usually within 2
m of the surface. In Puerto Rico, the species grows
in areas that receive from 750 to 1800 mm of mean
annual precipitation. White alling plants are
moderately intolerant of shade, generally starting in
openings. Seedlings and saplings survive in
relatively sunny understories. Adults flower and
General Description.—White alling, also known fruit in intermediate crown positions. The species
as wild olive, mangle bobo, aceituna Americana, is rare in uplands, probably because of competition.
olivier bard, and oliba, is a shrub or small tree It has been cultivated successfully at elevations up
usually 3 or 4 m in height, occasionally reaching 9 to 1,500 m (Little and others 1974). White alling
m and 15 cm in stem diameter. Although seedlings grows as rare to common components of stands but
have a tap and lateral root system, older plants rely does not form pure or nearly pure stands.
on shallow lateral roots that are stiff and brittle
with a brown, corky bark. Trunks of white alling Reproduction.—White alling flowers and fruits
are usually covered with a thick, light brown, throughout the year (Little and others 1974). Fruits
furrowed bark. The heartwood is light gray-brown, collected in Puerto Rico averaged 0.311 + 0.015
hard, heavy, and somewhat brittle; the sapwood is g/fruit. Air-dried seeds cleaned from the above
tan in color. Multiple stems commonly diverge collection averaged 0.0898 + 0.0033 g/seed or
from the base of the principal stem. The crowns are 11,000 seeds/kg. Sown in commercial potting mix,
vertical and narrow with a thin complement of 44 percent of the seeds germinated between 12 and
foliage. Leaves are oblong to linear-lanceolate, 3 to 68 days after sowing (author’s observation). Seeds
11 cm long by 1 to 2 cm broad, and pointed at both are dispersed by water and presumably by birds
ends. The usually solitary, axillary tubular flowers and mammals that eat the fruits. Fruit production is
are yellow blotched with purple and 2 cm long. usually good and seedlings are common in small
Drupes are ovoid, tapering to a point with a openings near seed sources. Although white alling
permanently attached style. Their corky-textured are often tipped in hurricanes, they sprout from the
flesh is yellowish-green to yellow at maturity, has a trunks and reform vertical crowns.
slightly bitter taste, and contains a single hard-
117
Growth and Management.—A small group of Liogier, H.A. 1997. Descriptive flora of Puerto
white alling seedlings grown in the nursery in Rico and adjacent islands. Vol. 5. Editorial de la
Puerto Rico ranged in height from 21 to 86 cm, 8 Universidad de Puerto Rico, San Juan, PR.
months after being pricked into containers. Sapling 436 p.
and sprout growth rate is moderate (about 0.5
m/year) and individual plants live 10 to 30 years. Little, E.L., Jr., R.O. Woodbury, and F.H.
Although no management experience has been Wadsworth. 1974. Trees of Puerto Rico and the
published, the species can probably be established Virgin Islands. Vol. 2. Agriculture Handbook
with nursery seedlings and be managed like other 449. U.S. Department of Agriculture,
short-statured species. Washington, DC. 1,024 p.
Benefits.—White alling helps protect the soil and Mahabir, D. and M.C. Gullifor. 1997. Use of
provides food and cover for wildlife. The wood is medicinal plants for diabetes in Trinidad and
useful for fuel and stakes. Ethanol extracts of white Tobago. Revista Panamericana de Salud Pública
alling showed promising insecticidal activity 1(3): 174-179.
against Boophilus microplus, Cylas formicarius,
and Tribolium confusum (Mansingh and Williams Mansingh, A. and L.A.D. Williams. 1998.
1998, Williams and Caleb-Williams 1997, Pesticidal potential of tropical plants II.
Williams and Mansingh 1993). White alling is Acaricidal activity of crude extracts of several
frequently used to control diabetes in Trinidad Jamaican plants. Insect Science and its
(Mahabir and Gullifor 1997). Extracts of the plant Application. 18(2): 149-155.
are used to control intestinal worms, treat herpes,
treat inflammation, insect bites, scarring, ulcers, Nelson, G. 1996. The shrubs and woody vines of
and wounds (Liogier 1990). Florida. Pineapple Press, Inc., Sarasota, FL.
391 p.
References
Sánchez deL.C., J. 2001. Plantas de la flora cubana
Hawaiian Ecosystems at Risk 2001. Mystery plant cultivadas en España. http://floraguide.es/
# 2, Bontia daphnoides Linn. (Myoporaceae). arboles/plantasdecuba.htm. 8 p.
http://hear.org/mysteryplants/autogendhtml/
mysteryplant2.htm. 3 p. Williams, L.A.D. and L. Caleb-Williams. 1997.
Insecticidally active sesquiterpene furan from
Howard, R. A. 1989. Flora of the Lesser Antilles, Bontia daphnoides L. Philippine Journal of
Leeward and Windward Islands. Vol. 6. Arnold Science 126(2): 155-162.
MA. 658 p. Williams, L.A.D. and A. Mansingh. 1993.
Pesticidal potential of tropical plants I.
Liogier, H.A. 1990. Plantas medicinales de Puerto Insecticidal activity in leaf extracts of sixty
Rico y del Caribe. Iberoamericana de Ediciones, plants. Insect Science and its Application. 14(5):
Inc., San Juan, PR. 566 p. 697-700.
118
Borrichia arborescens (L.) DC. sea ox-eye
ASTERACEAE
Synonyms: Buphthalmum arborescens L.

Borrichia argentea DC.
Borrichia glabrata Small.
John K. Francis
planted as an ornamental, it is not known to have

naturalized outside the native range. Borrichia x
cubana of southern Florida is believed to be a
natural hybrid of sea ox-eye and Borrichia
frutescens (L.) DC. (Nelson 1996).
Ecology.—Natural sea ox-eye grows near the

seashore, usually within the influence of salt spray.
Because the species is low in stature, grows slowly,
and is intolerant of shade, it must occupy areas
with low competition from other vegetation. It
finds these conditions on headlands, seaside rocks,
dunes, beach strands, low hammocks, and the
edges of mangroves and brackish marshes. Sea ox-
eye will grow in more favorable conditions if
artificially protected from competition. Almost any
soil will do, including both acid and alkaline soils.
Sea ox-eye tolerates both excessively drained and
General Description.—Sea ox-eye, also known as poorly drained conditions. Mean annual rainfall
tree oxeye, silver sea ox-eye, oxeye daisy, sea may range from 900 to 1500 mm. It is drought
daisy, seaside tansy, gull feed, clavelón de playa, resistant (Gillman 1999).
and fleur-sorleil bord de mar, is an evergreen, low
shrub usually 1 m or less in height with stem Reproduction.—Sea ox-eye blooms in spring and
diameters of 1 cm, which forms mounds and mats. summer in Florida (Long and Lakela 1976) and
Older plants have numerous stems. The plants are throughout the year in Puerto Rico (author’s
supported by short rhizomes and a moderate observation). It is pollinated by insects. Seeds
number of lateral roots. The stems are gray with a collected in Puerto Rico averaged 0.00115 g/seed
white, brittle wood and a 3-mm pith. Twigs bear or 870,000 seeds/kg. Only 16 percent of this
deep leaf scars. The foliage tends to be crowded at collection germinated when placed in moist potting
ends of twigs. The simple, opposite, sessile leaves mix. Workman (1980) recommends propagating
are fleshy and yellow-green, light-green, or gray- sea ox-eye by sowing in pots. Also, the plant can
green, oblanceolate to spatulate, entire, and 3 to 8 be propagated by cuttings (Gillman 1999). Seed
cm long. The foliage is resinous and aromatic. Sea dispersal appears to be by water (storm surges) and
ox-eye flowers are usually solitary, terminal heads strong winds. Once established, plants widen by
about 2.5 cm across on peduncles 2 to 5 cm long. constant layering of the semiprostrate stems. New
The corolla and the ray florets are bright yellow clumps probably are established by the rooting of
and the disk florets are orange-yellow. The black broken stems moved by storms.
achenes are 3 to 4 mm long with a pappus in the
form of a dentate cup less than 1 mm long. The Growth and Management.—Growth of sea ox-
chromosome number is 2n = 28 (Howard 1989, eye is slow. Branch extension is about 10 to 30
Liogier 1997, Long and Lakela 1976, Nelson 1996) cm/year. Although, stems and branches die, as long
as conditions remain favorable, clonal plants may
Range.—Sea ox-eye is native to southern Florida, live almost indefinitely. Newly established plants
the Bahamas, Bermuda, the West Indies, Mexico should be protected from overtopping by grass,
and Guatemala (Liogier 1997). Although it is brush, and trees. It is important not to regularly
119
irrigate sea ox-eye plants used in landscaping Arboretum, Harvard University, Jamaica Plain,
(Gillman 1999). MA. 658 p.
Benefits.—Sea ox-eye is available commercially Knox, G.W. 2002. Drought tolerant plants for
and is widely planted as an ornamental. It is North and Central Florida. Pinellas County
recommended for foundation plantings and as a Extension. http://coop.co.pinellas.fl.us/fyn/
border shrub (Workman 1980), particularly where publications/fyn_drought_tolerant.html. 12 p.
salt and salt spray are prevalent and in xeric
(unwatered) gardens (Knox 2002). It also attracts Liogier, H.A. 1997. Descriptive flora of Puerto
butterflies (Gilman 2002). A tea is prepared from Rico and adjacent islands. Vol. 5. Editorial de la
branch tips and leaves to relieve colds, coughs, and Universidad de Puerto Rico, San Juan, PR.
fish poisoning (Garland 2002). 436 p.
References Long, R.W., and O. Lakela. 1976. A flora of

tropical Florida. Banyon Books, Miami, FL.
Garland, K. 2002. Borrichia arborescens (Sea Ox- 962 p.
eye). University of the Virgin Islands.
http://rps.uvi.edu/VIMAS/seaoxeye.htm. 1 p. Nelson, G. 1996. The shrubs and woody vines of
Gillman, E.F. 1999. Borrichia arborescens. Fact 391 p.
Sheet FPS-68. Cooperative Extension Service,
University of Florida, Gainesville, FL. 3 p. Workman, R.W. 1980. Growing native. The
Sanibel-Captiva Conservation Foundation, Inc.,
Howard, R. A. 1989. Flora of the Lesser Antilles, Sanibel, FL. 137 p.
Leeward and Windward Islands. Vol. 6. Arnold
120
Bourreria virgata (Sw.) G. Don roble de guayo
BORAGINACEAE
Synonyms: Ehretia virgata Sw.
John K. Francis
outer surface deeply grooved (author’s
observations, Liogier 1995, Little and others
1974).
Range.—Roble de guayo is native to Cuba,

Hispaniola, and Puerto Rico, including the Island
of Vieques (Little and others 1974). It is not
elsewhere.
Ecology.—Roble de guayo grows in areas that

receive about 700 to 2000 mm of mean annual
precipitation. Elevations of habitat in Puerto Rico
range from 30 to 300 m (Little and others 1974). It
colonizes well- to excessively well-drained soils
with a wide range of textures and pH’s from about
5.5 to 8.0. Most of the hillside, ridge, and hilltop
soils are shallow and rocky. In Puerto Rico, roble
de guayo is locally common in the moist and dry
limestone hills and in dry foot hills of igneous and
metamorphic (including ultramafic) rocks
(Breckon and García 2001, Little and others 1974).
The species is an important component of the
cactus scrub vegetation type in Cuba (World
General Description.—Roble de guayo, also Wildlife Fund 2001). It is moderately intolerant of
known as palo de vaca, hoja menuda, guazumilla, shade, growing in the understory of low basal area
raspalengua, and cafecillo, is an evergreen shrub or forests but needs overhead sun to produce good
small tree sometimes reaching 9 m in height and fruit and seed crops. The species survives best in
15 cm in stem diameter at breast height. It is harsh environments where competition is minimal.
somewhat branchy and may have single or
multiple stems emerging from the ground. The Reproduction.—Roble de guayo flowers and
stem bark is gray and smoothish to slightly fruits throughout the year (Little and others 1974).
fissured. The wood is light brown, hard, heavy, Production of fruits and seeds is moderate. Two
and relatively brittle. Roble de guayo is deeply and collections of fresh fruits from Puerto Rico
solidly rooted with a tap and lateral root system of weighed an average of 0.453 + 0.093 g/fruit
brown, somewhat stiff roots. Twigs are slender, (Francis and Rodríguez 1993) and 0.423 + 0.012
hairless to densely hairy, green when young, g/fruit. Air-dried seeds separated from the
becoming gray. The coriaceous, shiny, green respective groups of fruits averaged 43,500
leaves are alternate but may be crowded near twig seeds/kg and 51,500 seeds/kg. Sown without
ends. Blades are elliptic, oblong, or obovate, 1.5 to pretreatment, the first sample of seed failed to
7 cm long by 0.7 to 2.5 cm broad, rounded to germinate and the second gave only 2 percent
slightly notched at the apex, and have a short germination (author’s observation). Germination is
petiole. The terminal inflorescence is a corymb or epigeal. The seeds are undoubtedly dispersed by
a cyme with two to 15 tubular, white flowers with birds that eat the fruits of this as well as other
five corolla lobes. Fruits are orange or red-orange, species of the genus. Seedlings are relatively
subglobose, 5 to 8 mm in diameter, often three- common.
lobed, drupes with persistant calyx and style. Each
fruit contains four or fewer light brown, hard Growth and Management.—Both seedlings and
nutlets. The nutlets are a quarter sphere with the adult plants grow slowly. Apparently, roble de
121
guayo lives for several decades. No management Puerto Rican trees and shrubs: second
experience has been published. The species is not installment. Research Note SO-374. U.S.
weedy in agricultural land or forest plantations and Department of Agriculture, Forest Service,
will rarely need control. Southern Forest Experiment Station, New
Orleans, LA. 5 p.
Benefits.—Roble de guayo helps protect the soil
and furnishes food and cover for wildlife. The Liogier, H.A. 1995. Descriptive flora of Puerto
stems are too small for use as lumber but should Rico and adjacent islands. Vol. 4. Editorial de
make excellent fuel. The species would probably la Universidad de Puerto Rico, San Juan, PR.
be a good ornamental for landscaping xeric 617 p.
gardens.
References Wadsworth. 1974. Trees of Puerto Rico and the
Virgin Islands. Vol. 2. Agriculture Handbook
Breckon, G.J. and R.G. García. 2001. Vascular 449. U.S. Department of Agriculture,
plants of Susúa Forest. University of Puerto Washington, DC. 1,024 p.
Rico, Mayaguez, PR. http://www.uprm.edu/
biology/profs/breckon/herbarium/florasusua. World Wildlife Fund. 2001. Cuban cactus scrub
htm. 52 p. (NT1306). http://www.worldwildlife.org/
wildworld/profiles/terrestrial/nt/nt1306_full.
Francis, J.K. and A. Rodríguez. 1993. Seeds of html. 8p.
122
Brickellia californica (Torr. & Gray) Gray California brickellbush
ASTERACEAE
Synonyms: Bulbostylis californica Torr. & Gray

Brickellia wrightii A. Gray
Coleosanthus californicus Kuntze
Coleosanthus albicaulis Rydb.
Brickellia tenera A. Gray
John K. Francis
supported by tap and lateral roots that are covered

by corky brown, finely-furrowed bark. The
alternate aromatic, gray-green leaves are deltoid-
ovate with a rounded or cordate base and a short-
acute tip and a crenate-dentate margin. They are 1
to 5 cm long and almost as wide with petioles 4 to
20 mm long. Inflorescences are heads in terminal
clusters on small lateral branches of leafy panicles.
The fragrant heads are 7 to 10 mm long and
contain eight to 18 greenish-cream florets. The
olive green achenes are 3 mm long tipped with a
white to brownish pappus. There is a base number
of n = 9 chromosomes (author’s observation,
Abrams and Ferris 1960, Harrington 1964, Munz
1974).
Range.—California brickellbush occurs in two

varieties. Variety californica grows in Oregon,
Idaho, California (including Santa Catalina,
Anacapa, and Santa Cruz Islands), Nevada, Utah,
Colorado, Arizona, New Mexico, Oklahoma,
Texas, and at least Chihuahua State in Mexico.
Variety jepsonii B.L. Robins is found only in
California (Abrams and Ferris 1960, Munz 1974,
Natural Resources Conservation Service 2003).
Although the species is planted as an ornamental,
it has not been reported to have naturalized outside
its native range.
General Description.—California brickellbush is Ecology.—California brickellbush is intolerant of

a suffruticose shrub 0.5 to 1.2 m in height. Stem shade and does not grow under tree or closed
bases reach a maximum diameter of about 2 cm, shrub canopies. Stands observed by the author
and clumps may reach a diameter of 1 m. Plants were along trails, intermittent streams, and steep,
usually have multiple stems arising from the root north-facing colluvial slopes. Abrams and Ferris
crown or short, near-surface horizontal stems. The (1960) say that California brickellbush grows in
branches are slender, upright, with even finer dry canyons and streambeds. It grows in soils
twigs. Current-year branches, which annually die derived from a wide range of parent materials,
back to a few cm from the ground-line, have a through the whole range of textures, and with pH’s
white to silver-gray bark, and older branches are of 5 to 8. California brickellbush grows in habitat
fine-furrowed gray or gray-brown. The wood is that receives annual rainfall amounts ranging from
yellowish white, medium density, and brittle with 220 to 640 cm (Las Pilitas Nursery 2003). The
annual rings. California brickellbush shrubs are species grows at between 915 and 2,290 m in
elevation in Arizona (Dittman 2003). It grows in
123
coastal and island vegetation types in California References
(Munz 1974) and therefore at low elevations.
Disturbance is probably required for Abrams, L., and R.S. Ferris. 1960. Illustrated flora
establishment. of the Pacific States. Stanford University Press,
Reproduction.—California brickellbush flowers
August through October (Tarweed Native Plants Dittman, L. 2003. Asteraceae: aster, daisy or
2003). The species flowers abundantly and composite family. http://www.nazflora.org/
potentially produces vast numbers of seeds. Air- asteraceae.htm. [not paged].
dried seeds collected in Utah averaged 3.525
million/kg. Hand-sorted seeds placed without any Harrington, H.D. 1964. Manual of the plants of
pretreatment on moist blotter paper germinated at Colorado. Sage Books, Denver, CO. 666 p.
71 percent beginning between 5 and 14 days after
sowing. Germination is epigeal (author’s Las Pilitas Nursery. 2003. Brickellia californica.
observation). Seed will have to be harvested by http://www.laspilitas.com/plants/123.htm. 3 p.
hand because wild plants are scattered. Stems layer
(root) when covered by soil or colluvial material. Moerman, D.E. 1986. Medicinal plants of Native
The seeds are wind-dispersed. Seedlings probably America. Technical Reports 19. University of
appear and survive only when conditions are Michigan Museum of Anthropology, Ann
nearly ideal. Arbor, MI. 534 p.
Growth and Management.—Save for a few Munz, P.A. 1974. A flora of Southern California.
centimeters of caudex, California brickellbush University of California Press, Berkeley, CA.
adds and loses its entire height (0.5 to 1.0 m) each 1,086 p.
growing season. The aerial portion of the caudex
also dies back about every 3 to 5 years while Natural Resources Conservation Service. 2003.
others are produced each year. Plant may live for Plants profile: Brickellia Ell., brickellbush.
one to several decades, renewing themselves with http://plants.usda.gov/cgi_bin/plant_profile.cgi?
suckers. The species is rarely abundant and symbol=BRICK. [not paged].
probably does not require control. Management
experience has not been published. Oklahoma Biological Survey. 2003. Brikellia
californica (Torr. & Gray) Gray.
Benefits.—California brickellbush helps protect http://www.biosurvey.ou.edu/shrub/brca3.htm.
the soil and furnishes food and cover for wildlife. 3 p.
It has been reported to be a larval food plant for
the tiger moth, Dysschema howardi Hy. Edwards Tarweed Native Plants. 2003. Current plant
(Walsh 2003). The species has been planted as an availability list. http://www.tarweednativeplants.
ornamental, particularly for the floral fragrance, in com/botanical.html. 5 p.
natural landscaping (Oklahoma Biological Survey
2003). Native Americans used infusions of the Walsh, B. 2003. Moths of southeastern Arizona:
leaves as a ceremonial emetic, as a lotion to treat Arctiidae (tiger moths). http://nitro.biosci.
infant skin sores, and for coughs and fevers Arizona.du/zeeb/butterflies/artic.html. 8 p.
(Moerman 1986).
124
Bromelia pinguin L. maya
BROMELIACEAE
Synonyms: none
John K. Francis
Ecology.—Maya is intermediate in shade

tolerance. Although it sometimes grows in open
areas, the most vigorous stands are found under
forest stands with moderate basal areas. In Puerto
Rico, natural stands occur in areas with from 850
to 2000 mm of rainfall and from near sea level to
600 m of elevation. All types of soils except very
poorly drained and saline soils are colonized.
Maya is sensitive to fire. Although many plants in
a colony will recover from a burn, they do so
slowly. Usually flowering is synchronized,
although an occasional plant flowers out of phase,
especially in moist habitat.
General Description.—Maya is a pineapple-like Reproduction.—Maya reproduces vegetatively

plant with large, sword-shaped, dark-green leaves and by seeds. The fruits, whose fresh weight
that have alternate curved spines about 5 mm long averaged 12.26 + 0.35 g in a Puerto Rican
on their edges. The name used is Spanish collection, contain 0 to over 100 seeds, depending
meaning “net” which derives from its use as a on their size. The fruits have a tough, fibrous rind.
barrier or possibly because its fibers may have In Puerto Rico, most of the fruits are eaten by
been used for making nets. Other common names fine-toothed animals (probably rats, mice, or fruit
are karatas, pingouin, bayonette, and pinguin bats) and it is assumed that the seeds are dispersed
(Howard 1979). Although it is mostly fleshy, by these small mammals. The black, teardrop-
maya is classed as a shrub because it is perennial, shaped seeds averaged 0.245 + 0.006 g (air dry).
of shrub size, and has a woody core at its base Seventy-five percent of these seeds germinated
and fibrous leaves. The plants are formed of a between 133 and 175 days after sowing. The
large basal rosette and rarely develop a plants are very fragilely rooted at first and develop
discernible stem. The roots are shallow, relatively at a moderate rate. After maya plants have reached
fine, all of a similar size, and radiate in all their full size and before flowering, most healthy
directions. At the start of a plant’s last year, it individuals produce one or sometimes two stiff
grows a stout scape inflorescence with many horizontal stolons about 0.5 m long. A new plant
wooly red-orange flowers. The new leaves forms at the terminus. The new plants grow
surrounding the inflorescence are also intensely rapidly and reach roughly half the parent plant’s
red-orange. A few months later, 3.5-cm-long height and diameter and become independent in
elliptical yellow berries ripen. After the fruits about a year. Consequently, (with the exception of
have withered, about 1 year after the start of those planted by humans) new colonies are started
fruiting, the plant dies. as seedlings from dispersed seeds and most plants
within colonies arise vegetatively. Maya has been
Range.—The native range of maya extends from propagated with apical bud explants with a high
Mexico through tropical South America and the degree of success (Mesa and Lajonchere 1996)
Caribbean islands (Howard 1979). In Puerto Rico
nearly all the stands are on or near abandoned Growth and Management.—Maya plants grow
farmland, which may indicate a relatively recent to full size (1 to 2 m in height and 2 to 3 m in
introduction. It has been planted and naturalized diameter) in 2 or 3 years. Mature stands can be
in Hawaii (Neal 1965) and many other tropical dense with interlacing crowns and little clear
areas. space. Five stands inventoried in Puerto Rico
ranged from 723 to 1,948 plants/ha. Four whole
125
plants averaged 81.6 percent water and ranged References
from 0.3 kg to 1.6 kg dry weight. The maya
portion of total stand dry biomass would run from Crane, J.C. 1945. Living fence posts in Cuba.
about 500 to 2,000 kg/ha. For the most part, maya Agriculture in America 5(2): 34-35, 38.
is undesirable in forests and pastures because it
takes up valuable space, restricts access, inhibits Guzman, D.J. 1975. Especies útiles de la flora
grazing, and provides nesting habitat and cover for Salvadoreña. Ministerio de Educación, Dirección
rats (Vélez and van Overbeek 1950). Maya stands de Publicaciones. San Salvador, El Salvador.
may be controlled by uprooting and piling the 703 p.
plants. If management objectives so dictate, maya
can be established by transplanting top-trimmed Howard, R.A. 1979. Flora of the Lesser Antilles,
wildlings. Leeward and Windward Islands. Vol. 3. Arnold
Benefits.—Historically, and to some extent MA. 586 p.
today, maya was used as a hedge or living fence
to inhibit entry into fields and homesteads (Crane Mesa, A.R. and G. Lajonchere. 1996.
1945). These hedges were never more than Micropropagation of Bromelia pinguin Lindl.
marginally effective—cattle and many species of Pastos y Forrajes 19(3): 217-223.
animals pass relatively easily and people can
ignore the spines or cut a path with a machete. Neal, M.C. 1965. In gardens of Hawaii. Special
The fruit, which has a white flesh, is bitingly acid, Publication 50. Bernice P. Bishop Museum
and tastes like pineapple, is eaten occasionally out Press, Honolulu, HI. 924 p.
of hand and used to make a refreshing drink (Neal
1965). The long, parallel leaf fibers were found to Vélez, I. and J. van Overbeek. 1950. Plantas
be of good quality and proposed for commercial indeseables en los cultivos tropicales. Editorial
production (Guzman 1975). Maya offers food and Universitario, Río Piedras, PR. 497 p.
cover for wildlife, such as small mammals and
birds.
126
Brunfelsia lactea Krug & Urban jasmín del monte
SOLANACEAE
Synonyms: none
John K. Francis
4-mm brown, elliptic seeds (Liogier 1995, Little
and others 1974).
Range.—Jasmín del monte is endemic to high-

elevation forests in eastern Puerto Rico (Liogier
1995, Little and others 1974) and is confined to
only a few locations. The species has been grown
widely in greenhouses and gardens, but there is no
record of it naturalizing.
Ecology.—Dwarf forests at the summits of

mountains are the habitat of jasmín del monte.
These areas lie between 980 and 1,100 m in
elevation (Little and others 1974) and receive more
than 3000 mm of mean annual precipitation. The
soils are clayey over igneous rocks with pH’s
between 5 and 6 and are saturated in their subsoils
most of the time. As an ornamental, jasmín del
monte blooms in partial shade (Gehrke 2000). In
natural habitat, the species can survive under forest
canopies but requires broken or full sunlight to
flower and fruit. Hurricane-force winds impact the
mountain tops where the species grows every one
to few decades, uprooting trees and defoliating the
General Description.—Jasmín del monte (a forest. This disturbance, plus occasional
Puerto Rican name), also known as aguacero, vega landslides, probably allows adult plants of jasmín
blanca, dama de noche, is an evergreen shrub or del monte to flower and fruit and seedlings to
occasionally a small tree. It has been recorded to 7 establish themselves. In recent years, the
m in height and 10 cm in stem diameter but disturbance of road cuts and other construction
usually measures only 1 to 3 m in height. Older have created additional habitat for the species. The
plants tend to have multiple stems and many, often minimum overwintering temperature for jasmín
intertwining branches. The bark is light brown, del monte as an ornamental is 10 °C
slightly furrowed, and usually covered with (Kuipplantenkwekerij Gommer 2003).
mosses and liverworts. The inner bark is white.
The wood is light brown, hard, and brittle. Plants Reproduction.—Jasmín del monte flowers
are supported by a tap and lateral root system of throughout the year (Little and others 1974). There
light brown, somewhat brittle roots. Roots are good fruit and seed production. A collection of
sometimes arise from the lower trunk. The leathery fresh fruits weighed from 2 to 8 g each. Air-dried
foliage is moderately abundant. Dark green leaves seed separated from those fruits averaged 0.0062 +
are elliptic, oval, or obovate, 5 to 15 cm long and 0.0002 g/seed. Sown without pretreatment in
2.5 to 6.5 cm broad, and have petioles 5 to 15 mm commercial potting mix, 13 percent germinated
long. The flowers are solitary at the terminus of between 1 and 3 months after sowing. Germination
twigs. They are strikingly fragrant, particularly at is epigeal. Seedlings are relatively common in
night. White or pale yellow, later tinged with disturbed patches near seed-bearing shrubs but rare
purple, the corolla is trumpet shaped, 5 to 7 cm elsewhere. Stems layer (root) whenever they come
long and 5 to 6 cm across, with five rounded lobes. in contact with the ground. Cut roots and stems
Purple to light brown, fleshy fruits (berries) are produce sprouts.
globose, 2 to 2.5 cm in diameter and contain many
127
Growth and Management.—Jasmín del monte http://www.tfts.org/Plant_Sale_May_2000.htm.
grows slowly in its natural habitat and appears to 12 p.
be long-lived. The author knows of no wildland
plantings or intentional management. Because the Kuipplantenkwekerij Gommer. 2003. Brunfelsia
species reproduces after disturbance, soil lactea. Kuipplantenkwekerij Gommer, Dalen,
unavoidably exposed during construction and road Netherlands. http://www.kwekerijgommer.com/
work should be left in good tilled condition to fotos/kuipplanten/Brunfelsia%20Lactea.htm.
encourage seedling establishment. 1 p.
Benefits.—Jasmín del monte helps hold the soil, Liogier, H.A. 1995. Descriptive flora of Puerto
provides cover and probably food for wildlife, and Rico and adjacent islands. Vol. 4. Editorial
adds to the aesthetics of the forest. The species is de la Universidad de Puerto Rico, Río Piedras,
planted as an ornamental in gardens and PR. 617 p.
greenhouses and is available for purchase from a
number of commercial nurseries. Little, E.L., Jr., R.O. Woodbury, and F.H.
Wadsworth. 1974. Trees of Puerto Rico and the
References Virgin Islands. Vol. 2. Agriculture Handbook
Gehrke, P. 2000. Sale plants 2000. Tropical Washington, DC. 1,024 p.
Flowering Tree Society, Miami, FL.
128
Caesalpinia bonduc (L.) Roxb. gray nicker bean
FABACEAE
Synonyms: Guilandina bonduc L.

Guilandina bonducella L.
Caesalpinia bonducella (L.) Fleming
Caesalpinia crista of Small and other authors
Guilandina crista (L.) Small
John K. Francis
Range.—Gray nicker bean is established in nearly

every tropical shore worldwide, including Florida,
Louisiana, Puerto Rico, the U.S. Virgin Islands,
Hawaii, Guam, and American Samoa (Howard
1988, Liogier 1988, Pacific Island Ecosystems at
Risk 2002, Vermillion 2001). It has apparently
arrived at these locations by natural means,
although unrecorded accidental or intentional
introductions cannot be ruled out.
Ecology—Gray nicker bean is intolerant of shade

and is usually open-grown or at least grows in
broken sunlight. It tolerates salt spray, salty soils,
and occasional flooding with seawater. The species
grows in all textures of mildly acid to alkaline soil.
General Description.—Gray nicker bean, also Annual rainfall in the areas where gray nicker bean
known as nicker nut, wait-a-bit, hold back, fever grows in Puerto Rico ranges from 750 mm to 1800
nut, hava de San Antonio, mato de playa, z’yeux à mm. The species grows from sea level to 850 m in
chatte, canique grise, and uri, is a spiny, elevation in India (Parrotta 2001). It grows most
scrambling shrub to 1.5 m in height (unsupported) frequently on the beach strand, on coastal dunes,
and 6 m or more in extension, and has stems up to and at the better-drained edge of mangroves. Gray
5 cm in diameter or more. Plants usually have a nicker bean also grows inland in disturbed areas. It
single stem arising from the ground but often competes well with grass and herbs and may
branch low on the stem. Seedlings form taproots ascend into the crowns of low trees.
and may retain them later; lateral roots are
extensive. The stems grow stiffly upright for 0.75 Reproduction.—Gray nicker bean flowers and
to 1 m before curving into a more horizontal fruits year-round in Florida (Nelson 1996). A
direction. The stems, twigs, and leaf rachises are collection of seeds from Puerto Rico weighed an
covered with straight or curved prickles. The 25- to average of 2.444 + 0.034 g/seed or 409 seeds/kg.
80-cm-long leaves are bipinnately compound with Twenty-five seeds each of weathered-scarified (by
four to nine pairs of pinnae, each with four to eight nicking), weathered-unscarified, fresh-scarified,
pairs of oblong to elliptic leaflets. Racemes of and fresh-unscarified treatments were incubated in
yellow flowers are lateral or terminal. Prickley, plastic bags of moist potting mix. Most of the
inflated legumes are flattened oval shaped, 5 to 10 scarified seed germinated within a few days and
cm long, and reddish-brown when dry. Within each had all germinated within 1 year. By 16 months, 23
pod are one to three (usually two) smooth, hard, 2- of the fresh, unscarified seeds had germinated and
cm long seeds that are olive drab in the pods and just one of the weathered, unscarified seeds had
remain so until exposure to the sun bleaches them germinated. Germination is hypogeal (author’s
to a light gray color. The pods open partially upon observation). Gray nicker bean seeds are
drying and eventually release the seeds (author’s apparently carried to sea by floods and storm
observation, Howard 1988, Liogier 1988, Nelson surges where they float until deposited on shore.
1996). There are 2n = 24 chromosomes (Long and The seeds are found on beaches in northern
Lakela 1976). Scotland and are known to be able to float in sea
129
water for as long as 19 years (Markland 2002). The References
scarifying action of sand, weathering, insects, or
rodents eventually allows water to enter the seeds Ajiwe, V.I.E., C.A. Okeke, H.U. Agbo, G.A.
and they germinate and establish themselves in the Ogunleye, and S.C. Ekwuozor. 1996. Extraction,
new environments. A seed collected at Macquarie characterization and industrial uses of velvet-
Island, Antarctica, a drift of not less than 1 year, tamarind, physic-nut and nicker-nut seed oils.
was scarified and grown successfully (Costin Bioresource Technology 57(3): 297-299.
1965). The stems layer (root) whenever they come
in contact with the ground. Stems sprout when cut Ali, M.S., S. Shameel, V.U. Ahmad, and K.
and lateral roots sometimes sucker. Clonal plants Usmanghanim. 1997. Chemical constituents of
tend to remain attached to one another. Caesalpinia bonduc. Pakistan Journal of
Scientific and Industrial Research 40(1-4): 20-
Growth and Management.—Growth of gray 22.
nicker beans is fairly rapid at all stages. Seedlings
reached 26 cm in 40 days after sowing. Older Amarsinghe, A.P.G., R.D. Sharma, C. Chaturvedi,
plants grow a meter or more per year. Individual and D.K. Agarwal. 1993. Anthelmintic effect of
stems live at least 4 years. The author knows of no Ayurvedic recipe Kuberakshadi yoga in
published management experience. Plantations intestinal worms among children.Journal of
undoubtedly could be established with potted Research and Education in Indian Medicine
seedlings or by direct seeding scarified seeds. 12(1): 27-31.
Control could probably be obtained by lopping
with a machete followed by herbicide treatment of Biswas, T.K., S. Bandyopadhyay, Biswapati-
the sprouts. Mukherjee, Bhaswar-Mukherjee, B.R. Sangupta,
and B. Mukherjee. 1997. Oral hypoglycemic
Benefits.—Gray nicker bean contributes to the effect of Caesalpinia bonducella. International
biodiversity of the forests where it grows, helps Journal of Pharmacognosy 35(4): 261-264.
protect the soil, and furnishes cover for wildlife.
The seeds have been used for centuries and are still Burkill, H.M. 1995. The useful plants of West
used as jewelry, prayer beads, good luck charms, Tropical Africa. Vol. 3. Royal Botanic Gardens,
and worry stones (Rancho Leona 2002, Workman Kew, UK. 857 p.
1980). They were anciently used as standards of
weight in India (Vijayanagara Coins 2002). The Costin, A.B. 1965. Long-distance seed dispersal to
ancient African game of mancala traditionally Macquarie Island. Nature 206: 317.
employs gray nicker beans as game pieces
(Driedges 1972). The species is sometimes planted Driedges, W. 1972. The game of Boa, or Mankala,
as a hedge to prevent undesired entry into property in East Africa. Mila 3(1): 7-19.
(Nelson 1996) and could be planted for dune
stabilization. Preparations of the seeds and other Howard R.A. 1988. Flora of the Lesser Antilles,
plant parts are used to treat a large range of Leeward and Windward Islands.
ailments (Burkill 1995, Parrotta 2001). Dicotyledoneae, Part 1. Vol. 4. Arnold
Triterpenoids, fatty acid triglicerides, and sterols Arboretum, Harvard University, Jamaica Plain,
isolated from seeds may possibly explain some of MA. 673 p.
the activity (Ali and others 1997, Rostogi and
others 1996). Seed extracts have been shown to Liogier, H.A. 1988. Descriptive flora of Puerto
lower blood sugar in laboratory animals (Biswas Rico and adjacent islands, Spermatophyta. Vol.
and others 1997, Sharma and others 1997) and 2. Editorial de la Universidad de Puerto Rico,
effectively suppress or cure infections of several Río Piedras, PR. 481 p.
species of round worms (Amarsinghe and others
1993, Rastogi and others 1996). The fat content of Long, R.W. and O. Lakela. 1976. A flora of
gray nicker nut seeds was measured at 34 + 0.83 Tropical Florida. Banyon Books, Miami, FL.
percent. The semidrying oil extracted would be 962 p.
useful for manufacture of high-quality alkyd resins,
polishes, and paint (Ajiwe and others 1996). Markland, J. 2002. Drift seeds. In: West Word,
Community paper for Glenfinnan, Lochailort,
Glenuig, Arisaig, Morar, Mallaig, Knoydart, and
the Small Isles. March 2002. http://road-to-the-
130
isles.org.uk/westword/march2002.html. p. 6-7. antifilarial triglyceride fraction from Caesalpinia
bonduc. Fitoterapia 67(1): 63-64.
Nelson, G. 1996. The shrubs and woody vines of
Florida. Pineapple Press, Inc., Sarasota, FL. Sharma, S.R., S.K. Dwivedi, and D. Swarup. 1997.
391 p. hypoglycaemic, antihyperglycaemic and
hypolipidemic activities of Caesalpinia
Pacific Island Ecosystems at Rick. 2002. bonducella seeds in rats. Journal of
Caesalpinia bonduc (L.) Roxb., Fabaceae. Ethnopharmacology. 58(1): 39-44.
http://www.hear.org/pier3/cabon.htm. 2 p.
Vermillion, W.G. 2001. Caesalpinia bonduc
Parrotta, J.A. 2001. Healing plants of Pininsular (Fabaceae) new to Louisiana. Sida 19(4): 1,181-
India. CABI Publishing, Wellingford, UK and 1,182.
New York. 917 p.
Vijayanagara Coins. 2002. Coinage. http://www.
Rancho Leona. 2002. Rancho Leona jewelry vijayanagaracoins.com/htm/coinage.htm. 7 p.
samples: rainforest seed chart.
http://rancholeona.com/seed.html. 2 p. Workman, R.W. 1980. Growing native. The
Sanibal-Captiva Conservation Foundation, Inc.,
Rastogi, S., A.K Shaw, and D.K. Kulshreshtha. Sanibal, FL. 137 p
1996. Characterization of fatty acids of .
131
Cajanus cajan (L.) Millsp. pigeon pea
FABACEAE
Synonyms: Cystisus cajan L.

Cajanus bicolor DC.
Cajanus flavus DC.
Cajanus indicus Spreng.
Cajanus luteus Bello
John K. Francis
which are flattened, somewhat constricted between

seeds, and 4 to 8 cm long, are mottled bronze-
purple when immature, drying to brown. They
contain two to nine mottled brown (white, red,
brown, gray, or black in improved varieties) seeds,
7 to 8 mm long by 6 mm broad. There are 2n = 22,
44, or 66 chromosomes (Liogier 1988, Long and
Lakela 1976).
Range.—Pigeon pea probably originated in India,

but may have come from Africa. Both are centers
of diversity for the genus Cajanus. It is clear that
the species has been under cultivation for a long
time and was spread by traders thousands of years
ago. The wild progenitor may be Cajanus
cajanifolius (Haines) van der Maesen of India and
Myanmar (van der Maesen 1990). Today, pigeon
pea is cultivated throughout the tropics and has
naturalized in many areas including Florida, Puerto
Rico, and the U.S. Virgin Islands (Liogier 1988,
Long and Lakela 1976).
Ecology.—Pigeon pea grows on a broad range of

General Description.—Pigeon pea, also known as well-drained soils, from sands to clays over
red gram, Congo pea, gungo pea, no eye pea, dhal, sedimentary, igneous, and metamorphic parent
gandul, gandure, frijol de árbol, and pois cajan, materials. It tolerates pH’s of from 4.5 to 8.4 and
occurs in several varieties. The old varieties in some varieties tolerate 6 to 12 mmhos/cm of
cultivation and semicultivation in the West Indies salinity. However, the species is sensitive to
treated here are semideciduous, short-lived shrubs waterlogging. Pigeon pea will grow in areas that
usually 1 to 4 m in height and 1 to 4 cm in basal receive from 530 to 4000 mm of mean annual
stem diameter. They are usually single stemmed, precipitation (Van Den Beldt 1988). Pigeon pea
freely branching, and become woody after a few grows best in temperatures of 18 to 30 °C. It is
months. The wood is moderately hard and brittle. subject to frost damage (Skerman and others 1988).
They have a deep taproot (to 3 m) with lateral roots The species is grown under cultivation from near
and nodulated fine roots. The branches and fine sea level to 3,000 m in elevation (Van Den Beldt
twigs support abundant light-green or yellow-green 1988). Although a large number of insects and
foliage. Spirally arranged, silky-pubescent, diseases affect it (American Phytopathological
trifoliolate leaves have narrowly elliptic, lanceolate Society 2002), the actual incidence of attack is low
or oblong leaflets, 2.5 to 9 cm long, the center (Center for New Crops and Plants Products 2002).
leaflet being slightly longer than the laterals. The Pigeon pea is intolerant of shade and tolerates only
five- to 12-flowered racemes are axillary. Flowers moderate competition. It does best in full sun on
are about 2 cm long, yellow, the standard often bare ground but can grow with side shade or
being orange to purple outside. The legumes, broken shade from trees and a low cover of grass
132
and forbs. Growth is moderately slow during the stems, the plant is damaged by browsing,
first 2 to 3 months of life during which time especially by cattle (Department of Primary
seedlings are not competitive with grass and Industries 2002). The species is planted as a green
weeds; afterwards pigeon pea competes well with manure crop, nurse crop, cover crop, a windbreak
vegetation equal or lower in height (van der hedge, as a host for lac insects, and as food for silk
Maesen 1990). In Puerto Rico, the species persists worms. The stalks are used for fuel, thatch, and
for a generation or two after cultivation and grows basketry (Center for New Crops and Plants
along roadsides and in waste places where seeds Products 2002). Pigeon pea forms root nodules in
have fallen, but the populations are not sustained association with Rhizobium sp. bacteria and is
indefinitely. Although seldom devastating, a large capable of fixing 41 to 280 kg/ha of nitrogen (Red
number of insects and diseases affect pigeon pea de Grupos de Agricultura de Cobertura 2002).
and can reduce yield or shorten a plant’s life Preparations of the leaves are used to treat
(American Phytopathological Society 2002). jaundice, inflammation, and sores of the mouth
(Parrotta 2001).
Reproduction.—Pigeon pea flowers nearly
throughout the year (Red de Grupos de Agricultura References
de Cobertura 2002). The flowers are self-
compatible and usually self-pollinated (Smartt American Phytopathological Society. 2002.
1976). Insects visit and pollinate the flowers (5 to Common names of plant diseases: Diseases of
40 percent cross pollination), but it is not known pigeonpea [Cajanus cajan (L.) Millsp.].
whether they increase seed yield (McGregor 2002). http://www.apsnet.org/online/common/names/
Upon drying, the pods spring and coil to release the pigeon.asp. 5 p.
seeds. Pigeon pea seeds weigh about 0.1g and
germinate in about 2 weeks (Center for New Crops Center for New Crops and Plants Products. 2002.
and Plants Products 2002). Cajanus cajan (L.) Millsp. Purdue University.
http://www.hort.purdue.edu/newcrop/duke_
Growth and Management.—Pigeon pea is energy/Cajanus_cajun.html. 6 p.
normally sown directly into prepared ground.
Seeding rates for pure stands are 12 to 25 kg of Department of Primary Industries. 2002. Legumes
seed/ha (Smartt 1976). Seeding depths of 2.5 to 5 for the tropics: pigeon pea (Cajanus cajan).
cm are recommended (Center for New Crops and Queensland Government. http://www.dpi.qld.
Plants Products 2002). No pregermination gov.au/pastures/4571.html. 2 p.
treatment of the seed is needed. Although some
varieties mature seed in 5 to 6 months, longer- Liogier, H.A. 1988. Descriptive flora of Puerto
lived, tall varieties including those that are more Rico and adjacent islands, Spermatophyta. Vol.
competitive in the wild take 10 to 12 months to 2. Editorial de la Universidad de Puerto Rico,
mature seed. These plants live about 5 years Río Piedras, PR. 481 p.
(Smartt 1976). Experimental yields of 50 dry
t/ha/year have been demonstrated; yields of 3 to 8 Long, R.W. and O. Lakela. 1976. A flora of
dry t/ha/year are obtained under normal Tropical Florida. Banyan Books, Miami, FL.
management (Van Den Beldt 1988). 962 p.
Benefits.—Pigeon peas are an important food in McGregor, S.E. 2002. Insect pollination of
developing tropical countries. An excellent source cultivated crop plants. Agriculture Research
of protein, the seeds (and sometimes the pods) are Service. http://gears.tucson.ars.ag.gov/book/
eaten as a vegetable, as a flour additive to other chap4/pig.html. [not paged].
foods, in soups, and with rice (Center for New
Crops and Plants Products 2002). Although they Nene, Y.L. and V.K. Sheila. 1990. Pigeonpea:
vary slightly, typical nutritional values for seeds geography and importance. In: Y.L. Nene, S.H.
are: moisture, 10.1 percent, protein 19.2 percent, Hall, and V.K. Sheila. The pigeonpea. CAB
fat, 1.5 percent, carbohydrates, 57.3 percent, fiber International, Wellingford, UK. p. 1-14.
8.1 percent, and ash, 3.8 percent (Smartt 1976).
About 3.4 million ha were under cultivation in Parrotta, J.A. 2001. Healing plants of Peninsular
1978-1988 period, 88 percent of it in India (Nene India. CABI Publishing, Wallingford, UK and
and Sheila 1990). Although pigeon pea makes New York. 917 p.
excellent forage, because of the brittleness of its
133
Red de Grupos de Agricultura de Cobertura. 2002. Smartt, J. 1976. Tropical pulses. Longman Group
Base de información sobre especies con Limited, London. 348 p.
potential de abonos verdes y cultivos de
cobertura. Rockefeller Foundation. http://www. Van Den Beldt, R.J. 1988. Cajanus cajan: it’s
rockfound.org.mx/cajanusbiesp.html. [not more than just a pulse crop. Nitrogen Fixing
paged]. Tree association. NFT Highlights 88-06. 5 p.
Skerman, P.J., D.G. Cameron, and F. Riveros. van der Maesen, L.J.G. 1990. Pigeonpea: origin,
1988. Tropical forage legumes. FAO Plant history, evolution, and taxonomy. In: Y.L.
Production and Protection Series 2. Food and Nene, S.H. Hill, and V.K. Sheila. The
Agriculture Organization of the United Nations. pigeonpea. CAB International. Wellingford,
692 p. UK. p. 15-46.
134
Callicarpa americana L. American beautyberry
VERBENACEAE (also placed in LAMIACEAE alt. Labiatae)
Synonyms: None
Kristina Connor
the West Indies (Bailey and Bailey 1976, Brickell

and Zuk 1996).
Ecology.—American beautyberry is a drought-

tolerant perennial shrub. Classified as a pioneer
species (Odenwald and Turner 1988), it is
common along forest edges and fence rows, and is
also a common understory plant in open pine
plantations in the southeastern United States. It is
found in openings after clear-cutting (Odenwald
and others 1996) and in moist thickets and
bordering swamps (Krüssman 1976). American
beautyberry is very tolerant of fire and resprouts
easily. However, it is intolerant of deep shade and
is only found in the better drained areas of
bottomland hardwood stands. It will grow in a
variety of soil textures and in a wide pH range
(USDA-NRCS 2001). Tolerant of temperatures
ranging from –17 oC to over 37.8 oC (Martin and
Mott 1997), it is cultivated for its fall display of
purple to violet fruits.
Illustration source: UDA [no date]
Reproduction.—The perfect, bluish white to
General Description.—American beautyberry, white axillary flowers first appear in early spring.
also known as beauty-berry, Bermuda mulberry, The bloom period can continue through late
French mulberry, sourbush, sow-berry, or Spanish summer. It is not uncommon to see both flowers
mulberry, is a shrub that can reach 2.4 to 2.7 m in and fruits on the same plant (Grelen and Duvall
height but is typically shorter, averaging 0.9 to 1.8 1966). The flowers are borne in dichotomous
m. The irregular, spreading bush has an open cymes 8 to 36 mm long (Bonner in press). The 3-
growth habit and grows as a single plant or in to 6-mm fruits are violet to purple berry-like
colonies. Bark is brown and raised lenticels are drupes, formed in clusters that encircle the stem.
found on older stems. Branches are scurfy or Each drupe contains four small seeds that
tomentose, gray to reddish brown in color. The germinate without scarification or stratification,
opposite, short-petioled, simple, deciduous leaves although germination can be slow. If seeds are
are ovate to elliptic in shape. They are pointed at sown in the fall, germination the following spring
both the base and the apex, and the margins are may be excellent (Dirr and Heuser 1987). Bonner
coarsely serrate. Leaves are glabrous or slightly (in press) reported that seeds can be easily cleaned
pubescent above and pubescent with stellate using any type of macerator, even a kitchen
trichomes below (Radford and others 1968, Martin blender for small lots. Seeds average 600 per
and Mott 1997). Leaf size is variable, ranging from gram. Longevity of seeds in storage is unknown,
7.6 to 20.3 cm in length and 5 to 12.7 cm in width. but they survive for at least 1 year in the soil seed
bank and are thought to be orthodox. Orthodox
Range.—American beautyberry ranges through the seeds can be dried to a moisture content of less
Southeastern United States from Maryland and than 12 percent and stored under refrigeration for
Virginia south to Florida and west to Arkansas, long periods (Roberts 1973). These seeds are
Oklahoma, and Texas. It also grows in the transported by animals and birds. American
Bahamas, Bermuda, Cuba, northern Mexico, and in beautyberry may also be propagated from
softwood stem cuttings taken in the summer and
135
fall that are treated with indol-3-butyric acid (IBA; Grelen, H.E. and V.L. Duvall. 1966. Common
1000 ppm) and put in a mist bed (Dirr and Heuser plants of longleaf pine-bluestem range. Research
1987). Paper SO-23. U.S. Department of Agriculture
Forest Service, Southern Forest Experiment
Growth and Management.—American Station, New Orleans, LA. 96 p.
beautyberry is a short-lived shrub that is easy to
grow and widely distributed in the Southeastern Krüssmann, G. 1976. Manual of Cultivated Broad-
United States. It can be used as a landscape border Leaved Trees and Shrubs. Volume I, A-D.
and makes a good mass planting. However, regular Timber Press, Beaverton, OR. 448 p.
pruning is required if it is to maintain its form and
produce masses of the fruits so attractive in the fall Martin, C.O. and S.P. Mott. 1997. American
of the year. The fruits occur only on new growth, Beautyberry (Callicarpa americana). Section
and plants should be heavily pruned in early spring 7.5.8, U.S. Army Corps of Engineers Wildlife
(Martin and Mott 1997). It will not do well in Resources Management Manual. Ecosystem
areas that experience flooding but can be used to Management and Restoration Research Program
restore surface-mined sites (Martin and Sick Technical Report EL- 97.15. Conservation
1995). It will resprout after fire (Martin and Mott Communications, Vicksburg, MS. 17 p.
1997) and can be easily transplanted.
Martin, H. and G. Sick. 1995. American
Benefits.—American beautyberry is a good source beautyberry for borrow pit reclamation in South
of food for deer and the fruits attract birds. It can Carolina Restoration and Management Notes
be used in reclamation work and for erosion 13(1): 90-96.
control (Brown 1945). It easily reseeds in nature.
Odenwald, N.G., C.F. Fryling, Jr., and T.E. Pope.
References 1996. Plants for American Landscapes. Louisiana
State Univ. Press, Baton Rouge, LA. 266 p.
Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third:
A Concise Dictionary of Plants Cultivated in the Odenwald, N.G. and J.R. Turner. 1988. Plants for
United States and Canada. McMillan Publishing the South: A guide for landscaping design.
Co., Inc., New York. 1,290 p. Claitor’s Publication Division, Baton Rouge, LA.
565 p.
Bonner, F.T. (In press). Callicarpa americana L.
American beautyberry. In: Seeds of Woody Radford, A.E., H.E. Ahles, and C.R. Bell. 1968.
Plants in the United States. Revised edition. Manual of the Vascular Flora of the Carolinas.
University of North Carolina Press, Chapel Hill.
Brickell, C. and J.D.. Zuk. 1996. The American 1,183 p.
Horticultural Society A-Z Encyclopedia of
Garden Plants. DK Publishing, Inc., New York. Roberts, E.H. 1973. Predicting the storage life of
1,092 p. seeds. Seed Science and Technology 1: 499-514.
Brown, C.A. 1945. Louisiana trees and shrubs. USDA, NRCS. 2001. The PLANTS database.
Bulletin 1, Louisiana Forestry Commission, National Plant Data Center, Baton Rouge,
Baton Rouge, LA. 262 p. http://plants.usda.gov. LA [not paged].
Dirr, M.A. and C.W. Heuser, Jr. 1987. The USDA. [no date]. Southern wetland floral field
reference manual of woody plant propagation: office guide to plant species. U.S. Department
from seed to tissue culture. Varsity Press, Athens, of Agriculture Soil Conservation Service, South
GA. 239 p. National Technical Center, Fort Worth, TX.
http://www.npwrc.usgs.gov. [not paged]
.
136
Calotropis procera (Ait.) Ait. f. giant milkweed
ASCLEPIADACEAE
Synonym: Asclepias procera Ait.
John K. Francis
cymes that grow at or near the ends of twigs. The

flowers are shallowly campanulate with five sepals
that are 4 to 5 mm long, fleshy and variable in
color from white to pink, often spotted or tinged
with purple. The fruits are inflated, obliquely ovoid
follicles that split and invert when mature to
release flat, brown seeds with a tuft of white hairs
at one end (Howard 1989, Liogier 1995, Little and
others 1974).
Range.—Giant milkweed is native to West Africa

as far south as Angola, North and East Africa,
Madagascar, the Arabian Peninsula, southern Asia,
and Indochina to Malaysia (Rahman and Wilcock
1991). The species is now naturalized in Australia,
many Pacific islands, Mexico, Central and South
America, and the Caribbean islands.
Ecology.—Giant milkweed favors open habitat

with little competition. This condition is most
completely met in overgrazed pastures and
rangeland. Other common habitats are beachfront
dunes, roadsides, and disturbed urban lots. The
species grows in dry habitat (150 to 1000 mm
General Description.—Giant milkweed is also precipitation) and sometimes in excessively
known as sodom apple, calotrope, French cotton, drained soils in areas with as much as 2000 mm of
small crown flower (English), algodón de seda, annual precipitation. Giant milkweed may be found
bomba (Spanish), cotton-france, arbre de soie, and in areas up to 1,000 m in elevation in India
bois canon (French) (Howard 1989, Liogier 1995, (Parrotta 2001). It roots very deeply and rarely
Neal 1965, Parrotta 2001). grows in soils that are shallow over unfractured
This plant is a soft-wooded, evergreen, rock. Soils of all textures and derived from most
perennial shrub. It has one or a few stems, few parent materials are tolerated, as well as soils with
branches, and relatively few leaves, mostly high sodium saturation. Beachfront salt spray is not
concentrated near the growing tip. The bark is detrimental. Competition with tall weeds, brush,
corky, furrowed, and light gray. A copious white and especially grass weakens existing plants, and
sap flows whenever stems or leaves are cut. Giant being overtopped and shaded by trees soon
milkweed has a very deep, stout taproot with few eliminates them. During droughts in Puerto Rico,
or no near-surface lateral roots. Giant milkweed giant milkweed is attacked by the orange aphid,
roots were found to have few branches and reach Aphis nerii Boyer de Fonscolombe, which causes
depths of 1.7 to 3.0 m in Indian sandy desert soils defoliation, death of branches, and aborted fruits
(Sharma 1968). The opposite leaves are oblong- (Little and others 1974).
obovate to nearly orbicular, short-pointed to blunt
at the apex and have very short petioles below a Reproduction.—Flowering and fruiting takes
nearly clasping, heart-shaped base. The leaf blades place throughout the year (Little and others 1974).
are light to dark green with nearly white veins. Hundreds to thousands of seeds may be produced
They are 7 to 18 cm long and 5 to 13 cm broad, per plant each year. The seeds in a Puerto Rican
slightly leathery, and have a fine coat of soft hairs collection averaged 0.0095 + 0.0027 g/seed or
that rub off. The flower clusters are umbelliform
137
about 100,000 seeds/kg (author’s observation). leaves are chopped and mixed with other feed,
Eighty-nine percent germination took place in consumption greatly increases with no ill effects.
potting mix between 7 and 64 days after sowing. (Abbas and others 1992, Nehra and others 1987).
Half the seed weight was found in the wing (silk). Shade-dried giant milkweed leaves contained 94.6
The seeds are dispersed by wind and may fly for percent dry matter, 20.9 percent ash, 19.6 percent
several hundred yards in gentle breezes. Seedlings crude protein, 2.2 percent fat, 43.6 percent acid
may arise in abundance after rainy periods, but detergent fiber, and 19.5 percent neutral detergent
only a few survive the first season. Using the fiber (Abbas and others 1992). Although it is
reserves in its large taproot, giant milkweed can lightweight, the wood is used in impoverished
resprout year after year when burned or cut. desert areas for a cooking fuel (Varshney and Bhoi
1988).
Growth and Management.—Giant milkweed
usually reaches heights of about 2 m, but may References
occasionally reach 5 m in height and stem
diameters of 25 cm (Little and others 1974). Abbas, B., A.E. El Tayeb, and Y.R. Sulleiman.
Growth is about 1 m the first year after sprouting 1992. Calotropis procera: feed potential for arid
in Puerto Rico. Senescence of individual stems zones. Veterinary Record 131(6): 132.
takes place in about 5 years, but plants often
resprout afterwards. Giant milkweed does not form Anver, S. and M.M. Alam. 1992. Effect of latex
dense stands, normally occurring as scattered seed dressing on interacting root-knot and
individuals. It can be a serious weed in pastures, reniform nematodes. Afro-Asian Journal of
overgrazed rangelands, and poorly managed hay Nematology 2: 1-2, 17-20.
fields. Although it probably is not possible to
eliminate existing stands through management Campolucci, P. and C. Paolini. 1990.
without ceasing grazing and harvest, it may be Desertification control in the Sahel regions—
possible to prevent their establishment. Some form low-cost large-scale afforestation techniques.
of chemical control would seem to be the only Note Tecniche 10. Centro di Sperimentazione
practical option for eliminating existing stands but Agricola e Forestale di Sperimentazione per la
no specific guidelines are available. Establishing Pioppicoltura. 24 p.
giant milkweed has been advocated for
environmental protection and as a nurse crop for Charu-Jain and P.C. Trivedi. 1997. Nematicidal
more valuable species (Campolucci and Paolini activity of certain plants against root-knot
1990). This can be done easily by planting nematode, Meloidogyne incognita, infecting
containerized seedlings or rooted cuttings. chickpea, Cicer arietinum. Annals of Plant
Protection Sciences 5(2): 171-174.
Benefits.—Giant milkweed tissues, especially the
root bark, are used to treat a variety of illness Howard, R.A. 1989. Flora of the Lesser Antilles,
including leprosy, fever, menorrhagia, malaria, Leeward and Windward Islands. Dicotyledoneae.
and snake bite (Parrotta 2001). The latex is toxic Part 3. Vol. 6. Arnold Arboretum, Harvard
and can cause blisters and rash in sensitive University, Jamaica Plain, MA. 658 p.
persons. The plant is occasionally grown as an
ornamental in dry or coastal areas because it is Liogier, H.A. 1995. Descriptive flora of Puerto
handsome, of a convenient size, and is easy to Rico and adjacent islands. Vol. 4. Editorial
propagate and manage. It is recommended as a de la Universidad de Puerto Rico, San Juan, PR.
host plant for butterflies (Mikula 2001). In the 617 p.
past, the silky hairs were used to stuff pillows
(Little and others 1974). Giant milkweed was Little, E.L., Jr., R.O. Woodbury, and F.H.
tested as a host for sandalwood, Santalum album Wadsworth. 1974. Trees of Puerto Rico and the
L., a partial root parasite. It resulted in greater Virgin Islands. Vol. 2. Agriculture Handbook
growth of sandalwood than all other species tested 449. U.S. Department of Agriculture,
(Shinde and others 1993). Extracts, chopped Washingtion, DC. 1,024 p.
leaves, and latex have shown great promise as
nematicides, in vitro and in vivo (Anver and Alam Mikula, R. 2001. Butterfly plants for your garden.
1992, Charu and Trivedi 1997). Sheep, goats, and www.butterflybreeders.com/pages/bflygdning/bu
camels will eat the leaves of giant milkweed tterflyplants.html. 5 p.
during droughts, but consumption is low. If the
138
Neal, M.C. 1965. In gardens of Hawaii. Special 11(3): 301-308.
Press, Honolulu, HI. 924 p. Sharma, B.M. 1968. Root systems of some desert
plants in Churu, Rajasthan. Indian Forester
Nehra, O.P., M.C. Oswal, and A.S. Faroda. 1987. 94(3): 240-246.
Management of fodder trees in Haryana. Indian
Farming 37(3): 31, 33. Shinde, S.R., R.D. Ghatge, and S.S. Mehetre. 1993.
Comparative studies on the growth and
Parrotta, J.A. 2001. Healing plants of Peninsular development of sandalwood tree in association
India. CAB International, Wallingford, UK and with different hosts. Indian Journal of Forestry
New York. 944 p. 16(2): 165-166.
Rahman, M.A. and C.C. Wilcock. 1991. A Varshney, A.C., and K.L. Bhoi. 1988. Cloth from
taxonomic revision of Calotropis bast fibre of the Calotropis procera (aak) plant.
(Asclepiadaceae). Nordic Journal of Botany Biological wastes 26(3): 229-232.
139
Capparis indica (L.) Druce sapo prieto
CAPPARACEAE
Synonyms: Capparis indica (L.) Rawc. & Rendle

Breynia indica L.
Capparis breynia L.
Capparis amygdalifolia Jacq.
Capparis amygdalina Lam.
Linnaeobreynia indica (L.) Hutch.
John K. Francis
narrowly elliptic, leathery blades 4 to 11 cm long

and 2 to 4.5 cm broad. The inflorescences
(corymbs) are near the ends of branches and
contain a few small, white flowers with long white
stamens and short yellow anthers. The fruits are
silvery-brown legume-like pods 4 to 26 cm long
that split along one side at maturity to expose a
bright-red interior and pulp-covered seeds. The
seeds are black, elliptic, and 5 to 7 mm in diameter
1974, Zamora 1989).
Range.—The range of sapo prieto includes

Jamaica, Hispaniola, Puerto Rico, the Virgin
Islands, the Lesser Antilles, the Dutch Antilles,
Trinidad, Venezuela, Southern Mexico to Costa
Rica and possibly Panama (Liogier 1985, Little and
others 1974, Zamora 1989).
Ecology.—Sapo prieto grows in a wide range of

well-drained soils derived from both sedimentary
(including limestone), igneous, and metamorphic
rocks. It may be found from near sea level to about
General Description.—Sapo prieto, a name used 600 m on steep as well as level slopes, and in areas
in Puerto Rico that means black toad in Spanish, is receiving from 750 to about 1700 mm of
also known as caper, linguam, burro, white willow, precipitation in Puerto Rico. In Costa Rica, sapo
colorín, vara prieta, palo zapo, taiche, endurece prieto grows from sea level to 300 m in elevation
maíz, curumo, guacoco, naranjuelo, pachaca, olivo in very rocky soils (Zamora 1989). The species is
macho, olivo, bois de mêche, bois-puant, bois-noir, reported to root deeply (Zoológico Virtual de
pois à mabou, paaloe pretae, and raba stokki (Little Barranquilla 2002). Sapo prieto may be found
and others 1974, Zamora 1989). It is a shrub or a growing in secondary and remnant forests. It is not
small tree, which under favorable circumstances a pioneer and will rarely be found in abandoned
may reach 8 m in height and 15 cm in diameter at fields or areas where all vegetation has been
breast height. Sapo prieto usually has a single stem removed. It is moderately intolerant of shade and
unless it has been damaged, but the species is very consequently does not grow under dense forest
limby and may have major branches emerging low canopies. It most frequently grows in an
on the trunk. The stem and branch bark is gray or intermediate crown position in relatively open dry
light brown and smooth with a bitter, yellow or forests or as a codominant in clumps of shrubs and
reddish inner bark. The twigs, petioles, and low trees in disturbed secondary forests.
undersides of leaves are covered with silvery, gray,
or golden scales. The alternate, simple leaves are Reproduction.—Little and others (1974) state that
supported by 6- to 10-mm petioles and have
140
sapo prieto flowers and fruits intermittently. References
Flowers were observed in Costa Rico in May and
September and fruits were present from May to Debrot, A.O., J.Y. Miller, L.D. Miller, and B.T.
September (Zamora 1989). A collection of sapo Leysner. 1999. The butterfly fauna of Curaçao,
prieto fruits from Puerto Rico weighed an average West Indies: 1996 status and long-term species
of 6.948 + 0.773 g/fruit. The air-dry seeds collected turnover. Caribbean Journal of Science 35(3-4):
from these fruits weighed an average of 0.0838 + 184-194.
0.0021 g/seed or 12,000 seeds/kg. Sown on
commercial potting mix without any pre-treatment, Howard R.A. 1988. Flora of the Lesser Antilles,
the seedlings began to emerge in 7 days and Leeward and Windward Islands.
reached a maximum germination of 49 percent in Dicotyledoneae, Part 1. Vol. 4. Arnold
64 days (author’s observation). The seeds are Arboretum, Harvard University, Jamaica Plain,
covered with a thin, pasty, scarlet pulp that is MA. 673 p.
presumably eaten by birds that transport the seeds.
Liogier H.A. 1985. Descriptive flora of Puerto
Growth and Management.—Sapo prieto has been Rico and adjacent islands, Spermatophyta. Vol.
referred to as slow growing (Zoológico Virtual de 1. Editorial de la Universidad de Puerto Rico,
Barranquilla 2002). Weaver (1990) measured one Río Piedras, PR. 352 p.
tree in St. John, U.S. Virgin Islands, as a part of a
larger study and found a 0.4 mm annual diameter Little, E.L., Jr., R.O. Woodbury, and F.H.
increment over a 5-year period. Increases in sapo Wadsworth. 1974. Trees of Puerto Rico and the
prieto density can probably be encouraged by Virgin Islands. Vol. 2. Agriculture Handbook
treatments to dry and moist forests near seed 449. U.S. Department of Agriculture,
sources to eliminate dense shade and create brush Washington, DC. 1,024 p.
patches and small openings. No wildland planting
experience has been reported.
Virgin Islands Wetlands Reserve. 2002. A list of
plants tagged along the trail at the reserve and
Benefits.—Sapo prieto makes a good ornamental
some of their uses. Virgin Islands Wetlands
that is evergreen, with attractive bicolor leaves. It
Reserve, University of the Virgin Islands, St.
has showy flowers and fruits, and a crown that can
Croix, U.S. Virgin Islands.
be shaped by pruning. However, the species is not
http://rps.uvi.edu/VIMAS/plantlist.htm. 5 p.
commonly used because it grows very slowly
(Virgin Islands Wetlands Reserve 2002, Zoológico
Weaver, P.L. 1990. Tree diameter growth rates in
Virtual de Barranquilla 2002). The wood is heavy
Cinnamon Bay Watershed, St. John, U.S. Virgin
and hard and useful for fuel, stakes, and small-
Islands. Caribbean Journal of Science 26(1-2):
diameter fence posts. Sapo prieto contributes to the
1-6.
biodiversity of the forests it inhabits, helps protect
those sites from erosion, and furnishes minor
Zamora, N. 1989. Flora arborescente de Costa
amounts of food and cover for wildlife. The
Rica. Editorial Techologica de Costa Rica, San
flowers are nectar sources for butterflies (Debrot
Jose, Costa Rica. 262 p.
and others 1999).
Zoológico Virtual de Barranquilla. 2002.
Resultados de busca por: Olivo macho.
http://zoobaq.org/formas/consultaveg2.php3?co
mun=Olivo. 1 p.
141
Capraria biflora L. goat-weed
SCROPHULARIACEAE
Synonyms: Capraria lanceolata Vahl
John K. Francis
south as Bolivia, and the Galapagos Islands (Flora

of Texas Consortium 2002, Grisebach 1963,
Pezzatti and others 1998). The species has
naturalized in Ghana, the Cape Verde Islands, and
Mauritius (Burkill 2000).
Ecology.—Goatweed is intolerant of shade. It is

usually open-grown but may be found in stands of
brush and in the understory of open forest. It will
die as the shade becomes dense. In Puerto Rico, it
grows in pastures, occasionally burned grasslands,
dry, early secondary forest, rocky coasts, and sandy
areas behind beaches. It is reported to grow on
beaches, sandy soils, and disturbed areas in Florida
(Long and Lakela 1976). The species grows on
soils derived from both sedimentary and igneous
rocks in a wide range of soil textures and fertility
but requires well-drained conditions. In Puerto
Rico, goatweed occurs in areas that receive from
750 to about 1700 mm of precipitation. It grows at
elevations from near sea level to 1,000 m in
Nicaragua (Stevens and others 2001). Disturbance
is probably required for establishment. Once
established, goatweed competes relatively well
with grass, weeds, and low brush. It is little
browsed by cattle.
General Description.—Goatweed, also known as
wild tea, savadilla, té del país, thé du pays, ditay Reproduction.—Goatweed blooms and fruits all
paye, and balsaminha, is an evergreen (or year (Long and Lakela 1976, Stevens and others
sometimes suffruticose) shrub to 1.5 m in height 2001). A collection of seeds from Puerto Rico
and 6 mm in basal stem diameter. The plant is averaged 42,300 seeds/g. Placed on moist filter
supported by a well-developed taproot with laterals paper, 21 percent germinated between 10 and 35
through its length. The shrub usually has multiple days after sowing. Germination is epigeal. The tiny
stems and many fine branches and twigs. The seeds are apparently dispersed by wind and water,
degree of hairiness is variable. Most plants have or by fortuitously sticking to animals or machinery.
foliage the full length of their stems. The leaves are Natural seedlings are not abundant. Plants readily
subsessile, narrowly lanceolate to obovate, 2 to 12 sprout from the lower stem or root when cut or
cm long, with entire to serrate margin. The white, burned.
five-lobed, campanulate, 5 mm-long flowers are
borne in groups of one to three in the leaf axils. Growth and Management.—Individual goatweed
Ovoid, 5-mm capsules contain many tiny, yellow stems live 2 or 3 years or more and are replaced by
seeds about 0.4 mm long (Howard 1989, Liogier other sprouts. The plant may live for many years.
1995, Stevens and others 2001). Sprouts grow about 0.5 m in their first year, much
less in succeeding years. Large numbers of seeds
Range.—Goatweed is native to Florida and Texas, can be collected by picking unopened capsules,
the Bahamas, the West Indies, Trinidad, Mexico drying them in a paper bag, and separating with a
through Central America, South America as far fine screen. Nursery procedures have not been
142
published. Goatweed is considered weedy in 789 p.
pastures. In the absence of tested procedures for its
control, standard techniques of grubbing or spot Guzmán, D.J. 1975. Especies útiles de la flora
spraying with broadleaf herbicides is Salvadoreña. Ministerio de Educación,
recommended. Mowing has little lasting effect on Dirección de Publicaciones, San Salvador, El
stands. Salvador. 703 p.
Benefits.—Goatweed helps revegetate disturbed Howard, R. A. 1989. Flora of the Lesser Antilles,
areas, protects the soil, and furnishes cover for Leeward and Windward Islands. Vol. 6. Arnold
wildlife. It is a nectar source for butterflies and is Arboretum, Harvard University, Jamaica Plain,
recommended in natural landscaping (University of MA. 658 p.
Florida Extension 2002). Goatweed has many uses
in herbal medicine. A tea prepared from the leaves Liogier, H.A. 1995. Descriptive flora of Puerto
is used as an eyewash (Burkill 2000), to soothe Rico and adjacent islands. Vol. 4. Editorial de la
skin itch (Secritaría de Medio Ambiente y Universidad de Puerto Rico, San Juan, PR.
Recursos Naturales 2002), and as a general tonic. 617 p.
However, overindulgence can result in stupor,
disorientation, and paralysis (Guzmán 1975). The Long, R.W. and O. Lakela. 1976. A flora of
infusion is also used to treat fever, flu, vomiting, Tropical Florida. Banyon Books, Miami, FL.
childbirth recovery, diarrhea, hemorrhoids, 962 p.
rheumatism, and swelling (Scofield 2002). The
alkaloid, biflorine, present in the leaves has Pezzatti, B., T. Irzan, and D. Cherix. 1998. Ants
antibiotic properties (Burkill 2000), and (Hymenoptera, Formicide): Lost paradise.
sesquiterpenoids, caprariolides A and B from aerial Darwin Foundation, Galapagos, Equador.
parts, have demonstrated strong insecticidal http://www.darwinfoundation.org/articles/n5900
activity against adult sweet potato weevils, Cylas 049802.html. 10 p.
formicarius elegantulus (Collins and others 2000).
Scofield, D. 2002. Medicinal usage: Capraria
References biflora L. (Scrophulariaceae). http://www.
cassiakeyensis.com/sofl_plants/med_caprariabifl
Burkill, H.M. 2000. The useful plants of West ora.html. 1 p.
Tropical Africa. Vol. 5. Royal Botanical Garden,
Kew, UK. 686 p. Secritaría de Medio Ambiente y Recursos
Naturales. 2002. Especies con usos do
Collins, D.O., W.A. Gillimore, W.R. Reynolds, maderables en bosques de encino, pino, y pino-
L.A.D. Williams, and P.B. Reese. 2000. New encino. http://www/semarnat.gob.mx/pfnm/
skeletal sesquiterpenoids, caprariolides A-D, CaprariaBiflora.html. 2 p.
from Capraria biflora and their insecticidal
activity. Journal of Natural Products 63: 1,515- Stevens, W.D., C. Ulloa-U., A. Pool, O.M.
1,518. Montiel, eds. 2001. Flora de Nicaragua.
Monographs in Systematic Botany Vol. 85, No.
Flora of Texas Consortium. 2002. Texas vascular 3. Missouri Botanical Garden, St. Louis, MO. p.
plant checklist: Scrophulariaceae. 1,911-2,666.
http://csdl/tamu.edu/FLORA/ftc/dft/ftc_scr.htm.
5 p. University of Florida Extension. 2002. Ask a
master gardener: List of native Florida
Grisebach, A.H.R. 1963. Flora of the British West specimens for butterfly gardening. http://
Indian Islands. J. Cramer-Weinheim. New York. volusia.org/extension_service/butterfly.htm. 2 p.
143
Capsicum annuum L. bird pepper
SOLANACEAE
Synonyms: Capsicum indicum microcarpum var. aviculare Dierb.

Capsicum bacatum of authors, not of L.
John K. Francis
1993), and Arizona (Tewksbury and other 1999). It

is felt that the species was introduced into Puerto
Rico by the aboriginal inhabitants (Barrett 1925).
Bird pepper is also widely cultivated and escaped
in the tropical and subtropical Eastern Hemisphere
(Bailey 1941). It is the wild ancestor of the
pimento, the bell pepper, and some of the hot
peppers (Bailey 1941). The domestication of these
pepper varieties from bird pepper is reported to
have taken place in Mexico or Central America
(Hawkens 1991).
Ecology.—In Puerto Rico, bird pepper inhabits the

subtropical moist forest and the wetter portion of
the subtropical dry forest (about 800 to 2000 mm
of annual precipitation). It is cold sensitive, and
hot, dry weather is desirable for fruit ripening
General Description.—Bird pepper is also known
(California Antilles Trading Consortium 2001). In
as red pepper, wild pepper, wild chili, pimiento,
favorable habitat, bird pepper can grow under a
and ají de gallina (Liogier 1995). The current
broken canopy or in disturbed areas without tree
growth is herbaceous, but it later hardens and
cover. However, in Arizona, the plants were found
becomes woody. The wood is brittle. Older plants
exclusively under partially shading (nurse) plants
are multistemmed and very branchy, with a thick,
(Tewksbury and other 1999). Under conditions of
leafy crown. The leaves are dark green, ovate to
low fertility, moisture, and light, bird pepper plants
lanciolate, 4 to 15 cm long, with petioles 0.5 to 3
do not become large and probably do not live more
cm long. The twigs are stiff and straight between
than 1 year. The species favors a near-neutral soil
nodes. The flowers are mostly solitary or in pairs.
reaction and a generous supply of bases. Well-
The corolla is greenish-white or yellowish-white, 1
drained soil with a sandy loam or silt loam texture
to 2 cm broad, with blue, violate, or yellow
is best (California Antilles Trading Consortium
anthers. The fruits, which are ellipsoidal berries
2001). After it is well established, bird pepper can
with thin flesh, are 0.4 to 3 cm long (usually about
survive dry seasons of 2 to 4 months. During this
0.8 cm in wild plants) and red or orange-red. The
period, it will partially defoliate and take on a
seeds are cream to yellow in color (Howard 1989,
wilted appearance, but will refoliate and grow
Liogier 1995). The variety growing as a shrub in
vigorously after the rains return.
wildland vegetation in Puerto Rico, Florida, and
elsewhere is C. annuum var. glabriusculum
Reproduction.—When conditions are favorable,
(Dunal) H. Pickersgill (Liogier 1995).
bird pepper begins flowering at about 3 months of
age (California Antilles Trading Consortium 2001)
Range.—Bird pepper ranges today from the
and flowers and fruits throughout the year. A
Southern United States to Argentina and
sample of fruits collected in Puerto Rico weighed
throughout the West Indies (Liogier 1995). The
an average of 0.0926 + 0.0057 g/fruit. Seeds from
species has been widely spread and cultivated by
the sample weighed an average of 0.0039 + 0.0001
Native Americans and European settlers. Escapes
g/seed or 260,000 seeds/kg. Bird pepper plants are
from cultivation have masked the original range.
prolific seed producers. Large plants may produce
Bird pepper may be found competing with native
hundreds of fruits per year. Fruits in the above
vegetation in Puerto Rico (author’s observations),
sample contained an average of 8.2 seeds/fruit.
Florida (Nelson 1996), Texas (Everitt and Drawe
144
Twenty-eight percent of the seeds in this collection References
germinated between 17 and 118 days after being
sown in potting mix. Seedlings may be established Bailey, L.H. 1941. The standard cyclopedia of
by sowing in prepared seed spots in the field or horticulture. Vol. 1. The MacMillan Co. New
grown in the nursery and transplanted. The seeds York, NY. 1,200 p.
are dispersed by birds that are immune to the
capsaicin the fruits contain. Rodents that would
Barrett, O.W. 1925. The food plants of Puerto
otherwise chew up and destroy the seeds will not
Rico. The Journal of the Department of
eat the fruits (UniSci 2001). Seedlings and plants
Agriculture of Puerto Rico 9(2): 168.
are common but scattered in early secondary forest
in Puerto Rico. However, bird pepper may
occasionally form small thickets.
California Antilles Trading Consortium. 2001.
Growth and Management.—Because the species Capsicum peppers. www.calantilles.com/
name “annuum” means annual and because capsicum_peppers.htm. 9 p.
cultivated forms are, or are treated as, annuals, bird
pepper has been assumed to be an annual (see
Liogier 1995). In its wild state, it is, in fact, a Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs
short-lived perennial (Floridata 2001), living 3 or 4 and cacti of South Texas. Texas Tech University
years, if conditions are favorable. Plants may Press. 213 p.
occasionally reach 5 m in height (Howard 1989).
Heights in Puerto Rico are commonly 2 to 3 m. Floridata. 2001. Capsicum spp.
The stems are slender, about 1 cm in diameter, in www.floridata.com/ref/c/caps_spp.cfm. 6 p.
larger plants (author’s observation).
Hawkens, J.G. 1991. The centers of plant diversity
Benefits.—Bird pepper has been cultivated for in Latin America. Diversity 7(1/2): 7-9
thousands of years and is the wild ancestor for
hundreds of named varieties of cultivated peppers. Hongo, H., and A.K. Karel. 1986. Effects of plant
However, the wild fruits are still harvested and extracts on insect pests of common beans.
used today. Bird pepper fruits are “hot” to very Journal of Applied Entomology 102(2): 164-
“hot” with a slight musky flavor. They are used to 169.
flavor food and make sauces. Capsicum
derivatives are used in a wide variety of medicinal Howard, R. A. 1989. Flora of the Lesser Antilles,
applications, mostly related to pain relief (eg: sore Leeward and Windward Islands. Vol. 6. Arnold
muscles, toothache, phantom limb pain). Arboretum, Harvard University, Jamaica Plain,
Capsaicin causes the brain to release endorphins MA. 658 p.
that promote a sense of well-being and deadens
pain receptors (Floridata 2001). Concentrated Krenzelok, E.P., and F.J. Provost. 1995. The ten
pepper sprays are used for riot control and most common plant exposures reported to
personal defense. Gardeners also use pepper poison information centers in the United States.
extracts to protect crops from insects and animals. Journal of Natural Toxins 4(2): 195-202.
Extracts of leaves and fruits of bird pepper have
shown insecticidal and fungicidal properties in Liogier, H.A. 1995. Descriptive flora of Puerto
laboratory tests (Hongo and Karel 1986, Patil and Rico and adjacent islands. Vol. 4. Editorial de la
Joi 1992, Williams and Mansingh 1993). Universidad de Puerto Rico, San Juan, PR.
However, to sensitive individuals, exposure to C. 617 p.
annuum fruits can be dangerous. This species
ranked fourth in plant poisoning incidents reported Nelson, Gil. 1996. The shrubs and woody vines of
to American poison information centers Florida. Pineapple Press, Inc. Sarasota, FL.
(Krenzelok and Provost 1995). In Texas and 391 p.
Arizona, several species of birds including the Rio
Grande turkey and the curve-billed thrasher eat the Patil, B.H. and M.B. Joi. 1992. Inhibition of
fruits (Everitt and Drawe 1993, UniSci 2001). tomato spotted wilt virus with leaf extracts of
some plants. Journal of Maharashtra
Agricultural University 17(2): 340-341.
145
Tewksbury, J.J., G.P. Nabhan, D. Norman, H. http://unisci.com/stories/20013/0726015.htm.
Suzan, J. Tuxill, and J. Donovan. 1999. In situ 3 p.
conservation of wild chilis and their biotic
associates. Conservation Biology 13(1): 98-107. Williams, L.A.D. and A. Mansingh. 1993.
Pesticidal potential of tropical plants—I.
UniSci 2001. Zing in the flavor helps chili pepper Insecticidal activity of leaf extracts of sixty
plants survive. Daily University Science News. plants. Insect Science and its Application 14(5):
697-700.
146
Capsicum frutescens L. red pepper
SOLANACEAE
Synonyms: Capsicum fastigiatum Blume

Capsicum annuum L. var. frutescens (L.) Kuntze
John K. Francis
rounded tip, 1.5 to 3.5 cm long and 0.5 to 1.2 cm

thick. The fruits are somewhat dry and contain few
to many (depending on fruit size) cream to yellow
lenticular seeds about 3 mm in diameter. The fruits,
especially the seeds and placenta, have a biting,
pungent taste. The species has 2n = 24
chromosomes (Bailey 1941, Bentley and Trimer
1880, Bosland and Votava 2000, Liogier 1995).
Range.—The original range of red pepper is

unknown, but it is believed to have been
domesticated in Central America, possibly Panama,
thousands of years ago. It spread throughout the
Neotropics before Columbus and has since become
almost pan-tropically cultivated and naturalized
(Bosland and Votava 2000). Unlike C. annuum L.,
which has many widely varying varieties,
domesticated red pepper has relatively few
varieties with a minimum of variation (Bentley and
Trimer 1880) and is not greatly different from the
wild type.
Ecology.—Red pepper grows on soils of all

textures in a wide range of fertilities. Moist, well-
drained conditions and loose structure is best for
rapid growth. Soil pH’s of 4.3 to 9.7 are tolerated
(Center for New Crops and Plant Products 2002).
The species is intolerant of shade: it will grow with
General Description.—Red pepper, also known as broken overhead shade and moderate competition
bird pepper, chili pepper, Cayenne pepper, Guinea from grass and forbs, but fruits best in full sun. Red
pepper, ají, and ají picante, is a short-lived peppers can be cultivated in areas that receive from
evergreen shrub usually 1 to 1.5 m in height and 1 30 to 430 cm of annual precipitation at elevations
to 3 cm in basal stem diameter. The shrub is from near sea level to more than 2,000 m. The
supported by a short to long taproot (depending on species is not frost tolerant and does not grow in
soil conditions), many spreading lateral roots, and temperatures below 7 °C (Center for New Crops
moderately abundant fibrous roots. The stem and and Plant Products 2002). Wild red pepper grows
larger branches of mature plants are woody but as individuals or groups in gardens, fields, vacant
moderately soft and weak. Bark of stems and older lots, river flood plains, abandoned fields, roadsides,
branches is light gray. The form is upright, the and early secondary forest. Although damping-off
abundant branching is often dichotomous, and the fungi, various species of insects, and nematodes
branches and twigs are slender. The ovate to ovate- sometimes damage or kill individual plants, serious
lanceolate leaves vary in size. The larger of them effects are rarely widespread.
are 4 to 12 cm long and 1 to 4.5 cm broad.
Greenish-white to yellowish-white flowers with Reproduction.—After about 3 months of growth,
blue, violet, or yellow anthers occur in groups of red pepper plants flower (Center for New Crops
two or more at the nodes. The berries are red or and Plant Products 2002) and fruit continuously as
red-orange at maturity, elongated with a pointed or
147
long as they live. The flowers are insect pollinated protection, law enforcement, and defense. Acetone
(Bosland and Votava 2000). Production of fruits and petroleum ether extracts caused complete
and seeds can be abundant. Fresh fruits from mortality of rice weevil (Sitophillus oryzae) in 15
several plants in a stand in Puerto Rico averaged days. Fruit powder was much less effective (El-
0.3668 + 0.0242 g/fruit. The variability was high Lakwah and others 1997). Red pepper has many
(CV = 39.5 percent). They ranged from 2 to 35 medicinal applications. Some of the most widely
seeds/fruit and averaged 16.6 + 2.9 seeds/fruit. Air- used and reliable are as a salve to relieve muscle,
dried seeds averaged 0.0041 + 0.0001 g/seed or joint, and toothache pain, to treat cough, asthma,
244,000 seeds/kg. Placed in moist potting mix, 72 and sore throat, as a stimulant, and to treat stomach
percent of the seeds germinated between 13 and 34 ache, seasickness, and flatulence. Anciently, it was
days after sowing. Germination is epigeous. The even used as an instrument of torture (Bentley and
seeds can be safely stored under refrigeration after Trimer 1880, Bosland and Votava 2000, Center for
air-drying and need no treatment before planting. New Crops and Plant Products 2002,
The seeds are dispersed by birds that eat the fruits Gardenguides.com 2002).
and are immune to the pungent chemicals they
contain. Young plants coppice when cut or broken. References
Rooting of cuttings and tissues has been
demonstrated for Capsicum (Bosland and Votava Bailey, L.H. 1941. The standard cyclopedia of
2000). horticulture. Vol. 1. The MacMillan Co. New
York, NY. 1,200 p.
Growth and Management.—Under continually
favorable conditions, red peppers live about 2 Bentley, R. and H. Trimer. 1880. Medicinal plants.
years. They grow rapidly during the first year, then Vol. 3. J.&A. Churchill, London. p. 147-227.
much more slowly, and finally dwindle and die.
Red peppers grown commercially are managed as Bosland, P.W. and E.J. Votava. 2000. Peppers:
annuals, much as other Capsicum peppers but for a Vegetable and spice capsicums. CABI
longer season. Plants that arise spontaneously in Publishing, Oxon, UK and New York. 204 p.
gardens and vacant land are usually allowed to
grow for occasional future harvest. Center for New Crops and Plant Products. 2002.
Capsicum pepper. Purdue University. http://
Benefits.—Red pepper is used and loved the world hort.purdue.edu/newcrop/med-aro/factsheets/
over as a condiment, added to food fresh, dried, CAPSICUM_PEPPER.html. 2 p.
refined, and ground (for Cayenne pepper and
curry), and as the principal or incidental ingredient
El-Lakwah, F.A., O.M. Khaled, M.M. Khattab,
in sauces. The source of the popular biting
and T.A. Abdel-Rahman. 1997. Toxic effects of
sensation are the capsaicinoids, principally
extracts and powders of certain plants against
capsaicin, which ranges from 600 to 13,000 ppm in
the rice weevil (Sitophilus oryzae). Annals of
the fruits (Center for New Crops and Plant
Agricultural Science (Moshtohor) 35(1): 553-
Products. 2002). The fruits are an excellent source
566.
of vitamins A and C (Bosland and Botava 2000). A
few thousand hectares are grown commercially,
Gardenguides.com. 2002. Cayenne. http://www.
and probably even more are grown in gardens or
gardenguides.com/herbs/cayenne.htm. 4 p.
harvested from the wild in rural areas of the
tropics. Coatings and powders are used with
varying degrees of success to deter browsing
animals and insects, and break children of thumb
Universidad de Puerto Rico, San Juan, PR.
sucking and nail biting. Pepper spray, whose active
617 p.
ingredient is capsaicin, is used widely for personal
148
Casearia decandra Jacq. tostado
FLACOURTIACEAE
Synonyms: Casearia parvifolia Jacq.

Casearia parvifolia Will.
Casearia albicaulis Risby
Samyda lancifolia Sessé & Moç.
John K. Francis
toothed margin. The small white, cream, or

greenish-white flowers are born in nearly sessile
clusters at the defoliate leaf nodes. The globose
fruits are capsules about 8 to 10 mm in diameter,
opening on three valves. The fruits are ripe when
they turn from greenish-white to cream with a
salmon-colored blush or light brown. The fleshy
part of the fruit is orange to red. There are one to
four, 4- to 5-mm seeds per fruit, depending on fruit
size (Howard 1989, Liogier 1994, Little and
Wadsworth 1964).
Range.—The native range of tostado covers the

islands of the West Indies from Hispaniola south
and extends from Honduras through Panama and
into South America as far south as Paraguay,
Bolivia, and Northern Argentina (Howard 1989,
Liogier 1994, Little and Wadsworth 1964).
Ecology.—Tostado grows on well-drained soils

ranging from coastal sands to upland clays, and on
soils derived from sedimentary (including
limestone), igneous, and metamorphic rocks.
Annual precipitation in habitat in Puerto Rico
ranges from about 900 mm to about 2200 mm. The
species grows to altitudes of at least 1,120 m in
General Description.—Tostado (a name used in Brazil (Silva and others 2002). In Bolivia it occurs
Puerto Rico) is also known by wild honey-tree, from 230 to 650 m in elevation (Killeen and others
wild-cherry, pipewood, biscuitwood, jumbie-apple, 1993), and in Puerto Rico, it grows from near sea
palo blanco, caracolillo, cerezo, corcho blanco, level to over 600 m. Tostado is moderately
cotorrerillo, duro-duro, guía mansa, machacomo, intolerant of shade. It grows in disturbed areas,
tapaculo, limoncaspi, bois jaune, coco-ravet, jaune forest openings, very rocky sites, and the
d’oeuf, and fortuga caspi. It is a shrub or small tree understories of lower density forests. In southern
usually 2 to 6 m in height and 2 to 8 cm in diameter Brazil, it occurs in the third canopy layer of
at breast height (d.b.h.). Tostado is supported by a Araucaria forests (Silva and others 2002), and in
strong taproot, somewhat finer lateral roots, and Bolivia in Amazon forests and savanna woodlands
abundant near-surface fine roots. The roots are stiff (Killeen and others 1993). In Venezuela, the
and ivory-colored. The species usually has a single species functions as a gap-filling species in bush
stem, unless disturbed, and smooth gray bark. The island savannas (San José and others 1991).
stemwood is moderately hard and moderately
heavy. The branches and twigs are numerous, Reproduction.—Tostado flowers and fruits
slender, and form a relatively dense crown. The irregularly throughout the year in Puerto Rico
alternate leaves have petioles 2 to 5 mm long, (Little and Wadsworth 1964), only a minority of
elliptic blades 3 to 9 cm long, with a finely saw- plants being in flower or fruit at any time. It is
149
reported to flower from August to October in References
Bolivia (Killeen and others 1993). Plants 1 m or
more in height in partial or full sunlight bear fruits. Francis, J.K. 2000. Estimating biomass and carbon
Shrubs and small trees bear from hundreds to content of saplings in Puerto Rican secondary
thousands of fruits each year. A collection of fruits forests. Caribbean Journal of Science 36(3-4):
from Puerto Rico weighed an average of 0.340 + 346-350.
0.031 g/fruit. They are extremely variable (CV =
69.9 percent), even within the same plant. Seeds Howard, R.A. 1989. Flora of the Lesser Antilles,
separated from the above collection weighed (air- Leeward and Windward Islands. Vol. 5. Arnold
dried) an average of 0.0406 + 0.0005 g/seed or Arboretum, Harvard University, Jamaica Plain,
24,600 seeds/kg. Without pre-treatment, these MA. 604 p.
seeds were sown on commercial potting mix and
91 percent germinated 7 days later (author’s Killeen, T., E. García E., and S.G. Beck. 1993.
observation). Tostado sprouts when cut or burned. Guía de árboles de Bolivia. Herbario Nacional
de Bolivia and the Missouri Botanical Garden.
Growth and Management.—Although tostado is St. Louis, MO. 958 p.
usually a shrub, if it survives several decades in
fertile sites with sufficient sunlight, it may Liogier, H.A. 1994. Descriptive flora of Puerto
occasionally become a tree 12 to 18 m in height Rico and adjacent Islands. Vol. 3. Editorial de la
(Howard 1989, Liogier 1994). Weaver (1990) Universidad de Puerto Rico. Río Piedras, PR.
reported an average 5-year annual diameter 461 p.
increment of only 0.05 cm for plants 4 to 9 cm
d.b.h. in St. John, U.S. Virgin Islands. The air- Little, E.L., Jr. and F.L. Wadsworth. 1964.
dried specific gravity of stemwood was measured Common trees of Puerto Rico and the Virgin
at 0.630 + 0.031. The above-ground average Islands. Agriculture Handbook 249. U.S.
carbon content was measured at 0.508. Total Department of Agriculture, Washington, DC.
above-ground dry weight in plants up to 5 cm 548 p.
d.b.h. can be estimated by the model: Wt =
34.356(D2S), where Wt is weight in grams, D is San José, J.J., M.R. Farinas, and J. Rosales. 1991.
diameter in cm at 30 cm above the ground-line, S is Spatial patterns of trees and structuring factors
total stem length in meters, and adjusted r-squared in a Trachypogon savanna of the Orinoco
equals 0.977 (Francis 2000). Llanos. Biotropica 23(2): 114-123.
Benefits.—The stems of tostado are sometimes Silva, D.W., E. Seitz, M. Burgo, S.M. Silva, and
used as fuel and fenceposts. This is an important J.J. Soares. 2002. Levantamento de espécies
honey plant. The small fruits are edible, but almost arbóreas no parque municipal das araucarias,
tasteless according to Little and Wadsworth (1964). Guapapuava, PR. Sociedad Botanica de São
However, the author found them sweet and Paulo, Brazil. http://www.ib.usp.br/sbsp/
pleasantly flavored. The species furnishes food and congresso/ec.htm. 1 p.
cover for wildlife, contributes to biodiversity, and
helps stabilize the soil. Weaver, P.L. 1990. Tree diameter growth rates in
Cinnamon Bay Watershed, St. John, U.S. Virgin
Islands. Caribbean Journal of Science 26(1-2):
1-6.
150
Casearia sylvestris Sw. wild-coffee
FLACOURTIACEAE
Synonyms: Samyda parviflora L.

Casearia parviflora L.
Anavinga samyda Gaertn. f.
John K. Francis
three valves to reveal usually three brown seeds

covered with a red or red-orange aril (author’s
observations, Croat 1978, Howard 1989, Liogier
1994, Little and Wadsworth 1964).
Range.—Wild-coffee is native to Cuba, Jamaica,

Lesser Antilles, Trinidad and Tobago, Mexico
through Central America, and South America as far
south as northern Argentina, Uruguay, and Bolivia
(Howard 1989, Instituto de Botánica Darwinion
2002, Killeen and others 1993, Little and
Wadsworth 1964).
Ecology.—Wild-coffee grows on soils of all

textures derived from nearly all parent materials.
However, it does not grow on swampy ground and
usually not on excessively drained sites. In Puerto
Rico, wild-coffee grows in areas receiving from
about 900 to over 3000 mm of annual precipitation
(author’s observation). However, in southern
Brazil, wild-coffee grows even in annually flooded
areas called várzeas (Universidad do Extremo Sul
General Description.—Wild-coffee, also known
Catarinense 1999). In Nicaragua, the species grows
as crack-open, cafeíllo, café silvestre, sarnilla,
from near sea level to 1,300 m in elevation
corta-lengua, guayabillo, guassatonga, and
(Stevens and others 2001). Wild-coffee
papelite, is a shrub or small tree usually 2 or 3 m in
demonstrates an intermediate tolerance to shade,
height, but occasionally reaching 10 m or more. On
being able to survive and grow slowly under a
clay soils, the plant relies for absorption and
forest canopy that is not too dense but requiring
support on extensive lateral roots that are white,
partial sunlight to reproduce. Plants growing in
moderately stiff, and have a corky bark. The plant
pine plantations and natural secondary forests in
may have a single or multiple stems with nearly
Puerto Rico held similar concentrations of N, P,
smooth, thin gray bark and a moderately branchy
and ash in their leaves, but K levels were nearly
habit. The sapwood is light brown and the
twice as high in natural forest (Lugo 1992). Wild-
heartwood is dark-brown, fine-textured, hard,
coffee is common in brushy pastures, early
heavy, and strong. The twigs are long and slender,
secondary forests, roadsides, fence rows, and
often horizontal or drooping. The previous year’s
gallery forests of stream bottoms (author’s
growth often has tufts in the leaf axils of what were
observation, Little and Wadsworth 1964). It is one
the fruit stalks. The alternate leaves have petioles 2
of the principal arborescent species of the Llanos
to 8 mm long and ovate to lanciolate blades 4 to 13
(plains) of the Orinoco area of Venezuela (van Os
cm long by 2 to 5 cm broad with a wavy edge and a
2000).
long-pointed tip. Tiny white, cream, or greenish
flowers are crowded (20 to 50) on short stalks in
Reproduction.—Under favorable conditions, wild-
the leaf axils. The flowers smell like a mixture of
coffee is able to flower and fruit at a young age,
honey and urine. The fruits are globose or slightly
about the second year as a sprout or at about 1 m in
flattened, 3 to 4 mm in diameter, splitting open on
151
height. Although individual plants do not flower 1990, Carvalho, and others 1999, Itokawa and
continuously, members of the population may be others 1990, Silva and others 1988). A commercial
seen flowering throughout the year (Little and perfume and a weight-loss product contain extracts
Wadsworth 1964, Stevens and others 2001). In of wild-coffee (Fontovit 2002, Multinível do Brasil
Panama, the species flowers principally in August S.A. 2002).
to December with sporadic flowering during the
dry season. The fruits mature in about 1 month References
(Croat 1978). A collection of fresh fruits (seed +
aril) from Puerto Rico weighed an average of Basile, A.C., J.A.A. Sertie, S. Panizza, T.T.
0.0286 + 0.0021 g/fruit. They vary considerably in Oshiro, and C.A. Azzolini. 1990.
size (coefficient of variation = 60.5). Air-dry seeds Pharmacological assey of Casearia sylvestris. I.
from the same collection averaged 0.0011 + 0.0001 Preventive anti-ulsur activity and toxicity of the
g/seed or 909,000 seeds/kg. Sown on peat, these leaf crude extract. Journal of
seeds germinated at 31 percent between 20 and 103 Ethnopharmacology 30(2): 185-197.
days after sowing (author’s observation). In
southern Brazil, 84,000 seeds/kg were reported and Carvalho, J.C.T., V.V. Vignoli, G.H.G. de Souza,
a low rate of germination (Universidad do Extremo K. Ujikawa, J.J. Neto, and G.H.B. de Souza.
Sul Catarinense 1999). Germination is epigeal. Just 1999. Antimicrobial activity of essential oils
one or two fruits per inflorescence are ripe at any from plants used in Brazilian popular medicine.
time. The only method of seed collection known to Acta Horticulturae 501: 77-81.
the author is to hand-pick the tiny ripe fruits
(capsules split exposing the red aril) and wet sieve Croat, T.B. 1978. Flora of Barro Colorado Island.
to separate the seeds. Birds disburse the seeds. Stanford University Press, Stanford, CA. 943 p.
Natural seedlings are rare to common. Plants past
the seedling stage have a high survival rate. Fontovit. 2002. Porangaba Fontovit.
Damaged plants coppice readily. http://fontovit.com.br/produtos/porangaba.htm.
1 p.
Growth and Management.—Wild-coffee has a
moderate growth rate and lives for at least 20 Howard, R.A. 1989. Flora of the Lesser Antilles,
years. Artificial reproduction is usually by seeds, Leeward and Windward Islands. Vol. 5. Arnold
but cuttings can also be rooted. Recommended Arboretum, Harvard University, Jamaica Plain,
spacing for plantings in Brazil is 4 by 4 m MA. 604 p.
(Universidad do Extremo Sul Catarinense 1999).
Because it becomes established as part of the early Instituto de Botánica Darwinion. 2002. Catálogo
secondary forest or in disturbed openings of more de las plantas vasculares de la Argentina:
advanced forest, management to promote Flacourtiaceae. http://www.darwin.edu.ar/
regeneration of the species might include the use of Catalogo/Flacourtiaceae.pdf. 3 p.
clearcuts or group selection in closed forest and
protection of open land from fires. Itokawa, H., N. Totsuka, H. Morita, K. Takeya, Y.
Iitaka, E.P. Schenkel, and M. Motidome. 1990.
Benefits.—The wood of wild-coffee is used for or New antitumor principles, casearines A-F, for
is suitable for fuel, fence posts, stakes, small poles, Casearia sylvestris Sw. (Flacourtiaceae).
rustic carpentry, and tool handles (Little and Chemical and Pharmaceutical Bulletin 38(12):
Wadsworth 1964). The species is recommended for 3,384-3,388.
urban planting as a food source for birds (Rio
Grande Energia 2002). It is a honey plant (Little Killeen, T.J., E. García E., and S.G. Beck. 1993.
and Wadsworth 1964). Leaves of wild-coffee Guía de árboles de Bolivia. Herberio Nacional
contains 2.5 percent essential oils with a pleasant de Bolivia and Missouri Botanical Garden, St.
aroma. They also contain caprionic acid, saponins, Louis, MO. 958 p.
alkaloids, flavonoids, and a host of other
chemicals. Extracts (particularly of the leaves) are Liogier, H.A. 1994. Descriptive flora of Puerto
used in traditional herbal medicine to treat a large Rico and adjacent Islands. Vol. 3. Editorial de la
number of different ailments (Universidad do Universidad de Puerto Rico. Río Piedras, PR.
Extremo Sul Catarinense 1999). Laboratory 461 p.
experiments have confirmed antiseptic, antitumor,
antiulser, and abortive activity (Basile and others
152
Little, E.L., Jr. and F.L. Wadsworth. 1964. rs.com.br/manual_poda/esp_recomendadas.asp.
Common trees of Puerto Rico and the Virgin 5 p.
Islands. Agriculture Handbook 249. U.S.
Department of Agriculture, Washington, DC. Silva, F.A., A.L.M. Baisch, B. Oliveira, A.M.
548 p. Battastini, F. Torres, G. Rocoski, E.S. Silva,
M.F. Alam, J.C.G. Apolinario, and A.J. Lapa.
Lugo, A.E. 1992. Comparison of tropical tree 1988. Preliminary pharmacological studies on
plantations with secondary forests of similar extracts from Casearia sylvestris Swartz. Acta
age. Ecological Monographs 62: 1-41. Amazonica Suplemento 18(1-2): 219-229.
Multinível do Brasil S.A. 2002. Uma nova geração Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
de colônias. http://www.multiniveldobrasil. Montiel, eds. 2001. Flora of Nicaragua.
com.br/bio_collection.html. 2 p. Monographs in Systematic Botany Vol. 85, No.
2. Missouri Botanical Garden Press. St. Louis,
van Os, M. 2000. The Llanos de Orinoco. MO. p. 945-1,910.
Wageningen Agricultural University,
Wageningen, Netherlands. http://www.wau.nl/ Universidad do Extremo Sul Catarinense. 1999.
rpv/ond_proj/dirjongm/Llanos/teksten/objectives Jornada Catarinense de plantas medicinais:
/chapter03.html. 8 p. Guaçatonga. http://www.unesc.rct-sc.br/
plantas_medicinais/guaca.htm. 4 p.
Río Grande Energia. 2002. Manual de arborização
e poda: espécies recomendadas. http://www.rge-
153
Cassine xylocarpa Vent. marble tree
CELASTRACEAE
Synonyms: Elaeodendron xylocarpum (Vent.) DC.

Freziera dioica Macfad.
Elaeodendron dioicum (Macfad.) Griseb.
Elaeodendron attenuatum A. Rich.
Elaeodendron xylocarpum (Vent.) DC.
Cassine rotundata (DC.) Kuntze
John K. Francis
They are entire or with a few teeth, elliptic to
obovate, 2.5 to 12.5 cm long, with a rounded to
spine-tipped apex. The inflorescences are tiny-
flowered, branched axillary cymes. The fruit is a
yellow (at maturity), rounded drupe, 1.5 to 3 cm
long with a thin, hard-fleshy exocarp and hard,
thick-walled stone with one to three cells and
elliptic seeds (Howard 1989, Liogier 1994, Little
and others 1974).
Range.—The range of marble tree consists of the

Bahamas, the Greater and Lesser Antilles, Mexico,
Belize, Costa Rica, Nicaragua, Panama, and
Venezuela (Howard 1989, Instituto Nacional de
Biodiversidad. 2000, Liogier 1994, Missouri
Botanical Garden 2002a, 2002b).
Ecology.—Marble tree is moderately intolerant of

shade. It can persist and grow slowly under low
forest but needs to be open-grown or at least have
broken sunlight to flower and fruit. The species is
drought-tolerant and usually grows on excessively
well-drained situations—beaches, bluffs, rocky
General Description.—Marble tree, also known as headlands, and rocky ridges. Marble tree often
spoon tree, poison tree, wild nutmeg, coscorrón, grows out of cracks in rocks. It is most often seen
aceituno, guayarote, mate prieto, pinipinche de in areas of limestone and ultramafic (serpentine)
sabana, bois-tan, bord-de-mer, and grosse-peau, is rocks. The species is reported to be a nickel
an evergreen shrub or small tree commonly 2 to 4 accumulator (Medina and others 1994). Colonized
m in height and 5 to 8 cm in diameter at breast areas in Puerto Rico range from 750 to about 1800
height. The plant is supported by an extensive mm of mean annual rainfall and at elevations from
shallow and deep lateral root system with sinkers near sea level to 300 m or more. The species may
and sometimes a discernable taproot. The roots are be found in remnant and late secondary forests.
orange and stiff. Marble tree often has several
stems emanating from the root crown or low on the Reproduction.—Marble tree flowers and fruits
trunk. The crowns are fairly dense (especially intermittently throughout the year (Little and
when open-grown), branchy, and often contorted. others 1974). A collection of air-dried seeds in
The bark is gray, smooth to finely fissured, with Puerto Rico averaged 3.19 + 0.076 g/seed or 313
inner bark that is purplish or pinkish. The twigs are seeds/kg. Sown in commercial potting mix, they
slender and green, becoming gray with age. Leaves began germinating in 26 days and completed 82
are variable, whitish-green to yellow-green, mostly percent germination. The species can also be
opposite, but sometimes alternate, thick and stiff. propagated by asexual means. Fifty-six and 21
154
percent, respectively, of air layers and misted MA. 604 p.
cuttings treated with IBA rooted. The newly
formed roots are fragile and easily broken off Instituto Nacional de Biodiversidad. 2000. Leaps
during transplanting (author’s observation). The and bounds. The Cutting Edge 7(2): 1-2,
seeds are probably dispersed by fruit bats. http://inbio.ac.cr/papers/manual_plantas?apr00le
Seedlings and saplings are relatively common in a.html. 2 p.
and around Puerto Rican stands of the species.
Liogier, H.A. 1990. Plantas medicinales de Puerto
Growth and Management.—Marble tree grows Rico y del Caribe. Iberoamericana de Edicones,
slowly. A 1-m tall sapling with a basal diameter of Inc. San Juan, PR. 566 p.
1.5 cm growing in the understory of low basal area
moist forest had 37 annual rings (author’s Liogier, H.A. 1994. Descriptive flora of Puerto
observation). Individuals as large as 20 cm in Rico and adjacent Islands. Vol. 3. Editorial de la
diameter at breast height (Little and others 1974) Universidad de Puerto Rico, San Juan, PR.
must be several decades old. Management of the 461 p.
species should include protection from fire and
from commercial development of habitat. Existing Little, E.L., Jr., R.O. Woodbury, and F.H.
understory seedlings and saplings could be Wadsworth. 1974. Trees of Puerto Rico and the
released from competition by thinning or Virgin Islands. Vol. 2. Agriculture Handbook
eliminating the overstory in patches. 449. U.S. Department of Agriculture,
Washingtion, DC. 1,024 p.
Benefits.—Marble tree is pretty enough to make a
good ornamental but has rarely been used as such. Medina, E., E. Cuevas, J. Figueroa, and A.E.
It is an important component of the plant Lugo. 1994. Mineral content of leaves from
community in the harsh environment where it is trees growing on serpentine soils under
most common and furnishes food and cover for contrasting rainfall regimes in Puerto Rico. Plant
animals. The fruits of marble tree are listed as a and Soil 158(1): 13-21.
food item for the endangered Anegada iguana
(Cyclura pinguis) (Zoological Society of San Missouri Botanical Garden. 2002a. Tropicos-Flora
Diego 2002). The wood is light-brown, hard, of Panama Checklist. http://mobot.org/cgi-
heavy, fine-textured, strong, and durable (Little bin/search_pick. 1 p.
and others 1974). It is certainly useful for fuel and
fence posts and possibly carving and turnery. The Missouri Botanical Garden. 2002b. W3-Specimen
fruits are reported to be edible, and extracts of the data base. http://mobot.org/cgi-bin/search_vast.
plant are a stimulant (Liogier 1990). 1 p.
References Zoological Society of San Diego. 2002.

Unraveling the relationships between Caribbean
Howard, R.A. 1989. Flora of the Lesser Antilles, iguanas and their ecosystems. http://www.
Leeward and Windward Islands. Vol. 5. Arnold sandiegozoo.org/conservation/fieldproject_
Arboretum, Harvard University, Jamaica Plain, anegada_plant.html. 2 p.
155
Castela erecta Turp. cockspur
SIMAROUBACEAE
Synonyms: Castela nicholsoni Hook.

Castelaria micholsoni (Hook.) Small
John K. Francis
elsewhere.
Ecology.—Cockspur is a coastal species. It grows

in beach strand vegetation, in sandy soils behind it,
and on rocky escarpments and hills somewhat
inland (Fundación La Salle de Ciencias Naturales
2002), a dominant to minor part of local xeric
scrub communities (Locklin 2002). It occurs to an
elevation of about 100 m in Puerto Rico (Little and
others 1974). These areas receive annual rainfall
totals of about 700 to 900 mm. Soil type does not
appear to be critical except that because of the
species’ short stature and slow growth, it tends to
be more successful in poor sites. Although
cockspur tolerates salt spray and mild salt in the
soil, it does not grow in very salty soils.
Reproduction.—In Puerto Rico, cockspur flowers

and fruits in the spring. The species bears male and
female flowers on different plants (dioecious).
Fleshy, red fruits are probably eaten and the seeds
dispersed by birds.
General Description.—Cockspur, also known as Growth and Management.—Cockspur is a slow-

goat-bush, retama, and urupagüita, is an evergreen, growing species. No management experience has
spiny shrub 1 to 4 m in height and up to 10 cm on been published.
stem diameter. The plant is multi-stemmed and
branchy. The twigs are stiff, sometimes zig-zag, Benefits.—Cockspur helps protect the soil and
whitish from fine hairs, and end in spines. There furnishes food and cover for wildlife. The sister
are also short spines at the leaf bases. The foliage species C. texana (T.&G.) Rose, once considered a
is sometimes dense, composed of alternate simple part of cockspur as C. erectas subsp. texana (T. &
oblong to elliptic, almost sessile leaves, 0.6 to 2.5 G.) Cronq., is considered an important browse
cm long by 0.3 to 1.2 cm broad, dark green and species (Secretaría de Medio Ambiente y Recursos
glabrous above, and hairy below. The foliage and Naturales. 2002). The common name, goat-bush,
twigs are bitter. Flowers are tiny, whitish to red suggests that it is browsed by goats.
and tightly clustered in the leaf axils. The fruits are
6- to 10-mm, red, fleshy drupes, one to four References
developing from a flower. Each fruit contains one
hard seed (Howard 1988, Liogier 1988, Little and Fundación La Salle de Ciencias Naturales. 2002.
others 1974). Playas de Aragua para uso turístico sustentable.
http://www.fundacite.arg.gov.ve/papelesf/docs/
Range.—Cockspur is native to Puerto Rico, the playas.pdf. 24 p.
Virgin Islands, Antigua, Barbuda, Anguilla,
Curacao, Aruba, northern Venezuela and northern Howard R.A. 1988. Flora of the Lesser Antilles,
Colombia (Howard 1988, Little and others 1974). Leeward and Windward Islands.
It is not known to have been planted or naturalized Dicotyledoneae, Part 1. Vol. 4. Arnold
156
Arboretum, Harvard University, Jamaica Plain, Locklin, C. 2002. Guajira-Barranquilla xeric scrub
MA. 673 p. (NT1308). World Wildlife Fund, Washington,
DC. http://www.worldwildlife.org/wildworld/
Liogier, H.A. 1988. Descriptive flora of Puerto profiles/terrestrial/nt/nt1308_full.html. 7 p.
2. Editorial de la Universidad de Puerto Rico, Secretaría de Medio Ambiente y Recursos
Río Piedras, PR. 481 p. Naturales. 2002. Especies forestales no
maderables y maderables no tradicionales de
Little, E.L., Jr., R.O. Woodbury, and F.H. zonas áricas y semiáridas: Castela texana
Wadsworth. 1974. Trees of Puerto Rico and the (T.&G.) Rose. http://www.semarnat.gob.mx/
Virgin Islands. Vol. 2. Agriculture Handbook pfnm3/fichas/castela_texana.htm. 2 p.
157
Ceanothus velutinus Dougl. ex Hook. snowbush ceanothus
RHAMNACEAE
Synonyms: none
John K. Francis
2001, Banner 2002, Brayshaw 1996, Harrington

1964, Nobs 1963, Pollock 2003, Welsh and others
1987).
Range.—The native range of snowbush ceanothus

includes British Columbia and Alberta south to
California and east to South Dakota and Colorado
(Abrams 1951, Anderson 2001, Natural Resources
Conservation Service 2003). There are two
varieties: hookeri and velutinus. The former, of the
Coast and Cascade ranges, in glaberous beneath
the leaves. The latter, occurring farther inland, is
finely hairy on the underside of the leaves
(Brayshaw 1996). The species is not known to
have naturalized outside its native range.
Ecology.—Snowbush ceanothus is a member of a

large number of plant associations (Anderson
2001). It favors well-drained soils and is
susceptible to root rot in more poorly drained
conditions (Hansen 2002). These are moderately
Drawing source: Britton and Brown 1913 acid to neutral, often rocky or gravely soils, poor
in nutrients, and of granitic parent material. The
General Description.—Snowbush ceanothus is taproot penetrates 1.8 to 2.4 m deep (Pollock
also known as shiny-leaf ceanothus, varnish-leaf 2003), and with the moisture-conserving leaf
ceanothus, sticky laurel, tobacco brush, mountain structure, impart the plant a great deal of drought
balm, buckbrush, deerbush, and snowbrush. It is resistance. Snowbush ceanothus grows in
an evergreen, upright to semiprostrate shrub 1 to 2 relatively low coastal habitat and in mountain sites
m (rarely to 4 m) in height. The shrub has one to up to 2,900 m in elevation (Welsh and others
many stems with reddish-brown bark. The twigs 1987). Although found on all aspects, it is more
are rigid, slightly flattened, and olive green. The likely to occur on southerly exposures (Anderson
plant is supported by a taproot and a deep, 2001). Precipitation and length of growing season
spreading root system. Simple, alternate leaves are vary a great deal because of the wide elevational
shiny dark green above, and pale beneath leathery, and latitudinal range of the species. Snowbush
and covered with a sticky resin. The 2- to 8-cm ceanothus is a pioneer species. Large numbers of
blades are broadly elliptical to ovate with rounded seedlings often appear after logging and especially
tips and rounded or subcordate bases. They have a after fires. The species generally grows in small to
fine serrate margin and three prominent veins large thickets, usually as the dominate species. It
proceeding from the base. The foliage is strongly develops best in full sun but can persist in open
aromatic, especially when crushed. Some have forest stands. It grows more rapidly than conifer
described it as being similar to cinnamon or seedlings and will suppress them for a time.
balsam. Inflorescences, which are borne on short However, after 10 to 75 years, conifers overtop it,
lateral branches on the previous year’s growth, and its vigor and abundance declines (Pollock
contain many small white, five-maris, perfect 2003).
flowers. Fruits are three-lobbed capsules 3 to 6
mm long. The seeds are shiny and tan to dark Reproduction.—Snowbush ceanothus flowers
brown. There are 2n = 24 chromosomes (Anderson from May through July and the fruits mature in
158
August and September (Banner 2002). The flowers round, although it makes up only a small portion
are pollinated by at least bees (Clark 1976). There of their diet. It is important fall and winter moose
is an average of 207,000 seeds/kg. When seeds browse in some areas. It has an in vitro dry matter
mature, beginning in August, they are ejected from digestibility of about 57 percent and a crude
the pod and dispersed a short distance (Anderson protein content of 7 to 18 percent (Anderson
2001). Good seed production begins when plants 2001). Snowbush ceanothus is important resting
are about 8 years old. As many as 1,250,000 and escape cover for many types of wildlife
seeds/ha/year can be produced. However, as much (Pollock 2003). Nitrogen is fixed by actinomycetes
as 99 percent of the annual seed crop can be of the genus Frankia in root nodules (Dalton
consumed by ants, birds, and rodents. (Pollock 1997). Snowbush ceanothus makes a beautiful and
2003). Seeds still accumulate in large numbers in hardy ornamental that can be pruned and shaped
the soil seed bank and can apparently remain (Hansen 2002). Native Americans used the leaves
viable for 200 to 300 years until conditions as a deodorant, smoke of the plant to kill bedbugs,
become favorable. Germination is epigeal (Reed and decoctions to treat flu, pain, and gonorrhea
1974). Heat, as from forest fires, appears to scarify (Moerman 1986). They also used the leaves as a
the seeds. Burned or damaged plants sprout from tobacco substitute (Pollock 2003).
the root crown. Layering also occurs when
branches come in contact with the soil (Anderson References
2001).
Abrams, L. 1951. Illustrated flora of the Pacific
Growth and Management.—The natural life States. Vol. 3. Stanford University Press,
span of snowbush ceanothus is longer than 25 Stanford, CA. 866 p.
years, and plants sometimes reach 50 years old.
However, stands sometimes begin to deteriorate Anderson, M.D. 2001. Ceanothus velutinus. In:
after 15 years (Anderson 2001). A 17-year-old U.S. Department of Agriculture, Forest Service,
stand in the Cascade Mountains in Oregon Rocky Mountain Research Station, Fire Sciences
contained over 4,000 stems/ha and averaged Laboratory, Fire Effects Information System.
34,000 kg/ha of above-ground biomass (Pollock http://www.fs.fed.us/database/feis/plants/shrub/
2003). Seeds may be sown in the late summer and ceavel/all.html. 73 p.
will germinate in the spring. Pretreatments are
necessary for good germination of spring-planted Banner, R. 2002. Deerbush. Utah State University,
seeds. A hot water soak (90 °C until the water Logan, UT. http://extension.usu.edu/coop/
cools) followed by cold stratification at 1 to 5 °C natres/range/Woody/deerbush.htm. 4 p.
for 30 to 90 days. Seeds should be sown in flats at
depths of twice the diameter of the seed in Brayshaw, T.C. 1996. Trees and shrubs of British
amended mineral soil. Germination may be as high Columbia. UBC Press, Vancouver, British
as 82 percent. Seedlings should be pricked into Colunbia, Canada. 373 p.
pots when they have gained several sets of true
leaves (Reed 1974). Plants can be propagated with Britton, N.L. and A. Brown. 1913. Illustrated flora
stem cuttings. Survival of transplanted seedlings is of the northern states, Canada, and the British
low (about 9 percent). Small seedlings survive possessions. Vol. 2, 2nd ed. Scribner, New York.
better than large ones (Pollock 2003). Snowbush 735 p.
ceanothus stands are difficult to control. Broadleaf
herbicides such as 2,4-D tend to top-kill rather Clark, L.J. 1976. Wild flowers of the Pacific
than eliminate it. Opening the brush canopy also Northwest. Gray’s Publishing Limited, Sidney,
allows seeds to germinate and grow. Glyphosate British Columbia, Canada. 604 p.
and hand slashing result in greater densities of the
species in the long run (Anderson 2001). Dalton, D. 1997. Nitrogen fixation: range of
organisms that can fix nitrogen. Reed College,
Benefits.—Snowbush ceanothus adds to the Portland, OR. http://academic.reed.edu/
beauty of the forest, helps protect the soil, biology/Nitrogen/Nfix1.html. 7 p.
furnishes food and cover for wildlife, and is a
useful ornamental and medicinal plant. The Hansen, W. 2002. Ceanothus. http://www.
species is little used by cattle and horses. Sheep nwplants.com/plants/shrubs/ceanothus/. 6 p.
browse it only when other forage is unavailable.
Deer and elk browse snowbush ceanothus year-
159
Harrington, H.D. 1964. Manual of the plants of Pollock, T. 2003. Ceanothus velutinus. University
Colorado. Sage Books, Denver, CO. 666 p. of University of Saskatchewan, Saskatoon,
Saskatchewan, Canada. http: //www.usask.ca/
Moerman, D.E. 1986. Medicinal plants of Native agriculture/plantsci/classes/range/ceanothus/
America. Technical Reports 19. University of html. 5 p.
Michigan, Museum of Anthropology, Ann
Arbor, MI. 524 p. Reed, M.J. 1974. Ceanothus L., ceanothus. In:
C.S. Schopmeyer, tech. coord. Seeds of woody
Natural Resources Conservation Service. 2003. plants in the United States. Agriculture
Plants profile: Ceanothus velutinus Dougl. ex Handbook 450. U.S. Department of Agriculture,
Hook. http://plants.usda.gov/cgi_bin/ Forest Service, Washington, DC. p. 284-290.
plant_profile.cgi?symbol=CEVE. 4 p.
Welsh, S.L., N.D. Atwood, S. Gooddrich, and L.C.
Nobs, M.A. 1963. Experimental studies on species Higgins. 1987. A Utah flora. Great Basin
relationships in Ceanothus. Publication 623. Naturalist Memoirs 9. Brigham Young
Carnegie Institution of Washington, University, Provo UT. 894 p.
160
Celastrus orbiculatus Thunb. oriental bittersweet
CELASTRACEAE
Synonyms: Celastrus articulatus Thunb.

Celastrus insularis Koidz.
Celastrus jeholensis Nakai
Celastrus lancifolius Nakai
Celastrus stephanotiifolius Makino
Celastrus strigulosus Nakai
Celastrus tatarinowii Rupr.
Celastrus versicolor Nakai
Jinshuang Ma and Gerry Moore

the petals 2.6 to 5.0 by 0.9 to 2.0 mm. The
functional male flowers have five stamens about as
long as the petals and a vestigial pistil. The
functional female flowers have a syncarpous pistil
with a three-lobed stigma, short columnar style,
and well-developed superior ovary. The fruit of
Oriental bittersweet is a bright orange, three-
valved, globose capsule, with each valve covering
one or two seeds that are enclosed in a fleshy,
crimson aril. The seeds (sans aril) are smooth, light
orange, oblong, 2.5 to 2.6 mm by 1.5 to 1.6 mm
(Clemants 2003). The genus name is derived from
the ancient Greek name Celastros (Kelastros),
Photo credit: Steven Clemants which in Greek was applied to a different
Mediteranean evergreen tree probably in the genus
General Description.—Oriental bittersweet is the now known as Phillyrea (Rehder 1940).
name used most often in North America to refer to
this nonnative vine. Other common names include Range.—Oriental bittersweet is native to east Asia
Asian or Asiatic bittersweet and round-leaved in Japan, Korea, and China, its southern limit
bittersweet. The woody roots can be up to 2 cm occurring along the Yangtze River watershed in
thick and are yellow to orange. Oriental China. In its native range it occurs in areas with
bittersweet is multistemmed with brown, round elevation ranges from 450 to 2200 m (Cheng and
twigs; the pith is solid and white. It can behave Huang 1999). The plant has become widely
like a shrub but it is usually a climbing woody vine established in the Eastern United States, occurring
that can reach up to 20 m in height. Oriental in 25 states (Shetler and Orli 2000, USDA 2003).
bittersweet is able to climb and effectively cover It is also known in Canada from Ontario and
other woody vegetation. The plant lacks tendrils or Quebec (Scoggan 1978).
aerial roots. The leaves of oriental bittersweet are
deciduous, alternate, spiral, stipulate, and measure Systematic Botany.—Oriental bittersweet is
(including the petiole) 2.8 to 13.7 by 2.9 to 7.6 cm, similar to American bittersweet (Celastrus
with the blades suborbicular to broadly obovate in scandens L.), and can be distinguished from it on
outline with crenulate to serrulate margins. The the basis of its leaves and inflorescences (Hou
leaf blades can have white hairs along the midvein 1955). The leaves of oriental bittersweet are
on the underside but are otherwise glabrous; the suborbicular to obovate and are usually less than
lateral veins are ascending. The leaf scars are flush twice as long as wide, whereas the leaves of
with the stem, not raised. The small, greenish- American bittersweet are elliptic to oblong and are
white, five-parted flowers are in axillary cymes of usually twice as long as wide. The flowers (and
three or four functionally unisexual flowers, with fruits) of oriental bittersweet are in short axillary
the partially fused sepals 3 to 7 by 1 to 2 mm and clusters; the flowers (and fruits) of American
161
bittersweet are in long terminal clusters. Both (Baker 1974, Newsome and Noble 1986, Sakai
species have the same chromosome number (n = and others 2001)–that made early horticulturists
23; Bowden 1945) and they can be hybridized prefer oriental bittersweet over American
(White and Bowden 1947, Pooler and others bittersweet (C. scandens L.) are the same
2002). The presence of some intermediate material characteristics that enable oriental bittersweet to be
in the wild with both axillary and terminal an extremely successful invasive plant.
inflorescences has led to the speculation that the
two species may hybridize in the wild (Dreyer and Reproduction.—Oriental bittersweet blooms in
others 1987, Mehrhoff in Pooler and others 2002). the spring and is pollinated by hymenopterous
Some spell the specific epithet “orbiculata” instead insects, especially bees, although wind may also
of “orbiculatus” used here. This is because be involved (Brizicky 1964). The fruit ripens in
botanists disagree as to whether the genus name the fall. The seeds are dispersed by birds and small
Celastrus L. should be treated as feminine mammals (Dreyer 1994). Humans are also
(“orbiculata”) or masculine (“orbiculatus”) (Paclt important dispersal agents since the fruiting stems
1998). In classical Greek, the generic name was are commonly used in dried flower arrangements
treated as feminine; however, Linnaeus treated the and subsequently discarded in compost and brush
genus as masculine. The matter is now being piles (Dreyer 1994). Oriental bittersweet can also
reviewed by a formal nomenclatural committee reproduce asexually through root suckering.
(Brummitt 2000).
Growth and Management.—Due to the plant’s
Ecology.—Oriental bittersweet grows in a wide highly invasive nature, it is not recommended that
variety of habitats, including, dune thickets, this plant be grown. Small infestations can be
fencerows, forests, forest edges, and roadsides. It controlled by cutting the stems and removing the
seems to do particularly well in disturbed habitats roots. Larger infestations require the application of
(Dreyer 1994, Steward and others in press). This glycophosphate herbicides (Miller 2002).
species was perhaps first introduced in the New
World during the 1860s and was later popularized References
for ornamental plantings (Patterson 1973, Rehder
1940). Oriental bittersweet is an aggressive Bailey, L.H. 1922. The standard cyclopedia of
invader that can dominate all vegetation levels of horticulture. vol 2. Macmillan Co., New York.
forested and open areas (Bergmann 2003). It 1,200 p
grows over other vegetation, completely covering
it. It can kill other plants by preventing them from Baker, H.G. 1974. The evolution of weeds. Annual
photosynthesizing and through girdling and Review of Ecology and Systematics 5: 1-24.
uprooting. Based on its native range and habitat
types, Oriental bittersweet can be expected to Bergmann, C. 2003. Oriental Bittersweet Celastrus
spread to additional areas in the United States and orbiculatus Thunb. http://www.nps.
Canada (J. Ma, personal observation). Oriental gov/plants/alien/fact/ceor1.htm. [not paged].
bittersweet is a shade tolerant species, able to
acclimate to low light levels under heavy shading Bowden, W.M. 1945. A list of chromosome
and grow explosively if shading is reduced. Once numbers in higher plants I. Acanthaceae to
large clones are established, it is difficult to Myrtaceae. American Journal of Botany 32: 81-
eliminate since it has developed such an extensive 92.
root system. While oriental bittersweet is
spreading and increasing in abundance, the native Brizicky, G.K. 1964. The genera of Celastrales in
American bittersweet is concurrently declining the southeastern United States. Journal of the
(Dreyer and others 1987, Forman and Stark 2001, Arnold Arboretum 45: 206-235.
Steward and others in press). Pooler and others
(2002) have hypothesized that the spread of Brummitt, R.K. 2000. Report of the Committee for
Oriental bittersweet and its hybridization with Spermatophyta: 49. Taxon 49: 261-278.
American bittersweet may be threatening the
genetic identity of the American bittersweet. The Cheng, C.Y., and P.H. Huang. 1999. Celastraceae.
very properties–faster growth (Bailey 1922, Flora Reipublicae Popularis Sinicae 45: 1-218
Rehder 1940), greater fecundity (Clemant and (in Chinese).
others 1991, Dreyer and others 1987, Hart 1928),
greater tolerance of environmental heterogeneity
162
Clemant, C., R. Warren, G. Dreyer, and P. Barnes. Patterson D. 1973 Distribution of Oriental
1991. Photosynthesis, water relations, and bittersweet in the United States. Journal of the
fecundity in the woody vines American and Elisha Mitchell Science Society 89: 245.
Oriental bittersweet (Celastrus scandens and
Celastrus orbiculatus). American Journal of Pooler, M.R., R.L. Dix, and J. Feely. 2002.
Botany 78 (Suppl.): 134-135. Interspecific hybridization between the native
bittersweet, Celastrus scandens and the
Clemants, S.E. 2003. Celastrus orbiculatus. New introduced invasive species, C. orbiculatus.
York Metropolitan Flora Project. Brooklyn Southeastern Naturalist 1: 69-76.
Botanic Garden http://www.bbg.org/sci/nymf/
encyclopedia/cel/cel0020b.htm#description. [not Rehder, A. 1940. Manual of cultivated trees and
paged]. shrubs. Macmillan Publishing Co., New York.
996 p.
Dreyer, G.D. 1994. Element stewardship abstract
for Celastrus orbiculata Asiatic bittersweet. The Saikai, A.K., F.W. Allendorf, J.S. Holt, D.M.
Nature Conservancy, Arlington, Virginia. Lodge, J. Molofsky, K.A. With, S. Baughman,
http://tncweeds.ucdavis.edu/esadocs/documnts/ R.J. Cabin, J.E. Cohen, N.C. Ellstrand, D.E.
celaorb.html.11p. McCauley, P.O’Neil, I.M. Parker, J.N.
Thompson, and S.G. Weller. 2001. The
Dreyer, G.D., L.M. Baird, and C. Fickler. 1987. population biology of invasive species. Annual
Celastrus scandens and Celastrus orbiculatus: Review of Ecology and Systematics 32: 305-
comparisons of reproductive potential between a 332.
native and an introduced woody vine. Bulletin of
the Torrey Botanical Club 114: 260-264. Scoggan, H.J. 1978. The Flora of Canada Part 3 –
Dicotyledoneae (Saururaceae to Violaceae).
Forman, J., and J. Stark. 2001. The introduction of National Museum of Natural Sciences, National
non-native plants into Massachusetts. Museums of Canada, Ottawa. 568 p.
http://omega.cc.umb.edu/~conne/jennjim/
celastrus.html. [not paged]. Shetler, S.G., and S.S. Orli. 2000. Annotated
checklist of the vascular plants of the
Hart, H.T. 1928. Delayed germination in seedsof Washington-Baltimore area. Part. 1. Ferns, fern
Peltandra virginica and Celastrus scandens. allies, gymnosperms, dicotyledons. Department
Publications of the Puget Sound Biological of Systematic Biology – Botany. National
Station of the University of Washington 6: 255- Museum of Natural History, Smithsonian
261. Institution, Washington, D.C. 186 p.
Hou, D. 1955. A revision of the genus Celastrus. Steward, A.M., S.E. Clemants, and G. Moore
Annals of the Missouri Botanical Garden 42: 2003. The concurrent decline of the native
215-302. Celastrus scandens and spread of the nonnative
Celastrus scandens in the New York City
Miller, J.H. 2002. Exotic pest plants and their metropolitan area. Journal of the Torrey
control. The Bugwood Network. Botanical Society 130: 143-146.
http://www.bugwood.org/weeds/forestexotics.ht
ml. Accessed 30 June 2003. [not paged]. USDA, NRCS. 2003. The PLANTS Database.
National Plant Data Center, Baton Rouge,
Newsome, A.E. and I.R. Noble. 1986. Ecological LA. http:plants.usda.gov. [not paged].
and physiological characters of invading species.
In: R.H. Groves and J.J. Burdon, eds. Ecology of White, O.E. and W.M. Bowden 1947. Oriental and
biological invasions. Cambridge University American bittersweet hybrids. Journal of
Press. p. 1-20. Heredity 38: 125-127.
Paclt, J. 1998. Proposal to conserve the name

Celastrus (Celastraceae) as being of feminine
gender. Taxon 47: 879-880.
163
Celastrus scandens L. American bittersweet
CELASTRACEAE
Synonyms: Celastrus bullata L.

Euonymoides scandens (L.) Medic.

of 14 to 50 functionally unisexual flowers, with
the partially fused sepals 3 to 7 by 1 to 2 mm and
the petals 1.8 to 4.0 by 0.9 to 1.2 mm. The
functional male flowers have five stamens about as
long as the petals and a vestigial pistil. The
functional female flowers have a syncarpous pistil
with a three-lobed stigma, short columnar style,
and well-developed superior ovary. The fruit of the
American bittersweet is a bright orange, three-
valved, globose to ellipsoid capsule, with each
valve covering one or two seeds that are enclosed
in a fleshy, crimson aril. The seeds (sans aril) are
smooth, light orange, ellipsoid, 3.5 to 4.5 by 1.8 to
2.0 mm (Clemants 2003). The genus name is
derived from the ancient Greek name Celastros
(Kelastros), which in Greek was applied to a
Mediterranean evergreen tree probably in the
genus now known as Phillyrea (Rehder 1940).
Range.—American bittersweet has been reported

throughout most of the United States, except
Florida and the Western States of Arizona,
California, Colorado, Idaho, Nevada, New
Mexico, Oregon, Utah and Washington (USDA.
2003). It is also known in Canada from Manitoba,
New Brunswick and Quebec (Scoggan 1978).
Illustration source: Britton and Brown 1897 Reports (Bailey 1949, Britton and Brown 1897) of
this plant from New Mexico (then a territory with
General Description.—American bittersweet is the same area as the present state) are probably in
the name used most often in North America to error. No voucher specimens exist for these reports
refer to this native vine. Other common names and there has been no other report of American
include climbing orange root, false bittersweet, bittersweet from New Mexico.
fever twig, staff-tree, waxwork, and yellow-root
(Dillingham 1907). The woody roots can be up to Systematic Botany.—American bittersweet is
2 cm thick and are yellow to orange. American similar to the Oriental bittersweet (Celastrus
bittersweet is multistemmed with brown, round orbiculatus Thunb.) and can be distinguished from
twigs; the pith is solid and white. It can behave it on the basis of its leaves and inflorescences
like a shrub but it is usually a climbing woody vine (Hou 1955). The leaves of American bittersweet
that can reach up to 10 m in height. The plant lacks are elliptic to oblong and are usually twice as long
tendrils or aerial roots. The leaves of American as wide, whereas the leaves of Oriental bittersweet
bittersweet are deciduous, glabrous, alternate, are suborbicular to obovate and are usually less
spiral, stipulate, and measure (including the than twice as long as wide. The flowers (and
petiole) 4.5 to 14.8 by 2.7 to 5.5 cm, with the fruits) of American bittersweet are in long terminal
blades elliptic to ovate in outline with serrulate clusters; the flowers (and fruits) of Oriental
margins. The lateral veins are ascending. The leaf bittersweet are in short axillary clusters. Both
scars are flush with the stem. The small, greenish- species have the same chromosome number (n =
white, five-parted flowers are in terminal racemes
164
23, Bowden 1945) and they have been hybridized reported as being poisonous to cattle (Brizicky
(White and Bowden 1947, Pooler and others 1964). Fruiting stems of the species are often used
2002). The presence of some intermediate material in dried arrangements.
in the wild with both lateral and terminal
inflorescences has led to the speculation that the References
two species may hybridize in the wild (Dreyer and
others 1987, Mehrhoff in Pooler and others 2002). Bailey, L.H. 1922. The standard cyclopedia of
horticulture Vol 2. Macmillan Co., New York.
Ecology.—American bittersweet grows in a wide 1,200 p.
variety of habitats, including dune thickets,
fencerows, forests, forest edges, and roadsides. Bailey, L.H. 1949. Manual of cultivated
The native American bittersweet is declining, plants.Macmillan Co., New York. 1,116 p.
while the nonnative Oriental bittersweet (Celastrus
orbiculatus Thunb.) is spreading and increasing in Baker, H.G. 1974. The evolution of weeds. Annual
abundance (Dreyer and others 1987, Forman and Review of Ecology and Systematics 5: 1-24.
Stark 2001, Steward and others in press). Pooler
and others (2002) have hypothesized that the Bowden, W.M. 1945. A list of chromosome
spread of Oriental bittersweet and its hybridization numbers in higher plants I. Acanthaceae to
with American bittersweet may be threatening the Myrtaceae. American Journal of Botany 32:
genetic identity of the American bittersweet. The 81-92.
very properties--faster growth (Bailey 1922,
Rehder 1940), greater fecundity (Clemant and Britton, N. L. and A. Brown 1897. An illustrated
others 1991, Dreyer and others 1987, Hart 1928), flora of the northern United States, Canada, and
greater tolerance of environmental heterogeneity the British possessions Vol 2. Charles Scribner’s
(Baker 1974, Newsome and Noble 1986, Sakai Sons, New York. 643 p.
and others 2001)--that made early horticulturists
prefer Oriental bittersweet over American Brizicky, G.K. 1964. The genera of Celastrales in
bittersweet are the same characteristics that enable the southeastern United States. Journal of the
Oriental bittersweet to outcompete American Arnold Arboretum 45: 206-235.
bittersweet.
Chittenden, F.J. and P.M. Synge. 1956. Dictionary
Reproduction.—American bittersweet blooms in of gardening. Vol. 1, 2nd Ed. Clarendon Press,
the spring and is pollinated by hymenopterous Oxford, UK. 512 p.
insects, especially bees, although wind may also
be involved (Brizicky 1964). The fruit ripens in Clemant, C., R. Warren, G. Dreyer, and P. Barnes.
the fall. The seeds are dispersed by birds and small 1991. Photosynthesis, water relations, and
mammals (Dreyer 1994). American bittersweet fecundity in the woody vines American and
can also reproduce asexually through root Oriental bittersweet (Celastrus scandens and
suckering. Celastrus orbiculatus). American Journal of
Botany 78 (Suppl.): 134-135.
Growth and Management.—American
bittersweet does well in many conditions. It is a Clemants, S.E. 2003. Celastrus orbiculatus. New
rapid grower, adding up to several meters in one York Metropolitan Flora Project. Brooklyn
growing season. Adequate sun is important for Botanic Garden http://www.bbg.org/sci/nymf/
fruit production. It can be planted in Hardiness encyclopedia/cel/cel0030b.htm#description. [not
Zones 3 to 8, in soils with pH ranges from 3.7 to paged].
6.5. It cannot tolerate saturated soils. Propagation
can be effected either through layering or seed Dillingham, F.T. 1907. The staff-tree, Celastrus
(Chittenden and Synge 1956). Seeds should be scandens, as a former food supply of starving
stratified in the dark at about 4 °C for 3 months Indians. American Naturalist 41: 391-393.
(Young and Young 1992)
Dreyer, G.D. 1994. Element stewardship abstract
Benefits.—The branches of American bittersweet for Celastrus orbiculata Asiatic bittersweet. The
were used by Native Americans, including the Nature Conservancy, Arlington, Virginia.
Chippewa, as a last resort food source (Dillingham http://tncweeds.ucdavis.edu/esadocs/documnts/c
1907, Palmer 1871). The leaves have been elaorb.html. 11p.
165
Dreyer, G.D., L.M. Baird, and C. Fickler, 1987. Rehder, A. 1940. Manual of cultivated trees and
Celastrus scandens and Celastrus orbiculatus: shrubs. Macmillan Publishing Co., New York.
comparisons of reproductive potential between a 996 p.
native and an introduced woody vine. Bulletin of
the Torrey Botanical Club 114: 260-264. Saikai, A.K., F.W. Allendorf, J.S. Holt, D.M.
Lodge, J. Molofsky, K.A. With, S. Baughman,
Forman, J. and J. Stark. 2001. The introduction of R.J. Cabin, J.E. Cohen, N.C. Ellstrand, D.E.
non-native plants into Massachusetts. http:// McCauley, P.O’Neil, I.M. Parker, J.N.
omega.cc.umb.edu/~conne/jennjim/celastrus. Thompson, and S.G. Weller. 2001. The
html. [not paged]. population biology of invasive species. Annual
Review of Ecology and Systematics 32: 305-
Hart, H.T. 1928. Delayed germination in seeds of 332.
Peltandra virginica and Celastrus scandens.
publications of the Puget Sound Biological Scoggan, H.J. 1978. The Flora of Canada Part 3 –
Station of the University of Washington 6: 255- Dicotyledoneae (Saururaceae to Violaceae).
261. National Museum of Natural Sciences, National
Museums of Canada, Ottawa. 568 p.
Hou, D. 1955. A revision of the genus Celastrus.
Annals of the Missouri Botanical Garden 42: Steward, A.M., S.E. Clemants, and G. Moore
215-302. 2003. The concurrent decline of the native
Celastrus scandens and spread of the nonnative
Newsome, A.E. and I.R. Noble. 1986. Ecological Celastrus scandens in the New York City
and physiological characters of invading species. Metropolitan area. Journal of the Torrey
In: R.H. Groves and J.J. Burdon, eds. Ecology of Botanical Society 130: 143-146.
biological invasions. Cambridge University
Press, Cambridge, MS. p. 1-20. USDA, NRCS. 2003. The PLANTS Database,
Baton Rouge, LA. http:plants.usda.gov. [not
Palmer, E. 1871. Food products of the North paged].
American Indians. In H. Capron, Report of the
Commissioner of Agriculture for the Year 1870, White, O.E. and W.M. Bowden 1947. Oriental and
Washinton, D.C. p. 404-428 American bittersweet hybrids. Journal of
Heredity 38: 125-127.
Pooler, M.R., R.L. Dix, and J. Feely. 2002.
Interspecific hybridization between the native Young, J.A. and C.G. Young. 1992. Seeds of the
bittersweet, Celastrus scandens and the woody plants of North America. Dioscorides
introduced invasive species, C. orbiculatus. Press, Portland, OR. 407 p.
Southeastern Naturalist 1: 69-76.
166
Celtis reticulata Torr. netleaf hackberry
ULMACEAE
Synonyms: Celtis douglasii Planch.

C. occidentalis L. var. reticulata (Torr.) Sarg.
C. laevigata Willd. var. reticulata (Torr.) L. Benson
Ann M. DeBolt
reticulate or netlike pattern of prominent venation

on the leaf underside (Lanner 1983). Twigs and
leaves are frequently infested by insect galls, but
vigor is not seriously affected (Hayward 1948).
The small, green, apetalous, polygamo-
monoecious flowers number from one to several in
the leaf axils. Pea-sized fruits are fleshy, reddish-
orange to purple globose drupes, each with a
single large seed. Some authorities consider the
highly variable netleaf hackberry to be a variety of
sugarberry (C. laevigata Willd.) (Natural
Resources Conservation Service 2003, Stephens
1973). Others recognize many intergrading forms,
and it is possible that future distribution maps of
C. reticulata, if the name is retained, will not
resemble those available today.
Range.—Netleaf hackberry is found east of the

Cascade and Sierra Nevada Mountains from
central Washington south to northern Mexico and
east to central Kansas, Texas, and Oklahoma
(Lanner 1983, Little 1976).
Ecology.—At elevations ranging from 100 to

2,000 m, netleaf hackberry grows in a variety of
habitats including mountain shrub, deciduous
riparian woodlands, live oak-mixed shrub and
wash scrub communities, rocky ravines, and as
scattered individuals in semi-desert grasslands,
Illustration credit: J.R. Janish in Hitchcock and pinyon-juniper, and Joshua tree woodlands (Albee
others 1964. and others 1988, Brown 1982, Carmichael and
others 1978, Plummer 1977). Populations can be
General Description.—Netleaf hackberry is also small or highly localized, particularly at the
known as western or Douglas hackberry. It is a northern latitudes (Daubenmire 1970), or they can
deciduous shrub to small tree from 3 to 20 m tall. comprise the dominant vegetation matrix. On the
Near the northern edge of its range in Idaho, its Edwards Plateau of Texas, netleaf hackberry
maximum height is 12 m (DeBolt and McCune grows as an overstory codominant in many
1995). This species’ growth form varies from a riparian woodlands (Van Auken and others 1979).
singular, reasonably straight-stemmed tree to a A hackberry association was presented by Dick-
gnarled shrub with multiple stems. The bark is Peddie and Hubbard (1977) for the mixed
gray, hard, and moderately thick, forming deciduous series of their riparian classification
protruding corky ridges with age (Elias 1980). scheme for New Mexico, and several ecologists
Leaves are 3 to 9 cm long, alternate, simple, ovate have recognized it as a vegetation type on the
to ovate lanceolate, and somewhat leathery with lower Snake and Salmon Rivers in Idaho and
scabrous surfaces and entire to serrate margins. Oregon (Huschle 1975, Tisdale 1986), where
The specific epithet reticulata comes from the warm canyons are adequate for its existence. At
167
northern latitudes stands are often associated with Bonner, F.T. 1974. Celtis L. Hackberry. In: C.S.
rock (DeBolt and McCune 1995), which provides Shopmeyer, tech. coord. Seeds of woody plants
extra moisture and some protection from wildfire. in the United States. Handbook 450. U.S.
Department of Agriculture, Agriculture
Reproduction.—Wind-pollinated, nonshowy Washington, DC. p. 298-300.
flowers appear in spring, just before or as the
leaves unfold, depending on latitude. Fruits ripen Brown, D.E. 1982. Great Basin montane
from September to November and persist on the scrubland. In: Biotic communities of the
trees until midwinter. They are dispersed by birds, American Southwest–United States and Mexico.
rodents, and other small mammals. Netleaf Desert Plants 4: 83-84.
hackberry exhibits low germination percentages
(37 percent) and high seed dormancy (Bonner Carmichael, R.S., O.D. Knipe, C.P. Pase, and
1974). Seeds are hard and thick-walled, probably W.W. Brady. 1978. Arizona chaparral: plant
the major impediment to rapid germination. associations and ecology. Agriculture Research
Germination is epigeal. Dormancy can be Paper RM-202. U.S. Department of Agriculture,
overcome with stratification at 5 oC in moist sand Ft. Collins, CO. 16 p.
or other suitable media for 120 days (Bonner
1974). Mechanical scarification enhances Daubenmire, R. 1970. Steppe vegetation of
germination after 45 days (DeBolt 1992). Netleaf Washington. Technical Bulletin 62. Agriculture
hackberry can reproduce vegetatively by sprouting Experiment Station, Pullman, WA. 131 p.
from the root crown, stem base, or lateral roots. It
can also be propagated by cuttings (Bonner 1974). DeBolt, A.M. 1992. The ecology of netleaf
hackberry in Idaho. Master’s thesis, Oregon
Growth and Management.—Seedlings establish State University, Corvallis, OR. 167 p.
from animal-, gravity-, or water-dispersed
propagules in sites free of herbaceous competition, DeBolt, A.M. and B. McCune. 1995. Ecology of
often in rocky areas or in the duff of an existing Celtis reticulata in Idaho. Great Basin Naturalist
stand. Plummer (1977) found netleaf hackberry 55: 237-248.
useful in stabilizing disturbed areas in pinyon-
juniper and mountain brush types of Utah. In Dick-Peddie, W.A. and J.P. Hubbard. 1977.
Texas and Arizona, it is increasingly being used Classification of riparian vegetation. Pages 85-
for riparian restoration and wildlife plantings. It 90 In: R.R. Johnson and A. Dale, eds.
currently receives limited use in artificial plantings Importance, preservation, and management of
in the northern portion of its range, though once riparian habitat. General Technical Report RM-
established, young plants are hardy and persistent. 43. U.S. Department of Agriculture, Ft. Collins,
Netleaf hackberry is generally slow-growing, but CO.
plants may live for 300 to 400 years (DeBolt and
McCune 1995). Elias, T.S. 1980. The complete trees of North
America. Times Mirror Magazines, Inc., New
Benefits.—Netleaf hackberry provides cover and York. 565 p.
food for a host of wildlife species. Its fruits are
particularly important for birds during winter Hayward, C.L. 1948. Biotic communities of the
(Hayward 1948, Lanner 1983), and its leaves and Wasatch chaparral, Utah. Ecological
twigs are browsed by bighorn sheep, mule deer, Monographs 18: 473-506.
whitetail deer, elk, and occasionally by livestock.
It could be used more widely for domestic Hitchcock, J.C., A. Cronquist, M. Ownbey, and
landscaping as it requires little maintenance and is J.W. Thompson. 1964. Vascular plants of the
drought tolerant. Its hard wood is still used today Pacific Northwest. Part 2: Salicaceae to
on a limited basis for tool handles and firewood. Saxifragaceae. University of Washington Press,
Seattle, WA. 597 p.
References
Huschle, G. 1975. Analysis of the vegetation along
Albee, B.J., L.M. Schultz, and S. Goodrich. 1988. the middle and lower Snake River. M.S. thesis,
Atlas of the vascular plants of Utah. Occasional University of Idaho, Moscow. 271 p.
Publication 7. Utah Museum of Natural History,
Salt Lake City, UT. 670 p.
168
Lanner, R.M. 1983. Trees of the Great Basin. Stephens, H.A. 1973. Woody plants of the North
University of Nevada Press, Reno, NV. 215 p. Central Plains. University Press of Kansas,
Lawrence, KS. 530 p.
Little, E.L., Jr. 1976. Atlas of United States trees.
Volume 3-Minor western hardwoods. Tisdale, E.W. 1986. Canyon grasslands and
Miscellaneous Publication 1,314. U.S. associated shrublands of west-central Idaho and
Department of Agriculture, Washington, DC. adjacent areas. Bulletin 40. Forestry, Wildlife,
215 p. and Range Experiment Station, University of
Idaho, Moscow. 42 p.
Natural Resources Conservation Service. 2003.
Plants profile: http://plants.usda.gov/ [not Van Auken, W.W., A.L. Ford, and A. Stein. 1979.
paged]. A comparison of some woody upland and
riparian plant communities of the southern
Plummer, A.P. 1977. Revegetation of disturbed Edwards Plateau. Southwestern Naturalist 24:
intermountain area sites. In: J.C. Thames, ed. 165-180.
Reclamation and use of disturbed lands of the
Southwest. University of Arizona Press, Tucson.
p. 302-337.
169
Cephalanthus occidentalis L. buttonbush
RUBIACEAE
Synonyms: Cephalanthus berlandieri Wernh.
Kristina Connor
Cuba, Central America, and in the West Indies

(Little 1979). Recognized varieties are C.
occidentalis L. var. californicus Benth., C.
occidentalis var. pubescens Raf., and C.
occidentalis var. angustifolius Dippel.
Ecology.—Buttonbush is a wetland species that

cannot tolerate drought. It commonly grows in
thickets in areas that have intermittent flooding.
The open, rangy plant is not particularly attractive
and is seldom found in cultivation today, although
Van Dersal (1938) and Vines (1960) reported that
it was cultivated as early as 1735 as a honey plant.
The creamy white summer flowers of buttonbush
attract butterflies, honeybees, and hummingbirds.
While it is common in natural environments, its
habitat is threatened in California and it grows
poorly along manmade waterways (Holstein
1984). Faber-Langendoen and Maycock (1989)
state that buttonbush abundance increases with
increasing depth of water and light levels, while
Holstein (1984) suggests distribution may be
limited by mean July temperatures of 20 oC. It is
General Description.—Buttonbush, also known classified as a pioneer species and grows best in
as buttonball, button willow, common buttonbush, wet areas that receive full sun. It is able to tolerate
honey-bells, globe-flower, and riverbush, is a some salinity that might result from hurricane
deciduous, wetland shrub or small tree that can storm surges but will not survive long-term
reach 6 m in height but generally averages 1 to 3 exposure to salt water (McCarron and others
m tall. The trunk base is often swollen (Snyder 1998).
1991). Branches are generally green when young
but darken upon maturity and have conspicuous, Reproduction.—The perfect, creamy white
raised lenticels (Brown and Kirkman 1990). The terminal and axillary flowers of buttonbush occur
short-petioled glossy green leaves are elliptic or in dense spherical heads 2 to 3.5 cm in diameter.
lanceolate-oblong; they are mostly opposite but, The flower balls stand on stalks that are 5 cm long.
on the same plant, can occur in whorls of three or The fragrant individual flowers have corollas 6 to
four. Leaves range in size from 7 to 18 cm in 10 mm long, with pubescent lobes, and four
length and are 4 to 10 cm wide. stamens that extend 5 to 8 mm beyond the
corollas. Flowers are produced over a long period,
Range.—Buttonbush is common along stream and from late spring, throughout the summer months,
pond borders, in swamps, floodplains and other and into early autumn. Flowers are thought to be
riparian areas throughout the eastern half of the self-incompatible (Imbert and Richards 1993). The
United States. It occurs naturally in southern Nova long-stemmed fruits are clusters of achenes. The
Scotia, New Brunswick, Quebec, and Ontario as hard nutlets are 4 to 7 mm long and turn reddish-
well as through the eastern half of the Great Plains brown when mature (August to November). Fruit
States; scattered populations and varieties are balls may persist throughout the winter. Bonner
found in Arizona, New Mexico, southern (1974a) reported an average of 295,000 seeds/kg.
California, and Texas. It also grows in Mexico, Seeds are high in carbohydrates (Bonner 1974b).
170
Longevity of seeds in storage is unknown but they Bonner, F.T. 1974b. Chemical components of
are thought to be orthodox. Thus, if seeds are dried some southern fruits and seeds. United States
to a moisture content of 12 percent or less, they Department of Agriculture Forest Service Res.
can be stored under refrigeration for long periods Note SO-183, New Orleans. 3 p.
(Roberts 1973). Germination is epigeal, cotyledons
forced above ground, and no pretreatment is Bramble, W.C. and M.K. Goddard. 1943. Seasonal
required. DuBerry (1963) and Bonner (1974a) browsing of woody plants by white-tailed deer
reported germinations from 78 to 86 percent while in the bear oak forest type. Journal of Forestry
Vines (1960) reported a low germination 41(7): 471-475.
percentage. Seeds may be transported by animals,
birds, or by water. Buttonbush may also be
Brown, C.L. and L.K. Kirkman. 1990. Trees of
propagated from tip cuttings in the spring or
Georgia and adjacent states. Timber Press,
mature-wood cuttings in the winter.
Portland, OR. 292 p.
Growth and Management.—Buttonbush is a fast
DuBerry, A.P., Jr. 1963. Germination of
growing but short-lived shrub. It can be used as a
bottomland tree seed while immersed in water.
landscape shrub border but if not planted in moist
Journal of Forestry 61: 225- 226.
soil, it must be watered frequently. It must also be
pruned to maintain a good form. In nature,
Faber-Langendoen, D. and P.F. Maycock. 1989.
buttonbush occurs in dense thickets. It will
Community patterns and environmental
resprout after fire (Vogl 1973, Wade and others
gradients of buttonbush, Cephalanthus
1980). It has no reported pest problems but is
occidentalis, ponds in lowland forests of
moderately sensitive to herbicides. Thickets can be
southern Ontario. Canadian Field-Naturalist
reduced by cutting.
103(4): 479-485.
Benefits.—Buttonbush seeds are an important
Holstein, G. 1984. California riparian forests:
food for water birds but can be toxic to other
deciduous islands in an evergreen sea. In: R.E.
animals (Snyder 1991). Dense buttonbush thickets
Warner and K.M. Hendris, eds. California
provide a safe nesting ground for many wetland
riparian systems: Ecology, conservation, and
birds. Buttonbush also serves as a source of honey
productive management Proceedings of a
for butterflies, bees, and hummingbirds. A
conference; 1981 Sept. 17-19, Davis CA.
decoction of the inner bark was used by Native
University of California Press, Berkeley, CA.
Americans as an emetic. The bark was also used as
p. 2-22.
a substitute for quinine. The wilted leaves, which
contain bitter glycosides, cephalin and
Imbert, F.M. and J.H. Richards. 1993. Protandry,
cephalanthin (ACES 2001), are reportedly toxic to
incompatibility, and secondary pollen
some animals, especially cattle that eat them when
presentation in Cephalanthus occidentalis
other foliage is scarce. However, buttonbush
(Rubiaceae). American Journal of Botany 80:
leaves are eaten by deer (Bramble and Goddard
395-404.
1943), apparently with no ill effects. Other plant
parts are less toxic.
Little, E.L., Jr. 1979. Checklist of United States
trees (native and naturalized). Agriculture
References
Handbook 541. United States Department of
Agriculture, Forest Service, Washington, D.C.
ACES, Alabama Cooperative Extension System.
375 p.
2001. Poisonous Plants of the Southeastern
United States.ANR-975 http://www.aces.edu/
McCarron, J.K., K.W. McLeod, and W.H. Conner.
dept/extcomm/ publications. 56 p.
1998. Flood and salinity stress of wetland
woody species buttonbush (Cephalanthus
Bonner, F.T. 1974a. Cephalanthus occidentalis L.,
occidentalis) and swamp tupelo (Nyssa sylvatica
common buttonbush. In: C.S Schopmeyere,
var. biflora). Wetlands 18: 165-175.
tech. coord. Seeds of woody plants in the United
States. United States Department of Agriculture
Roberts, E.H. 1973. Predicting the storage life of
Forest Service Agriculture Handbook 450,
seeds. Seed Science and Technology 1: 499-514.
Washington, D.C. 883 p.
171
Snyder, S.A. 1991. Cephalanthus occidentalis. In: Vines, R.A. 1960. Trees, shrubs, and woody vines
United States Department of Agriculture, Forest of the Southwest. University of Texas Press,
Service, Rocky Mountain Research Station Fire Austin, TX, 1,104 p.
Sciences Laboratory. Fire Effects Information
System, http://www.fs.fed.us/database/feis. 10 p. Vogl, R.J. 1973. Effects of fires on the plants and
animals of a Florida wetland. American Midland
Van Dersal, W.R. 1938. Native woody plants of Naturalist 89: 334-347.
the United States: their erosion-control and
wildlife values. United States Department of Wade, D., J. Ewel, and R. Hofstetter. 1980. Fire in
Agriculture Miscellaneous Publication 393, South Florida ecosystems. United States
Washington, D.C. 362 p. Department of Agriculture Forest Service
General Technical Report SE-17. Asheville, NC.
125 p.
172
Cercocarpus intricatus Wats. littleleaf mountain-mahogany
ROSACEAE
Synonyms: Cercocarpus ledifolius var. intricatus (Wats.) Jones

Cercocarpus arizonicus Jones
Stanley G. Kitchen
arising primarily from short spur branchlets on

second-year growth. Hard, light brown fruits are
cylindrical achenes, approximately 1 mm wide and
4 to 10 mm long with a persistent twisted or
spiraling tail (style) 1 to 3 cm long (Welsh and
others 1987). Fruit body and especially the tail are
covered with short (1 mm) hairs that extend from
the axis when dry and facilitate wind dispersal.
Taxonomy.—The evolutionary process by which

Littleleaf mountain-mahogany segregated from
curlleaf mountain-mahogany (C. ledifolius Nutt.
var. ledifolius) appears to still be in progress with
intermediates between the species widespread
(Stutz 1990). The shorter and more compact
littleleaf mountain-mahogany is more drought
tolerant, has fewer stamens and shorter tails on
fruits than does curlleaf mountain-mahogany
(Stutz 1990). Although hybridization occurs
between the two species, littleleaf mountain-
mahogany appears to exhibit much greater
reproductive isolation from true mountain
mahogany (C. montanus Raf.) than does its
ancestral species (Stutz 1990). Walker and Turley
(1999) cited examples of hybrids between littleleaf
and true mountain-mahoganies, suggesting that a
lack of hybridization may be due, at least in part,
to limited interspecies contact.
Range.—Littleleaf mountain-mahogany is found

throughout most of Nevada and Utah and in parts
of eastern California, northern Arizona, and
extreme western Colorado at elevation of 900 to
General Description.—Littleleaf mountain- 3,000 m (Davis 1990). Limited populations have
mahogany, also known as little leaf cercocarpus also been reported for New Mexico and Wyoming
and dwarf mountain-mahogany, is a long-lived, (NatureServe Explorer 2002).
unarmed, intricately branched and occasionally
tree-like, evergreen shrub. Height of mature plants Ecology.—Within desert shrub, pinyon-juniper,
varies from 0.5 to 2.5 m. Mature bark is smooth and mountain brush communities, littleleaf
and light gray. Dark brown heartwood is dense and mountain mahogany is typically found rooted in
extremely hard. Persistent leaves are 3 to 18 mm cracks and crevices of exposed sedimentary and
long, 0.8 to 1.4 mm wide (oblong to linear) and metamorphic rocks; commonly limestone,
leathery with strongly revolute margins (Welsh dolomite, and quartzite in the Great Basin and
and others 1987). Leaf pubescence varies from sandstone in the Colorado Plateau (Davis 1990). It
glabrous to villous. Perfect, apetalous flowers, 3 to is also found in shallow, poorly developed soils
9 mm long, occur singularly or in small clusters derived from these substrates. It grows best in full
173
sun. Soil pH of littleleaf mountain-mahogany Resources, Utah State University, Logan, UT.
habitat is neutral to slightly basic, (Fairchild and p. 1-13.
Brotherson 1980). Annual precipitation is 200 to
500 mm. Although undocumented, it is believed Fairchild, J.A. and J.D. Brotherson. 1980. Micro-
that this species possesses nitrogen fixing root habitat relationships of six major shrubs in
nodules similar to those reported for close relatives Navajo National Monument, Arizona. Journal of
(Davis 1990). Littleleaf mountain-mahogany is Range Management 33: 150-156.
browsed heavily by wild ungulates and domestic
sheep when it can be reached. Kitchen, S.G. and S.E. Meyer. 1990. Seed
dormancy in two species of mountain-mahogany
Reproduction.—Flowering occurs from mid-May (Cercocarpus ledifolius and Cercocarpus
to late June (Walker and Turley 1999). Fruits ripen montanus). In: K.L. Johnson, ed. Proceedings of
and disperse in July and August. Years of the Fifth Utah Shrub Ecology Workshop: The
abundant seed production are rare. A high Genus Cercocarpus; 1988 July 13-14; Logan,
percentage of empty fruits is common. Plummer UT. College of Natural Resources, Utah State
and others (1968) reported 112,000 cleaned seeds University, Logan, UT. p. 27-42.
(fruits with tails removed) per kg. Seed dispersal is
by wind. NatureServe Explorer. 2002. An online
encyclopedia of life [web application]. Version
Growth and Management.—Although 1.6. NatureServe. Arlington, VA, USA.
information on seed germination for this species is http://www.natureserve.org/explorer. [not
lacking in the literature, primary dormancy at seed paged].
dispersal is probably similar to that of curlleaf
mountain-mahogany. Kitchen and Meyer (1990) Plummer, A.P., D.R. Christensen, and S.B.
observed almost no germination for this species Monsen. 1968. Restoring big-game range in
without prechill (1 oC). They reported germination Utah. Publ. No. 68-3. Utah division of Fish and
of six seed collections after 6 weeks of prechill Game, Ephraim, UT. 183 p.
ranged from 53 to 96 percent of viable. Seedling
growth rate is slow, and plants require 10 to 30 Stutz, H.C. 1990. Taxonomy and evolution of
years to reach maturity. Management alternatives Cercocarpus in the western United States. In:
for littleleaf mountain-mahogany are limited due K.L. Johnson, ed. Proceedings of the Fifth Utah
to the nature of the sites it occupies. Shrub Ecology Workshop: The Genus
Cercocarpus, 1988 July 13-14, Logan, UT.
Benefits.—Littleleaf mountain-mahogany College of Natural Resources, Utah State
provides cover and year-round forage on harsh University, Logan, UT. p. 15-25.
landscapes. It is of particular value to native
ungulates that depend on rugged escape terrain Walker, S.C. and D. Turley. 1999. Characteristics
such as mule deer (Ococolius hemionus), bighorn of mountain mahogany (Cercocarpus) species
sheep (Ovis Canadensis), and mountain goat and hybrids in Utah hybrid zone. In: E.D.
(Oreamnos americanus). McArthur, W.K. Ostler, and C.L. Wambolt,
comps. Proceedings: Shrubland Ecotones; 1998
References. August 12-14; Ephraim, UT. Proceedings
RMRS-P-11. U.S. Department of Agriculture,
Davis, J.N. 1990. General ecology, wildlife use, Forest Service, Rocky Mountain Research
and management of the mountain mahoganies in Station, Ogden, UT. p. 32-34.
the Intermountain West. In: K.L. Johnson, ed.
Proceedings of the Fifth Utah Shrub Ecology Welsh, S.L., N.D. Atwood, L.C. Higgins, and S.
Workshop: The Genus Cercocarpus; 1988 July Goodrich. 1987. A Utah Flora. Brigham Young
13-14; Logan, UT. College of Natural University Print Services, Provo, UT. 894 p.
174
Cercocarpus ledifolius Nutt. curl-leaf mountain mahogany
ROSACEAE
Synonyms: None
Christopher Ross
pinyon/juniper, scrub oak/mountain brush, aspen,

ponderosa, jeffrey and other pines, and spruce/fir
(Martin 1950, Ross 1999). It occurs on a wide
variety of soils (Davis and Brotherson 1991),
including decomposed granite and limestone,
altered andesites (Billings 1976) and others. Curl-
leaf mountain mahogany often occurs in rock
outcrops and talus slopes with no apparent soil
development. In pine and spruce/fir forests it may
eventually be overshadowed by tall trees and
shaded out. It often forms dense, closed canopy
stands that permit little understory or interspecific
competition. Sapsuckers and other woodpeckers
may attack intermediate age class trees, girdling
the stems and causing extensive stand mortality in
some cases (Ross 1999). Other predators include
the bark beetle Renoci’s heterodoxus (Kraft 1960)
and the leaf defoliater Anacamtodes clivinavia
profanata (Furniss and Barr 1969). A variety of
other insects also attack the species at times.
Reproduction.—Flowers are one to three per

inflorescence, axillary, and more or less sessile,
with a straight stigma and 15 to 25 stamens. Fruits
General Description.—Curl-leaf mountain
are unusual for a member of the rose family, 4 to 8
mahogany is a large-sized shrub to small or
mm long, hard, with a 2.4 to 3.5 cm plume style
medium tree. Young stems are smooth and gray,
(Lis 1993). Seed production is episodic but may be
becoming rough and brown in mature specimens,
very heavy at times, with the seeds forming drifts
and often scaly in older trees. The evergreen leaves
beneath trees. Seed predation by insects in fall may
are narrowly lanceolate, often revolute, about 1 to
be nearly complete at times (Dealy 1975).
1.5 cm long with a 1- to 3-mm petiole. They are
Reproduction is sporadic, occurring usually on bare
densely white-wooly underneath. (Lis 1993). The
mineral soil, including fresh road cuts, but very
trees are extremely long lived in the absence of
uncommon in established plant communities. The
external sources of mortality and are often by far
increase in cheatgrass (Bromus tectorum L.) and
the oldest members of the communities in which
other annuals over its range has apparently reduced
they occur (Ross 1999).
reproduction in many areas (Ross 1999). Under
established stands, germination is common but the
Range.—Curl-leaf ountain mahogany is widely
seedlings usually damp off and die (Ross 1999).
distributed in North America, from Montana to
Episodic regeneration and mortality, especially
Baja California and from near the Pacific Coast in
from fire and sapsuckers, may create even-aged
southwestern Oregon to the Bighorn Mountains in
stands with almost no diversity of age class.
Wyoming and near Chadron, Nebraska. It occurs at
elevations of 610 to 1,372 m in the north, and to
3,000 m or higher in the south (USDA USFS 1937, Fire Effects.—Although curl-leaf mountain
Martin 1950). mahogany is sometimes referred to as a weak
resprouter after fire, this is very uncommon. In the
Ecology.—Curl-leaf mountain mahogany occurs in western Great Basin, it is invariably killed by fire
a variety of plant associations, including sagebrush, regardless of intensity, and never resprouts. Even
175
very light burns that do no apparent damage to References
mature trees result in full mortality within 1 year.
Closed mature stands may have insufficient Billings, W. D. 1976. Islands of sierran plants on
understory to carry fire, so that the fire and the arid slopes of Peavine Mountain. Mentzelia
resulting mortality are confined to the edges of the 6: 32-39.
stands. Regeneration by seed may occur after fire
if the soil is not colonized by other species. In Davis, J.N. and J. D. Brotherson. 1991. Ecological
some cases, the resulting stands occupy essentially relationships of curlleaf mountain mahogany
the same area as the burned stand. Historic fire (Cercocarpus ledifolius Nutt.) communities in
lines and burns are often clearly demarcated by the Utah and implications for management. Great
sharp boundaries of living stands. The wood is Basin Naturalist 51: 153-66.
rarely consumed by fire and often persists for
decades after, offering precise information about Dealy, J.E. 1975. Ecology of curlleaf mountain
past distribution and age classes. With changes in mahogany (Cercocarpus ledifolius Nutt.) in
fire intensity and return interval, usually related to eastern Oregon and adjacent areas. PhD.
increases in exotic annuals, curl-leaf mountain dissertation. Oregon State University, Corvallis,
mahogany in many areas is decreasing and is OR.
increasingly restricted to higher elevations, steeper
slopes, and cooler aspects (Ross 1999). Furniss, M.M. and W. F. Barr. 1969. Bionomics of
Anacamprodes clivinaria profanata on mountain
Growth and Management.—Although curl-leaf mahogany in Idaho. Agricultural Experiment
mountain mahogany is very long lived and Station Research Bulletin 73. University of
eventually assumes a tree form, young specimens Idaho, Moscow, ID. 24 p.
grow quickly as multistemmed shrubs. Stands are
typically composed of a single age class, perhaps Furniss M.M., D.C. Ferguson, K.W. Boget, J.W.
with younger specimens on the edges. It may be Burkhardt, A.R. Tiedemann, and J.L.
readily grown from seed for transplanting, although Oledmeyer. 1988. Taxonomy, life history, and
young transplants are attractive to rodents, ecology of a mountain mahogany defoliator,
lagomorphs, and large game. Seed collection is Stamnodes animata Pearsall in Nevada. Fish
easily done, although the hairy styles are a strong and Wildlife Research 3, USFWS, Washington,
respiratory and skin irritant. Management has D.C.
focused on increasing the forage production of old,
browsed stands via clipping and pruning (Davis Garrison, G.A. 1953. Effects of clipping on some
and Brotherson 1991), although this may result in range shrubs. Journal of Range Management 6:
tree mortality (Garrison 1953, Thompson 1970). 309-317.
Benefits.—Curl-leaf mountain mahogany has Kraft, G.F. 1960. Insects affecting bitterbrush and
found extensive use in the past as a fuel and other range plants. Office Report. Entomology
charcoal source. The very hard, beautiful wood Dept., Oregon State University, Corvallis, OR.
has been used for lathe work and parts for musical 55 p.
instruments (Ross 1999). Living stands provide
significant blue grouse nesting habitat (Stauffer Lis, R. 1993. Cercocarpus Mountain Mahogany.
and Petersen 1986). It is one of the few species that In: J.C Hickman, ed. 1993. The Jepson Manual:
meet protein requirements for wintering deer higher plants of California. U.C. Press, Berkeley
(Welch and McArthur 1979) and is heavily favored and Los Angeles, CA.
by bighorn sheep in summer (Rominger et al.
1988). Where present in even small amounts in Martin, F.L. 1950. A revision of Cercocarpus.
sagebrush, curl-leaf mountain mahogany allows Brittonia 7: 91-111.
greater wildlife diversity and abundance than
would otherwise occur (Furniss et al. 1988). It is Rominger, E.M., A.R. Dale, and J.A. Bailey. 1988.
also used increasingly as a commercial, residential, Shrubs in the summer diet of Rocky Mountain
and highway landscaping species. bighorn sheep. Journal of Wildlife Management
52: 47-50.
Ross, C. 1999. Population dynamics and changes

in curlleaf mountain mahogany (Cercocarpus
176
ledifolius Nutt.) in two adjacent Sierran and LaSal National Forest, USDA Forest Service
Great Basin mountain ranges. PhD. Dissertation, Range Improvement Notes, Vol. 15, No. 3.
University of Nevada, Reno.
USDA USFS. 1937. Range Plant Handbook.
Stauffer, D.F. and S.R. Petersen. 1986. Seasonal Washington D.C. U.S. Govt. Printing Office.
microhabitat relationships of blue grouse in p. 132–133.
southeastern Idaho. Great Basin Naturalist 46:
117-22. Welch, B.L. and E.D. McArthur. 1979. Nutritive
value of big sagebrush and other shrubs. In L.H.
Thompson, R.M. 1970. Experimental top pruning Stelter and others., eds. Proceedings, Shrub
of curlleaf mountain mahogany on the South establishment on disturbed arid and semi-arid
Horn Mountain, Ferron Ranger District, Manti- lands, Wyoming Fish and Game Dept. p. 9-22.
177
Cercocarpus montanus Raf. true mountain-mahogany
ROSACEAE
Synonyms: Cercocarpus betuloides Nutt. in T. & G.

Cercocarpus betulifolius Nutt. ex Hook.
Cercocarpus parvifolius Nutt.
Cercocarpus flabellifolius Rydb.
Stanley G. Kitchen
while the lower surface is generally pubescent

(Welsh and others 1987). Perfect, apetalous
flowers, 10 to 17 mm long, occur singularly or in
small clusters arising from axiles or on short spur
branchlets along upper portions of 2-year-old
stems. Hard, pale green to light brown fruits are
pubescent, cylindrical achenes, approximately 1 to
2 mm wide and 8 to 15 mm long with a persistent
twisted or spiraled style or tail 3 to 10 cm in
length. The numerous 1 to 2 mm long hairs present
on this appendage extend almost perpendicular to
the style axis when dry and facilitate wind
dispersal. Because of the reflective nature of these
hairs, abundant seed crops give plants a frosted
look in sunlight.
Taxonomy.—Nine varieties of true mountain

mahogany are recognized of which var. montanus
Raf. is most widespread (Marshall 1995,
NatureServe Explorer 2002). Varieties with
southwestern United States distributions include
vars. argenteus (Rydb) F.L. Martin, glaber (S.
Wats.) F.L. Martin, and the evergreen
paucidentatus (S. Wats) F.L. Martin. Closely
related species of the Southwest and Mexico
include: C. mexicanus Hendrickson, C. rzedowski
Hendrickson, and C. fothergilloides Kunth,
suggesting a southern North American origin
(Stutz 1990). Coastal California has varieties with
General Description.—True mountain- restricted distributions including the critically
mahogany, also known as mountain cercocarpus, imperiled (S1) Santa Catalina mountain-mahogany
birchleaf cercocarpus, birchleaf mountain- or C. montanus traskiae (Eastw.) F.L. Martin,
mahogany, alderleaf mountain-mahogany, which is sometimes treated as a separate species,
blackbrush, deerbush, tallowbrush, and lintisco and the vulnerable (S3) Island mountain-
(Spanish), is a long-lived, unarmed, mostly mahogany or C. montanus blancheae (Schneid.)
deciduous shrub, with few to many ascending F.L. Martin (NatureServe Explorer 2002). True
stems generally 1 to 4 m tall and 0.5 to 5 cm in mountain-mahogany is known to hybridize with
diameter at the base. Mature bark is smooth and other mountain-mahogany species where
light gray. Stout tap and lateral roots penetrate populations are in contact (Stutz 1990, Walker and
deep into available soil and rock crevices. Leaves Turley 1999). Hybrids derived from crosses with
are simple and alternate. Leaf length varies from 6 the more tree-like and evergreen curlleaf mountain
to 44 mm. Leaf blade shape is oblanceolate to mahogany (Cercocarpus ledifolius var.
roundish with shallow forward-pointing teeth on intermontanus N. Holmgren) are evergreen,
the terminal margin. The upper surface is glabrous intermediate in stature and leaf shape, and usually
178
sterile. In contrast, hybrids formed with C. possible significance of rodents as secondary
ledifolius Nutt. spp. ledifolius, are quite fertile and dispersers of mountain-mahogany seeds is
backcross with parent populations (Stutz 1990). unknown.
Range.—True mountain-mahogany, like other Growth and Management.—Although within-

members of the genus, is endemic to dry coastal plant flowering and fruit maturation are well
and interior foothills and mountains of the Western synchronized, some within-population asynchrony
United States and Mexico (Marshall 1995, Stutz is expected. Ripened fruits are easily dislodged
1990). In the United States, it is centrally located and collected by striking or shaking branches over
in Colorado, Utah, and Wyoming and to lesser hand-held hoppers. Optimal season for harvest
degrees in South Dakota, Nebraska, Kansas, varies from June to September. During harvest and
Oklahoma, Texas, New Mexico, Arizona, Nevada, handling, fruit hairs dislodge and can cause
California, Oregon, Idaho and Montana (Davis considerable discomfort to eyes and skin,
1990, NatureServe Explorer 2002). Within its revealing the motivation for the cowboy moniker,
range it grows at elevations of 1,000 and 3,000 m “hell feathers” (Plummer and others 1968).
(Davis 1990). Commercially available seed is typically cleaned
to 95 percent purity and 85 percent viability. Seeds
Ecology.—True mountain-mahogany grows in have been warehouse-stored for 7 years without
solid, sometimes extensive stands, and in mixtures significant loss in viability (Stevens and others
in sagebrush and mountain shrublands and pinyon- 1981). Although primary dormancy is apparently
juniper woodlands. This species is also found in absent for some collections of true mountain-
openings of ponderosa pine, mixed conifer, and mahogany seed (Deitschman and others 1974),
aspen forests. It does best in full sun. Annual moist chilling (stratification) treatments of 2 to 12
precipitation ranges from 250 to 600 mm. Soils are weeks are typically required to break dormancy
well drained, near-neutral to slightly basic, and (Kitchen and Meyer 1990). In one study, seeds
deep to absent (rock crevices). Nitrogen fixation in germinated in chilling (2 oC) between 7 and 15
root nodules and ectomycorrhizal facilitation of weeks of treatment (Kitchen and Meyer 1990). A
phosphorus uptake (Hoeppel and Wollum 1971) germination inhibitor may play a role in
are probably critical for the success of this species germination regulation (Moore 1963). Laboratory
on infertile soils. Plants often appear hedged from viability evaluations using standard TZ
heavy winter use by wildlife. Regeneration by (tetrazolium chloride) tests can be problematic due
crown sprouting is best after early season fires to difficulty in extracting undamaged embryos
(Marshall 1995). Ferguson (1983) suggested that from fruits (Kitchen and others 1989). Seeds are
the ability to sprout following fire might be used in restoration plantings of highly disturbed
ecotype dependent. Stand regeneration by seed is sites (e.g. surfacing mines) and for wildlife habitat
slow. improvement. Seedling vigor is fair and growth
rate is slow to moderate. Plants reach maturity in 5
Reproduction.—Anthesis for this wind pollinated to 15 years. Plants are tolerant of moderate to
species occurs between April and early July heavy use by browsers (Turley 2000) and shoots of
depending on elevation, latitude, and aspect. Fruits true mountain-mahogany provide adequate
ripen and disperse from July to October. A unique nutrition as winter forage for wildlife. Productivity
pattern of resource partitioning is commonly decreases when plants are protected from
observed in which branches with few seeds and herbivory over extended periods (Waugh 1990).
long leader growth occur concurrently with Decadent stands can be rejuvenated using light to
branches sporting many fruits and little leader moderate top removal techniques such as one-way
development (Walker and Turley 1999). Abundant chaining (Davis 1990). Both containerized and
fruit production may occur at intervals of 1 to 10 bare-root stock (1 and 2 year old) is available from
years (Plummer and others 1968). A high select nurseries.
percentage of nonviable (empty) fruit is common
for this species and the genus Cercocarpus in Benefits.—True mountain mahogany is an
general. Means of 90,000 and 130,000 cleaned important, often dominant shrub in many parts of
seeds per kg (fruits with tails removed) have been the Western United States. It provides valuable
reported in the literature (Kitchen and others 1989, winter forage for wildlife. It also provides
Deitschman and others 1974). Primary seed protection for, and increases the fertility of,
dispersal is by wind. However, seeds are also naturally occurring and human-caused (e.g. mine
gathered and likely cached by rodents. The spoils) infertile soils.
179
References
Moore, T.C. 1963. A germination inhibitor in
Davis, J.N. 1990. General ecology, wildlife use, achenes of Cercocarpus montanus. Ecol. 44:
and management of the mountain mahoganies in 406-409.
the Intermountain West. In: K.L. Johnson, ed.
Proceedings of the Fifth Utah Shrub Ecology NatureServe Explorer. 2002. An online
Workshop: The Genus Cercocarpus; 1988 July encyclopedia of life. Version 1.6. NatureServe,
13-14; Logan, UT. College of Natural Arlington, VA, USA. http://www.natureserve.
Resources, Utah State University, Logan, UT. org/explorer [not paged].
p. 1-13.
Plummer, A.P., D.R. Christensen, and S.B.
Deitschman, G.H., K.R. Jorgensen, and A.P. Monsen. 1968. Restoring big-game range in
Plummer. 1974. Cercocarpus. In: C.S. Utah. Publication 68-3. Utah Division of Fish
Schopmeyer, tech. coord.. Seeds of Woody and Game, Ephraim, UT. 183 p.
Plants of the United States. Agricultural
Handbook 450. U.S. Department of Agriculture, Stevens, R., K.R. Jorgensen, and J.N. Davis. 1981.
Forest Service, Washington, DC. p. 309-312. Viability of seed from thirty-two shrub and forb
species through fifteen years of warehouse
Ferguson, R.B. 1983. Use of wildland shrubs for storage. Great Basin Naturalist 4: 274-277.
wildland plantings in the Intermountain West.
In: S.B. Monsen and N. Shaw, comps. Managing Stutz, H.C. 1990. Taxonomy and evolution of
Intermountain Rangelands—Improvement of Cercocarpus in the western United States. In:
Range and Wildlife Habitats; Proceedings of K.L. Johnson, ed. Proceedings of the Fifth Utah
Symposia; 1981 September 15-17; Twin Falls, Shrub Ecology Workshop: The Genus
ID and 1982 June 22-24; Elko NV. General Cercocarpus; 1988 July 13-14; Logan, UT.
Technical Report INT-157. U.S. Department of College of Natural Resources, Utah State
Agriculture, Forest Service, Intermountain University, Logan, UT. p. 15-25.
Research Station, Ogden, UT. p. 136-147.
Turley, D. 2000. Effects of browsing on true
Hoeppel R.E. and A.G. Wollum. 1971. mountain mahogany (Cercocarpus montanus
Histological studies of ectomycorrhizae and root Raf.) MS Thesis. Brigham Young University.
nodules from Cercocarpus montanus and Provo, UT. 115 p.
Cercocarpus paucidentatus. Canadian Journal of
Botany 49: 1315-1318. Walker, S.C. and D. Turley. 1999. Characteristics
of mountain mahogany (Cercocarpus) species
Kitchen, S.G. and S.E. Meyer. 1990. Seed and hybrids in Utah hybrid zone. In: E.D.
dormancy in two species of mountain-mahogany McArthur, W.K. Ostler, and C.L. Wambolt,
(Cercocarpus ledifolius and Cercarpus comps. Proceedings: Shrubland Ecotones; 1998
montanus). In: K.L. Johnson, ed. Proceedings of August 12-14; Ephraim, UT. Proceedings
the Fifth Utah Shrub Ecology Workshop: The RMRS-P-11. U.S. Department of Agriculture,
Genus Cercocarpus; 1988 July 13-14; Logan, Forest Service, Rocky Mountain Research
UT. College of Natural Resources, Utah State Station, Ogden, UT. p. 32-34.
University, Logan, UT. p. 27-42.
Waugh, W.J. 1990. Stagnation and decadence of
Kitchen, S.G., S.E. Meyer, G.R. Wilson, and R. Cercocarpus montanus in a southeastern
Stevens. 1989. Addition of Cercocarpus Wyoming big game exclosure. In: K.L. Johnson,
montanus—true mountain-mahogany to the ed. Proceedings of the Fifth Utah Shrub Ecology
Rules. Association of Official Seed Analysts Workshop: The Genus Cercocarpus; 1988 July
Newsletter 63: 28-30. 13-14; Logan, UT. College of Natural
Resources, Utah State University, Logan, UT. p.
Marshall, K.A. 1995. Cercocarpus montanus. In: 89-95.
U.S. Department of Agriculture, Forest Service,
Rocky Mountain Research Station, Fire Sciences Welsh, S.L., N.D. Atwood, L.C. Higgins, and S.
Laboratory, Fire Effects Information System, Goodrich. 1987. A Utah Flora. Brigham Young
http://www.fs.fed.us/database/feis/ 17 p. University Print Services, Provo, UT. 894 p.
180
Cereus greggii Engelmann night-blooming cereus
CACTACEAE
Synonyms: Peniocereus greggii (Engelm.) Britt. & Rose

Cereus greggii Engelmann var. roseiflorus Kuntze
Cereus pottsii Salm-Dyck
typically less than 1.5 m. The gray-brown stems
are four to five ribbed and approximately 1.25 cm
in diameter. There are 11 to 13 dark spines that
tend to be swollen at the base, per areole. The
lower spines may have some white coloration. The
flower is particularly remarkable, having waxy
white, pointed “petals” with numerous white to
yellow-tipped stamens. It is fragrant and up to 5
cm in diameter and 15 cm long. The orange-red,
oblong fruit is up to 7.6 cm long and 3.8 cm in
diameter with short spines. The root is a fleshy
taproot and typically weighs 2.3 to 6.8 kg
(Weniger 1991, Earle 1963, Vines 1960). In old
plants it may be up to 0.6 m in diameter and weigh
125 lbs (Weniger 1991). In 1919 a specimen with
purplish flowers was collected in Organ, New
Mexico, but since then only plants with white
flowers have been recorded (Weniger 1969). There
are two varieties of night-blooming cereus, Cereus
greggii var. greggii and Cereus greggii var.
transmontanus. Variety transmontanus differs
from the typical variety in that the flowers are
larger, approximately 7.5 cm in diameter, and the
hypanthium is particularly spiny, and the spines
generally 1.5 to 3.0 cm long (Benson 1982, Martin
and Hutchins 1980). The hypanthium is the cup-
like enlargement of the floral axis below the calyx
Illustration credit: Tonia Masaood that surrounds the ovary. The hypanthium of
variety greggii is covered by minute,
General Description.—The common names for inconspicuous spines and the flowers are smaller.
Cereus greggii are night-blooming cereus, Texas These varieties are in need of further study
night-blooming cereus, Arizona queen-of-the- (NMNPPAC 1984). Some sources (Natural
night, reina-de-la-noche, sweet-potato cactus, and Resources Conservation Service 2003) maintain
deer horn cactus. It is a perennial, succulent shrub that the synonym, Peniocereus greggii (Engelm.)
(Vines 1960). Night-blooming cereus is a Federal Britt. & Rose is the correct name. Night-blooming
Species of Concern and is listed as endangered by cereus provides an interesting evolutionary link. It
the State of New Mexico (Sivinski and Lightfoot appears to possess anatomical characteristics that
1995, NMRPTC 2002). It is threatened by illegal are relictual from its nonsucculent, woody,
collection throughout its range. Although this plant Pereskia-like ancestor (Mauseth and others 1998).
is cryptic and may be more common than verified
documentation suggests, many occurrences of Range.—Night-blooming cereus grows in
night-blooming cereus are known to have been southern New Mexico, southeastern Arizona,
extirpated due to collection pressures, and its western Texas, eastern Chihuahua, northeastern
range-wide abundance is believed to be declining. Durango, northern Zacatecas, and Coahuila,
Night-blooming cereus has erect or sprawling Mexico (Kearney and others 1960, Powell 1998,
stems that may grow to 2.4 m in length but are
181
Vines 1960). Benson (1982) defined the range of Growth and Management.—The stems generally
variety transmontanus as the Arizona and Sonoran grow up through, and later become supported by,
Desert that possibly also includes Hildalgo County shrubs. It is likely that the shrubs provide
in New Mexico, and that of variety greggii as the protection and a suitable microclimate for
Chihuahuan Desert of New Mexico, western Texas germination and subsequent plant development.
and Mexico. Any activity that reduces shrub cover is a threat to
night-blooming cereus. Therefore, fire is a
Ecology.—Night-blooming Cereus grows in dry potential hazard, although the direct effect of fire
alluvial soils at elevations between 370 and 1,500 on night-blooming cereus is not well documented.
m. Again, the varieties are distinguished by variety The substantial tuberous root may provide the
transmontanus growing at the lower end of the plant the means to recover if the above-ground
elevation range (300 to 1,050 m) and variety stems are killed, and therefore, it is likely that
greggii growing above 1200 m (Benson 1982). individual plants can tolerate light fires that do not
Night-blooming cereus generally grows in slightly decimate the shrubs with which they are associated
broken to level terrain in desert grassland or (Thomas 1991). More severe fires that scorch and
Chihuahuan desert scrub. Typically it grows in heat the soil are likely to be more detrimental.
sandy to silty gravelly soils on upper to mid Herbivores such as white-throated woodrats
bajadas among, depending upon the desert region, (packrats) and cactus borers (Cactobrosis
creosotebush (Larrea tridentata), mesquite fernaldialis) eat the stems, but new stems soon
(Prosopsis glandulosa), paloverde (Cercidium sprout from the tuberous root (ASDM 2002).
species), and knife-leaf condalia (Condalia Many populations throughout its range have been
species). It is also commonly associated with extirpated due to private and commercial
Ocotillo (Fouquieria splendens), tarbush collection, and the time required for population re-
(Flourensia cernua) and dropseed (Sporobolus establishment is not known. However, although
species). Habitat alteration such as that caused by slow growing, it is easily propagated by short stem
the high impact trampling of heavy grazing is cuttings and from seed. Seeds of many cacti
suspected to reduce population size (NMNPPAC species, including Cereus species, are subject to
1984). physiological dormancy. For example, seeds of
Cereus griseus were dormant at maturity but
Reproduction.—As the common name suggests, germinated after 8 weeks in a dry environment
night-blooming cereus blooms only at night. Just (Baskin and Baskin 2001). Cereus griseus seeds
after dark the flowers open in a series of “jerks” have an absolute light requirement for germination
(Benson 1982). Individual flowers only bloom for (Baskin and Baskin 2001). If the plant is dug up
one night. The total flowering period of plants for transplantation it is actually difficult to keep
within a population is also brief and generally all alive (Weniger 1969). Apparently the root is
flowers bloom within a 2-day period. However, in susceptible to fungus. Damp soil has to be
unusual circumstances, flowers on different avoided, but even if root rot is prevented, the plant
individuals within a population may bloom tends to decline over a few years if kept in a pot or
periodically over a period of almost one month. in a greenhouse (Weniger 1969).
Flowering occurs in late May into July depending
on geographic area (Earle 1963, Epple 1995, Benefits.—Night-blooming Cereus plants have
Kearney and others 1960). The fragrance of the commercial value especially among cactus
flowers is pleasant, and humans can detect the collectors (Epple 1995). Not only is it an
perfume of the flowers up to 30 m away from the aesthetically desirable cactus, but it has
plant (Benson 1982, Epple 1995, Vines 1960). considerable medicinal value. The root is generally
Flowers are pollinated principally by hawk moths believed to be the organ that has medicinal
(ASDM 2002) and possibly other night feeding properties (Moerman 1998, Moore 1989, Powell
insects (Epple 1995) that search for nectar. The 1998). However, the stems also have the same
flowers are not self-fertile and must be cross- properties although “more feebly” (Moerman
pollinated. Therefore the hawk moths (Sphingidae) 1998, Moore 1989, Powell 1998). Active
must fly hundreds of yards between the sparsely ingredients include penicerol, viperidone, desoxy
distributed plants (ASDM 2002). In areas where viperidone, viperidinone, β-sitosterol and, most
pesticides are heavily used for agriculture, the likely, caffeine (Moore 1989). It is likely that
hawkmoth populations are devastated, and most of night-blooming cereus is not unique in its
the flowers on plants in adjacent natural habitat medicinal value as other species of Cereus contain
fail to fruit (ASDM 2002). similar active ingredients (Ecdybase 2002). The
182
commercial toll on this species has not been Mauseth, J.D., T. Terrazas, and S. Loza-Cornejo.
established, but wild-harvested material is 1998. Anatomy of relictual members of
available on the Internet (Pacific West Botanicals subfamily Cactoideae, IOS Group 1a
2002). It is called “pain in the heart” by the Death (Cactaceae). Bradleya 16: 31-43.
Valley Shoshones (Moore 1989). This tribe
presumably uses it in a manner similar to Native Moerman, D.E. 1998. Native American ethno-
Americans of Nevada who ingest an infusion of botany. Timber Press, Portland, OR. 927 p.
the roots as a cardiac stimulant (Moerman 1998).
Other Native Americans have used a decoction of Moore, M. 1989. Medicinal plants of the desert
the roots for diabetes, the seedpods mixed with and canyon west. Museum of New Mexico
deer fat as a salve for sores, and the cut slices of Press, Santa Fe, NM. 184 p.
root as an externally applied cure for chest colds
(Moerman 1998, Vines 1960). The fruits, flowers, Natural Resources Conservation Service. 2003.
young stalks, and roots have been eaten for food Plants profile: Peniocereus greggii (Engelm.)
(Moerman 1998). This use of the root may account Britt. & Rose. http://plants.usda.gov/ [not
for the common name “sweet potato cactus.” paged].
Chewing the raw root has been reported to quench
thirst. Birds eat the seed and contribute to seed New Mexico Native Plants Protection Advisory
dispersal (Vines 1960). Committee. 1984. A handbook of rare and
endemic plants of New Mexico. University of
References New Mexico Press, Albuquerque, NM. 291p.
Arizona Sonora Desert Museum. 2002. New Mexico Rare Plant Technical Council. 2002.
Peniocereus greggii. Internet web site: NM Rare Plants Web Page: http://nmrareplants.
http://www.desert unm.edu/reports/ pengre.htm.
museum.org/books/nhsd_peniocereus.html#top.
Pacific West Botanicals. 2002. Internet web site:
Baskin, C.C. and J.M. Baskin. 2001. Seeds, http://www.pacificwestbotanicals.com/
ecology, biogeography, and evolution of Night_Blooming_Cereus.html.
dormancy and germination. Academic Press,
New York. 666 p. Powell, A. M. 1998. Trees & shrubs of Trans-
Pecos and adjacent areas. University of Texas
Benson, L. 1982. The cacti of the United States University Press, Austin, TX. 498 p.
and Canada. Stanford University Press,
Stanford, CA. p. 588-593. Sivinski, R. and K. Lightfoot 1995. Inventory of
rare and endangered plants of New Mexico.
Earle, W.H. 1963. Cacti of the Southwest. Desert Miscellaneous Publication 4. New Mexico
Botanical Garden, Phoenix, AZ. 210 p. Forestry and Resources Conservation Division,
Energy, Mineral and Natural Resources
Ecdybase. 2002. Internet Web Page: http:// Department, Santa Fe, NM.
ecdybase.org/index.php?row=308&action=
browse. Thomas, P.A. 1991. Response of succulents to
fire: A review. International Journal of Wildland
Epple, A.O. 1995. A field guide to the plants of Fire 1(1): 11-22.
Arizona. Falcon Press Publishing Co., Helena,
MT. 347 p. Vines, R.A. 1960. Trees, shrubs, and woody vines
of the Southwest. Sixth printing 1986.
Kearney, T.H., R.H. Peebles, J.T. Howell, and E. University of Texas Press. Austin, TX. 1104 p.
McClintock. 1960. Arizona flora. 2d ed.
University of California Press. Berkeley, CA. Weniger, D. 1969. Cacti of the Southwest.
1,085 p. University of Texas Press, Austin, TX. 249 p.
Martin, W.C. and C.R. Hutchins. 1980. A Flora of Weniger, D. 1991. Cacti of Texas and neighboring
New Mexico. Strauss & Cramer, Hirschberg, states. University of Texas Press, Austin, TX.
Germany. 2,592 p. 356 p.
183
Cestrum diurnum L. day jasmine
SOLANACEAE
Synonyms: Cestrum fastigiatum Jacq.

Cestrum diurnum L. var. fastigiatum (Jacq.) Stehlé in Fournet
Cestrum diurnum portoricense O.E. Schulz in Urban
John K. Francis
from cultivation in at least Florida, southern Texas,
Hawaii, Puerto Rico, Guam, and American Samoa
(Florida Exotic Plants Council 2001, Institute of
Pacific Islands Forestry 2001).
Ecology.—Day jasmine grows in gardens,

roadsides, fencerows, pastures, vacant lots, and
abandoned farmland in Puerto Rico (Liogier 1995).
The species is particularly associated with pasture
fencerows because its seeds are deposited there by
birds, vegetative competition is controlled by
grazing, and day jasmine is rarely eaten by cattle.
There is no mention in the literature of the types of
natural forest stands where day jasmine grows
within its native range. However, because the
species is intolerant of heavy shade and quickly
disappears when overtopped by forest, it can be
inferred that day jasmine is an opportunist that
invades disturbed areas and quickly completes its
life cycle ahead of encroaching forest. The species
may grow as individual plants or in thickets. Day
jasmine will grow in areas receiving from about
General Description.—Day jasmine is a single or 800 to 3000 mm of precipitation, but most
multistemmed shrub or rarely a small tree that is aggressively colonizes areas that receive from
also known as day cestrum, wild jasmine, ink-bush, about 1400 to 2400 mm of precipitation. It tolerates
Chinese inkberry (English), dama de día, rufiana, soils of all textures, apparently from all parent
galán de día, and saúco tintóreo (Spanish) (Little materials, but is most common in limestone areas
and others 1974). The bark of day jasmine is nearly (Florida Exotic Plants Council 2001). Day jasmine
smooth and gray. The inner bark is yellow-brown does not prosper on exposed subsoil nor grow on
and slightly bitter. The twigs are slender, greenish swampy ground. The plant is salt tolerant if not
gray, and sometimes drooping (Liogier 1995, Little exposed to heavy salt spray or overwash from
and others 1974). The leaves of day jasmine are storms (Florida Exotic Plants Council 2001).
oblong or oblong-elliptic, and shiny green to
yellow green, and somewhat membranous. They Reproduction.—Day jasmine flowers and fruits
are about 15 by 6.5 cm and have petioles about 2.5 year-round (Florida Exotic Plants Council 2001).
cm long. The small, white flowers are grouped in The fruits from a Puerto Rican collection were
panicles attached at the bases of the last leaves in sweet and slightly bitter with little other flavor and
each branch. These develop into elliptical, weighed (fresh) an average of 0.208 + 0.010
purplish-black berries about 6.5 mm long (Howard g/fruit. The seeds weighed an average of 0.0021+
1989, Liogier 1995, Little and others 1974). 0.0005 g/seed (air dry). The fruits are much more
variable in size (CV = 49) than the seeds (CV =
Range.—Day jasmine is native to the Bahamas, 25). There are from four to 14 seeds/fruit (Florida
Cuba, Jamaica, Hispaniola, and the Cayman Exotic Plants Council 2001). Day jasmine seeds
Islands (Little and others 1974). It has been are dispersed by frugiverous birds (Neal 1965,
introduced as an ornamental into most of tropical Little and others 1974). Seventy-nine percent of the
and subtropical America. The species has escaped seeds from the Puerto Rican collection cited above
184
germinated between 7 and 30 days after sowing. Calcinogenic glycosides. http://www.ansci.
edu/plants/toxicagents/calglyco.htm. 2 p.
Growth and Management.—Day jasmine rarely
reaches more than 4.5 m in height and 8 cm in Florida Exotic Plants Council. 2001. Cestrum
trunk diameter (Little and others 1974). Early diurnum L., Solanaceae/nightshade family.
growth is relatively fast and life spans are www.fleppc.org/pdf/Cestrum%20diurnum.pdf.
relatively short (5 to 20 years). Although seeds are 2 p.
still sold by regional seed houses and ornamental
seedlings are offered by nurseries, nothing is Howard, R.A. 1989. Flora of the Lesser Antilles,
published about propagation and management. Leeward and Windward Islands. Dicotyledoneae.
With a plant so aggressive, few nursery and Part 3. Vol. 6. Arnold Arboretum, Harvard
planting problems seem likely. Because the species University, Jamaica Plain, MA. 658 p.
is a weed in many situations, control may be
desirable. In the absence of control guidelines, Institute of Pacific Islands Forestry. 2001. Plant
cutting followed by herbicide treatment of the threats to Pacific Ecosystems: Cestrum diurnum
sprouts is suggested. L., Solanaceae. www.hear.org/pier/cediu.htm.
2 p.
Benefits.—Day jasmine has long been planted as
an ornamental for its pleasing appearance, Island Resource Foundation. 2001. Threatened and
moderate size, ease of establishment, and fragrant endangered birds of the insular Caribbean: plain
flowers. The plant has the drawback of being pigeon. www.irf.org/binornat.htm. 14 p.
poisonous to livestock. The leaves contain a
calcinogenic glycoside called 1,25- Liogier, H.A. 1995. Descriptive flora of Puerto
dihydroxycholecalciferol that leads to a vitamin D Rico and adjacent islands. Vol. 4. Editorial de
toxicity that results in elevated serum calcium and la Universidad de Puerto Rico, San Juan, PR.
deposition of calcium in soft tissues. Fifteen to 30 617 p.
percent of day jasmine leaves in an animal’s diet is
sufficient to cause symptoms (Animal Science at Little, E.L., Jr., R.O. Woodbury, and F.H.
Cornell University 2001). However, the fruits of Wadsworth. 1974. Trees of Puerto Rico and the
day jasmine are one of the three foods that make up Virgin Islands. Vol. 2. Agriculture Handbook
the bulk of the diet of the endangered plain pigeon 449. U.S. Department of Agriculture,
(Columba inornata) in Puerto Rico (Island Washingtion, DC. 1,024 p.
Resource Foundation 2001).
Neal, M.C. 1965. In gardens of Hawaii. Special
References Publication 50. Bernice P. Bishop Museum
Press, Honolulu, HI. 924 p.
Animal Science at Cornell University. 2001.
185
Cestrum laurifolium L’Hér. dama de noche
SOLANACEAE
Synonyms: Cestrum citrifolium Ritz in Hoffm.

Cestrum alaternoides Poir. in Lam.
Cestrum lambertii Dunal in DC.
John K. Francis
one to three or more black seeds per fruit with an

ivory spot on the ventral side (author’s observation,
Howard 1989, Liogier 1995).
Range.—Dama de noche is native to Cuba,

Lesser Antilles, and Trinidad (Grisebach 1963,
Ecology.—Dama de noche is most common in

areas underlain by limestone rocks but also grows
in areas of igneous rocks. The soils are well
drained, often rocky with a wide range of textures.
In Puerto Rico, it may be found from a little above
sea level to about 500 m in elevation and in areas
that receive from about 900 to 2200 mm of annual
precipitation. Dama de noche has an intermediate
tolerance of shade. It usually grows in the
understory of medium- to low-density forest. The
species grows and fruits best when small openings
allow broken overhead sunlight to enter. Dama de
noche does not invade disturbed areas quickly and
so is mainly found in middle and late secondary
forests and remnant forests. It can be quite
common but does not form dense stands.
General Description.—Dama de noche, also
known as candlewood, torch, galán de monte, bois- Reproduction.—Dama de noche is a good fruit
flambeau, and citronnier, is an evergreen shrub and seed producer and may yield a few to 100 or
usually 1 to 2 m in height but sometimes reaching more fruits per season. Flowering and fruiting
4 m. Dama de noche is a Puerto Rican name seem to be synchronized, at least within a given
meaning lady of the night. It has one or a small area. Fruits were observed ripening in a forest in
number of stems arising from the root crown or the north of Puerto Rico during the dry season
low on the main stem. The plant tends to root (March). A collection of fruits from that area
shallowly with stout lateral roots that are hard and averaged 0.290 + 0.012 g/fruit. Air-dried seeds
woody with a corky or fleshy bark. The stems and separated from them averaged 0.0169 + 0.0004
branches are slender with gray bark and grow g/seed or 59,200 seeds/kg. Sown on commercial
vertically for 1 to 2 m and then curve horizontally. potting mix, 93 percent germinated between 14 and
The alternate, glabrous leaves are shiny green on 60 days after sowing. Seeds are apparently
the upper surface, narrowly elliptic to obovate, dispersed by birds, and seedlings are well scattered
with a short petiole and blades that are 4 to 15 cm and not abundant. Plants resprout when cut or
long. The inflorescences are short axillary racemes damaged.
with three to five yellow or greenish-yellow, five-
lobed, tubular flowers 10 to 20 mm long. The fruits Growth and Management.—Dama de noche
are purple to black, ovoid to subglobose berries, 6 appears to have a moderate growth rate, probably
to 10 mm long, with a persistent calyx. There are not exceeding 1 m of height growth per year. No
186
nursery, planting, or management experience has References
been published. Protection of forests from fires and
development is recommended to encourage the Grisebach, A.H.R. 1963. Flora of the British West
formation and development of natural stands. Indies. J. Cramer-Weinheim, New York. 789 p.
Benefits.—Dama de noche contributes to the Howard, R. A. 1989. Flora of the Lesser Antilles,
biodiversity of the forests where it grows, helps Leeward and Windward Islands. Vol. 6. Arnold
protect the soil, and furnishes food and cover for Arboretum, Harvard University, Jamaica Plain,
wildlife. The stems are generally too small to use MA. 658 p.
for fuel.
617 p.
187
Chamaebatiaria millefolium (Torr.) Maxim. fern bush
ROSACEAE
Synonyms: Spiraea millefolium Torr.

Sorbaria millefolium Focke
Basilima millefolium Greene
Chamaebatiaria glutinosa Rydb.
Spiraea glutinosa Fedde
Nancy L. Shaw
Hitchcock and others 1961, Welsh and others
1987). Chromosome number is x = n = 9
(McArthur and Sanderson 1985). Variability
within the species appears low and hybridization
with other species has not been reported.
Range.—Fern bush is endemic to the Great Basin,

Colorado Plateau, and surrounding areas. It occurs
east of the Cascade and Sierra Nevada Mountains
from central Oregon eastward into Wyoming and
south to southeastern California, Nevada, Utah,
northern Arizona, and northern New Mexico
(Hitchcock and others 1961, Phillips 1949, Welsh
and others 1987).
Ecology.—Fern bush grows in fissures in rock

outcrops and cliffs and on well-drained soils of
dry, rocky, gravelly canyons and mountain slopes
at elevations ranging from 900 to 3,400 m
(Hickman 1993). It is often an early successional
Drawing source: Hitchcock and others 1961 species on cinder cones and basalt lava flows, but
it also grows on soils derived from limestone and
General Description.—Fern bush or desert sweet, granite (Eggler 1941, Everett 1957, Merkle 1952).
the single species in its genus, is an upright, Fern bush occurs in isolated populations or as an
multistemmed, sweetly aromatic shrub 0.3 to 2 m associated species in Artemisia spp., Pinus spp.-
tall. Bark of young branches is brown, becoming Juniperus spp., northern Juniperus, mountain
smooth and gray with age. Leaves are leathery, brush, Populus tremuloides Michx., Pinus flexilis
alternate, simple, twice pinnately divided, stipulate, James, P. ponderosa Dougl., Picea spp. A.--Abies
and clustered near the branch tips. Foliage and spp., Pinus longaveva D. Bailey, and chaparrel
young branches are viscid and pubescent with communities (Hickman 1993, Welsh and others
simple and stellate, pointed and glandular-capitate 1987).
hairs. Southern populations are evergreen (Phillips
1949), while northern populations are largely Reproduction.—Plants flower from June to
deciduous, retaining a few leaves though winter September (Hitchcock and others 1961, Phillips
and initiating new leaf growth in early spring. The 1949) with irrigation prolonging the flowering
profuse, terminal, leafy-bracteate panicles are 5 to season (Shaw and Hurd 2001). Seeds ripen in
20 cm long and produce abundant flowers that are August to October. The follicles are hand
showy, creamy white, and insect-pollinated. collected. They open as they dry, and seeds are
Flowers are complete, regular, and five-merous. extracted by screening. Shaw and Hurd (2001)
Follicles are pubescent and coriaceous. Seeds are found an average of about 3.7 million seeds/kg for
yellowish to brownish and linear to narrowly two seed lots grown under irrigation. Fresh seeds
fusiform. Germination is epigeal (Hickmann 1993, are nondormant while stored seeds require a 1- to
188
3-month prechill (Phillips 1949, Shaw and Hurd in Cormophyta 15. New observations on
2001). The optimum temperature for germination cyanogenesis in Rosaceae. Planta Medica 41:
of southwestern populations is 18 to 26 °C 313-327.
(Phillips 1949).
Hawksworth, F.G. and R.L. Mathiasen. 1978.
Growth and Management.—Seedlings establish Hosts of juniper mistletoe at Walnut Canyon
from shallow seeding on well-drained soils where National Monument, Arizona, USA. Great Basin
vegetative competition is limited (Shaw and Hurd Naturalist 38: 89.
2001). Fall seeding or artificial prechilling is
required for seeds of northern populations. Hickman, J.C., ed. 1993. The Jepson manual of
Seedlings develop rapidly if adequate water is higher plants of California. University of
available. Bareroot seedlings can be lifted California Press, Berkeley, CA. 1,400 p.
following one growing season. Irrigated plants
begin flowering during the second year (Shaw and Hitchcock, C.L., A. Cronquist, M. Ownbey, and
Hurd 2001). Unlike its namesake genus, J.W. Thompson. 1961. Vascular plants of the
Chamaebatia Benth., fern bush is not nodulated by Pacific Northwest. Part 3: Saxifragaceae to
nitrogen-fixing actinomycetes (McArthur and Ericaceae. University of Washington Press,
Sanderson 1985). Fern bush is a rare host of Seattle, WA. 614 p.
Phoradendron juniperinum A. Gray, the juniper
mistletoe (Hawksworth and Mathiasen 1978). McArthur, E.D. and S.C. Sanderson 1985. A
cytotaxonomic contribution to the western North
Benefits.—Fern bush provides cover for many American rosaceous flora. Madroño 32: 24-28.
organisms. It is sometimes browsed by mule deer
(Odocoilus hemionus Rafinesque), sheep (Ovis Merkle, J. 1952. An analysis of a pinyon-juniper
spp.), and goats (Capra hircus), but it receives community at Grand Canyon, Arizona. Ecology
little use by cattle (Bos spp.). First cultivated in 33:385-384.
1878 (Rehder 1940), fern bush is a valued
ornamental due to its showy flowers, long Mozingo, H.N. 1987. Shrubs of the Great Basin.
flowering period, and fernlike foliage (Hitchcock University of Nevada Press, Reno, NV. 342 p.
and others 1961, Phillips 1949). Mass plantings,
xeriscapes, screens, hedges, and specimen plants Phillips, J. 1949. Southwestern landscaping with
thrive in full sun (Phillips 1949). Native native plants. Museum of New Mexico Press,
Americans used a tea made from its leaves for Santa Fe, NM. 140 p.
treatment of stomachaches (Mozingo 1987). Plants
produce cyanic acids in their tissue (Fikensher and Rehder, A. 1940. Manual of cultivated trees and
others 1981). shrubs. Macmillan Publishing Company, Inc.,
New York. 996 p.
References
Shaw, N.L. and E.G. Hurd. 2001 Chamaebatiaria
Eggler, W.A. 1941. Primary succession on millefolium (Porter) Maxim. fern bush, desert
volcanic deposits in southern Idaho. Ecological sweet. In: F.T. Bonner and R.G. Nisley, eds.
Monographs 3: 277-298. Woody plant seed manual. Agriculture
Handbook. U.S. Department of Agriculture,
Everett, P.C. 1957. A summary of the culture of Forest Service, Washington, D.C.
California plants at the Rancho Santa Ana http://wpsm.net/index.html. 6 p.
Botanic Garden. Rancho Santa Ana Botanic
Garden, Claremont, CA. 233 p. Welsh, S.L., N.D. Atwood, L.C. Higgins, and S.
Goodrich, eds. 1987. A Utah flora. Great Basin
Fikenscher, L.H., R. Hegnauer, and H.W.L. Naturalist Memoirs 9. Brigham Young
Ruijgrok. 1981. Distribution of hydrocyanic acid University, Provo, UT. 894 p.
189
Chenopodium oahuense (Meyen) Aellen. 'aheahea
CHENOPODIACEAE
Synonyms: Atriplex oahuensis Meyen,

Chenopodium oahuense var discosperma Fosb.,
Chenopodium pekeloi Degener, I. Degener & Aellen,
Chenopodium sandwicheum Moq.
Randy S. Senock and Sarah A. Taylor
Ecology.―‘Aheahea is dispersed across the full

range of dryland habitats in Hawaii, from coastal
to subalpine. It appears primarily as a shrub but
has been found on Mauna Kea on the island of
Hawaii 3.5 to 4.5 m tall (Lamb 1981). ‘Aheahea
behaves as a colonizer on old lava flows following
site disturbance. In many dry upper elevation
shrublands on the island of Hawaii, ‘aheahea
dominates woody plant density and may be found
sharing habitat with native naio, Myoporum
sandwicense Gray, and mamani, Sophora
chrysophylla (Salisb.) Seem, trees (Lamb 1981).
The US Army has documented 'aheahea as being
the most widespread woody plant species across
their 108,000 acre Pohakuloa Training Area.
General Description.―Chenopodium oahuense, Following wildfire in that same area, ‘aheahea
known most commonly as 'aheahea or 'aweoweo, reestablishment was delayed for approximately 1
is a nonscented to lightly scented shrub that year. Within 4 years, however, it had exceeded its
sometimes takes on the form of a small tree. Plants preburn density approximately 15-fold (Sherry and
may be erect, ascending or prostrate, branched, others 1999). 'Aheahea appears well adapted to
and total height 0.5 to 2 m. Older branches turn tolerate drought by dying-back during periods of
gray and woody with age, while younger tissue low moisture and then rapidly growing during
remains mealy pubescent, thickened and somewhat periods of higher available moisture.
fleshy. Leaves are rhombic to broadly deltate, 2 to
4.2 cm long, and 1.3 to 2.5 cm wide, with both Reproduction.―Seed production is extremely
surfaces pubescent, the upper surface less so and high for this species. The inflorescence is a large
greener. Leaves three-lobed, teeth rounded to terminal panicle composed of many small, dense
obtuse, base truncate to cuneate, petioles 1.3 to 2.5 glomerules of flowers. ‘Aheahea peak
cm long (Wagner and others 1990). The leaves are reproductive events appear correlated with
strongly scented when crushed (Lamb 1981). favorable environmental conditions such as
Flowers in small, dense clusters grouped into prolonged periods of available moisture. However,
terminal, almost leafless panicles. A calyx nearly 1 plants can be found in flower nearly any time of
mm long, entirely surrounding the fruit at year. Chromosome number 2n = 36 (Wagner and
maturity. Seeds are light to dark brown and may be others 1990).
horizontal or sometimes vertical, about 0.8 mm in
diameter, the surface covered with short, blunt, Growth and Management.―‘Aheahea grows
rounded projections (Wagner and others 1990). rapidly and remains fleshy during the first several
months of growth. In later stages, branches are
Range.―While the genus is pantropical in brittle and easily damaged. However larger, older
distribution, this species is indigenous to Hawaii. individuals become single-stemmed and develop
‘Aheahea occurs on Laysan, French Frigate thickened woody branches. ‘Aheahea has been
Shoals, Necker, Nihoa, Lisianski, and all of the used sparingly as an ornamental in native and
main Hawaiian Islands except Kaho‘olawe xeriscape gardens. The parent plant produces
(Wagner and others 1990). hundreds of seeds, which are easily propagated.
190
'Aheahea prefers partial to full sun with light
watering. The species may have potential in native References
habitat restoration as a nurse species or to enhance
Anonymous. 1996. Hawaiian use of native plants.
microclimatic understory conditions in favor of
http://www.k12.hi.us/~waianaeh.
woody plant establishment.
hawaiianstudies/nplants6.html
Benefits.―The leaves and plant tips are used for
Ching, F. 2002. Ulu Ka Hoi. http://kms.
greens, and may be eaten like spinach, wrapped in
kapalama.ksbe.edu/projects/yellow/default.
ti [Cordyline fruticosa (L.) A. Chev.] leaves and
html. [not paged].
cooked on hot coals. Hawaiians used the plant to
make a poultice for minor wounds during battle.
Kawaharada, D. 2002. Visiting Mokumanamana.
Leaves were pounded or crushed and applied like
http://explorers.bishopmuseum.org. [not paged].
rubbing alcohol, the healing effect due to the
presence of chlorophyll in the leaf (Ching 2002).
Kessing, J., L. Martin, and R.FL Mau. 1993.
Hawaiians also used the dark red bark pounded in
http://www.extento.hawaii.edu/kbase/crop/
conjunction with other ingredients to create a
Type/Nysius.htm
concoction that would enhance and beautify the
skin of a newborn baby when ingested by a
Lamb, S.H. 1981. Native trees and shrubs of the
pregnant or nursing mother (Anonymous 1996).
Hawaiian Islands. The Sunstone press, Santa Fe,
Two limbs of the shrub wood were connected to
New Mexico. 158 p.
create a "makua mano," or shark hook, and in this
fashion, 'aheahea was used to catch sharks (Ching
Sherry, K., J.M. Castillo, and R.B Shaw. 1999.
2002). The native shrub is also of great importance
Effects of wildfire on vegetation and rare plants
to a certain rare beetle, Rhyncogonus biformis, that
in arid montane shrublands. Pohakaloa training
lives on Necker Island. The beetle is nocturnal and
area, Hawaii. 1999 Hawaii Conservation
emerges at night where it feeds soley on the leaves
Conference, Honolulu, HI.
of the 'aheahea (Kawaharada 2002).
Wagner, W.L., D.R. Herbst, and S.H. Sohmer.
Detrimental Effects.―The flower heads are host
1990. Manual of the Flowering Plants of
plants for Nysius nemorivagus, a white Lygaeid
Hawaii. University of Hawaii Press, Honolulu,
bug that can become a nuisance pest for certain
HI. 1,854 p.
agricultural crops such as cabbage, cucumbers,
potatoes and all types of squash (Kessing and
others 1993).
191
Chiococca alba (L.) A.S. Hitchc. West Indian snow-berry
RUBIACEAE
Synonyms: Lonicera alba L.

Chiococca racemosa L.
John K. Francis
in Florida, the Bahamas, through the Greater and
Lesser Antilles, in Mexico, Central America, and
northern South America. The species is not known
to have naturalized outside its native range.
Ecology.—West Indian snow-berry grows in moist

and dry forests from near sea level to 700 m or
more in all types of topography. It will grow on
soils of all textures derived from both sedimentary
and igneous parent materials including ultramafic
rocks (serpentine). Very poorly drained and highly
saline conditions are not tolerated. The species may
be found in remnant forests in both natural
openings and the understory. It invades brushy
pastures and roadsides and appears early in the
reforestation process. It is intermediate in tolerance
to shade. West Indian snow-berry competes well
with weeds and brush, but does not prosper in
dense grass swards. The species does not appear to
be seriously affected by any insect or disease.
Reproduction.—West Indian snow-berry flowers

in Puerto Rico from June through October and
yields ripe fruits from June through December
General Description.—West Indian snow-berry, (Acevedo-Rodriguez and Woodbury 1985). The
is also known as David’s root, bejuco de berac, small white, cream, or yellow flowers are borne in
buenda, liane des sorciers and several other racemes that arise from the leaf axils. The globose
common names (Howard 1989). It is one of the but slightly flattened fruits are white and vary in
most common woody plants in Puerto Rican moist size (4 to 8 mm) and weight (by a factor of 3) in
forests. The species grows as a scrambling shrub the same inflorescence. The fruits are sweet with
or woody vine that often climbs taller vegetation little other flavor. The average fruit weight from a
and may reach 6 m of extension (Acevedo- Puerto Rico collection was 0.117 + 0.003 g. The
Rodríguez and Woodbury 1985). The slender, dark brown seeds, which averaged about two per
green stems and branches are four-angled to round fruit, weighed 0.004 + 0.000 g. Germination is
in cross section. Opposite branching occurs at the epigeal. For the above sample, germination was 48
nodes where one or two shoots develop. A vertical percent and occurred between 33 and 59 days after
shoot at each node commonly ascends from sowing. Stems root when they come in contact with
prostrate stems. The root system consists of the soil, which results in interconnected patches
shallow laterals and abundant fine roots. West and many independent plants. Asexual
Indian snow-berry has opposite, thin to coriaceous, reproduction by air layers and rooted cuttings
dark green leaves that are ovate or oblong and should be easy. Long distance dispersal appears to
pointed at the tip and pointed to rounded at the be principally done by birds.
base. The petioles are 2 to 10 mm long and tiny
lobbed stipules are present (Liogier 1997). Growth and Management.—West Indian snow-
berry plants under moderate canopy openings grow
Range.—West Indian snow-berry grows naturally up to 0.5 m per year in each of its many stems.
192
Plants develop slowly in the nursery, requiring 3 Technical Report SO-58. U.S. Department of
months to grow large enough to prick into pots and Agriculture, Forest Service, Southern Forest
about an additional year to reach outplanting size. Experiment Station, New Orleans, LA. 331 p.
Seedlings and sections of older plants can be dug
up in the forest and transplanted successfully Bailey, L.H. 1941. The standard cyclopedia of
(Workman 1980). Once established, West Indian horticulture. Vol. 1. The MacMillan Company,
snow-berry needs little maintenance. New York, NY. 1,200 p.
Benefits.—West Indian snow-berry is used to a Howard, R.A. 1989. Flora of the Lesser Antilles,
limited extent as an ornamental in naturalistic Leeward and Windward Islands. Dicotyledoneae.
landscaping (Workman 1980) and is sometimes Part 3. Vol. 6. Arnold Arboretum, Harvard
cultivated as a flowering vine in greenhouses University, Jamaica Plain, MA. 658 p.
(Bailey 1941). Being very common and
widespread, the species contributes to the Liogier, H.A. 1997. Descriptive flora of Puerto
biodiversity of many Neotropical forests. It is Rico and adjacent islands. Vol. 5. Editorial de la
grazed by cattle, goats and certainly wild ruminants Universidad de Puerto Rico, San Juan, PR.
and produces fruits that are eaten by birds and 436 p.
other animals. The roots of West Indian snow-
berry are used as a purgative, diuretic, vomitive, Liogier, H.A. 1990. Plantas medicinales de Puerto
and antidiarrhetic in herbal medicine (Liogier Rico y del Caribe. Iberoamericana de Ediciones,
1990). Inc., San Juan, PR. 566 p.
References Workman, R.W. 1980. Growing native. The
Acevedo-Rodriguez, P. and R.O. Woodbury. 1985. Sanibel, FL. 137 p.
Los bejucos de Puerto Rico. Vol. 1. General
193
Chromolaena geraniifolia (Urban) King & H.E. Robins. geranium-leaf eupatorium
ASTERACEAE
Synonyms: Osmia geraniifolia (Urban) Britt. & Wilson

Eupatorium geraniifolium Urban
John K. Francis
locally common in scattered populations. It may be

found on a wide variety of soils but most
frequently is found on well-drained, medium-
textured, often rocky soils that develop over
igneous, sedimentary, and metamorphic (including
ultramafic) rocks. The species is most common in
areas of volcanic rock. Geranium-leaf eupatorium
grows on sites that receive from about 1200 to
about 3000 mm of mean annual precipitation at
elevations of 400 to 1,000 m (Liogier 1997). The
species is moderately intolerant of shade; it grows
well in openings and under the canopy of low
basal-area forest. Geranium-leaf eupatorium occurs
in remnant and middle- to late-secondary forests,
old road cuts and fills, bluffs, and unstable slopes.
It grows as scattered individual plants and in small
clumps under favorable conditions.
Reproduction.—Geranium-leaf eupatorium was

observed in bloom in March by the author. It
General Description.—Geranium-leaf eupator- flowers and fruits abundantly. Seeds collected near
ium, a name assigned by the author in the absence Cayey, Puerto Rico, averaged 0.00033 g/seed or 31
of a local common name, is an evergreen shrub up million seeds/kg. These seeds were sown on moist
to 2 m in height and 1 cm in stem diameter. The filter paper and yielded 67 percent germination
plant usually has several stems arising from the between 7 and 24 days after sowing. The seeds are
root crown. The root system is composed of fibrous dispersed by the wind. Seedlings and successful
lateral and fine roots. Geranium-leaf eupatorium natural reproduction are not common.
produces relatively few branches. The foliage,
which has a chrysanthemum-like odor, tends to be Growth and Management.—Individual stems of
concentrated near the branch ends. The opposite geranium-leaf eupatorium live about 5 years. Plants
leaves, which have a short petiole, are densely live much longer because of sprouting that replaces
pubescent, 2 to 6 cm long by 1.5 to 5 cm broad, senescent stems. No management guidelines have
and deeply lobed or coarsely toothed. been published. Planting is probably the best way
Inflorescences are small terminal corymbs of of establishing new populations, and natural
oblong-cylindrical heads. The corolla of the small reproduction probably can be encouraged by
flowers is blue. The fruits are dark-colored creating openings with scattered patches of bare
achenes, 4 to 5 mm long with a pappus 4 mm long soil in forests near seed sources.
(author’s observation, Liogier 1997).
Benefits.—Geranium-leaf eupatorium is a pretty
Range.—Geranium-leaf eupatorium is endemic to plant and contributes to the aesthetic appeal of the
Puerto Rico. It occurs mainly in mountainous areas forest. It helps stabilize the soil and furnishes cover
of the central part of the island (Liogier 1997). It is for wildlife.
not known to have been planted or naturalized
elsewhere.
Ecology.—Geranium-leaf eupatorium can be
194
References

436 p.
195
Chromolaena odorata (L.) King & H.E. Robins. Christmas bush
ASTERACEAE
Synonyms: Eupatorium odoratum L.

Eupatorium conyzoides Vahl
Eupatorium brachiatum Sw. ex Wikstr.
Eupatorium atriplicifolium Vahl
Osmia odorata (L.) Schultz-Bip.
John K. Francis
Ecology.—Christmas bush grows from near sea

level to over 1,000 m in elevation (Binggeli 1999).
It thrives on all types of well-drained soil and can
grow on soils relatively low in fertility.
Disturbance is required before a site can be
colonized (Pacific Island Ecosystems at Risk
2001). Once established, Christmas bush competes
aggressively with herbs, grass, and shrubs in open
areas. In its native range, it is frequently seen on
roadsides, riverbanks, vacant lots, abandoned
farmland, and neglected pastures. Christmas bush
has found a particular niche in the slash-and-burn
General Description.—Christmas bush, also agriculture cycle. In Borneo, Christmas bush and
known as bitter bush, Siam weed, baby tea, other perennial grasses and shrubs invade within 3
cariaquillo, Santa María, and fleurit-Noël, is a years of abandonment and are gradually replaced
scrambling shrub (Howard 1989, Liogier 1997). It by trees (Ohtsuka 1999). The species is not shade-
may reach 1 m or more as a free standing shrub and tolerant and will not grow under a closed forest
4 m or more when climbing into trees or shrubs. stand. It is also intolerant of frost (Binggeli 1999)
Stems reach 2 cm in diameter. The plants are and is limited by drought (below about 900 mm of
maintained by a system of abundant, yellowish, mean annual precipitation).
fine lateral roots. Multiple sprouts arise from the
root crown and lower stems. The individual Reproduction.—Christmas bush blooms annually
branches are long with relatively few branches. and is an abundant producer of seeds. Flowering
Foliage occurs only on recent growth. The and fruiting begins after plants are 1 year old
opposite, three-nerved leaves are deltoid to ovate- (Binggeli 1999). The flowers are pollinated by
lanciolate, usually with a dentate margin and a long insects. The small fruits mature in about a month
pointed tip. The leaves are aromatic when crushed. (Binggeli 1999). One collection of seeds in Puerto
The inflorescences are corymbs of cylindrical Rico averaged 2,670,000 seeds/kg but did not
heads located on the terminals of lateral branches. germinate. A second collection averaged 1,560,000
There are 15 to 25 tubular florets per head, white, seeds/kg and gave 11 percent germination between
lavender, pink, or blue in color. The seeds are a 3 and 120 days after sowing. Germination is
brownish gray to black achene that is 4 mm long epigeal. The seeds are wind-dispersed, and
with a pale brown pappus 5 or 6 mm long (Howard transport by animals is possible because of small
1989, Liogier 1997). hooks on the seeds. In India, it was observed that
only about 1.4 percent of the first-year seedlings
Range.—Christmas bush is native from Florida survived into the second year (Binggeli 1999).
through the West Indies and from Texas through Stems root whenever they come in contact with the
Central and South America to Argentina (Howard ground.
1989, Liogier 1997). It has been accidentally or
deliberately introduced and has naturalized Growth and Management.—Individual stems last
throughout much of the tropics, including Guam about 2 years and die back to or near the base and
and Hawaii (Pacific Island Ecosystems at Risk are replaced by new sprouts. Plants easily survive
2001). cutting and fire. The best current control method is
196
mechanical or hand cutting followed by herbicide Eupatorium odoratum as a weed in teak
treatment. Partial control can be obtained through (Tectona grandis) forest. Lembaga Penelitian
the use of aggressive cover crops. Relatively good Hasil hutan 312. Lopora, Indonesia. 25 p.
biological control has been obtained with
Pareuchaetes pseudooinsulata Rego Barros Howard, R.A. 1989. Flora of the Lesser Antilles,
(Lepidoptera) in Guam and several other Pacific Leeward and Windward Islands. Vol. 6. Arnold
islands (Pacific Island Ecosystems at Risk 2001). Arboretum, Harvard University, Jamaica Plain,
MA. 658 p.
Detriments and Benefits.—Invasion of Christmas
bush has been disastrous by seriously suppressing Kaleta, D., S.R. Ghosh, and C.N. Saikia. 1999.
native species in disturbed forests and pastures in Medium density particle board from weeds.
the tropics outside its native range. The shrub is Journal of Scientific and Industrial Research
reported to be highly allelopathic to nearby 58(9): 705-710.
vegetation (Muniappan 1994), a fact that has been
demonstrated in controlled studies (Sahid and Liogier, H.A. 1990. Plantas medicinales de Puerto
Sugau 1993). Christmas bush reduces the diameter Rico y del Caribe. Iberoamericana de Ediciones,
growth of teak in infested plantations (Daryono and Inc., San Juan, PR. 566 p.
Hamzah 1979). It was thought to be useful in the
control of Imperata grass and for this reason was Liogier, H.A. 1997. Descriptive flora of Puerto
deliberately introduced into the Ivory Coast, but Rico and adjacent islands. Vol. 5. Editorial de la
the results were disappointing (Binggeli 1999). Universidad de Puerto Rico, San Juan, PR.
Because of the abundance of dead leaves and dry 436 p.
shoots, Christmas bush stands are a fire hazard
(Muniappan 1994). Cattle do not eat Christmas M’Boob, S.S. 1991. Preliminary results of a
bush; however, it is browsed by white-tailed deer survey and assessment of Chromolaena odorata
(Meyer and others 1984). In herbal medicine, leaf (Siam weed) in Africa. Biotropica Special Pub.
extracts with salt are used as a gargle for sore 44: 51-55.
throats and colds. It is also used to scent aromatic
baths (Liogier 1990). Extracts of Christmas bush Meyer, M.W., R.D. Brown, and M.W. Graham.
have been shown to inhibit or kill Neisseria 1984. Protein and energy content of white-tailed
gonorrhoeae (the organism that causes gonorrhoea) deer diets in the Texas coastal bend. Journal of
in vitro (Caceres and others 1995) and to accelerate Wildlife Management 48(2): 527-534.
blood clotting (Triratana and others 1991). A
satisfactory medium-density particleboard was Muniappan, R. 1994. Chromolaena odorata (L.)
prepared from Christmas bush stems (Kaleta and R.M. King and H. Robinson. In: R. Labrada,
others 1999). During fallows between cultivation, J.C. Caseley, and C. Parker, eds. Weed
Christmas bush adds copious amounts of organic management for developing countries. Plant
matter to the soil and may reduce the populations Production and Protection Paper 120. Food and
of nematodes (M’Boob, 1991). It is also useful as Agriculture Organization of the United Nations,
mulch for row crops (Swennen and Wilson 1984). Rome. 93-94.
References Ohtsuka, T. 1999. Early stages of secondary

succession on abandoned cropland in north-east
Binggeli, P. 1999. Chromolaena odorata (L.) King Borneo Island. Ecological Research 14(3): 281-
& Robinson (Asteraceae). http://members.tripod. 290.
co.uk/WoodyPlantEcology/docs/web-sp4.htm.
4 p. Pacific Island Ecosystems at Risk. 2001. Invasive
plant species: Chromolaena odorata (L.) King
Caceres, A., H. Menendez, E. Mendez, E. & Robinson, Asteraceae.
Cohobon, B.E. Samayoa, E. Jauregui, E. Peralta, http://www.hear.org/pier/chodo.htm. 3 p.
and G. Carrillo. 1995. Antigonorrheal activity of
plants used in Guatemala for the treatment of Sahid, I.B. and J.B. Sugau. 1993. Allelopathic
sexually transmitted diseases. Journal effects of lantana (Lantana camara) and siam
Ethnopharmacol 48(2): 85-88. weed (Chromolaena odorata) on selected crops.
Weed Science 41(2): 303-308.
Daryono, H. and Z. Hamzah. 1979. A study of
197
Swennen, R. and G.F. Wilson. 1984. In-situ mulch Triratana, T., R. Suwannuraks, and W.
production for plantain. Banana Newsletter 7: Naengchomnong. 1991. Effect of Eupatorium
20-22. odoratum on blood coagulation. Journal of
Medical Association of Tailand 74(5): 283-287.
198
Chrysobalanus icaco L. coco-plum
CHRYSOBALANACEAE
Synonyms: Chrysobalanus pellocarpus G.F.W. Meyer

C. icaco L. var. genuinus Stahlé & Quentin
C. icaco L. var. pillocarpa (G.F.W. Meyer) C. Martius
John K. Francis
disturbance and planting (Little and others 1974).

Chyrysobalanus orbicularis Schum., C. ellipticus
Soland. ex Sabine, and C. atacarensis A. Chev.
from Africa were all once considered to be
subspecies of coco-plum (Paradis 1987). Probably
all the African references to C. icacos are really
one of these species. There is disagreement among
taxonomists as to whether varieties exist among
coco-plumb (Howard 1988, Liogier 1985). At
most, variation among populations is minor.
Ecology.—Coco-plum is a coastal species. It

commonly grows as single plants or thickets on
dunes and rocky headlands. It may also be found
on shallow soils in moist areas up to a 450-m
elevation in Puerto Rico (Little and others 1974).
Although the species can survive a great deal of
stress from storms, salt spray, and flooding, it is
low in stature, relatively intolerant of shade, and
only persists where competing vegetation is short.
Scale insects and caterpillars sometimes damage
natural and ornamental plants in Florida (Vargas-
Simon and others 1997).
General Description.—Coco-plum, also known as Reproduction.—The flower clusters (cymes) are

icaco, icaque ponne, pork-fat-apple, zicate, and shorter than the leaves and borne at the bases of the
many other common names, is a medium-sized leaves near the ends of the branches. Several
coastal shrub or, rarely, a small tree. It generally greenish-white flowers less than 1 cm in diameter
has multiple brown or gray stems that are smooth compose the clusters. Coco-plum flowers and fruits
to scaly. The twigs are green and hairless when nearly throughout the year. The white to purple
young and turn reddish brown with raised lenticels. fruits are drupes that resemble plumbs. Their thin
The shrub’s habit may be creeping or erect. The flesh is spongy, whitish, and slightly sweet to
branches support many shiny, dark-green, leathery, almost tasteless when ripe. Immature fruits are
round, or elliptic leaves 3.8 to 8.2 cm in length and astringent (Little and others 1974). The fleshiness
2.5 to 5.7 cm in width. The under surfaces of the of the mesocarp of fruits varies considerably across
leaves are light green. The petiole is about 3 mm its range (Howard 1988). One large (2.5 m tall)
in length. The simple leaves are alternate in two coco-plum plant growing under a Pinus caribaea
rows, turned upward (Little and others 1974). Morelet plantation in Puerto Rico yielded 760 ripe
fruits in a single picking. Ripe fruits from another
Range.—Coco-plum is native to coastal areas of Puerto Rican collection weighed 4.36 + 1.17 g
southern Florida and the Bahamas through the (mean and standard deviation), and seeds averaged
Caribbean. It is also found along the coasts of 1,790/kg in one Puerto Rican collection. This seed
Mexico, through Central America and South lot gave 89 percent germination beginning 34 days
America, to Ecuador and northern Brazil. The after sowing sample (Francis and Rodríguez 1993).
range has been extended inland in those areas by No scarification or other seed treatment is needed.
199
Seed dispersion is presumed to be by gravity, hypoglycemic effects. Leandra 6-7: 7, 63-75.
water, birds, bats, domestic animals, and humans.
Natural reproduction may be sparse to abundant. Francis, J.K. and A. Rodríguez. 1993. Seeds of
Artificial reproduction is usually by seeds with Puerto Rican trees and shrubs: second
plants grown as potted seedlings. Apical and basal installment. Research Note SO-374. U.S.
semiwoody, leafy cuttings treated with hormones Department of Agriculture, Forest Service,
will root in 6 to 8 weeks in mist bed conditions Southern Forest Experiment Station. New
(Vargas-Simon and others 1997). The best Orleans, LA. 5 p.
treatment tested was Indol Acidic Acid (IAA) at
5,000 ppm applied to apical stem cuttings. Howard, R.A. 1988. Flora of Lesser Antilles. Vol.
Adventitious roots arose from the vascular 4. Arnold Arboretum, Harvard University,
cambium, and lateral roots arose from the Jamaica Plain, MA. 673 p.
paricycle.
Liogier, H.A. 1985. Discriptive flora of Puerto
Growth and Management.—Growth is slow in Rico and adjacent islands. Vol. 1. Editorial de la
the nursery and the field, so new plantings of coco- Universidad de Puerto Rico, San Juan, PR. 352
plum must be protected from both herbaceous and p.
woody competition.
Benefits.—The fruits are edible raw and can be Wadsworth. 1974. Trees of Puerto Rico and the
made into preserves. Coco-plum seeds, which have Virgin Islands. Vol. 2. Agriculture Handbook
a high oil content, are also edible. The wood is 449. U.S. Department of Agriculture,
light brown, hard, and heavy (specific gravity 0.8) Washington, DC. 1,024 p.
and is used for fuel and rustic construction (Little
and others 1974). Various parts of the plant have Paradis, G. 1987. Concerning the division of the
been used in folk medicine. The species is known genus Chrysobalanus into species and
to have hypoglycemic effects (Costa 1977). It is a subspecies in West Africa (Chrysobalanaceae).
honey plant and furnishes food for wildlife. Coco- A Sciences Naturelles 45(3,4): 246-254.
plum is used in the Eastern and Western
Hemespheres as an ornamental. Another important Vargas-Simon, G., G. Arellano-Ostoa, and E.
benefit from the species is for dune and soil García-Villanueva. 1997. Propagación por
stabilization. estacas con hojas de icaco (Chrysobalanus icaco
L.) y anatomía del enraizamiento. Proceedings of
References the International Society for Tropical
Horticulture 41: 264-269.
Costa, O. de A. 1977. Brazilian plants with
200
Chrysophyllum oliviforme L. satinleaf
SAPOTACEAE
Synonyms: Chrysophyllum microphyllum Jacq.

Chrysophyllum monopyrenum Sw.
Cynodendron oliviforme (L.) Baehni
John K. Francis
secretes a white milky sap when cut. The fruits are

sweet and edible. Satinleaf has 2n = 52
chromosomes (Gilman and Watson 1993, Liogier
1995, Little and Wadsworth 1964, Long and
Lakela 1976, Nelson 1996).
Range.—Satinleaf is native to Florida, the

Bahamas, the greater Antilles, and Belize (Liogier
1995). The species is planted in many tropical
countries as an ornamental and has become
naturalized in Hawaii and French Polynesia
(Pacific Island Ecosystems at Risk 2002). It has
been declared endangered in the wild in Florida
(Crowley 2002).
Ecology.—Satinleaf grows on a wide variety of

soils in all textures and in pH’s from about 5 to 8,
and has a moderate tolerance to salt (Institute of
Food and Agricultural Sciences 2002). It grows on
coastal sands and shallow clays over limestone in
Puerto Rico, in pinelands and hammocks of the
everglades and keys of Florida, and in low-
elevation moist, secondary forests and Eucalyptus
plantations in Florida (Little and Wadsworth 1964,
General Description.—Satinleaf, also known as Pacific Island Ecosystems at Risk 2002). Satinleaf
wild star-apple, damson plum, saffron-tree, occurs in remnant and middle secondary forests.
caimitillo, caimitillo de perro, camitillo cimarró, When young, it demonstrates an intermediate
teta de burra, macanabo, and caïmite marron, is a tolerance to shade and can grow and develop in the
beautiful, evergreen shrub or small tree usually 3 to understory of relatively low-density forests. To
5 m in height. Under favorable conditions, older bear fruit, it must have increased light, as in
individuals occasionally reach 10 m or more in intermediate and codominant crown positions.
height and 30 cm in diameter at breast height. The Satinleaf plants are severely damaged by
single or multiple stems have gray-brown bark with temperatures of 0 °C and below. They are resistant
fissures and plates. Twigs are slender and rusty or to drought and storm damage (Gilman and Watson
reddish-brown. The alternate leaves are elliptic, 1993).
ovate or oblong, 3 to 11 cm long and 2 to 5 cm
broad, with shiny, dark-green, hairless upper Reproduction.—Puerto Rican satinleaf plants
surfaces and coppery under-surfaces owing to a flower July to October and mature fruits in
dense covering of short hairs. The bell-shaped February (Little and Wadsworth 1964). They are
flowers are one to many on short pedicels in the reported to flower and fruit all year in Florida
leaf axils. The 3- to 5-mm, five-lobbed corolla is (Long and Lakela 1976). The flowers are insect
yellowish- or greenish-white. The fleshy fruits are pollinated. Seeds are dispersed by frugivorous
dark-purple, blue, or black, elliptical, one-seeded birds (Pacific Island Ecosystems at Risk 2002).
berries 1 to 2 cm long. They have whitish flesh Seedlings are relatively common in suitable habitat
surrounded by a gummy or rubbery skin that in Puerto Rico. Seed weights and treatment
201
information is not available. Misted hardwood capitata (Wiedemann). University of Florida.
cuttings and air layers, both treated with 0.3 http://creatures.ifas.ufl.edu/fruit/Mediterranean_
percent IBA (indol-butyric-acid), were tested as fruit-fly.htm. 19 p.
means of propagating the species vegetatively.
Thirty-five of 69 cuttings rooted in 7 months and Gilman, E.F. and D.G. Watson. 1993.
one of 39 air layers rooted in 6 months (author’s Chrysophyllum oliviforme, Satinleaf. Fact Sheet
observation). Ornamental production is by seed and ST-166. Environmental Horticulture
semihardwood cuttings under mist (Gilman and Department, Florida Cooperative Extension
Watson 1993). Service, University of Florida. 2 p.
Growth and Management.—Growth is relatively Institute of Food and Agricultural Sciences. 2002.
slow at all stages, and the plants are relatively Flowering and shade trees (A-E). University of
long-lived. Natural reproduction can probably be Florida Extension http://miami-dade.ifas.ufl.
encouraged by disturbance under low-density edu/programs/fyn/publications/dtplfloweringtree
secondary forests near seed sources. s-a-e.htm. 7 p.
Benefits.—Satinleaf contributes to the aesthetics of Liogier, H.A. 1995. Descriptive flora of Puerto
the forests where it grows, helps protect the soil, Rico and adjacent islands. Vol. 4. Editorial de la
and furnishes food and cover for wildlife. It was Universidad de Puerto Rico, San Juan, PR.
listed as an important source of nectar for 617 p.
honeybees in the Dominican Republic (Marcano
Fondeur 1973). The foliage is probably unpalatable Little, E.L., Jr. and F.L. Wadsworth. 1964.
to ungulates; at least the key deer (Odocoilleus Common trees of Puerto Rico and the Virgin
virginianus ssp. clavium) will not eat it (Schaus Islands. Agric. Handb. 249. U.S. Department of
and others 2002). The fruits are edible and Agriculture, Washington, DC. 548 p.
sometimes used to make jelly (Little and
Wadsworth 1964). In areas of infestation, satinleaf Long, R.W. and O. Lakela. 1976. A flora of
is usually heavily attacked by Mediterranean fruit Tropical Florida. Banyon Books, Miami, FL.
fly (Ceratitis capitata (Wiedemann) (Department 962 p.
of Entomology and Nematology 2002). The wood,
which has a specific gravity of 0.9, is hard, heavy, Marcano Fondeur, E. de J. 1973. La flora apícola
and strong. It is used for construction in Cuba de la República Dominicana. http.//
(Little and Wadsworth 1964). Satinleaf has been marcano.freeservers.com/nature/estudios/apicola
extensively, but not intensively, planted as an /dicotsp.htm. 11 p.
ornamental in Florida, Hawaii, and elsewhere. It
makes a pretty addition to natural landscaping, a Nelson, G. 1996. The shrubs and woody vines of
good foundation plant, and an attractive lawn, Florida. Pineapple Press, Inc., Sarasota, FL.
street, and parking lot tree. Pruning may be 391 p.
necessary to maintain a single stem and to remove
drooping branches (Gilman and Watson 1993). Pacific Island Ecosystems at Risk. 2002.
Chrysophyllum oliviforme L., Sapotaceae.
References http://www.hear.org/pier3/choli.htm. 2 p.
Crowley, M. 2002. Florida’s native plants— Schaus, C., S. Wade and J. Dunan. 2002. Key deer
protected. Native Florida Heritage Web Ring. and plants they won’t eat. Monroe County
http://nsis.org/garden/pr-plants-c.html. 5 p. Extension Service, University of Florida.
http://monroe.ifas.ufl.edu/key_deer_plants.htm.
Department of Entomology and Nematology. 4 p.
2002. Mediterranean fruit fly—Ceratitis
202
Chrysothamnus nauseosus (Pallas ex Pursh) Britt. rubber rabbitbrush
ASTERACEAE
Synonyms: Bigelovia nauseosa Gray

Chrysocoma nauseosa Pallas ex Pursh
Ericameria nauseosa (Pallas ex Pursh) Nesom & Baird

highly tomentose and number between 20 and 25.
Chrysothamnus nauseosus is uniformly diploid, x
= 9, 2n = 18 (Anderson 1966).
Taxonomy.—The International Plant Names

Index (IPNI) recognizes 30 subspecies of C.
nauseosus (IPNI. 2002). Some sytematists
promote the transfer of rubber rabbitbrush to the
genus Ericameria (Anderson 1995). Subspecies
may vary by morphology, color, and secondary
metabolic compounds. Each subspecies is likely
the result of reproductive isolation inbreeding,
although subspecific sympatry has been observed.
Some of the more common subspecies include ssp.
albicaulis, hololeucus, consimilis, graveolens, and
salicifolius (McArthur and others 1979). White
stem rubber rabbitbrush (C. n. ssp. albicaulis and
hololeucus) vary from other subspecies by their
dense white to grayish tomentum, the size and
arrangement of involucral bracts (five distinct
vertical ranks, 8 to 10 mm in length), and the
corolla lobe length (1 to 2 mm in ssp. albicaulis,
0.5 to 1 mm in ssp. hololeucus). These two
subspecies are segregated by elevation. Threadleaf
rubber rabbitbrush (C. n. consimilis) is a tall form,
up to 3.0 m, with green to yellow-green tomentum
and thread-like leaves. Ecologically, spp.
consimilis is more likely to be found in alkaline
valley bottoms. Green rubber rabbitbrush (C. n.
ssp. graveolens) has yellow-green to green stems
General Description.—Rubber rabbitbrush also and is less pubescent than other subspecies.
know as common rabbitbrush is a shrub typically Mountain rubber rabbitbrush (C. n. ssp.
0.3 to 2.0 m in height. Some types, however, are salicifolius) has larger leaves (4 to 8 cm long and 3
smaller while others exceed heights of 3.0 m. to 10 mm wide), less uniform involucaral bracts,
Usually several erect stems arise from the base and and is one of the more palatable subspecies to
branch to create a rounded form. Branches are livestock and big game.
covered with a felt-like tomentum, permeated by a
resinous gum, which can vary in color from green, Range.—Rubber rabbitbrush is common
yellow-green, gray-green, to white. Leaf shape can throughout the Intermountain West extending from
vary from filiform to broadly linear, and leaf British Colombia to Saskatchewan and south to
length can range from 18 to 63 mm. Like western Texas, Sonora, Baja California, and
branches, leaves are typically covered with eastern California (Anderson 1986). It inhabits
tomentum. The flower heads are generally plains, valley bottoms, and foothills and is often
arranged into a cymose infloresence. Involucral associated with sagebrush, pinyon-juniper, and
bracts in each head may vary from glabrous to ponderosa pine communities.
203
Ecology.—Rubber rabbitbrush establishes readily stable and disturbed plant communities. Its ability
on disturbed areas and can often become the to stabilize soil and reduce wind and water erosion
dominant vegetation on lands that have been preserves habitat for other shrubs, forbs, and
disturbed by fire, insects, vehicles, or heavy grasses. This shrub is an excellent plant for erosion
grazing. The species spans a broad elevation control because it has deep roots, produces heavy
gradient, extending from 150 to 2,700 m. with litter, and is able to establish on severe sites
annual precipitation generally in the 250 to 350 (Aldon and Pase 1981, McArthur and others 1995,
mm range. This shrub grows well in sandy, USDA 1937). Rubber rabbitbrush is browsed by
gravelly, or clay-alkaline soils, depending on the deer, pronghorn, and by domestic animals, but
subspecies or ecotype. Some subspecies have palatability varies greatly between subspecies and
broader ecological amplitudes than others, ecotypes. The white to grayish subspecies
allowing them to establish in a variety of habitats (albicaulis, hololoeucus, and salicifolius) are more
(McArthur and others 1979). Although it grows palatable than the green subspecies (gravelens and
vigorously, it does not exclude or inhibit consimilis) (Goodrich and others 1999 Hanks and
herbaceous species. On the contrary, production of others 1975). Unlike some forage shrubs, however,
herbaceous cover was greater where rubber rubber rabbitbrush is quite resilient to browsing.
rabbitbrush was present than when it was not In addition to food, C. nauseosus also provides
present (Frischknecht 1963). cover for small mammals, birds, and newly born
pronghorn (Yoakum 1986). In addition to its
Reproduction.—Rubber rabbitbrush blooms from ecological importance, people have also
August to October. Seeds mature beginning in recognized the chemical properties of C.
mid-October until late fall. In most conditions, nauseosus. Rubber rabbitbrush has been
rubber rabbitbrush is a heavy seed producer. Seed investigated as a possible source of rubber and
germination in rubber rabbitbrush is quite high, other chemicals (Hall and Goodspeed 1919;
and seed viability remains relatively high (65 Hegerhorst and others 1987). Some accessions, for
percent) for up to 3 year’s storage under ordinary example, produce up to 6 percent stem rubber and
warehouse storage conditions (Stevens and others 20 percent resin. Finally, some forms have
1981). When exposed to cool night (4.7 oC) and ornamental shrub potential (McArthur and others
warm day temperatures, seed germinates in about 1979, Weber and others 1985).
2 days (Weber and others 1985). Under
standardized single cool temperature conditions, References
however, seeds from lower elevations and latitudes
germinate more quickly (2 weeks as compared to Aldon, E.F. and C.P. Pase. 1981. Plant species
4) than those from higher elevations and latitudes adaptability on mine spoils, in the Southwest: a
(McArthur and others 1987, Meyer and others case study. Research Note RM-398. U.S.
1989). Seed (achene) size in rubber rabbitbrush Department of Agriculture, Forest Service,
appears to be correlated to habitat; subspecies Rocky Mountain Forest and Range Experiment
adapted to sandy sites have larger achenes (Meyer Station, Fort Collins, CO. 3 p.
1997). There are about 1,530 cleaned seeds per
gram (McArthur and others 1979). Anderson, L.C. 1966. Cytotaxonomic studies in
Chrysothamnus (Asterae, Compositae).
Growth and Management.—Once established, American Journal of Botany 53: 204-212.
wind-disseminated achenes allow rubber
rabbitbrush to spread easily and quickly (Meyer Anderson, L.C. 1986. Sympatric subspecies in
1997, Young and others 1984). It is fast growing, Chrysothamnus nauseosus. In: E. D. McArthur
reaching maturity in 2 to 4 years but relatively and B.L Welch, comps. Proceedings—
short-lived shrub, usually 5 to 20 years (personal symposium on the biology of Artemisia and
observation). It can be aerially seeded. Rubber Chrysothamnus; 1984 July 9-13, Provo, UT.
rabbitbrush can be troublesome on rangelands General Technical Report INT-200. U.S.
when it invades and occupies areas at high Department of Agriculture, Forest Service,
densities. It is a difficult to control because it is Intermountain Research Station, Ogden, UT: 98-
resistant to herbicides and has the ability to 103.
resprout from the crown (Cook and others 1965).
Anderson, L.C. 1995. The Chrysothamnus-
Benefits.—Rubber rabbitbrush can be an Ericameria connection (Asteraceae). Great
important structural and ecological component in Basin Naturalist 55: 84-88.
204
Cook, C.W., P.D. Leonard, and C.D. Bonham. McArthur, E.D., R. Stevens, and S.B. Monsen
1965. Rabbitbrush competition and control on 1995. Adaptation and success of big sagebrush
Utah rangelands. Bulletin 454. Utah and rubber rabbitbrush on disturbed sites. In:
Agricultural Experiment Station., Logan, UT. G.E. Schuman and G.F. Vance, eds. Decades
28 p. later: a time for reassessment, proceedings of the
12th Annual Meeting of the American Society
Frischknecht, N.C. 1963. Contrasting effects of big for Surface Mining and Reclamation; 1995 June
sagebrush and rubber rabbitbrush on production 3 -8, Gillette, WY. American Society for
of crested wheatgrass. Journal of Range Surface Mining and Reclamation, Princeton,
Management 16: 70-74. WV. p. 811-823.
Goodrich, S., E.D. McArthur, A. Huber, and J.E. Meyer, S.E. 1997. Ecological correlates of achene
Ott. 1999. Ungulate browsing of two mass variation in Chrysothamnus nauseosus
populations of rubber rabbitbrush. In: E.D. (Asteraceae). American Journal of Botany. 84:
McArthur, W.K. Ostler, and C.L. Wambolt, 471-477.
comps. Proceedings: shrubland ecotones; 1998
August 12-14; Ephraim, UT. Proc RMRS-P-11. Meyer, S.E., E.D. McArthur, and G.L. Jorgensen.
Ogden, UT: U.S. Department of Agriculture, 1989. Variation in germination response to
Forest Service, Rocky Mountain Research temperature in rubber rabbitbrush
Station: 196-199. (Chrysothamnus nauseosus:Asteraceae) and its
ecological implications. American Journal of
Hall, H.M. and T.H. Goodspeed. 1919. A rubber Botany 76: 981-991.
plant survey of Western North America. II.
Chrysil, a new rubber from Chrysothamnus Stevens, R., K.R. Jorgensen, and J.N. Davis. 1981.
nauseosus. University of California Publications Viability of seed from thirty-two shrub and forb
in Botany 7: 183-264. species through fifteen years of warehouse
storage. Great Basin Naturalist 41: 274-277.
Hanks, D.L., E.D. McArthur, A.P. Plummer, B.C.
Giunta, and A.C. Blauer. 1975. U.S. Department of Agriculture, Forest Service.
Chromatographic recognition of some palatable 1937. Range plant handbook. United States
and unpalatable subspecies of rubber rabbitbrush Government Printing Office, Washington, D.C.
in and around Utah. Journal of Range 816 p.
Management 28: 144-148.
Weber, D.J., T. Davis, E.D. McArthur, and N.
Hegerhorst, D., D.J. Weber, and E.D. McArthur. Sankhla. 1985. Chrysothamnus nauseosus
1987. Resin and rubber content in (rubber rabbitbrush): mutiple-use shrub of the
Chrysothamnus. Southwestern Naturalist 32: desert. Desert Plants 7: 172-209.
475-482.
Yoakum, J. 1986. Use of Aremisia and
International Plant Names Index. 2002. Chrysothamnus by pronghorns. In: McArthur, E.
International Plant Names Index. D. and B. L Welch, comps. Proceedings—
xhttp//www.ipni.org. [not paged]. symposium on the biology of Artemisia and
Chrysothamnus; 1984 July 9-13, Provo, UT.
McArthur, E.D., A.C. Blauer, A.P. Plummer, and General Technical Report INT-200. U.S.
R. Stevens 1979. Characteristics and Department of Agriculture, Forest Service,
hybridization of important intermountain shrubs. Intermountain Research Station, Ogden, UT.
III. Sunflower family. Research Paper INT-220. p. 176-180.
USDA Forest Service, Intermountain Forest and Young, J.A., R.A. Evans, and B.L. Kay. 1984.
Range Experiment Station, Ogden, UT. 82 p. Persistence and colonizing ability of rabbitbrush
collections in a common garden. Journal of
McArthur, E.D., S.E. Meyer, and D.J. Weber. Range Management 37: 373-377.
1987. Germination rate at low temperature:
rubber rabbitbrush population differences.
Journal of Range Management 40: 530-533.
205
Chrysothamnus parryi (Gray) Greene Parry rabbitbrush
ASTERACEAE
Synonyms: Bigelovia parryi Gray

Ericameria parryi (Gray) G.L. Nesom & G.I. Baird
Linosyris parryi Gray
to 11 mm long. Involucral bracts can vary in

length (9 to 14 mm) and shape (acuminate to very
attenuate) according to subspecies. Parry
rabbitbrush is uniformly diploid n = 9, 2x = 18
(Anderson 1966).
Taxonomy.—Parry rabbitbrush is a diverse group

with 12 subspecies (Anderson 1986, McArthur and
Meyer 1987). Subspecies vary one from another
by numerous characteristics including stature,
growth habit, leaf shape, leaf size, involucral bract
shape, number of flowers per head, pubescence,
and habitat. Neesom and Baird (1993) recently
suggested that Parry rabbitbrush would better be
placed in Ericameria based on molecular genetic
data. Anderson (1995) was not comfortable with
this change. Notable subspecies include asper,
attenuatus, howardii, monocephalus, nevadensis,
and parryi.
Subspecies asper is a low shrub 15 cm or
more high and inhabits mountainsides that border
the deserts in western Nevada and eastern
California (Anderson 1986, McArthur and others
1979). Leaves are covered with short-stalked resin
glands and flower heads contain five to ten disc
flowers.
Chrysothamnus parryi ssp. attenuatus has
small leaves up to 4 mm in length. Stems are
General Description.—Parry rabbitbrush is a mostly erect and shrubs can grow as high as 60
shrub, intermediate in height, growth habit, and cm. Flower heads contain five to seven disc
stem and leaf tomentum between rubber flowers and bracts are ranked into five vertical
rabbitbrush [Chrysothamnus nauseosus (Pallas ex rows. This subspecies is found in Utah,
Push) Britton] and low rabbitbrush southwestern Colorado, and northwestern New
(Chrysothamnus viscidiflorus Nutt.). It is usually Mexico (Anderson 1986, McArthur and others
short, being 20 to 60 cm tall, with numerous 1979).
spreading to erect flexible branches. Branches are Subspecies howardii is similar in size to
covered with a felt-like white to green tomentum. subspecies attenuatus. The uppermost leaves
Leaves are narrowly linear to elliptic. Leaf usually extend beyond the uppermost heads of the
dimensions range from 0.5 to 8 mm wide and 1 to inflorescence. Flower heads have five to seven
8 cm long. The leaf surface is somewhat viscid and pale yellow disc flowers and involucral bracts are
leaf pubescence can vary from glabrous to vertically ranked with spreading tips. This
tomentose. Flower heads are composed of four to subspecies is endemic to Utah, southern Wyoming,
twenty yellow disc flowers; heads are usually Colorado, New Mexico, and Nebraska (Anderson
arranged in terminal racemes that sometimes form 1986, McArthur and others 1979).
panicles. Corollas are tubular to funnelform and 8
206
In subspecies monocephalus, leaf shape common species, such as rubber and low
can vary from linear-oblanceolate to spatulate. rabbitbrushes.
Like subspecies howardii, the upper leaves usually
extend beyond the inflorescence. One unique Reproduction.—Parry rabbitbrush flowers from
characteristic is that flower heads occur singly or July to September and seed matures in October
in pairs rather than forming racemes. Flower heads and November. Achenes are 5 to 6 mm long and
contain five or six disc flowers subtended by covered with long, shaggy, oppressed hairs. We
obscurely ranked involucral bracts. This presume there are about 1,600 cleaned achenes per
subspecies occurs in high mountains between 790 gram and that they have good fertility as is the
and 3,400 m in eastern California and western case for other Chrysothamnus species
Nevada. (Deitschmann and others 1974). Achenes are wind
Subspecies nevadensis is comprised of disseminated in late fall or winter. In addition to
small shrubs up to 60 cm in height. Leaves are sexual reproduction, Parry rabbitbrush has been
linear to linear-oblanceolate and covered with observed to spread through underground roots
resin or gray tomentum. The uppermost leaves (Paulsen and Miller 1968, McArthur and others
rarely extend beyond the inflorescence. Flower 1979). We are unaware of seed germination data.
heads contain four to six yellow disk flowers and
are subtended by ranked involucral bracts with Growth and Management.—Seed germinates in
slender recurved tips. Subspecies nevadensis the early spring and seedlings become established
occurs on dry mountainsides from eastern during the following spring and summer. Plants
California to eastern Nevada, southwestern Utah, generally mature within 5 years and typically live
and Northern Arizona, being most common on the 15 to 20 years. Paulsen and Miller (1968) reported
eastern slopes of the Sierra Nevada (Anderson that, when necessary, Parry rabbitbrush can be
1986, McArthur and others 1979). controlled by the herbicide Tordon®.1
Subspecies C. parryi ssp. parryi consists
of low shrubs with erect branches and can be as Benefits.—Although Parry rabbitbrush is a
tall as 76 cm. The uppermost leaves usually extend component of western uplands, it is spotty in
beyond the inflorescence. Flower heads include 10 distribution and usually sparse in population
to 20 disc flowers and are subtended by obscurely density. It has some importance as a browse source
ranked bracts with attenuate tips. This subspecies in those areas where it is locally abundant. Some
grows on dry hillsides and plains in central subspecies of Parry rabbibrush, including C. parryi
Nevada, southern Utah, south-central Wyoming, ssp. asper and C. parryi ssp. monocephalus, are
western Colorado, and northern New Mexico particularly attractive plants and have horticultural
(Anderson 1986, McArthur and others 1979). potential as ornamentals.
Anderson (1970) made a case for this taxon being
the most primitive of all Chrysothamnus. References
Parry rabbitbrush in the form of C. parryi
ssp. attenuatus has been implicated in the Anderson, L.C. 1966. Cytotaxonomic studies in
parentage of the putative stabilized hybrid species Chrysothamnus (Asterae, Compositae). American
C. nauseosus ssp. uintahensis Anderson 1984). Journal of Botany 53: 204-212.
Range.—The range of Parry rabbitbrush extends Anderson, L.C. 1970. The karyotype of
from California in the west, to western Nebraska Chrysothamnus parryi ssp. parryi and its
in the east, and from Wyoming in the north, to implication. Transactions of the Kansas
New Mexico and Arizona in the south. Subspecies, Academy of Science 72: 399-401.
however, may be more geographically restricted.
Within its range, Parry rabbitbrush may be found Anderson, L.C. 1984. Chrysothamnus nauseosus
at elevations between 790 and 3,400 m. ssp. uintahensis: a stabilized hybrid. Great Basin
Naturalist 44:416-420.
Ecology.—Parry rabbitbrush grows on dry, open
foothills and mountains. Precipitation throughout Anderson, L.C. 1986. An overview of the genus
its range generally exceeds 380 mm. Parry Chrysothamnus. In: E.D. McArthur and B.L
rabbitbrush tends to increase on overgrazed or Welch, comps. Proceedings—symposium on the
otherwise disturbed areas. Populations are usually
smaller and more scattered than those of more 1
Use of trade names does not imply endorsement
of the U. S. Department of Agriculture
207
biology of Artemisia and Chrysothamnus; 1984 U.S. Department of Agriculture, Forest Service,
July 9-13, Provo, UT. General Technical Report Intermountain Forest and Range Experiment
INT-200. U.S. Department of Agriculture, Forest Station, Ogden, UT. 82 p.
Service, Intermountain Research Station, Ogden,
UT: 29-45. McArthur, E.D. and S.E. Meyer. 1987. A review
of the taxonomy and distribution of
Anderson, L.C. 1995. The Chrysothamnus- Chrysothamnus. In: K.L. Johnson, ed.
Ericameria connection (Asteraceae). Great Basin Proceedings of the Fourth Utah Ecology
Naturalist 55: 84-88. Workshop; 1986 September 17-18, Cedar City,
UT. College of Natural Resources, Utah State
Deitschman, G.H., K.R. Jorgensen, and A.P. University, Logan, UT. p. 9-18.
Plummer. 1974. Chrysothamnus. In: C.S.
Schopmeyer, tech. coord., Seeds of Woody Nesom, G.L. and G.I. Baird. 1993. Completion of
Plants of the United States. Agriculture Ericameria (Asteraceae: Asterae), diminution of
Handbook 450. Washington, DC, U.S. Chrysothamnus. Phytologia 75: 74-93.
Department of Agriculture, Forest Service.
p. 326-328. Paulsen, H.A., Jr. and J.C. Miller. 1968. Control of
Parry rabbitbrush on mountain grasslands of
McArthur, E.D., A.C. Blauer, A.P. Plummer, and Western Colorado. Journal of Range
R. Stevens. 1979. Characteristics and Management 21: 165-177
hybridization of important intermountain shrubs. .
208
Chrysothamnus viscidiflorus Nutt. low rabbitbrush
ASTERACEAE
Synonyms: Bigelovia douglasi Gray

Chrysothamnus douglasi Clements & Clements
Crinitaria viscidiflora Hook.
Ericameria viscidiflora (Hook) L. C. Anderson
perfect, fertile disc flowers, which are arranged in

compact terminal cymes. Each head includes
approximately 15 involucral bracts that are
arranged in poorly to well-defined vertical ranks.
The bracts of some subspecies have a greenish to
brownish thickened spot near their apex
(McArthur and others 1979). Low rabbitbrush is
the only species within the genus with frequent
polyploidy; diploids, tetraploids, and hexaploids
have been documented (x = 9, 2n = 18, 36, and 54,
respectively) (Anderson 1966). In addition,
occasional triploids and pentaploids are known
(Anderson 1971).
Taxonomy.—This species includes five

subspecies and several ecotypes within subspecies
(Anderson 1980, McArthur and Meyer 1987). The
most common subspecies include two glabrous
subspecies, viscidiflorus and axillaris, and two
pubescent subspecies lanceolatus and puberlus.
Abrams and Ferris (1960) describe low rabbitbrush
as a highly polymorphic species with several
freely integrating subspecies with overlapping
distributions. Stickyleaf low rabbitbrush or green
rabbitbrush (ssp. viscidiflorus) is the largest
subspecies of low rabbitbrush. Mature shrubs are
generally more than 50 cm tall whereas mature
shrubs of other subspecies are generally less than
50 cm tall. In addition, the leaves, stems, and
inflorescences of ssp. viscidiflorus are viscid
(sticky). Both mountain low rabbitbrush or
General Description.—Low rabbitbrush, also yellowbrush (ssp. lanceolatus) and hairy low
known as yellowbrush, green rabbitbrush, little rabbitbrush (ssp. puberlus) are shorter at maturity
rabbitbrush, or Douglas rabbitbrush, is an erect than ssp. viscidiflorus, and both have finely
shrub, with many branches stemming from a pubescent stems. Mountain low rabbitbrush has
simple base. It is typically 0.3 to 1.1 m tall. gray or straw-colored stems, whereas hairy low
Brittle, erect branches can be glaborous or rabbitbrush has yellow to green stems. All four of
puberulent with pale green or white bark. Leaves these subspecies are fairly widespread throughout
are often twisted. Leaf shape can vary between the Great Basin. Some previously recognized
narrowly linear to oblong or lanceolate, with leaf subspecies, such as ssp. elegans and pumulis, have
length ranging from 1 to 6 cm. Leaves may be been consolidated; chromatographic work
glaborous or pubescent, the degree of which may substantiates the consolidations (Anderson 1980;
vary by subspecies. Flowerheads include five McArthur and others 1978). There is a difference
of opinion on the taxonomic affinity of C.
209
viscidiflorus with C. nauseosus and C. parryi viscidiflorus were 37 and 58 percent, respectively.
based on molecular genetic techniques (Nesom Seeds from plants adapted to higher elevations
and Baird 1993; Anderson 1995) (where seeds ripen earlier in the year) are more
dormant than those collected from lower
Range.—Low rabbitbrush is one of the most elevations and may be more chill responsive
widely distributed shrubs in Western North (Meyer 1996). There are about 1,720 cleaned seeds
America (McArthur and others 1979). One reason per gram (Deitschman and others 1974).
for its broad distribution is the species’ great
ecological amplitude. It is found growing on dry, Growth and Management.—Seedlings emerge in
open areas from British Columbia and Montana, spring and grow quickly. After a period of growth
south to New Mexico, Arizona, and eastern and establishment, shrubs often produce seed after
California. their second growing season (Meyer 1996). Low
rabbitbrush can dominate rangelands that are
Ecology.—Due to its adaptability, low rabbitbrush grazed heavily or cleared. This species has a strong
grows in a variety of habitats. Plants generally, tendency to sprout from the base (Wasser 1982),
however, inhabit drier sites, such as foothills, which may make it more difficult to remove and
mountains, dry plains, and valleys. Annual difficult to control with herbicides (Cook and
precipitation within its range varies between 250 others 1965). Fall is the preferred time to seed;
and 535 mm, with notable exceptions at higher seed can be drilled or broadcast and should remain
elevations. Anderson (1986) has shown that within the top 3 mm of the soil surface. Seeds can
polyploid races, in subspecies where polypoidy be harvested by hand and by vacuum harvesting
occurs (ssp. lanceolatus, puberulus, and techniques.
viscidiflorus), are adapted to lower and drier sites
than their diploid counterparts. Interestingly, Benefits.—Low rabbitbrush is an important native
Anderson has also observed a correlation within component of Western North American
certain subspecies between plant stature and leaf shrublands. It can provide important browse to
size with altitude and precipitation (personal both game and livestock, especially during late fall
communication). In addition to its ability to and winter months after more desirable forage has
tolerate xeric conditions, some variants of low been consumed. As a forage shrub, the palatability
rabbitbrush can grow well in areas with poor or of low rabbitbrush varies significantly among
disturbed soils. Low rabbitbrush can be found at subspecies, varieties, locality, and season. Some
elevations ranging from 790 to 3,400 m. It is subspecies, e.g., ssp. lanceolatus, may be used
commonly associated with big sagebrush heavily by domestic livestock and wildlife,
(Artemisia tridentata), broom snakeweed whereas others are consumed little, if at all (USDA
(Gutierrezia sarothrae), and other rabbitbrushes 1937, Goodrich and others 2001). In addition to its
Chrysothamnus sp.), although it can be associated value as a browse species, low rabbitbrush has
with shadscale (Atriplex confertifolia), winterfat successfully been used to revegetate depleted
(Ceratoides lanata), fourwing saltbush (Atriplex rangelands and other disturbed sites (Plummer
canescens), greasewood (Sarcobatus 1977).
vermiculatus), and occasionally pinyon (Pinus sp.)
and juniper (Juniperus sp.). Goodrich and others References
(2001) note that C. viscidiflorus does not
commonly overtop associated herbaceous Abrams, R. and R.S. Ferris. 1960. Illustrated flora
vegetation. Elevation tends to partition the of the Pacific States, Vol. 4, Bignoniaceae to
subspecies; mountain low rabbitbrush (ssp. Compositae. Stanford University Press,
lanceolatus) especially and stickyleaf low Stanford, CA. 732 p.
rabbitbrush (ssp. viscidiflorus) grow well at higher
elevations, while hairy low rabbitbrush (ssp. Anderson, L.C. 1966. Cytotaxonomic studies in
puberlus) does best on lower desert foothills. Chrysothamnus (Asterae, Compositae).
Reproduction.—Shrubs flower from August
through October; seed matures from October until Anderson, L.C. 1971. Additional chromosome
December. Like other members of the genus, low numbers in Chrysothamnus (Asteraceae).
rabbitbrush seeds (or achenes) are wind Bulletin of the Torrey Botanical Club 98: 222-
disseminated. Meyer (1996) found that 225.
germination rates at 15 oC for ssp. lanceolatus and
210
Anderson, L.C. 1980. Identity of narrow-leaved U. S. Department of Agriculture, Forest Service,
Chrysothamnus viscidiflorus (Asteraceae). Intermountain Forest and Range Experiment
Great Basin Naturalist 40: 117-120. Station, Ogden, UT. 82 p.
Anderson, L.C. 1986. Cytogeography of McArthur, E.D., D.L. Hanks, A.P. Plummer, and
Chrysothamnus viscidiflorus. In: E.D. McArthur A.C. Blauer. 1978. Contributions to the
and B.L. Welch, comps. Proceedings— taxonomy of Chrysothamnus visicidiflorus
symposium on the biology of Artemisia and (Astereae, Compositae) and other
Chrysothamnus; 1984 July 9-13, Provo, UT. Chrysothamnus species using paper
General Technical Report INT-200. Ogden, UT: chromatography. Journal of Range Management
U.S. Department of Agriculture, Forest Service, 31: 216-223.
Intermountain Research Station. p. 93-97.
McArthur, E.D. and S.E. Meyer. 1987. A review
Anderson, L.C. 1995. The Chrysothamnus- of the taxonomy and distribution of
Ericameria connection (Asteraceae). Great Chrysothamnus. In: K.L. Johnson, ed.
Basin Naturalist 55: 84-88. Proceedings of the Fourth Utah Ecology
Workshop; 1986 September 17-18, Cedar City,
Cook, C.W., P.D. Leonard, and C.D. Bonham. UT. College of Natural Resources, Utah State
1965. Rabbitbrush competition and control on University, Logan, UT. p. 9-18.
Utah rangelands. Bulletin 454. Utah
Agricultural Experiment Station., Logan, UT. Meyer, S.E. 1996. Chrysothamnus Nutt. In:
28 p. Woody Plant Seed Manual,
http://www.wpsm.net/Chrysothamnus.pdf. 11 p.
Deitschman, G.H., K.R. Jorgensen, and A.P.
Plummer. 1974. Chrysothamnus. In: C.S. Nesom, G.L., and G.I. Baird. 1993. Completion of
Schopmeyer, tech. coord., Seeds of Woody Ericameria (Asteraceae: Asterae), diminution of
Plants of the United States. Agriculture Chrysothamnus. Phytologia 75: 74-93.
Handbook 450. U.S. Department of Agriculture,
Forest Service. p. 326-328. Plummer, A.P. 1977. Revegetation of disturbed
intermountain area sites. In: J. L. Thames,
Goodrich, S., R.M. Thompson, and A. Huber. editor, Reclamation and use of disturbed land in
2001. A yellowbrush/grass community type the Southwest. University of Arizona Press,
from the Uinta Mountains and Utah plateaus. Tucson, AZ: 302-339.
In: E.D. McArthur and D.J. Fairbanks, comps.
Shrubland ecosystem genetics and biodiversity: U.S. Department of Agriculture, Forest Service.
proceedings; 2000 June 13-15, Provo, UT. 1937. Range plant handbook. United States
Proceedings RMRS-P-21. U. S. Department of Government Printing Office, Washington, DC.
Agriculture, Forest Service, Rocky Mountain 816 p.
Research Station, Ogden, UT. p. 344-346.
Wasser, C.H. 1982. Ecology and culture of
McArthur, E.D., A.C. Blauer, A. P. Plummer, and selected species useful in revegetating disturbed
R. Stevens. 1979. Characteristics and lands in the West. U.S. Department of the
hybridization of important intermountain shrubs. Interior, Fish and Wildlife Service, Washington,
III. Sunflower family. Research Paper INT-220. DC. 347 p.
211
Cissampelos pareira L. velvet leaf
MENISPERMACEAE
Synonyms: Cissampelos caapeba L.

Cissampelos microcarpa DC.
John K. Francis
Range.—Velvet leaf is native from Mexico to

Argentina and Peru on the New World mainland
and in the West Indies (Instituto Botánico
Darwinian 2002, Secretaría de Medio Ambiente y
Recursos Naturales 2002, Stevens and others
2001). It is native to Florida, although rare or
possibly locally extinct (Nelson 1996). It is also
found throughout tropical Asia and Africa (Long
and Lakela 1976, Parrotta 2001), although it is not
clear whether it is native or naturalized there.
Ecology.—Velvet leaf is a “good site” species. It

usually does not grow on exposed clay subsoils,
compacted soils, excessively drained, or very
poorly drained soils. The species occurs on a wide
variety of soil textures, pH levels, and on soils
derived from most parent materials including
limestone and ultramafic rocks (serpentine). In
Puerto Rico, velvet leaf grows in areas that receive
from 750 to about 2400 mm of annual precipitation
General Description.—Velvet leaf is also known at elevations from near sea level to about 1,500 m
as abuta, pareira, alcotá, bejuco de mona, oreja de (Stevens and others 2001). Velvet leaf may be
raton, hierba de peso, curarina, liane amère, liane- found on roadsides, fencerows, river banks,
cordé, and liane molle. It is a shrubby climber (or hammocks, brushy pastures, and secondary and
liana) that reaches 3 to 6 m along the ground or remnant forests. It is moderately intolerant of shade
into the crowns of trees. The stem is woody, and does not grow under the closed canopy of high
flexible, and slender (to 1 cm), and twines for forest. Openings, and therefore disturbance, appear
support. The pith and rays of the stem cross section necessary for the maintenance of the species.
resemble a wagon wheel with spokes; annual rings
are not visible. The root system consists of flexible, Reproduction.—Velvet leaf flowers and fruits all
light-brown lateral roots with sinkers and year in the Americas (Acevedo-Rodríguez 1985,
moderately abundant fine roots. There are Stevens and others 2001). In India it flowers
relatively few branches. Alternate leaves are between July and October and fruits from October
usually softly pubescent on both surfaces. The to December (Parrotta 2001). Fruits collected in
petioles are 3 to 7 cm long. Venation is palmate in Puerto Rico weighed an average of 0.1925 +
widely oval or nearly round 4- to 10-cm blades. 0.0008 g/fruit. Seeds from them averaged 0.0109
The small staminate and pistillate inflorescences g/fruit (air-dried). When planted in commercial
(cymes) are borne in leaf axils. The fruits are juicy potting mix without pretreatment, 26 percent of the
red, red-orange, or yellow, hairy drupes 4 to 5 mm seeds germinated between 28 and 61 days after
in diameter. Each fruit is partially covered by a sowing (author’s observation). Birds presumably
rounded bract. The seeds have a croissant shape disperse the seeds. Fruit and seed production are
(Acevedo-Rodríguez 1985, Howard 1988, Liogier generally moderate, and seedlings are rarely
1985, Stevens and others 2001). The species’ abundant. Stems layer (root) wherever they contact
chromosome number is 2n = 24 (Long and Lakela the soil.
1976).
Growth and Management.—No information is
212
available on the growth rate of seedlings. Growth International BioPark Foundation. 2002. Abuta,
of sprouts is rapid (at least 3 m in the first year). family Menispermaceae. http://www.biopark.
Individual stems are not long-lived, but by org/peru/abuta.html. 3 p.
sprouting and layering, plants or clones may last
many years. The creation of small forest openings
and protection against heavy grazing are probably
good strategies to promote natural establishment of
new plants.
Benefits.—Velvet leaf adds to biodiversity and
Long, R.W. and O. Lakela. 1976. A flora of
biomass, helps stabilize the soil, and furnishes food
tropical Florida. Banyan Books. Miami, FL.
and cover for wildlife. A major interest in the
962 p.
species arises from the natural medicinal benefits
of the plant’s chemical contents. Known as the
“midwife’s herb,” it has been used for centuries by Morita, H., K. Matsumoto, K. Takeya, and H.
native peoples of South America to treat menstrual Itokawa. 1993a. Azafluoranthene alkaloids from
cramps, prevent threatened miscarriage, control Cissampelos pareira. Chemical and
uterine hemorrhages, and ease childbirth and Pharmaceutical Bulletin 41(7): 1,307-1,308.
postpartum pain (Rain-tree 2002). The list of other
natural medicinal applications to which the herb is
applied is large: urinary infections, kidney stones, Morita, H., K. Matsumoto, K. Takeya, H. Itokawa,
and Y. Iitaka. 1993b. Structures and solid state
arthritis, snakebite, cough, dysentery, piles, ulcers,
pain, indigestion, colic, skin irritations, stings, tautomeric forms of two novel antileukemic
intestinal worms, and wounds (Parrotta 2001, Rain- tropoloisoquinoline alkaloids, pareirubrines A
and B, from Cissampelos pareira. Chemical and
tree 2002). Ground tissues and preparations of
velvet leaf are sold throughout the world in Pharmaceutical Bulletin 41(8): 1,418-1,422.
markets, shops, and mail-order companies. The
physiological effects are apparently derived from a Nelson, G. 1996. The shrubs and woody vines of
number of alkaloids found in the tissues of the Florida. Pineapple Press, Inc., Sarasota, FL.
plant (International BioPark Foundation 2002, 391 p.
Morita and others 1993a, Morita and others
1993b). Parrotta, J.A. 2001. Healing plants of Peninsular
India. CABI Publishing, Wallingford, UK and
References New York. 917 p.
Acevedo-Rodríguez, P. 1985. Los bejucos de Rain-tree. 2002. Abuta. http://www.rain-

Puerto Rico. Vol. 1. General Technical Report tree.com/abuta.htm. 7 p.
SO-58. U.S. Department of Agriculture, Forest
Service, Southern Forest Experiment Station, Secretaría de Medio Ambiente y Recursos
New Orleans, LA. 331 p. Naturales. 2002. Especies con usos no
maderables in bosques tropicales y
Howard R.A. 1988. Flora of the Lesser Antilles, subtropicales: Cissampelos pareira L.
Leeward and Windward Islands. http://www.semarnat.gob.mx.pfnm2/fichas/cissa
Dicotyledoneae, Part 1. Vol. 4. Arnold mpelos_pareira.htm. 2 p.
MA. 673 p. Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
Montiel (eds.). 2001. Flora de Nicaragua.
Instituto de Botánica Darwinian. 2002. Catálogo Monographs in Systematic Botany Vol. 85, No.
de las plantas vasculares de la Argentina. 2. Missouri Botanical Garden, St. Louis, MO.
http://www.darwin.edu.ar/Catalogo/indicevascul p. 945-1,910.
ares.htm. [not paged].
213
Citrus aurantiifolia (Christm.) Swingle Key lime
RUTACEAE
Synonyms: Limonia aurantifolia Christm.

Limon spinosum Mill.
Citrus limonia Osbeck
Citrus lima Luman
Citrus spinosissima G.F.W. Meyer
Citrus acida Roxb.
John K. Francis
axillary clusters. The fruits (hesperidiums) are

ellipsoidal, 3 to 5 cm in diameter, have juicy,
greenish-yellow flesh, and are yellow at maturity.
They contain a few white, pointed seeds about 1
cm long (Liogier 1988, Little and Wadsworth
1964).
Range.—Key lime is believed to be native of

eastern Malaysia. It was introduced to the Asian
mainland early in historical times and carried by
Arab traders to the Middle East and eventually
came to Europe during the Crusades (Burkill
1997). The species was introduced to the West
Indies by Columbus during his second voyage
(Ehler 2002). Key lime has been planted
throughout the tropics and has naturalized in at
least Puerto Rico, the Virgin Islands (Little and
Wadsworth 1964), and the Florida Keys (Nelson
1996).
Ecology.—Key lime is most competitive in areas

that receive from about 700 to 1000 mm of mean
annual precipitation. It tolerates drought better than
any of the other citrus fruit species (Morton 1987).
General Description.—Key lime is the name used If planted, it will grow but becomes increasingly
most often to refer to a primitive race of Citrus susceptible to disease in areas that receive up to
arantiifolia cultivated and naturalized in the West about 2000 mm of annual precipitation. Most well-
Indies. It is also referred to as Mexican lime, West drained soils are suitable, particularly those rich in
Indian lime, lima, limón criollo, limón agria, limón calcium (Morton 1987). Elevations below 900 m
boba, and citron (Little and Wadsworth 1964). are best (Secritaría del Medio Ambiente y Recursos
Key lime is an evergreen, spiny shrub or small tree Naturales 2002). Key lime is intolerant of shade
to 6 m in height. The plant has single or multiple and will not survive long under a closed forest
stems and irregular branches covered with canopy. Cattle grazing encourages it somewhat by
smoothish brown to gray bark. The twigs are eliminating some of the competition without
quadrangular (when young), green, and bare sharp damaging it. Plants are top-killed by fires and hard
axillary spines 3 to 17 mm long. The leaves are frosts, but will sprout and survive if the disturbance
yellow-green to dark green, with 5- to 28-mm is not repeated frequently. Naturally reproduced
winged petioles and elliptic to oval leathery 4- to trees may be seen on roadsides, fencerows,
13-cm long blades with edges that have minute abandoned farms, secondary forests, and coastal
rounded teeth. The crushed foliage has a strong, hammocks.
distinct, spicy (citrus) odor and taste. The four- to
five-petaled white flowers occur in few-flowered
214
Reproduction.—Key lime flowers in late spring citrus fruits cause phytophotodermatosis in
and fruits in fall to spring in Florida (Nelson 1996). sensitive individuals (Bruneton 1999). The wood is
In Puerto Rico, it flowers in spring to early summer hard and heavy, but is used for little other than
and fruits in summer and fall (Little and fuel. It is a good honey plant. With pruning, the
Wadsworth 1964). The fruits ripen and fall from plant can be used for a living fence post (Little and
the trees 5 to 6 months after flowering (Morton Wadsworth 1964) and can be formed into a hedge
1987). Fruits in one collection in Puerto Rico (Burkill 1997).
ranged from 3.0 to 4.5 cm in diameter and weighed
from 19 to 51 g. Air-dried seeds separated from References
them averaged 0.0702 + 0.0037 g/seed or 14,000
seeds/kg. Placed on moist blotter paper, 84 percent Bruneton, J. 1999. Toxic plants dangerous to
germinated between 15 and 49 day after sowing. humans and animals. Lavoisier Publishing,
Germination is epygeal. Many of the seeds are Paris. 545 p.
polyembryonic. Seedlings begin to fruit in 3 to 6
years (Morton 1987). In the West Indies, the seeds Burkill, H.M. 1997. The useful plants of West
are mainly dispersed by gravity and humans. There Tropical Africa. Vol. 4. Royal Botanic Gardens,
are undoubtedly animal vectors in the original Kew, UK. 969 p.
native habitat. Key lime can be air-layered using
indol buteric acid (IBA) with a high degree of Ehler, J.T. 2002. Key lime (Citrus aurantifolia).
success (Morton 1987). Food Reference, Key West, FL.
http://foodreference.com/html/artkeylimes.html.
Growth and Management.—Growth of Key lime 2 p.
is slow, less than 0.5 m/year. Plants may live for 25
years or more. Although the species is not managed Katzer, G. 2002. Lime [Citrus aurantifolia
in the wild, existing trees are usually protected by (Christm. & Panz.) Swengle]. Gernet Katzer’s
local inhabitants, whenever possible. Plantations spice pages. http://www-ang.kfunigraz.
are established using potted material that is set at ac.at/~katzer/engl/Citr_aur.html. 4 p.
7.5- by 7.5-m spacing. Pruning is usually not
necessary (Morton 1987). Liogier H.A. 1988. Descriptive flora of Puerto
Benefits.—Key lime has been under cultivation or 2. Editorial de la Universidad de Puerto Rico,
semicultivation for thousands of years. Improved Río Piedras, PR. 481 p.
horticultural varieties, which have bigger, generally
seedless fruits and thornless plants, now dominate Liogier, H.A. 1990. Plantas medicinales de Puerto
the local and international fruit markets. The Rico y del Caribe. Iberoamericana de Ediciones,
unimproved type is still managed commercially Inc., San Juan, PR 566 p.
and is the basis for a juice and lime oil industry
(Morton 1987). It is also planted for a dooryard Little, E.L., Jr. and F.L. Wadsworth. 1964.
fruit tree, and fruits are harvested from the wild. Common trees of Puerto Rico and the Virgin
The principal use is still for food, refreshing drinks, Islands. Agriculture Handbook 249. U.S.
tasty disserts, and for seasoning meats, vegetables, Department of Agriculture, Washington, DC.
salads, sauces, and casseroles (Ehler 2002, Katzer 548 p.
2002). The wild type is superior in flavor to the
improved varieties because it has a stronger flavor Morton, J.F. 1987. Citrus aurantifolia Swingle.
and a higher acid content (Ehler 2002). The from: Fruits of warm climates.
pericarp (rind) contains 7 percent essential oil with http://www.hort.perdue.edu/newcrop/morton/me
principal constituants, citral, limonene, and xican_lime.html. 8 p.
fenchon as well as terpineol, bisabolene, and other
terpenoids (Katzer 2002). Key lime is used to treat Nelson, Gil. 1996. The shrubs and woody vines of
a huge number of ailments (Burkill 1997, Liogier Florida. Pineapple Press, Inc. Sarasota, FL.
1990). The author recommends a tea prepared from 391 p.
juice, fruit rind, or leaves as an expectorant and to
relieve catarrh brought on by colds and flu. The
fresh fruits and bottled juice are an excellent source
of vitamin C and were once relied upon to prevent
scurvy. Essential oils of Key lime and some other
215
Secritaría del Medio Ambiente y Recursos
Naturales. 2002. Especies con usos no
maderables en bosques tropicales y
subtropicales: Lantana involucrata L.
http://www.semarnat.gob.mx.pfnm2.fichas/
citrus_aurantifolia.htm. 2 p.
216
Clematis orientalis L. oriental virginsbower
RANUNCULACEAE
Synonyms: Clematis aurea A. Nels. & J.F. Macbr.

Viticella orientalis (L.) W.A. Weber
John K. Francis
Range.—Oriental virginsbower is native from

Turkey through Asia to the Korean Peninsula and
south to Iran and Northwest India (Griffiths 1994).
The species has been widely planted as an
ornamental and has naturalized and escaped in
Idaho, Nevada, Utah, Colorado, and New Mexico
(Natural Resources Conservation Service 2003)
and probably other places in the world. There are
two varieties, orientalis and robusta, separated on
the basis of pedicel length and thickness (Wu and
others 2001).
Ecology.—Oriental virginsbower grows in

shrublands, riverbanks, gullies, sand depressions,
and riparian forests in hot, dry valleys and desert
and semidesert areas. In Utah, oriental
virginsbower is mostly found along rivers, creeks,
and intermittent streams at the mouth of canyons.
It is not present in cooler sites farther up the
canyons. It requires well-drained soils, but is not
particular about soil texture, and tolerates acid and
alkaline soils from a wide range of parent
General Description.—Oriental virginsbower, materials. It does well on chalky soils (Plants for a
also known as orange peel or orange peel clematis, Future 2003). The species inhabits sites with
is a deciduous, sometimes suffruticose vine or elevations between 400 and 3,800 m (Wu and
scrambling shrub. The multiple stems may reach 8 others 2001). It is hardy to about –15 °C (Plants
m in length and as much as 7 cm in thickness. For a Future 2003) and presumably survives from
Stems are covered with longitudinally fissured roots after top damage. Oriental virginsbower will
gray-brown bark. Inner bark is green. Vines cling climb taller vegetation, fences, and rocks, but
to rocks and other plants by twining as they climb. forms mounds and mats when objects to climb are
The roots are weak and flexible. Fine roots are tan not available.
or orange-tan. The leaves are pinnately compound
with three to seven coarsely-toothed to entire, Reproduction.—Oriental virginsbower flowers
ovate to linear-lanceolate leaflets 1 to 5 cm long. between August and October. Both flowers and
The foliage has a gray-green color. Axillary fruits are often present on the plant at the same
cymes, which may have one to many flowers, are time. The flowers are pollinated by bees and flies
borne on the current season’s growth. Four yellow, (Plants for a Future 2003). A group of air-dried
yellow-orange, or yellowish-green sepals give the seeds collected in Utah averaged 911 seeds/g.
flowers their color. Flowers have a delicate Oriental virginsbower seeds are dispersed by the
fragrance. The feather-like style, 3 to 10 cm long, wind. Seedlings are not abundant. It reproduces
remains attached to the achene and functions as a vegetatively by sprouting from the root crown and
wing. There are 2n = 16, 32 chromosomes (Plants by layering.
for a Future 2003, Welsh and others 1987, Wu and
others 2001). Growth and Management.—Growth of oriental
virginsbower is rapid, at least 1 m/year from
sprouts or existing stems. Although the risk of
217
environmental damage is not severe, because it is References
an alien, the species should not be used in
environmental restoration plantings. No specific Griffiths, M. 1994. Index of garden plants. Timber
recommendations for control are available, but the Press, Portland, OR. 1,234 p.
general procedure of cutting plants and spot
spraying the sprouts with broadleaf herbicide Natural Resources Conservation Service. 2003.
would probably be effective. The use of potted Plants profiles: Clematis orientalis L., oriental
seedlings or rooted cuttings for ornamentals is virginsbower. http://plants.gov/cgi_bin/
recommended. They should be outplanted in the plant_profile.cgi?symbol=CLOR. 5 p
spring after the frost danger has passed. Pruning
should take place in the spring (Plants for a Future Plants For a Future. 2003. Database search results:
2003). Clematis orientalis. http://www.scs.leeds.ac.uk/
cgi-bin/pfaf/arr_html?Clematis+orientalis. 7 p.
Benefits.—Oriental virginsbower is planted as an
ornamental and ground cover. Plants escaped into Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
the wild help protect the soil, furnish cover for Hinnins, eds. 1987. A Utah flora. Great Basin
wildlife, and add beauty to the landscape, Naturalist Memoirs 9. Brigham Young
especially during the fall when it produces flower University, Provo, UT. 894 p.
and seed heads. Oriental virginsbower is used in
landscaping as a flowering ground cover. Infusions Wu Z., P.H. Raven, and Hong D. (eds.). 2001.
are used as a gargle for sore and ulcerated throats Flora of China. Vol. 6. Science Press, Beijing
and to treat dog bites (Plants for a future 2003). and Missouri Botanical Garden Press, St. Louis,
Mo. 512 p.
218
Clibadium erosum (Sw.) DC. carruzo
ASTERACEAE
Synonyms: Trixis erosa Sw.
John K. Francis
Vincent, Costa Rica, and Panama (Howard 1989,

Missouri Botanical Garden 2003). The species is
elsewhere.
Ecology.—Carruzo is a species of moist and wet

mountains. It grows at elevations above about 400
m in Puerto Rico and it is reported from 800 and
1,300 to 1,500 m in Costa Rica (Instituto Nacional
de Biodiversidad 1997, Missouri Botanic Garden
2003). Mean annual rainfall of its habitat in Puerto
Rico ranges from 2000 to over 4000 mm/year. The
soils are mostly clayey, derived from a variety of
rock types, anaerobic in their subsoils, and have
pH’s between 5 and 6. Because the species is light-
demanding, it grows mainly in disturbed areas
(Little and Wadsworth 1964) such as roadsides,
cuts and fills, landslides, tree-tip mounds, logged
areas, plantations, fields, and abandoned
construction sites. Once established, carruzo
General Description.—Carruzo, also known as competes aggressively with grass, herbs, and
jackass breadnut, cachimbo, tuchima, turma de shrubs of equal or lesser height. Overtopped, it
toro, bois énivrant, and medsinnyé benní, is a soon dies.
shrub or small tree up to 6 m in height and 10 cm
in basal stem diameter. These plants have a single Reproduction.—Carruzo flowers and fruits nearly
stem from the ground, unless damaged, and are throughout the year, at least in Puerto Rico (Little
much branched in the crown. The trunk bark is and Wadsworth 1964). Fruits are produced in great
gray to greenish brown, smooth to slightly numbers. A group of akenes harvested in Puerto
fissured. The white wood is soft to medium in Rico averaged 0.001867 g/fruit or 536,000
hardness, brittle, without annual rings, and has a 3- fruits/kg. The author could not germinate the hard
mm pith. The plant often breaks at forked stems. akenes on moist blotter paper. Some sort of
Carruzo plants are supported by a root system of pretreatment is probably necessary. Fruits can be
tan, flexible laterals and may or may not retain a collected in quantity by hand. Means of dispersal
taproot. The twigs are ringed at the nodes by is unknown. Seedlings are common on disturbed
opposite, three-lobbed leaf scars and densely ground near seed sources. Carruzo shrubs will
pubescent. The leaves are broadly ovate, with sprout if disturbed and will layer if intact stems
long, 2- to 8.5-cm petioles, coarsely and finely come in contact with the ground.
incised-dentate, and have an elongated tip. The
inflorescences are many-branched clusters Growth and Management.—Carruzo grows
(corymbs) containing many heads with white to rapidly, adding 1 to 1.5 m/year to its height. It
rose-purple flowers. Several black, 2- by 1.5-mm, begins flowering and fruiting after about 1 year
seed-like fruits (akenes) are produced per flower and probably lives 2 to 5 years. There is no
(Howard 1989, Liogier 1997, Little and published record of planting or propagation
Wadsworth 1964). techniques. Because it grows rapidly and thickly in
disturbed areas, carruzo sometimes becomes
Range.—Carruzo is native to Puerto Rico, St. weedy in fields, plantations, and along roads.
Martin, Saba, St. Kitts, Nevis, Montserrat,
Guadeloupe, Dominica, Martinique, St. Lucia, St.
219
Stands along roads are routinely controlled by http://www.inbio.ac.cr/bins/k03/p13/c045/o0144
mowing. /f01362/g007091/s020661.htm. 1 p.
Benefits.—Carruzo helps protect the soil in Liogier, H.A. 1997. Descriptive flora of Puerto
disturbed areas, serves as a nurse species for later Rico and adjacent islands. Vol. 5. Editorial de la
successional species, and provides cover and Universidad de Puerto Rico, Río Piedras, PR.
possibly food for wildlife. 436 p.
References Little, E.L., Jr. and F.H. Wadsworth. 1964.

Common trees of Puerto Rico and the Virgin
Howard, R.A. 1989. Flora of the Lesser Antilles, Islands. Agriculture Handbook 249. U.S.
Leeward and Windward Islands. Dicotyledoneae. Department of Agriculture, Forest Service,
Vol. 6. Arnold Arboretum, Harvard University, Washington, DC. 548 p.
Jamaica Plain, MA. 658 p.
Missouri Botanical Garden. 2003. Flora
Instituto Nacional de Biodiversidad. 1997. Lista de Mesoamericana: lista anotada—03 Feb 2003.
especímenes de Clibadium erosum. http://mobot.org/cgi-bin/search_vast. [not
paged].
220
Clidemia cymosa (Wendl. ex Spreng.) Alain camasey peludo
MELASTOMATACEAE
Synonyms: Heterotrichum cymosum (Wendl.) Urban

Melastoma cymosa Wendl. in Spring.
Heterotrichum eggersii Cogn.
John K. Francis
to hundreds of tiny, light brown seeds (Liogier
1995, Little and Wadsworth 1964).
Range.—Camasey peludo is endemic to Puerto

Rico (Liogier 1995). It grows mainly in the
Luquillo Mountains, the Cordillera Central, and
moist foothills in the eastern, northern, and western
districts (Little and Wadsworth 1964).
Ecology.—Camasey peludo is common and

widespread but never abundant. It is confined to
disturbed, open areas, such as roadsides, landslides,
recently abandoned fields, and large tree-fall gaps.
Camasey peludo grows in areas receiving from
about 1600 to over 3000 mm of annual
precipitation on soils derived from igneous and
sedimentary rocks. These upland soils are mostly
highly weathered clays with slow internal drainage.
The species grows from a few meters above sea
level to nearly 1,000 m in elevation. Camasey
peludo are occasionally attacked (sometimes
General Description.—Camasey peludo, which lethally) by white flies (Aleyrodidae) and mealy
means hairy melastome, is also known locally as bugs (Pseudocoddidae, author’s observation).
camasey colorado, camasey de paloma, pelúa, and Cyrtopeltis modesta Distant (Hemiptera),
terciopelo. It is a medium to large shrub to 4.5 m in Paurocephala heterotrichii Caldwell (Homoptera),
height and 7.5 cm in trunk diameter. Shrubs 1.5 to and Bleparomastix ebulealis Guenée (Lepidoptera)
2.5 m in height are typical. The stems may be have also been observed attacking camasey paludo
single or multiple. The bark is thin, gray-brown, (Martorell 1975).
and smooth or slightly fissured. There are
relatively few branches. The twigs are red or Reproduction.—Camasey peludo flowers and
orange near the tips due to a thick covering of fruits nearly throughout the year (Little and
reddish hairs, greenish below, becoming brown as Wadsworth 1964). A collection of fruits averaged
they grow older. The young leaves are bright red or 1.298 + 0.046 g/fruit and ranged from 0.650 to
orange. As the leaves approach full size, the color 1.998 g/fruit. Seeds separated from them averaged
fades to green on the upper surface with reddish 98 million seeds/kg (air dry). Sown on moist filter
accents at the edges and other areas where hairs are paper, 85 percent germinated between 11 and 49
concentrated and yellow-green on the lower days after sowing. No data on nursery management
surface. The leaves are ovate with a cordate base of seedlings is available. Forty-four branches of
and an elongated tip and have seven veins radiating camasey peludo were air layered by girdling and
from the base. The petioles are 4 to 9 cm long and treating with 0.3 percent IBA in talcum, then
the blades are 8 to 18 cm long and 7 to 12 cm covered with commercial potting mix and wrapped
broad. The 1.9-cm, white or pinkish flowers are with aluminum foil. Eighty-nine percent rooted.
grouped in terminal panicles. The purple berries However, IBA-treated cuttings placed in a mist bed
that develop are about 12 mm in diameter. They entirely failed to root and grow. Natural layering is
are juicy and edible with a delightful flavor common.
somewhat like blueberries. Each berry contains up
221
Growth and Management.—Camasey peludo used because of the poor form and small size of the
plants are relatively short-lived, lasting about 3 to 6 plants (Little and Wadsworth 1964).
years. Resprouting and layering may enable them
to persist somewhat longer. There is no reported References
planting or management experience for this
species. Plants cloned by layering should be used Liogier, H.A. 1995. Descriptive flora of Puerto
for planting until methods of growing the plants Rico and adjacent islands. Vol. 4. Editorial de la
from seed are developed. Natural reproduction in Universidad de Puerto Rico, San Juan, PR.
areas of abundant rainfall may be encouraged by 617 p.
cultivation or scalping to expose a bare seedbed
during wet portions of the year. Little, E.L., Jr. and F.L. Wadsworth. 1964.
Benefits.—Camasey peludo is an attractive Islands. Agriculture Handbook 249. U.S.
roadside plant and should be encouraged whenever Department of Agriculture, Washington, DC.
possible. Its beauty also suggests real potential as 548 p.
an ornamental, although its life is short and it has a
tendency to unsightly decline in its last year. The Martorell, L.F. 1975. Annotated food plant catalog
fruit, currently almost unknown to the public, is of the insects of Puerto Rico. Department of
very tasty and deserves attention. The fruits are a Entomology, Agricultural Experiment Station,
food source for birds. The light-brown, moderately University of Puerto Rico, Río Piedras, PR.
hard wood has a specific gravity of 0.6 but is not 303 p.
222
Clidemia hirta (L.) D. Don Koster’s curse
MELASTOMATACEAE
Synonyms: Melastoma hirta L.

Clidemia crenata DC.
Melastoma elegans Aublet
Clidemia elegans (Aublet) D. Don
John K. Francis
coffee-colored seeds (author’s observation,

Howard 1989, Liogier 1995, Smith 1992, Stevens
and others 2001).
Range.—Koster’s curse is native to the Greater

and Lesser Antilles, southern Mexico, Central
America, and South America as far south as
Northern Argentina and Bolivia (Howard 1989,
Liogier 1995, Stevens and others 2001). It has
naturalized in Hawaii, American Samoa, many of
the Pacific and Indian Ocean islands, Australia,
Southern Asia, Sri Lanka, India, and East Africa
(Binggeli 1997, Pacific Island Ecosystems at Risk
2002).
Ecology.—In its native range, Koster’s curse

colonizes disturbed habitat—landslides, river
banks, burned areas, tree tip mounds, pastures, old
fields, plantations, fence rows, and roadsides. It
occurs as scattered plants, occasionally as thickets,
which flourish for a few years and succumb to
competition or disease and insects. It tolerates a
General Description.—Koster’s curse (a wide range of soil properties as long as moisture is
Hawaiian name) is also known as clidemia, soap adequate. Annual rainfall ranges from about 1200
bush, camasey, camasey peludo, nigua, kak mél, to 4000 mm. Elevation may vary from near sea
bon bon mél, canot-macaque, and bonbon bleu. It level to 1,500 m. The species grows well in full sun
is a weak upright or scrambling shrub that varies in but is moderately tolerant of shade. In some non-
mature size from less than 1 m to 5 m in height, native habitat in the absence of the usual insects,
depending on habitat. Koster’s curse is supported fungi, and fast-growing competitors, Koster’s curse
by shallow lateral roots and abundant fine roots. forms dense and tall, monotypic thickets under
There are usually multiple stems arising as sprouts forest canopies that shade out virtually all
from the rootstalk and base of the principal stem, vegetation below them (Binggeli 1997, Smith
especially in older and disturbed plants, resulting in 1992).
dense shrubs. The branches and twigs are slender.
The 5- to 18-cm, opposite leaves have a short Reproduction.—Koster’s curse flowers and fruits
petiole, five to seven principal nerves radiating year-round in continuously moist habitat. After dry
from the base, an ovate blade with a serrated to seasons or droughts, in which it may defoliate or
entire edge, usually subcordate base and pointed die back, it recovers and begins flowering again. A
tip. The whole plant is covered with brown hairs. large plant may produce more than 500 fruits/year
Inflorescences are branched axillary cymes with (Smith 1992). A collection of fresh fruits from
five-merous white flowers. The fruits are Puerto Rico weighed an average of 0.2171 +
ellipsoidal, purple-to-black berries 6 to 8 mm long 0.0081 g/fruit. Air-dried seeds separated from them
with a flavor similar to blueberries, but weaker. averaged 0.00383 g/seed or 260,000 seeds/kg
Each fruit contains over 100, 0.5- to 0.7-mm-long, (author’s observation). The seeds are dispersed by
223
birds, feral pigs, other animals, and humans. The Franca, F., E.L. Lago, and P.D. Marsden. 1996.
seeds can remain viable in the soil for up to 4 Plants used in the treatment of leishmanial
years. The stems root on contact with moist soil; ulcers due to Leishmania (Vannia) braziliensis
even detached leaves have been observed to root in an endemic area of Bahia, Brazil. Revista da
(Smith 1992). Sociedade Brasileira de Medicina Tropical
29(3): 229-232.
Growth and Management.—Koster’s curse
grows slowly in the first months and faster Howard, R.A. 1989. Flora of the Lesser Antilles,
thereafter. It often dominates at the end of the first Leeward and Windward Islands. Vol. 5. Arnold
year (Binggeli 1997). Individual stems live 2 or 3 Arboretum, Harvard University, Jamaica Plain,
years in Puerto Rico, although plants may live MA. 604 p.
longer by sprouting. There is little motivation to
encourage Koster’s curse and often a need to Liogier, H.A. 1995. Descriptive flora of Puerto
control it. Single-pass mowing is usually Rico and adjacent islands. Vol. 4. Editorial de la
ineffective. Spraying with broadleaf herbicides is Universidad de Puerto Rico, San Juan, PR.
the most widely used control method, 617 p.
supplemented by hand pulling (Rubber Research
Institute of Malaya 1973). At best, such measures Murdiati, T.B., C.S. McSweeney, R.S.F.
are temporary. Some success has been achieved Campbell, and D.S. Stoltz. 1990. Prevention of
with biocontrol by the introduction of Liothrips hydrolysable tannin toxicity in goats fed
urichi, a thrips (Binggeli 1997). A number of other Clidemia hirta by calcium hydroxide
natural controls are being studied (State of Hawaii supplementation. Journal of Applied Toxicology
Department of Agriculture 2002). 10(5): 325-331.
Benefits and Detriments.—Desired or not, Pacific Island Ecosystems at Risk. 2002. Clidemia
Koster’s curse helps revegetate disturbed areas and hirta (L.) D. Don, Melosomataceae. http://www.
provides a reliable food source for wildlife. The hear.org/pier3clhir.htm. 5 p.
species is used in Brazil to treat Leishmania
braziliensis skin infections (Franca and others Rubber Research Institute of Malaya. 1973.
1996). On the other hand, Koster’s curse is a Clidemia hirta in South Johore. Planter’s
serious threat to understory plant species in fragile Bulletin 128: 140-144.
tropical island ecosystems (Pacific Island
Ecosystems at Risk 2002). It is a serious weed in Smith, C.W. 1992. Distribution, status, phenology,
tropical plantations such as rubber (Hevea rate of spread, and management of clidemia in
brasiliensis Muell. Arg.). Sheep have been shown Hawai’i. In: C.P. Stone, C.W. Smith, and J.T.
to control most weeds in plantations but will not Tunison, eds. Alien plant invasions in native
eat Koster’s curse (Chee and Faiz 2002). Goats ecosystems of Hawaii, management and
suffer toxicity from hydrolysable tannin when fed research. University of Hawaii Cooperative
the plant (Murdiati and others 1990). National Park Resources Studies Unit,
Honolulu, HI. p. 241-253.
References
State of Hawaii Department of Agriculture. 2002.
Binggeli, P. 1997. Clidemia hirta (L.) D. Don Plant pest control, persisting biological control
(Melastomataceae). http://members.lycos.co.uk/ problems. http://www.hawaiiag.org/hdoa/
WoodyPlantEcology/docs/web-sp5.htm. 4 p. pi_ppc_bioprob.htm. 6 p.
Chee, Y.K. and A. Faiz. 2002. Sheep grazing Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
reduces chemical weed control in rubber. Montiel, eds. 2001. A Flora de Nicaragua.
Australian Centre for International Agriculture Monographs in Systematic Botany Vol. 85, No.
Research. http://www.aciar.gov.au/publications/ 2. Missouri Botanical Garden Press, St. Louis,
proceedings/32/paper26.pdf. 4 p. MO. p. 945-1,910.
224
Coccoloba krugii Lindau wild-grape
POLYGONACEAE
Synonyms: none
John K. Francis

Islands, St. Martin, Anguilla, Barbuda, and
Antigua. A natural hybrid, Coccoloba krugii x
uvifera, with features intermediate between the two
parents is found in Puerto Rico, the Virgin Islands,
Anguilla, and Barbuda (Howard 1988, Liogier
1985, Little and others 1974).
Ecology.—Wild-grape grows on well-drained to

excessively drained, mostly slightly acid to mildly
alkaline soils in the whole range of soil textures. It
is found in Puerto Rico and the Virgin Islands from
near sea level to 125 m in elevation (Little and
others 1974) in areas that receive from 750 to 950
mm of annual precipitation. Wild-grape is
intolerant of shade. Because it grows slowly, in
order to compete it must grow on difficult sites
such as beach strands, dry coastal forests, shallow
rocky headlands, hillsides, and ridges. The species
is mostly found in remnant forests and sometimes
highly disturbed galleries. Wild-grape may occur
as individual plants, in groups, or in thickets.
General Description.—Wild-grape, also known Reproduction.—Wild-grape flowers in the fall

as white wood, crabwood, and bow-pigeon, is an and bears fruits during the winter (Little and others
evergreen shrub or small tree 1.5 to 6 m in height 1974). A collection of fruits from Puerto Rico
and stem diameters up to 10 cm. The plant is averaged 0.0985 + 0.0021 g/fruit. Air-dried seeds
supported by an extensive lateral root system of separated from them weighed an average of 0.0148
black roots. Older plants usually have multiple + 0.003 g/seed or 68,000 seeds/kg. However,
stems arising from the root crown. The smooth or sowing them in commercial potting mix with no
slightly fissured bark is gray. A dense, branchy pretreatment resulted in only 5 percent
crown is formed of trident or bifurcate branches germination. The seeds are dispersed by birds,
and slender twigs. Leaves have a reddish petiole 5 iguanas, and possibly mammals. Seedlings are not
to 6 mm-long, and reddish midvein. The blades are common. Plants sprout when cut or burned.
coriaceous, entire, ovate to suborbicular, and 2 to 6
cm long. This species is dioecious (male and Growth and Management.—Growth of wild-
female flowers on different trees). The flower grape is slow, perhaps 10 to 20 cm of height per
clusters are terminal spikes. Flowers are tiny, year. Plants are relatively long-lived and may
greenish-white, five-merous, and nearly stalkless. renew themselves by coppicing and continue
The fruits are fleshy and black, ovoid and clonally through more than one generation of
somewhat triangular in cross section, 4 to 5 mm crowns. No management experience has been
long, and have a mildly sweet flavor somewhat published. Planting any slow-growing species in
like grapes. The fruit juice stains skin and clothing. wildlands is a challenge; protection of remnant
Each fruit contains one dark brown akene (Howard forests is probably the best approach to promoting
1988, Liogier 1985, Little and others 1974). wild-grape.
Range.—Wild-grape is native to the Bahamas, Benefits.—Wild-grape wood is heavy and hard and
225
useful for at least fuel. The fruits are edible and International Reptile Conservation Foundation.
tasty, though small. The fruits are counted among 2002. Effects of introduced ungulates on the
the food sources of the endangered Anegada iguana iguana on Anegada. http://cyclura.com/
(Cyclura pinguis) (International Reptile article/articleview/248/1/6/. 6 p.
Conservation Foundation 2002).
References Rico and adjacent islands, Spermatophyta. Vol.
Howard R.A. 1988. Flora of the Lesser Antilles, Río Piedras, PR. 352 p.
Leeward and Windward Islands.
Dicotyledoneae, Part 1. Vol. 4. Arnold Little, E.L., Jr. R.O. Woodbury, and F.H.
Arboretum, Harvard University, Jamaica Plain, Wadsworth. 1974. Trees of Puerto Rico and the
MA. 673 p. Virgin Islands. Vol. 2. Agric. Handb. 449. U.S.
1,024 p.
226
Coccoloba microstachya Willd. pockhout
POLYGONACEAE
Synonyms: Coccoloba parvifolia Poir. in Lam.

Coccoloba klotzschiana Meisn. in DC.
Coccoloba microstachya var. ovalifolia Miesn. in DC.
Coccoloba microstachya var. lanceolata Meins. in DC.
Coccoloba microstachya var. rotundifolia Urb. ex Lindau
Coccoloba microstachya var. obtusifolia Lindau
John K. Francis
and generally rounded at both ends. Inflorescences

are terminal spikes. Male and female flowers are
borne on different shrubs (dioecious). The flowers
are greenish white and tiny. The egg-shaped fruits
are sessile, 6 mm long, nearly black or dark brown
when ripe, and contain one blackish seed (Liogier
Range.—Pockhout is native to Hispaniola, Puerto

Rico and its offshore islands, and the Virgin
Islands (Liogier 1985). Little and others (1974)
also list Anguilla as having a population. The
species in not known to have been planted or
Ecology.—Pockhout grows on a wide variety of

well- to excessively-drained soils over
sedimentary (especially limestone), igneous, and
metamorphic (especially ultramaphic) rocks. The
sites are usually hill tops, ridges, or upper slopes
with shallow and rocky soils. In Puerto Rico, this
habitat varies from 750 to about 1700 mm of mean
annual precipitation at elevations from near sea
General Description.—Pockhout, a name used in level to 760 m. The species is normally dispersed
the Virgin Islands, is known as uvillo, uverillo, throughout the secondary forest and does not form
and negra loca in Puerto Rico and Hispaniola. It is pure thickets (Weaver and others 1999). Pockhout
an evergreen or semideciduous shrub or small tree is moderately intolerant of shade. It grows in
usually 2 to 6 m in height and 2 to 7 cm in intermediate and codominant positions in open,
diameter at breast height. It is limby and twiggy low forests as well as dominating in more difficult
with a contorted form and often develops multiple sites.
stems from suckers on the lower stem. The wood
is light brown and hard. The plants are supported Reproduction.—Pockhout flowers during the
by a weak taproot and many, extensive, reddish- summer and fruits during the winter (Little and
brown, tough and flexible lateral roots. The fine others 1974). Fruit and seed production can be
roots have clearly visible mycorrhizal sheaths. The relatively heavy in open-grown trees. Fresh fruits
stem bark is gray, smooth, becoming deeply collected in Puerto Rico averaged 0.0517 + 0.0075
furrowed into plates in old individuals. The twigs g/fruit. Air-dried seeds separated from them
are brown to gray, smooth with lenticels. The averaged 70,400 seeds/kg. Sown without
light-green, alternate leaves have short petioles, 3 pretreatment on moist filter paper, 98 percent
to 6 mm long, have broadly elliptic to ovate- germinated beginning 16 days after sowing
lanceolate blades 3.5 to 7 cm long, entire edges (Francis and Rodríguez 1993). Seedlings are
227
uncommon.
Growth and Management.—Pockhout is slow Rico and adjacent islands, Spermatophyta. Vol.
growing and long lived. No published planting or 1. Editorial de la Universidad de Puerto Rico,
management experience is known to the author. Río Piedras, PR. 352 p.
Benefits.—Pockhout contributes to the Little, E.L., Jr., R.O. Woodbury, and F.H.
biodiversity of secondary and remnant forests, Wadsworth. 1974. Trees of Puerto Rico and the
helps protect the soil, and furnishes food and cover Virgin Islands. Vol. 2. Agriculture Handbook
for wildlife. The wood is good for fuel. 449. U.S. Department of Agriculture,
Washington, DC. 1,024 p.
References
Weaver, P.L., J.L. Ramirez, and J.L. Coll-Rivera.
Francis, J.K. and A. Rodríguez. 1993. Seeds of 1999. Las Cabezas de San Juan Nature Reserve
Puerto Rican trees and shrubs: second (El Faro). General Technical Report IITF-5.
installment. Res. Note SO-374. Southern Forest International Institute of Tropical Forestry, U.S.
Experiment Station, U.S. Department of Department of Agriculture, Forest Service, Río
Agriculture, Forest Service, New Orleans, LA. Piedras, PR. 58 p.
5 p.
228
Coccoloba uvifera (L.) L. seagrape
POLYGONACEAE
Synonyms: Coccolobis uvifera Jacq.

Polygonum uvifera L.
John A. Parrotta
Ecology.—Seagrape is one of the first species to

colonize sandy and rocky shores within its natural
range, and is rarely found at any great distance
from sea coasts (Little and Wadsworth 1964,
Adams 1972). It is a drought-hardy, salt-tolerant
species that grows in areas receiving between 500
and 1500 mm of rainfall with a dry season of up to
8 months (Von Carlowitz 1986). Its best growth
occurs on well-drained loamy sands that are
slightly to moderately alkaline in reaction. It can,
however, tolerate rocky, calcareous soils and soils
derived from igneous rocks, provided they are
freely drained (FAO 1982). A light-demanding
species, seagrape often forms dense thickets on
sand dunes, and is typically associated with a
number of other shrub, vine and tree species.
While most commonly found in dune shrub
formations, it does occur farther inland in coastal
woodland formations up to 150 m in elevation in
the Caribbean and elsewhere in its tropical and
subtropical American range (Stoffers, 1956, Craig
1984, Gooding 1974). Throughout its range, it is
General Description.—Seagrape, also known as susceptible to attack by a number of defoliating
seaside grape, bay grape, uva de playa, uvero, insect species, as well as leaf, stem, and root
raisin de la mer, and several other common names, pathogens, though none appear to pose a serious
is a large, low-branching shrub or small tree with threat to the species (USDA 1960, Spaulding
thick, smooth branches and a stout trunk. It is a 1961, Chellman 1978, Martorell 1975, National
coastal, tropical species easily recognized by its Research Council 1983, Parrotta 1994).
large, thick, nearly circular leaves and long,
grapelike clusters of edible, purple fruits. Reproduction.— Flowering and fruiting in
seagrape usually begins when trees are 6 to 8 years
Range.—Seagrape is widely distributed along the old. Its small (5 mm across), fragrant, greenish-
Atlantic, Caribbean and Pacific coasts of the white, male and female flowers, borne on separate
American tropics and subtropics between 25° N. trees, form terminal and lateral clusters 10 to 23
and 10° S. latitudes (Parrotta 1994). It is believed cm in length. Flowering and fruiting may occur
to be native to southern Florida, the Bahamas, and throughout the year, although flowering is
the West Indies, and is currently naturalized on generally concentrated between January and
both coasts of Mexico from Sinaloa and Veracruz August and fruiting between March and October in
southward through Central America and South its native Caribbean range (Little and Wadsworth
America to northern Peru and northeastern Brazil. 1964, Parrotta 1994). The flowers are pollinated
It has been introduced as an ornamental and by bees and other insects. The elliptic or egg-
seashore windbreak species in Hawaii and several shaped fruits, measuring about 2 cm in diameter,
tropical countries in the Asia-Pacific region are grouped in clusters resembling bunches of
(National Research Council 1983). grapes. The fruits ripen approximately 2 months
229
after flowering. The ripe, purple fruits contain a and dyeing (Little and Wadsworth 1964, Uphof
single, hard, elliptic seed about 1 cm long 1968). The astingent roots and bark have been
surrounded by an edible, tartly sweet pulp and a used in traditional medicine in Puerto Rico and
thin, fleshy covering. Fresh seed weights range elsewhere in the Caribbean (Liogier 1990).
from 1,000 to 1,400 seeds/kg, based on collections Seagrape’s tolerance of saline soils and sea spray
in Puerto Rico (Parrotta 1994). The fruits are has made it an excellent and popular choice for
consumed by a variety of birds that help to ornamental plantings and coastal windbreaks. It
disperse the seeds. Natural reproduction is prunes well and makes an attractive hedge.
generally good on open coastal sites, although
seedlings do not tolerate shading or competition References
from grasses and herbs. In the nursery, seagrape
can be propagated from seed in light-textured, Adams, C.D. 1972. Flowering plants of Jamaica.
well-drained soils; no pretreatment is required to University of the West Indies, Mona, Jamaica.
break seed dormancy. Seeds should be sown on or 848 p.
near the soil surface. Germination takes place
between 18 and 50 days after sowing, and Chellman, C.W. 1978. Pests and problems of
seedlings reach plantable size (15 cm tall) 4 to 5 South Florida trees and palms. Florida
months after sowing. It can also be propagated Department of Agriculture and Consumer
vegetatively by air-layering, ground-layering, and Services, Division of Forestry, Talahassee, FL.
grafting, though mature wood cuttings are most 103 p.
commonly used to produce rooted cuttings for
mass propagation of selected cultivars (FAO 1982, Craig, R.M. 1984. Plants for coastal dunes of the
Parrotta 1994). Gulf and South Atlantic coasts and Puerto Rico.
Information Bulletin 460. U.S. Department of
Growth and Management.—Growth rates in Agriculture, Soil Conservation Service,
seagrape are strongly influenced by wind exposure; Washington, DC. 41 p.
on exposed sites, heights typically increase from
the windward to leeward edges of stands. On a less FAO. 1982. Fruit-bearing forest trees: technical
exposed coastal site with sandy soils receiving an notes. FAO For. Pap. 34. Food and Agriculture
annual rainfall of approximately 1500 mm in Organization of the United Nations, Rome.
northeastern Puerto Rico, average heights of 0.9 177 p.
and 2.9 m were recorded for 1- and 3-year-old
plantings (Parrotta 1994). No information is Gooding, E.G.B. 1974. The plant communities of
available on growth rates of older trees. Under Barbados. Government Printing Offic,e
optimal conditions seagrape can attain a height of 8 Bridgetown, Barbados. 177 p.
to 18 m and a stem diameter of 45 to 60 cm
(Record and Hess 1943). Being shade-intolerant
and susceptible to competition from grasses, herbs
Rico y del Caribe. Iberoamericana de Ediciones,
and other woody species, planted stands should be
Inc., San Juan, PR. 566 p.
regularly weeded until they are taller than
competing vegetation.
Little, E.L., Jr. and F.H. Wadsworth. 1964.
Benefits.—Throughout its range, the wood is used Common trees of Puerto Rico and the Virgin
as firewood and for making charcoal. The reddish- Islands. Agriculture Handbook 249. U.S.
to dark-brown heartwood, distinguished from the Department of Agriculture, Forest Service,
light-brown sapwood in larger trees, is hard, close- Washington, DC. 548 p.
grained, and used in turning and occasionally for
furniture, inlay work and cabinetry (Gooding Martorell, L.F. 1975. Annotated food plant catalog
1974). The edible fruits can be eaten raw, used in of the insects of Puerto Rico. Agricultural
preserves, or fermented like grapes to make wine. Experiment Station, University of Puerto Rico,
The flowers yield abundant nectar, and the Río Piedras, PR. 303 p.
resulting honey is of good quality, light amber in
color, with a spicy flavor (National Research
Council 1983). The astringent red sap exuded from National Research Council. 1983. Firewood crops:
the cut bark, known commercially as West Indian shrub and tree species for energy production,
or Jamaican kino, was formerly used for tanning
230
Vol. 2. BOSTID Report 40. National Academy Stoffers, A.L. 1956. The vegetation of the
of Sciences, Washington, DC. 92 p. Netherlands Antilles. 15: Studies on the flora of
Curaçao and other Caribbean islands. Vol.
1.Nartinus Nijhoff, The Hague. 142 p.
Parrotta, J.A. 1994. Coccoloba uvifera (L.) L. –
Sea grape, uva de playa. Research Note SO-ITF-
Uphof, J.T.C. 1968. Dictionary of economic
SM-74. U.S. Department of Agriculture, Forest
plants. Verlag von J. Cramer, New York.
Service, Southern Forest Experiment Station,
591 p.
New Orleans, LA. 5 p.
USDA. 1960. Index of plant diseases in the United
Record, S.J. and R.W. Hess. 1943. Timbers of the
States. Agriculture Handbook 165. U.S.
New World. Yale University Press, New Haven,
CT. 640 p.
531 p.
Spaulding, P. 1961. Foreign diseases of forest trees

Von Carlowitz, P.G. 1986. Multipurpose tree and
of the world. Agriculture Handbook 191. U.S.
shrub seed directory. International Council for
Research on Agroforestry, Nairobi. 265 p.
361 p.
231
Coffea arabica L. coffee
RUBIACEAE
Synonyms: Coffea vulgaris Moench

Coffea laurifolis Salisb.
Coffea moka Hort. ex Heynh.
John K. Francis
and Wadsworth 1964, Wrigley 1988).
Range.—The original native population of coffee

was in the highlands of Ethiopia with possible
disjunct populations in nearby highland areas of
Sudan and Kenya. All of those areas have been
altered and under semicultivation for many years
(Charrier and Berthaud 1985). Coffee was first
cultivated by Arabs during the 14th century and
introduced into the New World and much of the
rest of the tropics during the 17th century (Smith
1985, Wrigley 1988). Today it is cultivated
throughout the moist subtropics and high-altitude,
moist tropics and has naturalized in many of these
areas including Puerto Rico, the Virgin Islands
(Little and Wadsworth 1964), Guam, and Samoa
(Pacific Island Ecosystems at Risk 2002).
Ecology.—The native range of coffee lies at 1,370

to 1,830 m in elevation (Wrigley 1988). In higher
latitudes, altitude becomes less critical. Optimum
temperatures range from 15 to 24 °C. Growth is
impaired above 25 °C. Frosts destroy both leaves
General Description.—Coffee, also known as and fruits (Willson 1985). The species requires a
café, is an upright, evergreen shrub or small tree up minimum of 1200 to 1500 mm of annual
to 5 m in height and 7 cm in diameter at breast precipitation. Precipitation in excess of 2500 to
height. The plant may grow with a single stem, but 3000 mm begins to be detrimental (Wrigley 1988).
often develops multiple stems by branching at the Although the species tolerates soil pH’s from 4 to
base or on the lower stem. The bark is light gray, 8, pH’s of 5.2 to 6.2 are preferred. Good drainage
thin, and becomes fissured and rough when old. is essential, and soil textures lighter than clays are
The wood is light-colored, hard, heavy, and tough. best (Willson 1985). Coffee is tolerant of shade and
The root system consists of a short, stout central usually grows in the forest understory. This
root, secondary roots radiating at all angles, and environment is simulated when coffee is grown
abundant fine “feeder” roots. The glabrous, shiny, under Inga vera Willd. and other shade species.
dark-green, opposite leaves have petioles 4 to 12 Also, it will grow well and is often cultivated in
mm long and ovate to elliptic blades 7 to 20 cm full sun. There are a number of serious pests and
long, with entire edges, and pointed at both ends. diseases of coffee in plantations; the effects appear
The fragrant, white flowers are in axillary clusters to be negligible in dispersed naturalized
of two to nine. The 1.0- to 1.8-cm drupes are populations.
ovoid, fleshy, green turning red and finally blue-
black. The fruits usually contain two greenish Reproduction.—Coffee usually flowers and fruits
seeds, 8 to 12 mm long, that are rounded and once per season, but in some areas it flowers twice.
flattened on one site with a medial groove. Coffee In Puerto Rico, flowering occurs January to April
is an allotetraploid with 2n = 44 chromosomes (Barrett 1925). It is reported to flower from
(Bailey 1941, Howard 1989, Liogier 1997, Little February through May in Nicaragua (Stevens and
232
others 2001). The flowers are insect pollinated 0.04 percent, stem 0.01 percent, and central root,
(Wrigley 1988). The fruits take from 6½ to 7 0.01 percent. Other alkaloids, xanthine, guanine,
months to mature (Bailey 1941). Seeds of the and trigonelline, which have stimulant and diuretic
varieties naturalized in Puerto Rico vary from properties, are also present (Burkill 1997). Caffeine
0.139 to 0.147 g/seed (Barrett 1925). Seeds 8 protects vegetative plant parts from insect and
weeks old germinate (95 percent in 32 days) better fungal attack and inhibits the growth of plants and
than fresh seed (Wrigley 1988). However, coffee bacteria near germinating seeds (Steiman 1997).
seeds stored more than 21 weeks at ambient Purified caffeine is widely sold as a medicinal
temperature begin to rapidly lose their viability stimulant, dietary aid, and headache remedy. Leaf
(Harrer 1963). The seeds are dispersed by birds, poultices are used to treat sores in Trinidad, and
and seedlings can be abundant. Artificial root sap or root infusions are drunk to relieve
propagation is usually by seed, but the species can scorpion stings (Burkill 1997). Coffee is also
be reproduced by budding and cuttings (Center for employed in folk medicine to treat asthma, flu,
New Crops and Plants Products 1996). headache, jaundice, nephrosis, malaria, sores, and
vertigo (Center for New Crops and Plants Products
Growth and Management.—Coffee has a 1996).
moderate growth rate. Plants begin bearing in 3 to
4 years and are in full fruit production in 6 to 8 References
years. Coffee plants may live over 100 years
(Center for New Crops and Plants Products 1996). Bailey, L.H. 1941. The standard cyclopedia of
Nursery plants are maintained in containers under horticulture. MacMillan, New York. 3,639 p.
shade for 6 to 12 months before outplanting
(Wrigley 1988). Also, plantations are established Barrett, O.W. 1925. The food plants of Puerto
by direct seeding into prepared seed spots (Center Rico. Journal of the Department of Agriculture
for New Crops and Plants Products 1996). Weed of Puerto Rico 9(2): 61-208.
control is necessary in the first years after
plantation establishment. Naturalized and escaped Burkill, H.M. 1997. The useful plants of West
coffee is usually not common enough or does not Tropical Africa. Vol. 4. Royal Botanic Gardens,
grow fast enough to warrant control. Kew, UK. 969 p.
Benefits.—As an exotic species in New World Center for New Crops and Plants Products. 1996.
forests, coffee has had a gentle impact on Coffea arabica L. Perdue University. http://hort.
biodiversity and contributes to wildlife food and perdue.edu/newcrop/duke_energy/Coffea_
cover, and soil stability. The wood is used mainly arabica.html. 7 p.
for fuel in the New World but is turned into chairs
and other types of furniture in Africa (Cheney Charrier, A. and J. Berthaud. 1985. Botanical
1925). Coffee is a good honey plant and yields a classification of coffee. In: M.N. Clifford and
light-colored honey (Little and Wadsworth 1964). K.C. Willson, eds. Coffee: botany, biochemistry,
Coffee berries, edible and slightly sweet, are eaten and production of beans and beverage. The AVI
occasionally by children and field workers. The Publishing Company, Inc., Westport, CN. p. 13-
fruit pulp, which is removed during processing, is 47.
sometimes fed to livestock but more often is
composted for fertilizer and mulch (Center for New Cheney, R.H. 1925. Coffee. The New York
Crops and Plants Products 1996). Coffee seeds University Press, New York. 244 p.
have been chewed as a stimulant in East Africa
from ancient times (Center for New Crops and Harrer, A.E. 1963. Coffee growing. Oxford
Plants Products 1996). The hot drink “coffee” is University Press, London. 127 p.
brewed from the roasted and ground seeds (or
“beans”) and is one of the world’s most popular Howard, R.A. 1989. Flora of the Lesser Antilles,
beverages. It is used to flavor candies, liquors, and Leeward and Windward Islands. Dicotyledoneae.
pastries. Probably the principal reason for its Part 3. Vol. 6. Arnold Arboretum, Harvard
popularity is the addictive stimulant alkaloid, University, Jamaica Plain, MA. 658 p.
caffeine (1,3,7-trimethylxanthine), 1.1 to 1.3
percent in the beans, but varying greatly in the Liogier, H.A. 1997. Descriptive flora of Puerto
beverage due to different brewing practices. The Rico and adjacent islands. Vol. 5. Editorial de la
alkaloid is present in the leaf at 0.30 percent, twig, Universidad de Puerto Rico, San Juan, PR.
233
436 p. Steiman, S. 1997. The effects of caffeine in Coffea
arabica L. http://www.grayskies.net/honeybear/
Little, E.L., Jr. and F.L. Wadsworth. 1964. arabica.htm. 2 p.
Islands. Agriculture Handbook 249. U.S. Stevens, W.D., C. Ulloa-U., A. Pool, and
Department of Agriculture, Washington, DC. O.M. Montiel, eds. 2001. Flora de Nicaragua.
548 p. Monographs in Systematic Botany Vol. 85,
No. 3. Missouri Botanic Garden Press, St.
Pacific Island Ecosystems at Risk. 2002. Coffea Louis, MO. p. 1,911-2,666.
arabica L., Rubiaceae. http://www.hear.org/
pier_v3.3/coara.htm. 3 p.
Willson, K.C. 1985. Climate and soil. In: M.N.
Clifford and K.C. Willson, eds. Coffee: botany,
Smith, R.F. 1985. History of coffee. In: M.N.
biochemistry, and production of beans and
Clifford and K.C. Willson, eds. Coffee: botany,
beverage. The AVI Publishing Company, Inc.,
biochemistry, and production of beans and
Westport, CN. p. 97-107.
beverage. The AVI Publishing Company, Inc.,
Westport, CN. p. 1-12.
Wrigley, G. 1988. Coffee. Longman Scientific
Technical and John Wiley & Sons, Inc. New
York. 639 p.
234
Colubrina arborescens (P. Mill.) Sarg. coffee colubrina
RHAMNACEAE
Synonyms: Colubrina colubrina (Jacq.) Millsp.

Colubrina ferruginosa Brongn.
Rhamnus colubrinus Jacq.
Ceanothus arborescens Mill.
Ceanothus colubrina (Jacq.) Lam.
John K. Francis
Range.—Coffee colubrina is native to southern

Florida, the Bahamas, the West Indies, and from
southern Mexico to Panama (Liogier 1994, Little
and Wadsworth 1964, Stevens and others 2001).
Although the species is planted as an ornamental,
it is not known to have become naturalized outside
its native range.
Ecology.—Coffee colubrina grows in a wide

variety of well-drained soils in its native range,
including soils of all textures, pHs ranging from
about 5.0 to 8.0, and soils derived from
sedimentary (including limestone), igneous, and
metamorphic (including ultramaphic) rocks. The
General Description.—Coffee colubrina, also species grows in areas of Puerto Rico that receive
known as greenheart, wild coffee, snakebark, from about 750 to 2500 mm of mean annual
blackbead tree, abeyelo, corazón de paloma, precipitation at altitudes from near sea level to
bijáguara, cascalata, sonzonate, and bois de fer, is about 600 m. Although the species is evergreen in
an evergreen or semideciduous shrub or small tree. moist sites, in dry areas, especially in excessively
It grows as a shrub with multiple stems 3 to 6 m in drained sites, it defoliates in response to prolonged
height in dry areas on beach fronts, deep sands, drought. Coffee colubrina grows on hammocks in
and porous limestone with little soil, and as a tree Florida (Nelson 1996) that have sandy soils
to 25 m in height on moist sites and fertile soils. (Gilman 1999), and in beach strand vegetation,
Bark on trunks is gray or brown, smoothish, coastal sands, dry to moist foothills and mountains
fissured, or platy. Inner bark is pink, brown, or in remnant and secondary forests in the West
reddish-brown. The roots are dark brown with Indies (author’s observation). The species tolerates
reddish inner bark and are somewhat brittle. The salt spray and moderate amounts of salt in the soil
twigs, flower clusters, young leaves, petioles, and (Gilman 1999). Coffee colubrina is moderately
underside of older leaves are covered with a rusty intolerant; it grows in full sun and partial shade of
pubescence. Leaves, which grow alternately in two low forest or broken high forest.
rows on twigs, are papery to leathery, ovate to
elliptic, 5 to 18 cm long, pointed at the tip and Reproduction.—Coffee colubrina is reported to
rounded at the base, and have prominent, curved bloom from spring to fall in Puerto Rico (Little
veins. The tiny yellow or yellow-green flowers and Wadsworth 1964) and throughout the year in
grow in short-stalked cymes in the leaf axils. Florida (Long and Lakela 1976). The flowers are
Fruits are globose capsules 6 to 10 mm in diameter insect pollinated. Seeds collected by the author in
that split into three parts and release globose, 3- to Puerto Rico averaged 70,400 seeds/kg (air-dried).
3.5-mm, hard, black seeds (Liogier 1990, Little Besides minor movement by gravity, wind, and
and Wadsworth 1964, Sargent 1923). water, the fruits pop open when dry and fling the
seeds a short distance (Gilman 1999).
Growth and Management.—Growth is slow in

dry forest areas and generally moderate in moist,
235
fertile soils. Four experimental plantings of coffee org/viewing/inyourbackyard/nativeplants.htm.
colubrina were made in Puerto Rico. Complete 11 p.
mortality occurred in the two drier, poorer sites,
but the trees grew 1 m/year in height for the first Francis, J.K. 1998. Tree species for planting in
few years on the better sites (Francis 1998). forest, rural, and urban areas of Puerto Rico.
Mortality probably occurred on the drier sites General Technical Report IITF-3. U.S.
because they were overcome by faster-growing Department of Agriculture, Forest Service,
competition. To ensure quick establishment, International Institute of Tropical Forestry, Río
planting of containerized stock for ornamental and Piedras, PR. 82 p.
wildland plantings is recommended. Protection
from weeds, vines, and faster-growing trees for 2 Gilman, E.F. 1999. Colubrina arborescens. Fact
or more years is imperative. Ornamental plants can Sheet FPS-137. University of Florida,
be forced into tree shape by continual pruning of Cooperative Estension Service, Gainesville, FL.
the lower branches (Gillman 1999). 3 p.
Benefits.—Coffee colubrina helps protect the soil, Liogier, H.A. 1990. Plantas medicinales de Puerto
contributes to the aesthetics of the forest, and Rico y del Caribe. Iberoamericana de Ediciones,
furnishes food and cover for wildlife. The plant Inc. 566 p.
attracts abundant insects including bees, wasps,
butterflies, and diurnal moths, which in turn Liogier, H.A. 1994. Descriptive flora of Puerto
provide food for warblers (Dendroica spp.), Rico and adjacent Islands. Vol. 3. Editorial de la
gnatchtchers (Polioptila spp.), kingbirds (Tyrannus Universidad de Puerto Rico. Río Piedras, PR.
spp.), and vireos (Vireo spp.) (Florida Fish and 461 p.
Wildlife Conservation Commission 2003). The
species is considered a honey plant, furnishing Little, E.L., Jr. and F.L. Wadsworth. 1964.
both nectar and pollen to honeybees (Apis Common trees of Puerto Rico and the Virgin
mellifera L.) May through September in the Islands. Agriculture Handbook 249. U.S.
Dominican Republic (Rivas-Laureano 2003). Department of Agriculture, Washington, DC.
Coffee colubrina is planted as a shade tree and 548 p.
ornamental in Florida, Guatemala, Nicaragua, and
El Salvador (Little and Wadsworth 1964, Stevens Long, R.W. and O. Lakela. 1976. A flora of
and others 2001). It is recommended for planting Tropical Florida. Banyon Books, Miami, FL.
in parking lot islands, in large containers, and 962 p.
small lawns, and for borders, screens, and hedges,
as well as for reclamation plantings (Gilman Nelson, G. 1996. The shrubs and woody vines of
1999). A tea made from the leaves and the wood is Florida. Pineapple Press, Inc., Sarasota, FL.
used as a remedy for rheumatism. Extracts are also 391 p.
used for antiseptic baths (Liogier 1990). The shiny
black seeds are used in necklaces in Jamaica. The Rivas-Laureano, S. 2003. Flora apícola
wood is hard and heavy with a specific gravity of Dominicana. http://www.apinetla.com.ar/rdom/
0.7. The sapwood is ivory or light brown, and the editorial-rdo/editorial1.htm. 2 p.
heartwood is yellowish brown. Because it is
resistant to decay, it is used for fence posts and Sargent, C.S. 1923. Manual of the trees of North
was formerly used for marine pilings (Little and America (exclusive of Mexico). Houghton
Wadsworth 1964). Mifflin, Boston, MA. 910 p.

Montiel, eds. 2001. Flora de Nicaragua,
Florida Fish and Wildlife Conservation Angiospermas. Monographs in Systematic
Commission. 2003. Native plants for backyard Botany Vol. 85, No. 3. Missouri Botanical
Florida habitats. http://www.floridaconservation. Garden Press, St. Louis, MO. p. 1,911-2,666.
236
Comptonia peregrina (L.) Coult. sweet fern
MYRICACEAE
Synonyms: Comptonia asplenifolia L.

Myrica asplenifolia (L.) Ait.
Paula M. Pijut
up to 0.7 cm long, with the ovary surrounded by

eight persistent bracts (Gleason and Cronquist
1991, Hall and others 1976). The fruit is an ovoid
nut, olive-brown, 3 to 5 mm long, enclosed in a
burr-like cluster of bracts (Dirr 1998, Gleason and
Cronquist 1991). The chromosome number of
sweet fern is 2n = 32 (Gleason and Cronquist
1991). Two varieties are recognized: var.
asplenifolia (L.) Fernald and var. tomentosa
A.Chev. (Krüssmann 1984, Missouri Botanical
Garden 2002).
Range.—Sweet fern occurs from Nova Scotia to

North Carolina, western South Carolina, and
northern Georgia, and west to Saskatchewan,
Minnesota, Illinois, and Tennessee (Gleason and
Cronquist 1991). Comptonia peregrina var.
asplenifolia occurs only from Long Island, New
York, to Virginia (Hall and others 1976). In
Canada, sweet fern is found in all three Maritime
Provinces, Ontario, and certain parts of Quebec
(Hall and others 1976).
General Description.—Sweet fern is a densely
branched, deciduous, rhizomatous shrub that Ecology.—Sweet fern prefers well-drained, dry,
grows 0.5 to 1.5 m tall and 1.2 to 2.5 m wide or acid, sandy or gravely soils with full exposure to
more (Dirr 1998, Gleason and Cronquist 1991). the sun (Del Tredici 1996, Schwintzer 1989,
Sweet fern is a dioecious or seldom monoecious Snyder 1993). These sites include dry piney
(Gleason and Cronquist 1991), actinorhizal woods, exposed mountain slopes, abandoned
nitrogen-fixing shrub (Del Tredici 1996, Ziegler pastures, barrens, highway embankments, gravel
and Huser 1963) with fern-like leaves and stems pits, weathered mine tailings, and cut-over forested
that are aromatic when crushed. Leaves are land (Del Tredici 1996). Sweet fern is shade
alternate, oblong-linear, deeply pinnately lobed, intolerant, adaptable to infertile soils, fixes
stipulate, dark green, pubescent, 5 to 12 cm long, 1 nitrogen in a symbiotic association with Frankia, a
to 1.5 cm wide, with petioles 0.3 to 0.6 cm long filamentous bacterium (actinomycete) (Callaham
(Dirr 1998, Gleason and Cronquist 1991, Hall and and others 1978), has cluster roots that aid in
others 1976). Stems are green, yellowish, or phosphorus uptake (Hurd and Schwintzer 1997),
reddish-brown when young, turning reddish-purple and is a pioneer colonizer of sites disturbed by fire,
or coppery-brown with age. Buds are globular, logging, clear-cutting, and road construction (Del
minute, solitary, and sessile, with two to four Tredici 1996, Lynham and others 1998, Snyder
exposed scales (Dirr 1998). The flowers are 1993). Sweet fern is drought and salt tolerant.
inconspicuous. Staminate catkins are olive green, Sweet fern colonizes areas via lateral roots that
cylindrical, clustered (usually three to four) on the form endogenous root buds from which shoot
tip of the previous year’s wood, 1.5 to 4 cm long, sprouts are readily formed (Louis and Torrey
and drooping (Gleason and Cronquist 1991, Hall 1991). The shrub is a serious weed problem in the
and others 1976). Pistillate catkins are dark red at commercial lowbush blueberry fields in Canada
the beginning of anthesis, subglobose at maturity, and New England (Hall and others 1976, Snyder
237
1993). The nymphs of the Saratoga spittlebug are readily formed (Louis and Torrey 1991). These
(Aphrophora saratogensis Fitch.) feed on sweet sprouts can spread extensively over large areas
fern but do not cause serious damage (Johnson and forming thickets. Juvenile stems (7.6 cm or less in
Lyon 1991). Sweet fern is susceptible to attack by length) root when treated with 3 g/l indole-3-
Botryosphaeria dothidea (Moug.:Fr.) Ces. & butyric acid and placed under mist (Dirr 1998).
DeNot. (causing dieback) and Gymnosporangium Root pieces (10 cm long x 0.2 cm wide, or 5 cm x
ellisii (Berk.) Ellis (alternate host for aecia stage) 1.3 cm) dug in late winter or early spring, placed
(Sinclair and others 1987). Sweet fern is the horizontally at a depth of 1.3 cm in a mixture of
alternate host of the fungus, Cronartium fine sand and sphagnum peat, will develop shoots
comptoniae Arth., that causes sweet fern blister and additional roots (Dirr 1998). Sweet fern can be
rust on hard pines (Farr and others 1989, Sinclair grown in containers and successfully transplanted
and others 1987). The fungus is not seriously (Dirr 1998). Micropropagation of sweet fern has
harmful to sweet fern, but it can reduce the growth been achieved by the induction of root buds in
and even cause death of susceptible pines, such as excised root culture (Louis and Torrey 1991).
jack (Pinus banksiana Lamb.), pitch (P. rigida
Mill.), shortleaf (P. echinata Mill.), and loblolly Growth and Management.—Sweet fern is a slow
(P. taeda L.) (Del Tredici 1996, Snyder 1993). to medium growing shrub that develops a broad,
flat-topped to rounded habit as it spreads and
Reproduction.—Sweet fern flowers are small, colonizes (Dirr 1998). It is hardy in zones two to
inconspicuous, wind-pollinated, and shed large six (USDA Plant Hardiness) and is sexually
amounts of pollen (Hall and others 1976). In the mature in 2 to 3 years (Snyder 1993). Burning and
United States the shrub blossoms in April or early disking following logging will stimulate the
May with fruit maturation by fall. In Canada, growth of sweet fern (Snyder 1993). Sweet fern is
sweet fern blossoms mid-May to mid-June with used for erosion control, low maintenance
fruit maturation from July through September plantings along highway embankments, and as
(Hall and others 1976). Hall and others (1976) naturalistic plantings under power company rights-
determined the mean number of staminate catkins of-way (Del Tredici 1996, Snyder 1993). Sweet
per shoot based on 42 shoots to be 6.43 + 0.48 fern can become weedy in pastures, old fields, and
(SE) and for the pistillate catkins 0.60 + 0.01 (SE). open woods, but can be controlled with herbicides,
The fruit of sweet fern is borne in a bur-like cluster such as dicamba, 2,4-D, and glyphosate (Hall and
of bracts containing on average 5.52 + 0.43 (SE) others 1976, Heyd and others 1987, Snyder 1993).
nuts per bur (Hall and others 1976). The pericarp
is composed of a thin, fleshy outer layer and a Benefits.—Sweet fern is planted as a landscape
hard, thick inner layer (Del Tredici and Torrey plant because of its fern-like, aromatic foliage and
1976). Seeds do not germinate readily but can stems. Because it fixes nitrogen, sweet fern is
remain viable (for as long as 70 years) buried in useful for rehabilitation of disturbed sites. Sweet
the soil (Del Tredici 1977). Sweet fern seeds are in fern has limited use as a food source and cover for
deep primary dormancy when they are shed from wildlife. The fruits are consumed by flickers,
the plant and become incorporated into the soil moose and deer browse sweet fern, and prairie
(Dow and Schwintzer 1999). They enter seasonal chickens and sharp-tailed grouse use it for nesting
cycles of secondary dormancy over time in which cover (Snyder 1993). The leaves can be used for
dormancy is induced during the summer. The potpourri and tea (Stokes 1981). The chemical
seeds are then released from this dormancy by betulin occurs in the leaves, root, and stem of
chilling during the winter. However, most buried sweet fern, and its biological activities have been
seeds still require exposure to strong daily described (Duke 1996).
temperature fluctuations before they germinate
(Dow and Schwintzer 1999). Strong temperature References
fluctuations indicate that shade cover has been
removed, which stimulates the seeds to germinate Callaham, D., P. Del Tredici, and J.G. Torrey.
in an open situation favorable for seedling 1978. Isolation and cultivation in vitro of the
establishment. Fresh seed will germinate (80 actinomycete causing root nodulation in
percent) after scarification and treatment with 500 Comptonia. Science 199: 899-902.
mg/l gibberellic acid (GA3) for 24 hours (Del
Tredici and Torrey 1976). Sweet fern propagates Del Tredici, P. 1977. The buried seeds of
itself vegetatively via lateral roots that form Comptonia peregrina, the sweet fern. Bulletin of
endogenous root buds from which shoot sprouts the Torrey Botanical Club 104: 270-275.
238
Del Tredici, P. 1996. Bulldozers and bacteria: the Johnson, W.T. and H.H. Lyon. 1991. Insects that
ecology of sweet fern. Arnoldia 56: 2-11. feed on trees and shrubs. Cornell University
Press, Ithaca, NY. 560 p.
Del Tredici, P. and J.G. Torrey. 1976. On the
germination of seeds of Comptonia peregrina, Krüssmann, G. 1984. Manual of cultivated broad-
the sweet fern. Botanical Gazette 137: 262-268. leaved trees and shrubs, Volume I, A-D. Timber
Press, Beaverton, OR. 448 p.
Dirr, M.A. 1998. Manual of woody landscape
plants, Their identification, ornamental Louis, I. and J.G. Torrey. 1991. In vitro clonal
characteristics, culture, propagation, and uses. multiplication of the actinorhizal plant
Stipes Publishing, Champaign, IL. 1,187 p. Comptonia peregrina. Plant Cell, Tissue and
Organ Culture 26: 89-96.
Dow, M.A. and C.R. Schwintzer. 1999. Seed
germination, seedling emergence, and seed bank Lynham, T.J., G.M. Wickware, J.A. Mason, and
ecology of sweet fern (Comptonia peregrina H. Krause. 1998. Soil chemical changes and
(L.) Coult.). Canadian Journal of Botany 77: plant succession following experimental burning
1,378-1,386. in immature jack pine. Canadian Journal of Soil
Science 78: 93-104.
Duke, J.A. 1996. Phytochemical Database, U.S.
Department of Agriculture, Agricultural Missouri Botanical Garden. 2002. w3TROPICOS,
Research Service, Beltsville, Maryland. Nomenclatural database. http://mobot.mobot.
http://www.ars-grin.gov/cgi- org/W3T/Search/vast.html. [not paged].
bin/duke/farmacy2.pl.
Schwintzer, C.R. 1989. All field-collected
Farr, D.F., G.F. Bills, G.P. Chamuris, and A.Y. actinorhizae examined on Comptonia peregrina
Rossman. 1989. Fungi on plants and plant and Myrica pensylvanica in Maine are spore
products in the United States. The American negative. Canadian Journal of Botany 67: 1,460-
Phytopathological Society, St. Paul, MN. 1,252 1,464.
p.
Sinclair, W.A., H.H. Lyon, and W.T. Johnson.
Gleason, H.A. and A. Cronquist. 1991. Manual of 1987. Diseases of trees and shrubs. Cornell
vascular plants of northeastern United States and University Press, Ithaca, NY. 575 p.
adjacent Canada. 2nd Edition. The New York
Botanical Garden, Bronx, NY. 910 p. Snyder, S.A. 1993. Comptonia peregrina. In: U.S.
Department of Agriculture, Forest Service,
Hall, I.V., L.E. Aalders, and C.F. Everett. 1976. Rocky Mountain Research Station, Fire Sciences
The biology of Canadian weeds. 16. Comptonia Laboratory. Fire Effects Information System.
peregrina (L.) Coult. Canadian Journal of Plant http://www.fs.fed.us/database/feis/plants/shrub/c
Science 56: 147-156. omper/index.html.
Heyd, R.L., R.L. Murray, and L.F. Wilson. 1987. Stokes, D.W. 1981. The natural history of wild
Managing Saratoga spittlebug in pine shrubs and vines, eastern and central North
plantations by suppressing sweet fern. Northern America. Harper and Row Publishers, New
Journal of Applied Forestry 4: 16-17. York. 246 p.
Hurd, T.M. and C.R. Schwintzer. 1997. Formation Ziegler, H. and R. Huser. 1963. Fixation of
of cluster roots and mycorrhizal status of atmospheric nitrogen by root nodules of
Comptonia peregrina and Myrica pensylvanica Comptonia peregrina. Nature 199: 508.
in Maine, USA. Physiologia Plantarum 99: 680-
689.
239
Conocarpus erectus L. button mangrove
COMBRETACEAE
Synonyms: Conocarpus procumbens L.

Conocarpus erectus var. procumbens (L.) DC.
Conocarpus erectus var. sericeus DC.
Conocarpus sericeus Jiménez
Conocarpus supinus Crantz
Terminalia erecta (L.) Baill.
John K. Francis
Inflorescences are terminal or axillary panicles of

tiny greenish-white flowers grouped in spheroidal
heads 3 to 5 mm in diameter. The thin, dry, 5- to
15-mm, two-winged seeds are densely packed into
globose clusters (Howard 1989, Liogier 1994,
Little and Wadsworth 1964, Nelson 1996,
Pennington and Sarukhan 1968, Stevens and others
2001).
Range.—Button mangrove is native to Bermuda,

both coasts of southern Florida, the Bahamas, the
West Indies, both coasts of Mexico, Central
America, South America (through Ecuador and
Brazil), and the Galapagos Islands. It is also native
to coastal areas of Tropical West Africa (Howard
1989, Little and Wadsworth 1964). The species has
been planted widely as an ornamental and has
naturalized in at least Hawaii (Pacific Island
Ecology.—Button mangrove is intolerant of shade.

It is almost always open-grown or in codominant
stands. It will tolerate competition by plants of
General Description.—Button mangrove is also
similar size but will not grow under the canopy of
known as buttonwood, sea mulberry, botoncillo,
taller trees. The species grows above the high tide
mangle prieto, mangle negro, palétuvier, and
line, along beaches and just landward of
mangue. The species is usually a shrub 1.5 to 4 m
Laguncularia racemosa (L.) Gaertn. and other
in height but can become a tree up to 20 m or more
mangroves. The soils in these situations are usually
in height. The root system consists mainly of
sandy or marly but sometimes loamy or clayey.
laterals and fine roots that are dark brown, weak
Button mangrove is particularly adaptable as an
and brittle, and have a corky bark. The plant
ornamental because it tolerates compacted soil, air
usually has an erect trunk or multiple trunks, but it
pollution, poor drainage, and drought (Gilman and
may assume a prostrate form and have limbs that
Watson 1993). It also tolerates salt water overwash
layer and become new individuals. The bark is gray
from storm surges and heavy salt spray. The
or brown, furrowed, fibrous, and moderately thin
species grows, although much less commonly, on
(about 8 mm). The inner bark is dark cream in
the lower floodplains of rivers and in upland
color. Stemwood (specific gravity of 1.0) is hard,
forests. Herbarium samples have been collected at
heavy, and strong. Branches are brittle. The twigs
elevations of 745 m in Costa Rica (Instituto
are slender, yellow-green, angled, flattened, or
Nacional de Biodiversidad 2002).
winged. The spirally arranged, elliptic to lanceolate
leaves are cartaceous to somewhat fleshy, 2 to 10
cm long, with petioles 3 to 9 mm long. Reproduction.—Button mangrove blooms
240
throughout the year in México (Pennington and Puerto Rican trees and shrubs: second
Sarukhan 1968) and from March to September in installment. Research Note SO-374. Southern
Florida (Nelson 1996). Tomlinson (1986) Forest Experiment Station, U.S. Department of
maintains that this species is dioecious. It is a Agriculture, Forest Service, New Orleans, LA.
consistent and abundant seed producer throughout 5 p.
most of its range. The fruit heads may contain from
35 to 56 fruits each (Masís and others 1999). Seeds Gilman, E.F. and D.G. Watson. 1993. Conocarpus
collected in Puerto Rico averaged 250,000 erectus, buttonwood. Fact Sheet ST-179. U.S.
seeds/kg. Seeds from this collection began Forest Service and Southern Group of State
germinating on moist filter paper in 9 days but Foresters, Gainesville, FL. 3 p.
were only 12 percent viable (Francis and
Rodríguez 1993). The seeds are dispersed by water Howard, R.A. 1989. Flora of the Lesser Antilles,
(Pacific Island Ecosystems at Risk 2002). Button Leeward and Windward Islands. Vol. 5. Arnold
mangrove can be propagated by partially burying Arboretum, Harvard University, Jamaica Plain,
large stakes in moist ground for living fence posts MA. 604 p.
(Little and Wadsworth 1964).
Instituto Nacional de Biodiversidad. 2002. Lista de
Growth and Management.—Button mangrove especímenes de Conocarpus erectus. http://
has a medium growth rate (Gilman and Watson www.inbio.ac.cr/bims/k03/p13/c045/o0264/f016
1993) and may live for several decades. 17/g007434/s021856.htm. 3 p.
Information is not available on wildland plantation
establishment. The planting of potted or Liogier, H.A. 1990. Plantas medicinales de Puerto
containerized seedlings seems the safest approach. Rico y del Caribe. Iberoamericana de Ediciones,
The species is not aggressive or invasive and only Inc., San Juan, PR 566 p.
requires control when land is converted to other
uses. Liogier, H. A. 1994. Descriptive flora of Puerto
Rico and adjacent Islands. Vol. 3. Editorial de la
Benefits.—Button mangrove provides food and Universidad de Puerto Rico, Río Piedras, PR.
cover for wildlife (various species of crabs and 461 p.
insects). It protects the soil during storm surges and
helps “fix” dunes (Popp and others 1989). Both Little, E.L., Jr. and F.H. Wadsworth. 1964.
button mangrove and silver buttonwood (C. erectus Common trees of Puerto Rico and the Virgin
var. sericeus Griseb.) are widely planted as Islands. Agriculture Handbook 249. U.S.
ornamentals in yards, parking lots, streets and Department of Agriculture, Forest Service,
parks (Gilman and Watson 1993, Nelson 1996). It Washington, DC. 548 p.
can be trained to form hedges. Potted plants are
used to form bonsai (Gilman and Watson 1993). Masís, A, D. Pérez, F. Chavarría, R. Espinoza, A.
Button mangrove wood is reported to be ideal for Guadamuz. 1999. Species page de Conocarpus
smoking fish and meat (Gilman and Watson 1993). erectus (Combretaceae), 14 April 1999. Species
The wood is durable but susceptible to dry-wood Home Pages. Area de Conservación Guanacaste,
termites [Cryptotermes brevis (Walker)] and is Costa Rica. http://www.acguanacaste.ac.cr/
used to make railroad ties, posts, for turnery, boat paginas_especie/plantae_online/magnoliophyta/
building, fuel, and charcoal. The bark and leaves combretaceae/conocarpus_erectus/c_erectus_14’
have been used in tannery (Little and Wadsworth ... 6 p.
1964). Bark from Belize contains 18 percent tannin
(Burkill 1985). Extracts of the bark are used to Nelson, G. 1996. The shrubs and woody vines of
treat bleeding gums, vaginal bleeding, colic, and Florida. Pineapple Press, Inc., Sarasota, FL.
skin ulcers (Liogier 1990). 391 p.
References Pacific Island Ecosystems at Risk. 2002.

Conocarpus erectus L., Combretaceae.
Burkill, H.M. 1985. The useful plants of West http://www.hear.org/pier_v3.3/coere.htm. 2 p.
Tropical Africa. Ed. 2. Vol. 1. Royal Botanic
Gardens, Kew, UK. 960 p. Pennington, T.D. and J. Sarukhan. 1968. Arboles
tropicales de México. Instituto Nacional de
Francis, J.K. and A. Rodríguez. 1993. Seeds of Investigaciones Forestales, Secretaría de
241
Agricultura y Ganadaría, México D.F., México. Stevens, W.D., C. Ulloa-U., A. Pool, and O.H.
413 p. Montiel, eds. 2001. Flora de Nicaragua.
Monographs of Systematic Botany Vol. 85, No.
Popp, M., U. Lüttge, W.J. Cram, M. Díaz, H. 1. Missouri Botanical Garden Press. p. 1-943.
Griffiths, H.J.S. Lee, E. Medina, C. Schäfer, K.-
H. Stimmel, and B. Thonke 1989. Water Tomlinson, P.B. 1986. The botany of mangroves.
relations and gas exchange of mangroves. New Cambridge University Press, Cambridge, UK.
Phytologist 111: 293-307. 419 p.
242
Corchorus hirsutus L. jack-switch
TILIACEAE
Synonyms: none
John K. Francis
Ecology.—Jack-switch is intolerant of shade and
grows in open areas or openings in low forest. It
cannot endure severe competition. The species
usually grows near the coast or inland on dry, often
excessively-drained sites. These are usually coastal
sands or rocky ridges and hillsides over limestone,
igneous, and metamorphic (including ultramaphic)
rocks. Jack-switch occurs as scattered plants or as
open stands with other species of similar size.
Because cattle do not eat or rarely eat the foliage,
the species benefits from over-grazing. It also
invades eroded and physically disturbed soils.
Reproduction.—Jack-switch blooms
continuously, except during periods of drought.
The species is insect pollinated (Marcano-Fondeur
1973). Capsules collected in Puerto Rico averaged
General Description.—Jack-switch, also known 28.0 seeds/capsule. Air-dried seeds from that
as wooly corchorus, mallet, cadillo, and collection averaged 1.109 million seeds/kg. Sown
malvavisco, is an upright woody to semiprostrate without pretreatment on the surface of wet peat, 10
shrub usually about 1 m in height but sometimes percent germinated over a 4-month period
reaching 2 m and 1.5 cm in basal diameter. It (author’s observation). Dispersal undoubtedly
usually has a single main stem and a somewhat occurs by wind and water; specialized means of
sparse, open crown. Plants examined by the author seed dispersal are unknown. Seedlings in wildland
had a deep taproot and fine lateral roots at all sites in Puerto Rico vary from common to rare.
levels. All parts except older branches and stems Plants apparently do not renew themselves by
are densely pale scruffy tomentulose with stellate resprouting after stem death. Reaction to fire is
hairs. The alternate leaves are ovate to oblong- unknown.
lanceolate, 2 to 6 cm long, have crenate to serrate
margins and obtuse to acute tips, and have petioles Growth and Management.—Jack-switch plants
2 to 7 mm long. The leaves, which have a gray- have a moderate to slow growth rate and live about
green color, tend to wilt rather than defoliate 5 years. Mechanical soil disturbance near seed
during dry seasons. Yellow flowers occur in groups sources and heavy grazing would probably lead to
of two to eight on axillary peduncles opposite the natural establishment of the shrubs. Jack-switch is
leaves. The four-chambered capsules are seldom common enough to warrant control, but if
ellipsoidal, about 12 mm long, and contain a needed, mowing may be sufficient.
number of 1.5- to 2-mm black seeds (Britton and
Millspaugh 1962, Howard 1989, Liogier 1994). Benefits.—Jack-switch furnishes cover for wildlife
and helps protect the soil. The plant is attractive
Range.—Jack-switch is native throughout the enough to be used as an ornamental in natural
West Indies and has been reported from Mexico landscaping, although that use is not yet reported.
through Central and South America and in It is a honey plant (Marcano-Fondeur 1973). The
northeastern Africa (Britton and Millspaugh 1962, leaves are cooked and eaten, and made into tea in
Howard 1989, Liogier 1994). Herbarium St. Croix, U.S. Virgin Islands. Branches were
specimens exist for Bolivia and Paraguay formerly used to make brooms (Garland 2002).
(Missouri Botanical Garden 2002). The species is
present as an exotic in the southern tip of Florida
(Institute of Systematic Botany 2002).
243
References http://www.plantatlas.usf.edu/main.asp?plantID
=4142. [not paged].
Britton, N.L. and C.F. Millspaugh. 1962. The
Bahama flora. The New York Botanical Garden, Liogier, H.A. 1994. Descriptive flora of Puerto
New York. 695 p. Rico and adjacent Islands. Vol. 3. Editorial de la
Universidad de Puerto Rico. Río Piedras, PR.
Garland, K. 2002. UVI Wetland Reserve plants: 461 p.
Corchorus hirsutus (jack-switch).
http://rps/uvi/edu/VIMAS/jackswitch.htm. 1 p. Marcano-Fondeur, E.J. 1973. La flora apícola de la
Republica Dominicana. http://marcano.
Howard, R.A. 1989. Flora of the Lesser Antilles, freeservers.com/nature/studios/apicola/dicotsp.
Leeward and Windward Islands. Vol. 5. Arnold html. 11 p.
MA. 604 p. Missouri Botanical Garden. 2002. W3-Specimen
Data Base: Current specimen list for Corchorus
Institute of Systematic Botany. 2002. Atlas of hirsutus. http://www.mobot.mobot.org/cgi-
Florida Vascular Plants: Corchorus hirsutus. bin/search_vast. [not paged].
University of South Florida, Tampa, FL.
244
Cordia lima (Desv.) Roemer & J.A. Schultes lija
BORAGINACEAE
Synonyms: Varronia lima Desv.

Cordia radula Spreng.
Cordia asperrima DC.
John K. Francis
precipitation) at elevations from near sea level to at

least 600 m. Colonized soils are variable in texture
with pH’s from about 5.0 to 7.5, well drained, and
usually rocky, on ridges, hillsides, and in draws.
Although the species grows over other rock types,
it is most common in limestone and ultramafic
(serpentine) areas. Lija is moderately intolerant of
shade and competes well with herbs and brush. It
grows under and at the edges of remnant forests
and middle to late secondary low forests with
relatively low basal areas. The species varies from
uncommon to relatively common in Puerto Rico.
Individual plants are usually widely scattered.
Reproduction.—A collection of fresh fruits from

Puerto Rico averaged 0.261 + 0.010 g/fruit. Air-
General Description.—Lija, which means dried seeds separated from them averaged 0.0515 +
sandpaper in Spanish, is an evergreen, vine-like, 0.0013 g/seed or 19,000 seeds/kg. Placed to
arching shrub 1 to 2.5 m in height and 1 to 2 cm in germinate on moist filter paper, 16 percent
basal diameter. Lija plants are supported by weak germinated within 6 months. Other germination
taproots, slender lateral roots, and abundant fine tests are needed to determine whether this is low
roots. The roots are dark brown. Older plants for the species. Pretreatments do not appear
usually have a number of stems that grow necessary. Apparently, the seeds are dispersed by
vertically for 1 to 2 m and then arch horizontally. birds. Seedlings in natural forests are uncommon.
Supported on other vegetation without twining or Established plants sprout when disturbed.
tendrils, they sometimes extend 4 m or more
laterally. The relatively few branches and twigs are Growth and Management.— Lija is a slow-
slender. The leaves are 2 to 6 cm long, stiff and growing shrub. Adult plants add only 10 to 30 cm
almost woody, ovate or elliptic, with toothed edges to their stem length annually. Individual stems
and a rough texture imparted by short, stiff hairs. appear to live 1 or 2 decades while plants, by
The inflorescences are terminal or lateral sprouting, may live much longer. No planting or
glomerules of small white subsessile flowers. management experience has been published for the
These produce globose one-seeded fruits (drupes) species. Protection from fire and development are
that are red when ripe and 4 to 5 mm in diameter. probably all that is required in relatively
The fruits have a mild, pleasantly sweet flavor undisturbed sites.
(author’s observation, Grisebach 1963, Liogier
1995). Benefits.—Lija contributes to the biodiversity in
forests where it lives, helps protect the soil, and
Range.—Lija is native to Hispaniola, Puerto Rico, furnishes food and cover for wildlife. The fruits are
and the Bahamas (Grisebach 1963, Liogier 1995). tasty but too small and too infrequently found to
It is not known to have been planted or naturalized justify collection.
elsewhere.
Ecology.—Lija grows in dry to moist habitat

(about 700 to 1800 mm of mean annual
245
References Liogier, H.A. 1995. Descriptive flora of Puerto
Grisebach, A.H.R. 1963. Flora of the British West Universidad de Puerto Rico, San Juan, PR.
Indian Islands. J. Cramer-Weinheim, New York. 617 p.
789 p.
246
Cordia polycephala (Lam.) I.M. Johnston black sage
BORAGINACEAE
Synonyms: Varronia polycephala Lam.

Varronia paniculata Wakström
Cordia wickstroemii Steudel
Cordia sulfurata Krause
Cordia ulmifolia Juss. ex Dumont
Lantana corymbosa L.
John K. Francis
have a serrate margin and are dark green above and

yellow green below. Whitish-green flowers are
grouped in tightly packed cymes arising near the
branch ends. The fruits (drupes) are red, globose,
2.5 to 5 cm in diameter, and have little flavor. The
seed has an irregular surface (Acevedo-Rodríguez
1985, Howard 1989, Liogier 1995).
Range.—Black sage is native to Hispaniola, Puerto

Rico, Vieques Island (Puerto Rico), the Virgin
Islands, St. Barts, St. Kitts, Guadeloupe, Dominica,
Martinique, St. Lucia, St. Vincent, Granada, and
South America as far south as Bahia, Brazil
(Centro Nordestino de Informações Sobre Plantas
2002, Howard 1989). It is not known to have been
planted or naturalized elsewhere.
Ecology.—Black sage most frequently grows in

abandoned fields and pastures, along roads, in early
secondary forest, in open forest on difficult terrain,
and in disturbed openings in old-growth forest. It
occurs in areas receiving from 1000 to 2500 mm of
annual precipitation, in soils of all textures and
parent materials at elevations from near sea level to
at least 600 m. The species does not occur in very
poorly drained soils. Black sage is not among the
early pioneers to colonize disturbed sites but enters
with brush that follows the weed and grass stage. It
competes well by scrambling across low vegetation
and climbing into the crowns of other shrubs and
General Description.—Black sage, also known as low trees. It is moderately intolerant to shade and is
basora, palo de perico, saraguero, saragüero, Santa able to grow under the canopy of low basal-area
María (Spanish), petit mahot, and mahot fin forest.
(French), is a scrambling shrub or woody vine
(Acevedo-Rodríguez 1985, Howard 1989, Liogier Reproduction.—As a population, black sage
1995). The single or multiple stems are gray, blooms and fruits all year, particularly during the
cylindrical, and strong with relatively few, slender summer (Acevedo-Rodríguez 1985). However,
branches. There may or may not be a weak taproot; individual plants usually flower at one time and
the plants rely on shallow lateral root systems with then rest for several months before flowering
abundant fine roots. Alternate scabrous leaves are again. The fresh fruits in one Puerto Rican
ovate to lanciolate and 8 to 15 cm long. The leaves collection averaged 0.12 g/fruit. The fruits are
247
consumed by birds that disperse the seeds. SO-58. U.S. Department of Agriculture, Forest
Seedlings are common but rarely abundant. Service, Southern Forest Experiment Station,
Growth and Management.—Black sage achieves
1 m or more of stem extension per year. Stems may Centro Nordestino de Informações Sobre Plantas.
grow to 1.5 cm of diameter and reach 5 m in height 2002. Base de dados CNIP, Checklist das
or extension from the point of rooting (Acevedo- plantas do Nordeste-Nomes provisórios-Versão
Rodríguez 1985). One 1.5-cm stem was observed 12: Cordia polycephala (Lam.) Johnst.
to have five growth rings. Black sage can live at Universidade Federal de Pernambuco.
least 5 years and perhaps much longer by http:150.161.125.13/pnechk/taxa/1057.shtml.
sprouting, which it readily does when injured. No 2 p.
planting, management, or control experience has
been published. In the absence of other control Howard, R.A. 1989. Flora of the Lesser Antilles.
information, grubbing out individual plants or spot Vol. 6. Arnold Arboretum, Harvard University,
treating with broad-leaf herbicides is Jamaica Plain, MA. 658 p.
recommended.
Benefits.—Black sage helps protect the soil and Rico and adjacent islands. Vol. 4. Editorial de la
furnishes food and cover for wildlife. The fruits are Universidad de Puerto Rico, San Juan, PR.
edible. It is a minor weed in pastures (Vélez and 617 p.
van Overbeek 1950).
Vélez, I. and J. van Overbeek. 1950. Plantas
References indeseables en los cultivos tropicales. Editorial
Universitaria, Río Piedras, PR. 497 p.
Acevedo-Rodríguez, P. 1985. Los bejucos de
Puerto Rico. Vol. 1. General Technical Report
248
Cornus sericea L. redosier dogwood
CORNACEAE
Synonyms: Cornus stolonifera Michx.
Paula M. Pijut
Fosberg and f. stolonifera (Michx.) Fosberg
(Missouri Botanical Garden 2002).
Range.— Redosier dogwood occurs from Alaska

to Newfoundland, south to Virginia in the east, to
Kansas in the Great Plains, to northern Mexico, in
the Rocky Mountains, and through California on
the West Coast (Crane 1989).
Ecology.—Redosier dogwood is extremely

adaptable to a wide range of soil and climatic
conditions. It prefers rich, moist, poorly drained
soils with high levels of mineral nutrients, and
moderate to full sunlight (Crane 1989). These sites
include swamps, low meadows, riparian zones,
wetlands, floodplains, forest openings and
understories. Redosier dogwood is suppressed in
shade, a dominant understory shrub in the early
successional willow and balsam poplar
communities, not a primary invader of wet prairies
and meadows (enters the stand later in succession),
and enters a stand after fire (Crane 1989). It is
Illustration source: USDA—Forest Service Collection, tolerant of fluctuating water tables and extremely
Hunt Institute cold temperatures. Redosier dogwood can spread
and form dense thickets (by layering) when stems
General Description.—Redosier dogwood is a touch the ground and roots form at the nodes. The
multi-stemmed, deciduous, stoloniferous shrub larvae (instars) of the dogwood sawfly
that grows 1 to 3 m tall and 3 m wide or more, [Macremphytus tarsatus (Say)] feed on redosier
often forming dense thickets. Leaves are opposite, dogwood causing defoliation (Johnson and Lyon
simple, ovate to oblong-lanceolate, acuminate, 1991). Pandemis pyrusana Kearfott (leafroller)
rounded at the base, 5 to 12 cm long, 2.5 to 6.4 cm attacks redosier dogwood in British Columbia and
wide, with petioles 1.3 to 2.5 cm long (Dirr 1998, California (Johnson and Lyon 1991). Redosier
Gleason and Cronquist 1991). Leaves are medium dogwood is susceptible to attack by Discula
to dark green in summer, purplish to reddish in the destructiva Redlin. sp. nov. (dogwood
fall, distinctly glaucous beneath, with five to seven anthracnose), Botryosphaeria dothidea
lateral vein pairs. Stems are slender, upright, bright (Moug.:Fr.) Ces. & De Not. (canker and dieback),
red to dark blood red, appressed pubescence on Septoria cornicola Desmaz. (leaf spot), and
younger stems, with prominent lenticels, and a Phyllactinia guttata (Wallr.:Fr.) Lev. (powdery
white pith (Dirr 1998). Buds are valvate, mildew) (Brown and others 1996, Jones and
appressed, and elongated. The inflorescence is a Benson 2001, Sinclair and others 1987). Tobacco
cyme (3.8 to 6.4 cm in diameter) with small (2 to 3 ringspot virus has been detected in naturally
mm), white to cream-colored flowers (Gleason and infected redosier dogwood (Sinclair and others
Cronquist 1991). The fruit is a globose, white 1987). Mycoplasma-like organisms have been
drupe (stone), 7 to 9 mm in diameter (Gleason and detected in witches-brooms of redosier dogwood
Cronquist 1991). The chromosome number of (Sinclair and others 1987).
redosier dogwood is 2n = 22 (Gleason and
Cronquist 1991). Two forms of C. sericea are Reproduction.—Redosier dogwood flowers are
recognized: f. baileyi (J.M. Coult and W.H. Evans) small, self-sterile with obligate outcrossing, and
249
are insect-pollinated (Crane 1989). In the United thick, extensive root system (Crane 1989).
States the shrub blossoms from late May to June Redosier dogwood is relatively tolerant to high
(June to August in the Northern United States) salinity tailings waters produced as a result of oil
with fruit maturation by fall. The fruit is a drupe and sand mining, making it useful for reclamation
containing a single two-celled and usually two- of sites (Renault and others 1999, 2001). The bark,
seeded stone, but only one seed is fully developed stems, leaves, roots, and fruit of redosier dogwood
(Brinkman 1974). Redosier dogwood seeds have a were used by Native Americans for a variety of
hard seed coat and dormant embryos (Crane 1989). uses including basket weaving, dyes, and tonics
Seeds do not germinate readily (can remain viable (Stevens and Dozier 2002). Redosier dogwood
in seedbanks) and require cold stratification and thickets provide cover for birds and other small
sometimes scarification in order for germination to wildlife. It is an important food source and winter
occur (Crane 1989). Fresh seed can be fall sown or browse for deer, elk, moose, bear, cottontail
stratified for 60 to 90 days at 5 ºC (Brinkman rabbits, snowshoe hares, grouse, quail, and other
1974). Acharya and others (1992) found that small birds and mammals (Crane 1989).
native (Alberta, Canada) redosier dogwood seed
germination was strongly influenced by the References
population from which seeds were collected, and
the yearly precipitation in those areas. Redosier Acharya, S.N., C.B. Chu, R. Hermesh, and G.B.
dogwood propagates itself vegetatively via stolons Schaalje. 1992. Factors affecting red-osier
(stems just on or under the ground) and by dogwood seed germination. Canadian Journal of
layering forming dense thickets. Plants are also Botany 70: 1,012-1,016.
produced from the roots, the bases of dying
branches, and injured stems (Crane 1989). Brinkman, K.A. 1974. Cornus L. dogwood. In:
Cuttings (with leaves) root readily when treated C.S. Schopmeyer, tech. coord. Seeds of woody
with 1 g/l indole-3-butyric acid (Dirr 1998). plants in the United States. Agriculture
Hardwood cuttings root (90 to 100 percent) when Handbook 450, U.S. Department of Agriculture,
placed in the field in late winter without a rooting Forest Service, Washington, D.C. p. 336-342.
treatment (Dirr 1998). Redosier dogwood can be
grown as bare-root, containerized, or balled-and- Brown, D.A., M.T. Windham, and R.N. Trigiano.
burlapped plants (Dirr 1998). 1996. Resistance to dogwood anthracnose
among Cornus species. Journal of Arboriculture
Growth and Management.—Redosier dogwood 22(2): 83-86.
is a fast (vigorous) growing shrub that develops a
loose, broad-spreading, rounded habit with Crane, M.F. 1989. Cornus sericea. In: U.S.
horizontal branches at the base (Dirr 1998). It is Department of Agriculture, Forest Service,
hardy in zones 2 to 7 (USDA Plant Hardiness) and Rocky Mountain Research Station, Fire Sciences
is sexually mature in 3 to 4 years (Crane 1989). Laboratory. Fire Effects Information System.
Redosier dogwood generally increases following http://www.fs.fed.us/database/feis/plants/shrub/c
fire (Crane 1989). Coppicing stimulates new, omper/index.html.
vigorous stem growth. Pruning and burning
redosier dogwood helps produce long straight Dirr, M.A. 1998. Manual of woody landscape
stems. Redosier dogwood is used for soil plants, their identification, ornamental
stabilization, reclamation sites, residential characteristics, culture, propagation, and uses.
landscape plantings, mass plantings along highway Stipes Publishing, Champaign, IL. 1,187 p.
banks, parks, and golf courses (Crane 1989, Dirr
1998, Renault and others 2001). Redosier Gleason, H.A. and A. Cronquist. 1991. Manual of
dogwood can be controlled by spraying with vascular plants of Northeastern United States
mixtures of 2, 4-D and dicamba (Crane 1989). and adjacent Canada. 2nd Edition. The New York
Botanical Garden, Bronx, NY. 910 p.
Benefits.—Redosier dogwood is planted as a
landscape plant because of its bright red stems, Johnson, W.T. and H.H. Lyon. 1991. Insects that
which are especially beautiful contrasting against a feed on trees and shrubs. Cornell University
snowy landscape. Because it is easy to establish Press, Ithaca, NY. 560 p.
and grows rapidly, redosier dogwood is useful for
rehabilitation of disturbed sites. It is an excellent Jones, R.K. and D.M. Benson. 2001. Diseases of
soil stabilizer along stream banks because of its woody ornamentals and trees in nurseries. The
250
American Phytopathological Society, St. Paul, plants to high salinity oil sands tailings. Journal
MN. 482 p. of Environmental Quality 28: 1957-1962.
Missouri Botanical Garden. 2002. w3TROPICOS, Sinclair, W.A., H.H. Lyon, and W.T. Johnson.
Nomenclatural database. http://mobot. 1987. Diseases of trees and shrubs. Cornell
mobot.org/W3T/Search/vast.html University Press, Ithaca, NY. 575 p.
Renault, S., C. Croser, J.A. Franklin, and J.J. Stevens, M. and I. Dozier. 2002. Redosier
Zwiazek. 2001. Effects of NaCl and Na2SO4 on dogwood. In: U.S. Department of Agriculture,
red-osier dogwood (Cornus stolonifera Michx) Natural Resources Conservation Service, The
seedlings. Plant and Soil 233: 261-268. PLANTS Database, National Plant Data Center,
Baton Rouge, LA. http://plants.usda.gov. [not
Renault, S., E. Paton, G. Nilsson, J.J. Zwiazek, and paged].
M.D. MacKinnon. 1999. Responses of boreal
251
Coursetia caribaea (Jacq.) Lavin añil falso
FABACEAE
Synonyms: Cracca caribaea (Jacq.) Lavin

Galega caribaea Jacq.
Tephrosia caribaea (Jacq.) DC.
Benthamantha caribaea (Jacq.) Kuntze
Brittonamra caribaea (Jacq.) Kuntze
Tephrosia aniloides Bello
John K. Francis
yellow-green. The racemes contain one to six pea-

like flowers that are white or yellowish, sometimes
lined with pink or purple. The pods are linear, 4 to
6 cm long and somewhat compressed between
seeds. The 10 to 25 seeds are about 2 mm long and
have squared ends (Howard 1988, Liogier 1988).
The diversity of the species across its range has
resulted in the description of nine varieties
(International Plant Index 2001).
Range.—Añil falso is native to northern Mexico

through Central America and South America to
Paraguay, and to Jamaica, Hispaniola, Puerto Rico,
the Virgin Islands, the Lesser Antilles, and
Trinidad in the Caribbean (Liogier and Martorell
2000).
Ecology.—Añil falso grows at elevations from

near sea level to over 400 m in Puerto Rico. These
areas receive from about 750 mm to about 1200
mm of annual precipitation. The species grows in a
wide variety of well-drained soils that originate
from igneous and sedimentary (including
limestone) rocks. Añil falso may be found in brush
lands, secondary forests, and remnant forests. It is
General Description.—Añil falso is a low shrub, moderately intolerant of shade. While it normally
usually about 0.6 m in height, but sometimes grows in the understory, it requires plenty of
reaching 2 m in height and 1 cm or more in basal filtered light. It can tolerate moderate competition
diameter. The common name is Spanish from from shrubs, herbs, and grass, but cannot survive in
Puerto Rico and means false indigo. Añil falso is heavy grass swards. The leaves are shed during the
supported by a tap and lateral root system with a dry season. In relatively undisturbed habitat in
moderate amount of fine roots. The species Puerto Rico, plants may grow as densely as one per
produces round to elongated root nodules (Dubey m2. Añil falso disappears from areas that are
and others 1972). The plant has a slender single overgrazed.
stem that usually resprouts with a single stem
when the plant is damaged. Plants grazed by cattle Reproduction.—Of a collection made in Puerto
may be stunted or contorted. The current year’s Rico, air-dry pods of añil falso weighed an average
mature growth is tender, but the stem is woody and of 0.073 + 0.005 g/pod. Eighteen pods were
normally without branches. The leaves are 4 to 12 threshed and yielded an average of 12.1 seeds/pod
cm long, have short petioles, and 9 to 18 opposite, with a minimum of two and a maximum of 19.
elliptic leaflets, 0.6 to 3.0 cm long. The foliage is These seeds weighed an average of 0.0028 +
252
0.0000 g/seed. They were sown without References
pretreatment on filter paper and yielded only 10
percent germination in 206 days. After mechanical Dubey, H.D., R. Woodbury, and R.L. Rodríguez.
scarification, 98 percent of the remainder of the 1972. New records of tropical legume
seeds germinated within 12 days. Grazing animals nodulation. Botany Gazette 133(1): 35-38.
are the principal seed vectors. Seedlings are
common and well-disbursed. Large plants are Howard R.A. 1988. Flora of the Lesser Antilles,
prolific seed producers. Even small and suppressed Leeward and Windward Islands.
plants produce a few seeds. Dicotyledoneae, Part 1. Vol. 4. Arnold
Arboretum, Harvard University. Jamaica Plain,
Growth and Management.—Añil falso grows up MA. 673 p.
to about 0.5 m per year in height. Plants live from 2
to several years. Wise management of the species International Plant Index. 2001. International plant
must preclude all but light grazing. Plantations for names index query: Coursetia caribaea
range improvement or environmental restoration www.ipni.org/ipni/query-ipni.html. 17 p.
could probably be established by sowing scarified
seed into cultivated understory sites before or at the Liogier, H.A. 1988. Descriptive flora of Puerto
beginning of the rainy season. Rico and adjacent islands, Spermatophyta. Vol.
Benefits.—Añil falso, when available, is grazed Río Piedras, PR. 481 p.
heavily by cattle. The species improves the soil by
fixing nitrogen. Liogier, H.A. and L.F. Martorell. 2000. Flora of
Puerto Rico and adjacent islands. Second ed.,
Rev. Editorial fe la Universidad de Puerto Rico,
253
Crataegus douglasii Lindl. black hawthorn
ROSACEAE
Synonyms: Crataegus brevispina Doug. ex Steud.

Crataegus brockwayae Sarg.
John K. Francis
Ontario, Washington, Oregon, California, Nevada,

Utah, Idaho, Wyoming, Montana, North Dakota,
South Dakota, Minnesota, Wisconsin, and
Michigan (Borialforest.org 2003, Natural
Resources Conservation Service 2003, Soper and
Heimburger 1982). There are two recognized
varieties. Crataegus douglasii var. douglasii
occupies nearly all the range (Natural Resources
Conservation Service 2003) and var. duchenensis
is found in a small area in Utah (Welsh and others
1987). Former varieties rivularis and suksdorfii are
now recognized as separate species (Natural
Resources Conservation Service 2003).
Ecology.—Black hawthorn grows mostly in forest

understories but also occurs in the open, often in
pure thickets. It is both a riparian species and
grows in upland sites where adequate moisture is
available. Slopes may be level to steep. Elevations
vary from 670 to 1,646 m. Soils vary greatly but
typically are deep and fine textured. The species
usually does not occupy recently disturbed sites.
One exception is that it is common in avalanche
General Description.—Black hawthorn, also tracks in Glacier National Park. Although the
known as Douglas hawthorn, river hawthorn, species grows along streams, it seldom grows in
western thorn apple, and Douglas thorntree, is a flood-prone areas. Black hawthorn burns readily
thorny deciduous shrub or small tree 1.5 to 9 m in but is somewhat fire tolerant due to sprouting and
height. It may have multiple stems from the base suckering. However, the extent of its stands and
or a single stem that begins branching just above thickets in eastern Washington has been limited by
the ground. The bark of older stems is smooth to frequent fires (Habeck 1991).
scaly and gray to brown; twigs are hairless, shiny,
slender, and reddish, and support stout, slightly Reproduction.—Black hawthorn flowers open in
curved spines up to 3 cm long. Leaves are May (Sargent 1923) or June (Soper and
alternate, thin, lanceolate to obovate, serrate to Heimburger 1982). The flowers are insect
doubly serrate, and 1.5 to 9.5 cm long and almost pollinated (Hansen 2003). The fruits ripen and fall
as wide. Showy, five-petaled, white flowers about in August through September (Sargent 1923).
12 mm wide with 10 pink stamens are borne in Fruits in southeastern Washington averaged 4.78
terminal corymbs. The flowers emit a somewhat seeds/fruit (Habeck 1991). Seeds make up about
fishy odor. Fruits are small (12 mm in diameter), 15 percent of the weight of ripe fruits. There are
apple-like black pomes with thick, light-yellow from 47,000 to 52,000 cleaned seeds/kg
pulp and three to five nutlets (Borialforest.com (Brinkman 1974). The species is difficult to
2003, Hansen 2003, Sargent 1923, Viereck and reproduce from cuttings (Hansen 2003). Black
Little 1972, Welsh 1974). hawthorn sprouts readily when cut and may sucker
from the roots (Habeck 1991).
Range.—Black hawthorn is native to southern
Alaska, British Colombia, Alberta, Saskatchewan,
254
Growth and Management.—Growth of newly References
established seedlings is slow (Habeck 1991).
Particulars on natural stand management are Borialforest.com 2003. Shrub species of the
lacking. Fruits are ripe when they are black and World’s boreal forests: Crataegus douglasii,
lustrous (Habeck 1991) and must be picked by black hawthorn. http://www.borealforest.org/
hand or clipped with a pruning pole. Pulp should world/herbs_shrubs/black_hawthorn.htm. 2 p.
be removed from the seeds by maceration and then
seeds should be air dried before sowing or storage. Brinkman, K.A. 1974. Crataegus L., hawthorn. In:
Seeds should be scarified with H2SO4 and C.S. Schopmeyer, tech. coord. Seeds of woody
stratified at 1 °C for 84 to 112 days before sowing. plants in the United States. Agriculture
Brinkman (1974) found that germination reached Handbook 450. U.S. Department of Agriculture,
about 30 percent in 35 to 45 days. Another source Forest Service, Washington, DC. p. 356-360.
cites 50 to 80 percent germination (Habeck 1991).
Dumroese and others (1997) report germination Dumroese, R.K., K.M. Hutton, and D.L. Wenny.
from 20 to 80 percent for lots treated identically. 1997. Propagating woody riparian plants in
Seedlings of most hawthorn species develop long nurseries. In: T.D. Landis and J.R. Thompson,
taproots and should not be kept in the nursery for tech. coords. National proceedings, Forest and
more than 1 year (Habeck 1991). Pruning should Conservation Nursery Associations. Gen. Tech.
be avoided and final target height of seedlings Rep. PNW-GTR-419. Pacific Northwest
should be 25 cm (Dumroese and others 1997). Research Station, U.S. Department of
Agriculture, Forest Service, Portland, OR. p. 71-
Benefits.—Black hawthorn contributes to the 76.
beauty of the riparian vegetation, helps protect the
soil from streambank and overflow erosion, and Habeck, R.J. 1991. Crataegus douglasii. In: U.S.
offers benefits to fauna. Although not as “showy” Department of Agriculture, Forest Service,
as some other hawthorns, the species is hardy and Rocky Mountain Research Station, Fire Sciences
has been used to a limited extent as an ornamental. Laboratory, Fire Effects Information System.
It makes excellent hedges and natural barriers, and http://www.fs.fed.us/database/feis/plants/shrub/
is planted as a wildlife cover and food plant, and cradou/all.html. 15 p.
for stream bank stabilization (Hansen 2003).
Livestock readily eat black hawthorn foliage but Hansen, W.W. 2003. Native plants of the
the thorny structure of shrubs and small trees Northwest: black hawthorn, Douglas hawthorn,
makes it difficult for them to utilize it well. They western thorn apple, and Douglas thorntree
prefer shrubs less than 1 m in height which they (Crataegus douglasii). http://www.nwplants.
may hedge. Mule deer also browse it. With 3.7 com/plants/trees/rosaceae/crataegus_douglasii/
percent protein, 3.8 percent fat, 19.3 percent fiber, index. html. 5 p.
and 4.0 percent ash, the species is moderately
palatable to browsing animals. Deer, upland game Moerman, D.E. 1986. Medicinal plants of Native
birds, song birds, and rodents eat the fruits. Black America. Technical Reports 19. University of
hawthorn makes excellent escape and nesting Michigan Museum of Anthropology, Ann
cover for wildlife (Habeck 1991). The fruits are Arbor, MI. 534 p.
made into pies, jellies, and jams (Hansen 2003).
Native Americans ate the fruits, sometimes dried Natural Resources Conservation Service. 2003.
and mixed with fat and other ingredients. The Plant profile: Crataegus douglasii Lindl., black
wood is hard and heavy and was used for tools and hawthorn. http://plants.usda.gov/cgi_bin/
weapons (Washington State Department of plant_profile.cgi?symbol=CRDO2. 5 p.
Transportation 2003). Native Americans used the
thorns for piercing ears, lancing boils, and making Sargent, C.S. 1923. Manual of the trees of North
fish hooks (Hansen 2003). They prepared a America (exclusive of Mexico). Houghton
poultice of chewed leaves to relieve swelling and Mifflin, Boston, MA. 910 p.
took decoctions of sap, bark, wood, and root for
stomach problems (Moerman 1986). Soper, J.H. and M.L. Heimburger. 1982. Shrubs of
Ontario. Royal Ontario Museum, Toronto,
Ontario, Canada. 495 p.
255
Viereck, L.A. and E.L. Little, Jr. 1972. Alaska
trees and shrubs. Agriculture Handbook 410. Welsh, S.L. 1974. Anderson’s flora of Alaska.
U.S. Department of Agriculture, Forest Service, Brigham Young University Press, Provo, Utah.
Washington, DC. 265 p. 724 p.
Washington State Department of Transportation. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
2003. Crataegus douglasii, black hawthorne. Higgins. 1987. A Utah flora. Great Basin
http://www.wsdot.wa.gov/environment/eao/ Naturalist Memoirs 9. Brigham Young
culres/ethbot/a-c/Crataegus.htm. 2 p. University Press, Provo, UT. 894 p.
256
Critonia portoricense (Urban) Britt. & Wilson guerrero
ASTERACEAE
Synonyms: Eupatorium portoricense Urban
John K. Francis
and the adjacent island of Vieques (Liogier 1997,

Little and others 1974). It is not known to have
been planted or naturalized elsewhere.
Ecology.—Guerrero grows on nearly all well-

drained soil types derived from sedimentary and
igneous rocks (including limestone and
ultramafics). Rooting is deep when soil permits,
but the typical sites where the species grows are
shallow over bedrock. The species is also found on
poor and eroded soils. It grows from near sea level
to 900 m in elevation (Little and others 1974).
Mean annual rainfall ranges from about 850 to
2200 mm. Although present in openings and rock
outcrops in late secondary and remnant forests,
guerrero is frequently common in early and middle
secondary forests. It invades neglected pastures.
Minor soil disturbance such as trampling by cows
appears necessary for establishment.
Reproduction.—Guerrero mainly flowers from

November to February and matures fruits from
December to March (Little and others 1974). Plants
General Description.—Guerrero, which means receiving full or nearly full sun are heavy seed
warrior in Spanish, is a large shrub or small tree to producers. The seeds are wind-disbursed and may
6 m in height and 15 cm in diameter at breast travel considerable distances. A collection of seeds
height. Older plants usually have multiple stems, averaged 0.00025 g/seed or about 4,000,000
arising from the root crown or lower stem. The seeds/kg. Seedlings are rare to common, but never
wood is brittle and has a central pith 2 or 3 mm abundant. Resprouting occurs after adult plants are
thick. Plants may or may not have well-defined burned or cut.
taproots. Lateral roots are robust and send down
sinker roots. The roots are tan-colored and stiff. Growth and Management.—Guerrero grows at a
Stem bark is thin, gray, and nearly smooth. The similar rate as associated species in the difficult
inner bark of guerrero is light green and bitter. habitat where it occurs. Sprouts add about 0.5 m of
Branches and twigs are not numerous, resulting in height per year for the first 4 or 5 years. Life span
thin crowns. The hairless leaves are opposite, 7 to appears to be 2 to 3 decades. No planting
16 cm long with short petioles, ovate-elliptic experience has been published.
blades with serrated or wavy-toothed edges, and
long-pointed tips. The foliage is fragrant. The Benefits.—Guerrero is browsed by goats but
inflorescences are heads within large, terminal mostly ignored by cattle. The species is sometimes
corymbs with tiny white flowers that produce, at planted as an ornamental for its fragrant foliage
maturity, tiny (3 mm long) achenes tipped with 3- (Little and others 1974). Three pyrrolizidine
mm white hairs (pappus) (Liogier 1997, Little and alkaloids were isolated from the aerial parts of the
others 1974). shrub (Wiedenfeld and others 1995). As a
medicinal plant, extracts from its leaves are said to
Range.—Guerrero is found only in Puerto Rico relieve arthritis, and to relieve pain when applied as
257
a topical alcoholic rub (Liogier 1990). Guerrero Liogier, H.A. 1990. Plantas medicinales de Puerto
contributes to biodiversity of the forests in which it Rico y del Caribe. Iberoamericana de Ediciones,
grows, furnishes cover and nesting sites for birds, Inc., San Juan, PR. 566 p.
and helps protect and stabilize soil in disturbed
areas. Little, E.L., Jr., R.O. Woodbury, and F.H.
Liogier, H.A. 1997. Descriptive flora of Puerto Washington, DC. 1,024 p.
Universidad de Puerto Rico, San Juan, PR. Wiedenfeld, H., R. Guerrero, and E. Roeder. 1995.
436 p. Pyrrolizidine alkaloids from Eupatorium
portoricense. Planta Medica 61(4): 380-381.
258
Crossopetalum rhacoma Crantz maidenberry
CELASTRACEAE
Synonyms: Rhacoma crossopetalum L.

Myginda rhacoma Sw.
Myginda rotundata Lam.
Myginda pallens J.E. Sm. in Rees
Crossopetalum pallena (J.E. Sm.) Kuntze
Rhacoma pallens (J.E. Sm.) G. Maza
Myginda latifolia M. Vahl
John K. Francis
contains a stone and one or two seeds (Howard

1989, Liogier 1994, Little and others 1974).
Range.—Maidenberry is native to Southern

Florida, the West Indies, and Venezuela (Howard
1989, Liogier 1994, Little and others 1974,
Alarcón 2001). Howard (1989) and Liogier (1994)
state that is found in Mexico and Central America,
but the species is not listed in the current check
lists for countries in the region.
Ecology.—Maidenberry is usually found in rocky

and dry or excessively drained sites. It tolerates a
wide range of well-drained soil types derived from
metamorphic (including ultramafic) rocks. In
Puerto Rico, maidenberry grows from near sea
level to 400 m in elevation in areas that receive
from 750 to about 1600 mm of precipitation. It is
moderately intolerant of shade and withstands only
moderate competition from grasses and shrubs. The
species will survive but not prosper under low
General Description.—Maidenberry is also basal-area dry forest. The roots support vascular-
known as Florida crossopetalum, rhacoma, poison- arbuscular mycorrhizae (TreeGuide Inc. 2002).
cherry, coral, manto, maravedí, palo de paloma, Maidenberry is listed as endangered in Florida
pico de paloma, limonejo, membrillo, ti bonbon, (Florida Fish and Wildlife Conservation
bonbon rouge, and placa chiquitu. It is an Commission 1997); however, it is relatively
evergreen shrub sometimes becoming a small tree. common in Puerto Rico and several West Indian
It commonly measures 1 to 3 m in height but may islands (Little and others 1974). In Florida, it is
reach 8 m. The shrub is supported by a major found in pinelands and occasionally hammocks
taproot and significant lateral roots with yellow (Nelson 1996). In the West Indies, maidenberry
bark. Maidenberry usually has multiple stems from grows in remnant and secondary forests, most often
the root crown and just above, and many branches. on rocky ridges and hillsides. The species’
Three shoots form at each node, one of which continued presence in overgrazed areas suggests
usually dies. The opposite or ternate leaves have a that it is not palatable to cattle.
petiole about 1 mm long, lanceolate to obovate
leathery blades usually with wavy-toothed edges. Reproduction.—Maidenberry flowers and fruits
The tiny greenish-red flowers are borne in axillary year-round as a process associated with new stem
cymes. The fruits that follow are fleshy, egg- growth (Nelson 1996). Birds disperse the seeds.
shaped drupes that ripen to a bright red. Each Despite almost constant fruit production, seedlings
259
are not common. Plants readily sprout when cut or Howard, R.A. 1989. Flora of the Lesser Antilles,
burned. Leeward and Windward Islands. Vol. 5. Arnold
Growth and Management.—Maidenberry grows MA. 604 p.
slowly and may live for several decades. No
management experience has been published.
International Reptile Conservation Foundation.
However, cattle grazing to reduce competition
2002. Effects of introduced ungulates on the
from grass, herbs, and vines will probably
iguana of Anegada. http://www.cyclura.com/
encourage reproduction and survival.
article/articleview/248/1/6/. 12 p.
Benefits.—Maidenberry furnishes food and cover
for wildlife and protects the soil. It is listed as one
of the food plants for the endangered Anegada
Inc., San Juan, PR 566 p.
iguana, Cyclura pinguis (International Reptile
Conservation Foundation 2002). The wood of
maidenberry is light brown and hard, but is used
only for fuel. All the plant tissues, especially the
Universidad de Puerto Rico. 461 p.
roots, are diuretic and are used to promote
urination and to treat kidney and bladder infections
(Liogier 1990).
References
Alarcón, C. 2001. Paraguana xeric scrub
(NT1313). World Wildlife Fund.
http://worldwildlife.org/wildworld/profiles/terre
strial/nt1313_full.html. 14 p.
391 p.
Florida Fish and Wildlife Conservation
TreeGuide Inc. 2002. Florida crossopetalum,
Commission. 1997. Florida’s endangered
Crossopetalum rhacoma Crantz, Celestraceae.
species, threatened species and species of
http://www.treeguide.com/nn/Species.asp?
special concern. http://floridaconservation.org/
SpeciesID=345&Region=NorthAmerican. 2 p.
pubs/endanger.html. [not paged].
260
Crotalaria lotifolia L. cascabelillo axilar
FABACEAE
Synonyms: Crotalaria lotifolia L. var. eggersii Senn

Crotalaria lotifolia L. var. grandiflora Urban
Crotalaria lotifolia L. var. grandifolia Urban
John K. Francis
(pods) are 2 to 3 cm long and 6 mm wide and

contain a few tan seeds that rattle when dry
suggesting the Spanish name cascabelillo, which
means little rattlesnake (author’s observations,
Range.—Cascabelillo axilar is native to the

Bahamas, the Greater Antilles, St. Barts, Dominica,
Martinique, the Grenadines, Barbados, Mexico,
and Honduras (Howard 1988, Liogier 1988).
Ecology.—In Puerto Rico, cascabelillo axilar

grows from a few meters above sea level to about
500 m elevation on the southern slopes of the
Sierra Central. These areas receive from 750 to
about 2200 mm of mean annual precipitation. It
grows on a variety of soils derived from igneous,
sedimentary (including limestone), and
metamorphic (including ultramaphic) rocks
(Breckon 2002). Cascabelillo axilar is moderately
intolerant of shade. It grows in openings and in the
understory of low basal-area forest, often in rocky
and excessively drained areas. The forests are
remnants and middle to late secondary. The species
does not tolerate competition from tall grass swards
and thick herb and brush growth.
Reproduction.—Cascabelillo axilar probably

General Description.—Cascabelillo axilar is a flowers near the end of the wet season. Plants with
suffruticose or relatively short-lived shrub usually mature pods were observed by the author in Puerto
1 m in height and 6 mm in basal diameter but Rico in March. Seed production can be abundant.
sometimes reaching 2 m in height and more than 1 Eighteen pods collected in Puerto Rico averaged
cm of basal diameter. It usually has a single stem, 6.7 seeds per intact pod. An unknown seed insect
stiff and straight in its lower portions, which had attacked some of the pods. Seeds collected
branches sparingly. The wood is moderately hard from the same area averaged 0.0086 + 0.0008
and tough. The branches are slender. Stem bark is g/seed or 117,000 seeds/kg. Sown on moist blotter
yellowish-green and striated. The plant is paper without pretreatment, 35 percent germinated
supported by a tap and lateral root system with in 5 months. Seeds from the same batch,
moderately stiff roots with yellow bark. Foliage is mechanically scarified, all germinated in 12 days.
concentrated near the top of the plant. Leaves are Germination is epigeal. Seed pods spring open
trifoliate on long (2 to 6 cm) petioles. Leaflets are upon drying flinging the seeds a short distance.
light green or yellow-green, oblong to elliptic, and Grazing animals probably also move seeds.
1 to 5 cm long. The inflorescences are few- Seedlings are relatively common on sites where the
flowered, axillary racemes whose unequal flowers species is present.
have a yellow corolla 5 to 7 mm long. The legumes
261
Growth and Management.—Once established, References
cascabelillo axilar adds about 0.5 m in height each
year and loses most of it to die-back the following Breckon, G.J. 2002. Preliminary checklist for
dry season. Individual plants probably live 1 to 3 Sierra Bermeja. http://www.uprm.edu/biology/
years or more. No management experience has profs/breckon/herbarium/FLORABERMEJA.
been published. Scarified seed sown and htm. [not paged].
incorporated into prepared ground or seed spots at
the start of the wet season may offer a good chance Howard R.A. 1988. Flora of the Lesser Antilles,
of success. Leeward and Windward Islands.
Dicotyledoneae, Part 1. Vol. 4. Arnold
Benefits.—Cascabelillo axilar contributes in a Arboretum, Harvard University. Jamaica Plain,
minor way to standing biomass, cover for wildlife, MA. 673 p.
the aesthetics of the forest understory, and it helps
protect the soil. The species is browsed by cattle Liogier, H.A. 1988. Descriptive flora of Puerto
and consequently is more common in protected Rico and adjacent islands, Spermatophyta. Vol.
areas. Cascabelillo axilar is present in moderately 2. Editorial de la Universidad de Puerto Rico,
grazed areas but absent from seriously overgrazed Río Piedras, PR. 481 p.
areas.
262
Crotalaria spectabilis Roth showy rattlebox
FABACEAE
Synonyms: Crotalaria leschenaultii DC.

Crotalaria macrophylla Weinm.
Crotalaria retzii Hitchc.
Crotalaria sericea Retz.
John K. Francis
5 to 17 cm long, entire, and have a 2-mm petiole

and broad, triangular stipules. The inflorescences
are terminal or subterminal racemes with 20 to 25
flowers with linear-triangular bracts. The bright
yellow unequal flowers are 1.5 to 2 cm long.
Inflated brown to black legumes are 3 to 5 cm long
and 1.8 to 2 cm thick. They contain several hard,
shiny brown to black seeds 4.5 mm long (author’s
observation, Damron and Jacob 2001. Howard
1988, Liogier 1988, Stevens and others 2001).
Range.—Showy rattlebox is native to the Indo-

Malaysia area (Parrotta 2001). It has been planted
widely and has naturalized in many tropical
countries including the Southern United States,
Hawaii, and Puerto Rico (International Legume
Database and Information Service 2002).
Ecology.—Not much is known about showy

rattlebox in its natural habitat. It grows on
roadsides and disturbed areas in Florida (Long and
Lakela 1976), and in Puerto Rican coastal sands
and limestone rubble in areas receiving about 1650
mm of annual precipitation (author’s observation)
and at the edge of mangroves (Liogier 1988).
Parrotta (2001) reports that it is common along
stream banks in deciduous forests in India. The
General Description.—Showy rattlebox, also species is a weed of agricultural crops in the
known as showy crotalaria and rattlebox, is an Southern United States (Bradley and Hagood
annual semiwoody herb in temperate areas, and a 2002). It is intolerant of shade and does not grow
short-lived shrub in its frost-free subtropical and under a forest canopy. The species seems to favor
tropical range. In Puerto Rico, it grows up to 2 m in sandy soils (McGregor 1976), grows well in
height and 3.5 cm in basal diameter. It normally alkaline soils, and at least tolerates moderate soil
has a single stem. The lower stems are covered salinity. Plants are often parasitized by dodder
with whorled, short, shelf-like old leaf bases and (Cuscuta spp.) in Puerto Rico. Showy rattlebox is
stipule remnants. The leaves and fruiting branches well nodulated with nitrogen-fixing bacteria
are shed progressively upward after they mature so (McGregor 1976).
that foliage and branches are only found on the
upper portion of the stem. The mid- and lower stem Reproduction.—Showy rattlebox blooms all year
has a white, brittle, medium-hard wood with a 1.5- in Florida (Long and Lakela 1976) but is reported
mm pith. The upper stem, branches, and foliage are to flower in April and May and fruit in October in
green to yellow-green. The plant is supported by a Nicaragua (Stevens and others 2001) and flower
tap and lateral system of stiff, tan roots. The between November and January and fruit from
simple, whorled leaves are oblanceolate to elliptic, December to February in India (Parrotta 2001).
263
The flowers are visited by honey bees and other Crotalaria spectabilis. http://ppws.vt.edu/scott/
insects (McGregor 1976). A collection of seeds weed_id/cvtsp.htm. 2 p.
from Puerto Rico averaged 0.0154 + 0.0000 or
65,000 seeds/kg. Only 3 percent of unscarified seed Burkill, H.M. 1995. The useful plants of West
placed on moist blotter paper germinated in 6 Tropical Africa. Vol. 3. Royal Botanic Garden,
months. However, 99 percent of scarified seed Kew, UK. 857 p.
Germination is epigeal. When sufficiently dry, Damron, B.L. and J.P. Jacob. 2001. Toxicity to
pods spring open, flinging the seeds a short poultry of common weed seeds. University of
distance. Florida, Cooperative Extension Service.
http://edis.ifas.ufl.edu/BODY_PS052. 7 p.
Growth and Management.—Showy rattlebox has
a moderate to fast growth rate and may reach a Howard R.A. 1988. Flora of the Lesser Antilles,
meter of height by 1 year old. Plants appear to live Leeward and Windward Islands.
2 or 3 years in Puerto Rico. Parrotta (2001) refers Dicotyledoneae, Part 1. Vol. 4. Arnold
to the species as biennial. Plantations are Arboretum, Harvard University. Jamaica Plain,
established by sowing scarified seed into prepared MA. 673 p.
ground.
International Legume Database and Information
Benefits and Detriments.—Showy rattlebox is a Service. 2002. ILDIS Legumes of the world:
beautiful plant and contributes to the aesthetics of Crotalaria spectabilis Roth. http://ildis.org/
wildlands and gardens where it is planted. It serves LegumeWeb/6.00/taxa/4077.shtml. 4 p.
as a nurse species during early reforestation and
helps protect the soil. Showy rattlebox is used as a Jain, S.K. and R.A. De Filipps. 1991. Medical
green manure crop, particularly where rotation for plants of India. Vol. 1. Reference Publications,
control of nematodes is important (Zago and others Inc., Algonac, MI. 408 p.
2002). It has also been grown as a fodder plant.
This has largely been abandoned because of the Liogier, H.A. 1988. Descriptive flora of Puerto
toxic alkaloids (principally monocrotaline) it Rico and adjacent islands, Spermatophyta. Vol.
contains. Concentrations are reported as: leaves 2. Editorial de la Universidad de Puerto Rico,
0.008 percent, seedpods 0.366 percent, and seeds Río Piedras, PR. 481 p.
1.958 percent (Burkill 1995). Chickens, turkeys,
and quail are very sensitive. Concentrations of Long, R.W. and O. Lakela. 1976. A flora of
crotalaria alkaloids between 0.01 and 0.1 percent of Tropical Florida. Banyan Books, Miami, FL.
the diet have adverse effects and 0.3 percent is fatal 962 p.
(Damron and Jacob 2001). Horses, cattle, and
swine are also sensitive. Sheep, goats, and dogs are McGregor, S.E. 1976. Insect pollination of
more resistant. Effects can be both acute and cultivated crop plants. Internet printing by the
chronic and include bloody diarrhea, anemia, USDA Agriculture Research Service, http://
jaundice, hair loss, and unthriftiness. There is no gears.tucson.ars.ag.gov/book/chap9/crotalaria.
known treatment (Alabama Cooperative Extension html. [not paged].
Service 2002). In herbal medicine, extracts of the
whole plant are used to treat impetigo and scabies Parrotta, J.A. 2001. Healing plants of Peninsular
(Jain and De Filipps 1991), as an antiseptic for India. CABI Publishing, Oxon, UK and New
cuts, and to treat intestinal worms (Parrotta 2001). York. 917 p.

Alabama Cooperative Extension Service. 2002. Monographs in Systematic Botany Vol. 85, No.
Poisonous plants of the Southeastern United 1. Missouri Botanical Garden Press, St. Louis,
States: Showy crotalaria Crotalaria spectabilis. MO. p. 945-1,910.
http://www.aces.edu/dept/extcomm/publications
/anr/anr-975/Showy%20crotalaria.pdf. 1 p.
Bradley, K. and S. Hagood. 2002. Virginia Tech

weed identification guide: Showy crotalaria:
264
Zago, C., V.P. de Melo and W.R. Maluf. 2002. A
aplicação da adubação verde em hortaliças.
Boletim Técnico de Hortaliças 16.
Departamento de Agricultura, Universidade
Federal de Lavras. http://www2.ufla.br/
~wrmaluf/bth016/bth016.html. 5 p.
265
Cuphea hyssopifolia Kunth false heather
LYTHRACEAE
Synonyms: Parsonia petiolata (L.) Rusby

Cuphia petiolata Koehne
John K. Francis
Range.—False heather is native from Mexico to

Panama (Howard 1989, Liogier 1994). It has been
planted widely and has naturalized and escaped in
at least Hawaii (Pope 1968), Puerto Rico (Liogier
1994), Montserrat, and St. Lucia (Howard 1989).
Ecology.—A moderate amount of disturbance is

required for false heather to establish and maintain
itself. Mowing, animal and vehicular traffic, and
light cultivation are typical enabling disturbances.
A moist climate (above 1400 mm of annual
precipitation) is required for natural stands. It does
not tolerate frost or salt (Watkins and Sheehan
1975). Partial shade from trees and light
competition from grass and weeds are tolerated,
but tall grass and heavy weeds will eliminate it. In
Hawaii, false heather is most abundant in moist and
partially shaded places above 240 m elevation
(Pope 1968). Clayey and medium textured soils
with good drainage are best. Japanese beetles
(Popillia japonica Newman) sometimes defoliate it
in Hawaii (Pope 1968).
Reproduction.—Flowering and seed production

General Description.—False heather is sometimes takes place constantly. The seeds are tiny (1.3
called cuphia, clammy cuphia, tarweed, elfin herb, million per kg) and difficult to collect. The period
or simply, heather (Griffiths 1992, Pope 1968). It is between flowering and seed formation is short, and
a woody, much branched shrub, although the seeds are expelled from the fruits at maturity. A
diminutive in size. While Howard (1989) states germination test of 94 seeds on filter paper resulted
that the species may reach 70 cm in height and in only two seeds germinating, over a period of 14
Turner and Wasson cite 60 cm as maximum height, days. Seedlings are not uncommon on moist,
25 to 30 cm are more typical (Liogier 1994). A disturbed soil. False heather also reproduces by
definite leader is maintained, but many of the layering whenever horizontal branches touch the
lateral branches grow along the ground surface. ground. Both seedlings and layers are important in
The bark is reddish brown and stringy. The root the maintenance of natural stands. Obtaining new
system consists of a short taproot and many laterals plants from seed is difficult. Nursery plants are
of equal thickness. The tertiary roots are fibrous. reproduced by cuttings; tips will root any time of
The leaves are subsessile, 1 to 3 cm long and year (Watkins and Sheehan 1975). Artificial
linear, lanceolate, or oblanceolate. Young stems reproduction by air and ground layers is also
and the underside of leaves are pubescent. The possible.
small white, pink, violate, or reddish purple
flowers appear in the axils of new leaves (Turner Growth and Management.—False heather is a
and Wasson 1997). There 5 to 8 ovules in each short-lived plant except when layering prolongs its
flower, and generally six 1-mm seeds per fruit life. After about 1 year, most plants begin a decline
(Howard 1989). in vigor and eventually die. Renewal of decadent
266
older plants can be promoted by mowing at about 5 Howard, R.A. 1989. Flora of the Lesser Antilles,
cm height, which cuts away the old plant structure Leeward and Windward Islands. Vol. 5. Arnold
from the newly layered branches. Nursery plants Arboretum, Harvard University, Jamaica Plain,
should be watered during dry periods or as needed MA. 604 p.
and given frequent light fertilizer applications.
False heather is susceptible to nematodes. Use of Liogier, H.A. 1994. Descriptive flora of Puerto
nematode-free media is recommended (Watkins Rico and adjacent Islands. Vol. 3. Editorial de la
and Sheehan 1975). Universidad de Puerto Rico. Río Piedras, PR.
461 p.
Benefits.—False heather has been cultivated
widely as an ornamental. It is used as a potted Pope, W.T. 1968. Manual of wayside plants of
plant, for low borders, and as ground cover in small Hawaii. Charles E. Tuttle Co, Publishers,
beds. False heather may become a weed in warm Rutland, VT. 289 p.
climates (Turner and Wasson 1997) and has
become a serious weed in some parts of Hawaii Turner, R.G., Jr. and Wasson, E. 1997. Botanica.
(Pope 1968). Barns and Noble, Inc. New York. 1,020 p.
References Watkins, J.V. and T.J. Sheehan. 1975. Florida

landscape plants, native and exotic. The
Griffiths, M. 1992. Index of garden plants. Timber University Press of Florida, Gainsville, FL.
Press, Portland, OR. 1,234 p. 420 p.
267
Curatella americana L. chaparro
DILLENIACEAE
Synonyms: none
Ariel E. Lugo and Ernesto Medina
Leaf morphology contributes to reduction

of heat loads and prevents overheating (Medina
1982). Stomata are slightly elevated and the leaf
epidermis is heavily cutinized (Mérida and Medina
1967). Leaves of chaparro are scleromorphic, with
low specific leaf area (<100cm2/g) and low
nutrient concentrations (Mérida and Medina 1967,
Medina 1977, 1982, Sarmiento and Monasterio
1983).
Throughout its range, chaparro is most
commonly associated with the following species:
trees such as Bowdichia virgilioides and
Byrsonima crassifolia; and grasses and sedges
such as Leptocoryphium lanatum and
General Description.—Chaparro in English is Trachypogon plumosus; and species of
known as the rough-leaf tree and it is also known Andropogon, Axonopus, Bulbostylis, Paspalum,
as careicillo, peralejo macho, vacabuey, aperalejo, and Rhynchospora (Sarmiento 1983). In Belize,
curatela, peralejo, peralejo de sabana, pomme chaparro grows in pine forests (Standley and
torche, raspa viejo, tlachicón, hoja man, yaha, Record 1936). Pollen from chaparro appeared in
lengua de vaca, malcajaco, hoja chigüe, raspa, Suriname and Colombia in strata dated at 45,000
raspa guacal, chumico de palo, hoja chigüe, yahal, years before present at the beginning of the
saha, chumico, azufre, curata, paricá, chaparro de Pleistocene and during glacial times (Van der
sabana, chaparro sabanero, chaparro colorado, Hammen 1983). Chaparro pollen was associated
parica, feuille rude, curatahie, carne de fiambre, with grass pollen and dry periods and has shown
caimbe, cajueiro bravo, sambaiba, caimbahiba, cycles of abundance and disappearance in response
cambaiba, sobro, sabaibinha, and chaáco (Liogier to climate change. The presence of chaparro pollen
1983, 1990, Standley and Record 1936, Salas in lake sediments established the evolution of
Estrada 1993, Siegel and Paguaga 1991, Vargas savanna vegetation in the Neotropics before the
Ulate 2001, Standley 1920, 1937, Pérez Arbelaez presence of humans (Sarmiento 1983). Chaparro
1978, Record and Mell 1924, Bastos 1984, Kileen pollen also established the boundary between rain
and others 1993). Curatélla means “plants being forests and savannas in Bolivia and the southern
used or worked”, for example, for polishing expansion of the rain forest boundary over the past
weapons and metal (Bailey 1941). It is an 3,000 years in response to climate change (Mayle
evergreen woody shrub less than 50 cm tall to a and others 2000), and vegetation changes in
tree 6 to 10 m in height (Lundell 1937, Liogier southeastern Brazil and Colombia since the
1990, Killeen and others 1993). Its most frequent Holocene and late Quaternary (Behling 1995,
habit is that of a gnarled shrub or small tree Behling and Hooghiemstra 1999).
(Sarmiento and Monasterio 1983).
In a family dominated by lianas, chaparro Range.—Chaparro has a large geographic range
is a twisted, thick-barked shrub/tree with that includes Cuba (Grisebach 1963); Hispaniola;
pubescent young branches (Schnee 1960). Leaves the states of Tepic, Tabasco, Chiapas, Veracruz,
are simple, alternate, oval to elliptic oval, 5 to 12 Campeche, Oaxaca, and Guerrero of México
cm wide and 10 to 15 cm long, rounded to (Standley 1920); Honduras; Belize (Standley and
emarginated, pinnately veined, hard, pubescent Record 1936); Nicaragua; Costa Rica; Panamá
(when young), and with very scabrous surfaces (Woodson and Schery 1965); Guatemala; El
and sinuous margins (Pittier 1926, Gentry 1993, Salvador; French Guiana; Suriname; Guyana;
Killeen and others 1993). Leaf borders are dentate. Trinidad; Venezuela; Colombia (Pérez Arbelaez
The apex is retuse and the base is decurrent on a 1978); Perú (Macbride 1956); Brazil; and Bolivia
short petiole (Schnee 1960). Stipules are (Beard 1953, Sarmiento 1983, Liogier 1990, Salas
deciduous and form small scars on the stem. Estrada 1993).
268
Genetics.—Comparative flavonoid chemistry of chaparro is one of the most common woody
chaparro and other members of the Dilleniaceae species.
family resulted in reticulate distribution of the Chaparro grows in many soil types. The
substances identified and did not permit most favorable are well-drained soils. Examples of
recognition of taxa beyond the level of genus and soils are red sands, red sandy loams, yellow sands,
family (Gurni and Kubitzki 1981). Woodson and well drained latosols, hard laterite crusts, ill-
Schery (1965) reported pentagyna as a variety of drained latosols, deeply weathered quartz gravel
chaparro in Panamá. and sands, serpentine-derived soils, entisols on
well drained sites, entisols on young alluvial
Ecology.—Chaparro is a savanna plant with a surfaces to oxisols on more ancient ones, alfisols,
uniform spatial distribution (San Jose and others and vertisols (Sarmiento 1983). Vargas Ulate
1991). It is adapted to fire, high air temperature, (2001) attributed the presence of chaparro in
and harsh soil conditions (Pittier 1926, Lundell Guanacaste, Costa Rica, to acid and infertile soils.
1937). Chaparro is among the first invaders after a In Colombia, the loss of the A and B horizons of
fire (Salas Estrada 1993). Medina and Silva (1990) granitic soils in sierra Nevada have led to
describe the savannas where chaparro grows as savannization and the establishment of chaparro in
“steady state (s) regulated by water-fire lands where it normally does not grow (Cavalier
interactions on a background of low nutrient and others 1998).
availability.” However, during the early phases of Chaparro has a deep root system reaching
growth, chaparro is susceptible to fire disturbance the water table in its adult stage. The species is a
(San José and Medina 1975). A low allocation of phreatophyte, so it grows new leaves during the
N to leaves--even young ones--reduces dry season (Sarmiento and Monasterio 1983,
photosynthetic capacity, but it is adaptive to fire as Medina and Francisco 1994).
only a small amount of N is lost when leaves burn Competition between chaparro and other
(Medina and Francisco 1994). plants in the savanna occurs in two stages:
Chaparro grows up to 500 m elevation in establishment and subsequent growth. During
Bolivia (Killeen and others 1993), and Venezuela establishment, grasses may hinder the growth of
(Sarmiento 1983), and it has extended its range in saplings and constitute a risk because of the high
Colombia to up to 500 to 1,500 m due to fire frequency. Chaparro roots face water shortages
savannization of forest lands (Cavalier and others as they grow through the grass root layer, so
1998). Frost appears to limit its northern and establishment is favored by sequences of wet years
southern distribution, and flooding or soil when fire frequency diminishes and soil moisture
saturation also limits its distribution (Sarmiento increases. Eventually, chaparro will out-grow
1983). grasses and shade them out (for example in tree
High acidity, high clay content, low groves).
mineral nutrient concentration, high exchangeable The chlorophyll content of chaparro is
Al, poor drainage, and poor aeration in the deeper low and correlated with low N and high ash
soil horizons favor chaparro (Lundell 1937, concentrations (Medina and Francisco 1994). Leaf
Gottsberger and Morawetz 1986). Chaparro will photosynthesis rate reached maximum values in
grow in depressions. In some sites, chaparro mature leaves during the humid season and
dominates the community and forms dense groves. declined in younger and older leaves and during
For example, within a matrix of flooded savannas the dry season (Sobrado 1996b). Young leaves are
in Bolivia, chaparro forms groves on ant and more efficient in water use than mature leaves.
termite hills and on hummocks (Killeen and others Stomatal conductance, and water loss,
1993). Sahales is the name used to describe groves decline during the dry season. Chaparro leaves do
of chaparro in northern Petén (Lundell 1937). not show photoinhibition. Stomatal conductance
Chaparro grows in a wide range of moisture differences between dry and wet season appear to
conditions from dry to wet savannas. In the be responsible for maintaining a homeostatic
Venezuelan Llanos, San José and Medina (1975) balance of leaf water and thus maintain xylem
report a mean annual rainfall of 1335 mm and a integrity in spite of the species vulnerability to
range of 580 to 1990 mm. A “normal” year has a 4 embolisms (Sobrado 1996a). Predawn water
to 6 month dry season. Beard (1953), Sarmiento potential, minimum leaf water potential, and
and Monasterio (1975), Sarmiento (1983), Medina hydraulic resistance of chaparro, did not show
and Silva (1990), and Medina and Huber (1992) significant differences throughout the year. In spite
report details of the climatic and edaphic of the scleromorphic appearance of leaves,
conditions of tropical American savannas where chaparro appears adapted only to short-term
269
diurnal deficits (Meinzer and others 1983). The capacity is reduced due to high mesophyll
transpiration curves for chaparro during the dry resistance to CO2 transfer (Medina 1982). Growth
season are always bimodal (Medina 1977, 1982) as is seasonal and its flowering is tardy (Sarmiento
plants close their stomata at noontime, and Monasterio 1983). It flowers and produces
independently of water availability (Medina 1967, new leaves during the dry season (Medina 1982).
1983). Transpiration rates are also low. The Leaf development takes 45 days (Nascimento and
dependency of leaf dark respiration on air others 1990), and leaf longevity is about 11
temperature changes seasonally (Medina 1982). months (Montes and Medina 1977). However,
The steepness of the relationship decreases from young, mature, and old leaves overlap in an
December, to February, to March, and to individual (Medina and Francisco 1994). Fruiting
September through November, when the and seed dispersal may continue for a short time
relationship is flat. during the rainy season. Fire does not interrupt this
The fungus Pseudocercospora curatellae cycle as burned leaves fall and within days there is
was isolated from leaves of chaparro (Furlanetto a new flush.
and Dianese 1999). The earthworm Pontoscolex Two hundred randomly selected trees of
roraimensis was associated with chaparro (Guerra chaparro growing in the seasonal savannas of
Torquemada 1994). Epiphytes are not abundant on western Venezuela had a mean stem diameter at
chaparro bark, probably due to its flaky deciduous ground level of 8.3 cm. The largest recorded
nature (Gottsberger and Morawetz 1993). Leaf diameter was 39.8 cm (Sarmiento and Monasterio
herbivory on chaparro was 4.6 percent near 1983). Chaparro wood has no growth rings and its
Cuiaba, Mato Grosso, Brazil (Nascimento and sapwood is not clearly defined (Record and Mell
others 1990). 1924).
Under a regime of annual burning,
Reproduction.—The white flowers have a chaparro will experience die back of all its annual
“disagreeable smell” (Grisebach 1963), occur in growth shoots, and growth will start the next
lateral panicles on defoliated old branches, and are season from adventitious buds that develop from
rarely terminal. Flowers are actinomorphic and vascular cambium of older branches (Sarmiento
hermaphroditic. Four to five petals are white or and Monasterio 1983). As a result of this
pinkish, 5 to 6 mm, free, and imbricate (Liogier response, “the tree behaves as a mechanical
1990). Four to five green sepals are rounded, 5 structure supporting a crop of annual branches”
mm long, persistent, and externally pilose (Schnee (Sarmiento and Monasterio 1983). The vascular
1960). Stamens are numerous, free, and persistent. cambium assumes the role of apical meristem as
The gynoecium has two hirsute carpels of 6 to 7 the main replacement tissue. Chaparro can
mm (Liogier 1990) with longitudinal dehiscence maintain itself as a hemixyle--with annual shoots
(Schnee 1960). They are coned at the base and and woody underground structures--and even
covered by long and abundant hairs. Follicles are complete the reproductive process in this growth
globose, hispid, coned in the lower half, with two form (Sarmiento and Monasterio 1983). Chaparro
seeds (Killeen and others 1993). Flowers and fruits can develop new individuals from injured roots
of chaparro are fasciculate and ramiflorous (Foldats and Rutkis 1975). Usually, saplings that
(Gentry 1993). Chaparro flowers between July and develop from injured roots have higher water
November in Bolivia (Killeen and others 1993) potentials than normal saplings because they are
and December to February in Venezuela (Montes already connected to the water table. Injury to
and Medina 1977). Seeds are oblong, bright, roots will stimulate vegetative regrowth, which in
chestnut brown to black, 3 mm long, with a small turn also facilitates formation of groves (San Jose
membranous hilum (Schnee 1960, Liogier 1983). and others 1991).
Seeds are long-lived and remain buried until the The mineralization of nitrogen, soil
proper conditions for germination occur (Medina organic matter, soil exchangeable Ca, Mg, and K,
and Francisco 1994). Fire induces germination and and total soil nitrogen were higher in soils under
sprouting of stems that are too thick to sprout chaparro trees than soils under savanna grasses.
under normal conditions (Foldats and Rutkis For soil organic matter, K, and cation exchange
1969). Seeds are dispersed by birds (Salas Estrada capacity, chaparro exceeded values measured
1993), including parrots and parakeets who eat under deciduous trees in savannas (García
chaparro fruits (Pérez Arbelaez 1978). Miragaya and others 1994, Sánchez and others
1997). Although chaparro is not an aluminum
Growth and Management.—Chaparro accumulating plant, it exhibits seasonal variation
assimilates carbon all year, but its photosynthetic in its nutrient concentration (Villela and de
270
Lacerda 1992). Potassium, Fe, Cu, and Zn were anti-inflammatory and analgesic activity of
highest in August, Ca and Mg in November, and Curatella americana L. Journal of
Al and Mn in May. The Al and Mn concentration Ethnopharmacology 67:171-177.
of mature chaparro leaves is 171 and 105 parts per
million, respectively (Medina 1977). Bailey, L.H. 1941. The standard cyclopedia of
horticulture. The Macmillan Co. New York.
Benefits.—Chaparro produces good wood with a
reddish hue and specific gravity of 0.85. Record Bastos, M.D.N.D.C. 1984. Floristic survey of
and Mell (1924) and de Matos Araujo and de fields of the state of Para Brazil. 1. The field of
Matos Filho (1977) contain wood anatomy Joannes Marajo Island. Boletim do Museu
information for chaparro. Small stem sizes limit Paraense Emilio Goeldi Serie Botanica 1:67-86.
wood utilization of shrubs and trees. However, the
wood is durable and used for charcoal, fuelwood, Beard, J.S. 1953. The savanna vegetation of
posts, small cabinetwork, and for turning (Record northern tropical America. Ecological
and Mell 1924, Standley and Record 1936, Killeen Monographs 23:149-215.
and others 1993, Salas Estrada 1993). Termites do
not attack chaparro wood (Pérez Arbelaez 1978). Behling, H. 1995. A high-resolution Holocene
Wood utilization increases where trees or shrubs pollen record from Lago do Pires, SE Brazil-
become locally abundant and grow in groves vegetation, climate and fire history. Journal of
(Pittier 1926). People use dry leaves as substitutes Paleolimnology 14:253-268.
for sand paper and for cleaning kitchen utensils or
polishing metals (Killeen and others 1993). Behling, H. and H. Hooghiemstra. 1999.
Chaparro bark produces tannins that provide a Environmental history of the Colombian
gray color when used for dying pelts (Killeen and savannas of the llanos Orientales since the last
others 1993). Ground seeds mixed with chocolate glacial maximum from lake records El Pinal and
flavors the drink in Oaxaca, México (Standley Carimagua. Journal of Paleolimnology 21:461-
1920). Chaparro was one of the most abundant 476.
pollens in honey samples during the months of
January, February, and March from an apiary Cavalier, J., T.M. Aide, C. Santos, A.M. Eusse,
north of Roraima State, Brazil (da Silva and Absy and J.M. Dupy. 1998. The savannization of
2000). Siegel and Paguaga (1991) suggested using moist forests in the Sierra Nevada de Santa
chaparro twigs and pods for prospecting for gold Marta, Colombia. Journal of Biogeography
in Costa Rica. In Venezuela, Clamens and others 25:901-912.
(1999) evaluated chaparro for industrial gum
production and found the species promising. Clamens, C., Leon de Pinto G., F. Rincon, V.
Antonio, and O. Beltran. 1999. The behavior of
Pharmaceuticals.—Chaparro is a medicinal plant Samanea saman and Curatella americana as
used extensively in folk medicine throughout its gum producers. Boletín del Centro de
range. Infusions from leaves and stems are used Investigaciones Biológicas Universidad del Zulia
for arthritis, diabetes, and to lower blood pressure 32(2): 67-78.
(Killeen and others 1993, Liogier 1990). Guerrero
and others (2002) found that a dose of 20 mg/kg of da Silva, S.J.R. and M.L. Absy. 2000. Analyses of
ethanolic extracts of chaparro showed significant pollen found in honey samples of Apis mellifera
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274
Cytisus scoparius (L.) Link Scotch broom
FABACEAE
Synonyms: Sarothamnus scoparius (L.) Wimmer ex Koch

Genista scoparius Lam.
Spartium scoparium L.
John K. Francis
10 mm long. It’s perfect flowers, which are

solitary in leaf axils or in clusters of two or three,
are about 2 cm long, brilliant yellow, and have the
typical form of flowers in the pea family. Orange,
red, pink, purple, and lilac flowers are available in
ornamental varieties. The fruits are dark brown,
linear pods 4 to 5 cm long that are ciliate at the
margins and contain five or six seeds each. The
seeds are 3 to 4 mm long, greenish brown, and in
the form of a flattened oval. There are 2n = 46 or
48 chromosomes (Abrams 1944, Meyer 2003,
New England Wild Flower Society 2003
University of Connecticut 2003, Welsh and others
1987).
Range.—Scotch broom is native to Western and

Eastern Europe, including the Mediterranean
islands, the British Isles, and western Scandinavia.
It has been planted and naturalized in Eastern and
Western United States, Hawaii, Canada, the
Azores, the Canary Islands, Australia, New
Zealand, South Africa, India, China, and parts of
southern South America (International Legume
Database and Information Service 2003, Natural
Resources Conservation Service 2003). Two
varieties of Scotch broom have naturalized in the
United States. Variety andreanus occurs only in
California and var. scoparius accounts for the rest
Drawing source: Britton and Brown 1913. of the naturalized range (Natural Resources
Conservation Service 2003).
General Description.—Scotch broom is also
known as broom, common broom, Scot’s broom, Ecology.—Scotch broom grows as scattered
English broom, green broom, broom tops, Irish plants, small clumps, and pure stands, and
tops, witch’s broom, basam, escoba negra, and competes fiercely with forbs, grasses, and tree
retama de escobas. It is a shrub, usually 1 to 2, seedlings. It requires disturbance, especially
occasionally 3 m in height, which sheds its leaves disturbance resulting in bare mineral soil to
early and remains leafless for most of the year. become established. It prefers sandy soils, but will
The green twigs are the principal photosynthetic grow in a wide variety of well-drained soils
organs. Branches are upright to arching. The (Prasad 2003). Scotch broom is drought tolerant.
angular, flexible twigs often occur in moderately The species is intolerant of shade and cannot
long sprays. Old stems become gray-brown. The survive under a closed forest stand. Nitrogen
plant grows a deep root system with a taproot. fixation occurs in root nodules, and fixed nitrogen
Leaves are trifoliate low on the twigs and simple is released to the environment in the litter (Parker
and sessile near the tips. The obovate to lanceolate and others 1994). Top injury from subfreezing
leaflets are 4 to 8 mm long, and petioles are 2 to temperatures may occur in cold climates, but
275
plants usually survive (Michigan State University stands reproduce prolifically. The application of
Extension 1998). Scotch broom is commonly seen herbicides has sometimes been effective (Prasad
along roads, in vacant lots and inactive 2003). A number of insect enemies of Scotch
construction sites, in old logging areas, in broom have arrived accidentally or have been
neglected pastures and orchards, and along fence- introduced into the United States. They may
rows. already be having a minor impact on the
competitive ability of the species (Syrett and
Reproduction.—Scotch broom flowers in May or others 1999). Research into biological control
June (New England Wild Flower Society 2003). continues, particularly in Australia and New
The flowers are pollinated by honey bees and Zealand (CSIRO 2003).
bumblebees (Meyer 2003). Seed production starts
when plants are 2 years old in Tasmania with Benefits.—Scotch broom, which produces a
around 300 seeds/year and increases to 12,000 to spectacular annual floral display and possesses a
30,000 seeds/shrub/year. The seeds are spread by unique form, has been planted throughout the
explosive release from the pods, which can throw temperate regions of the world as border and
the seeds 1 to 3 m (Alchemy-works 2003), as well accent plants. It is particularly useful in this regard
as by water, soil movement, and machinery because it succeeds on poor soils with little care. It
(National Haritage Trust 2003). Ants also disperse has also been used successfully to protect exposed
seeds. Seeds average 125 seeds/g. (Meyer 2003). soils and stabilize dunes. Scotch broom furnishes
Seeds can remain viable in the soil seed bank for food (mainly from the seeds) and cover for
as long as 80 years (National Haritage Trust 2003). wildlife. The leaves and twigs are generally
The species can also be propagated with cuttings unpalatable to domestic livestock and wild
(University of Connecticut 2003). mammals. The tissues contain the alkaloids
cytisin, sparteine, and isosparteine, and the
Growth and Management.—Scotch broom glycoside scoparin. However, large amounts must
grows from 30 to 46 cm/year (Michigan State be ingested before illness or death results (Everest
University Extension 1998) and usually lives from and others 2003). The stems and twigs have been
10 to 15 years (Prasad 2003). Fruits can be picked used from ancient times to make brooms and were
by hand after they are mature but before they burst used for ritual brooms called besoms. The species
open. They should be spread to dry, threshed, and was also used to make baskets, thatching, wattle
screened. These seeds are orthodox and can be fencing, and a green dye, and to flavor beer before
stored for many years. Because they are hard- the introduction of hops (Alchemy-works 2003).
coated, scarification is needed to ensure adequate Decoctions of the twigs and leaves were and are
and uniform germination. This can be done by still occasionally used to treat gout, jaundice,
nicking or sanding the seed coat or by applying edema, and other conditions (Grieve 2003).
dry heat for 1 minute at 130 °C or 15 minutes at 70
°C. Viability is normally about 80 percent. References
Because roots are delicate, the use of containerized
stock is recommended for planting rather than Abrams, L. 1944. Illustrated flora of the Pacific
bare-root stock (Meyer 2003). However, because States. Vol. 2. Stanford University Press,
the species is invasive, planting for conservation or Stanford, CA. 635 p.
ornamental purposes is usually discouraged. In
fact, the species has been declared a noxious weed Alchemy-works. 2003. Cytisus scoparius-Scotch
in five Western States where planting is prohibited broom. http://www.alchemy-works.com/
(Natural Resources Conservation Service 2003). cytisus_scoparius.html. 1 p.
Scotch broom is not a problem in cultivated land
or in land fully timbered, only in pastures, Britton, N.L. and A. Brown. 1913. Illustrated flora
roadsides, sites normally occupied by low of the northern states, Canada, and the British
vegetation, and in regenerating timberlands. possessions. Vol. 2, 2nd Ed. Scribner, New York.
Control of Scotch broom is difficult. Grubbing 735 p.
large plants and pulling seedlings, or power
cultivation, effectively kills the plants, but new CSIRO-European Laboratory. 2003. Our research:
seedlings continually appear from the soil seed Scotch broom-Cytisus scoparius Link, Fabaceae.
bank. Mowing only encourages branching near the http://www.csiro-europe.org/cytisus.html. 2 p.
ground. Stands of Scotch broom do not carry a fire
readily (Parker and others 1994), and burned
276
Everest, J.W., T.A. Powe, Jr., and J.D. Freeman. New England Wild Flower Society. 2003. Invasive
2003. Poisonous plants of the southwestern plant atlas of New England: Cytisus scoparius
United States. Alabama Cooperative Extension (Scotch broom). http://webapps.lib.uconn.edu/
System. http://www.aces.edu/dept/extcomm/ ipane/browsing.cfm?descriptionid=48. 6 p.
publications/anr/anr-975.pdf. 56 p.
Parker, B., G. Miller, L.C. Burrell. 1994. Weeds:
Grieve, M. 2003. Broom. http://www.botanical. Scotch broom. PNW 130. Pacific Northwest
com/botanical/mgmh/b/broom-70.html. 9 p. Extension, Oregon State University, Corvallis,
OR. 10 p.
International Legume Database and Information
Service. 2003. Cytisus scoparius (L.) Link. http: Prasad, R. 2003. Scotch broom, Cytisus scoparius
//www.ildis.org/LegumeWeb/6.00/taxa/6046. L. in British Columbia. Canadian Forest Service,
shtml. 4 p. Ottawa, Ontario, Canada. http://www.pfc.
forestry.ca/biodiversity/broom_e.html. 9 p.
Meyer, S.E. 2003. Cytisus scoparius (L.) Link. In:
F.T. Bonner and R.G. Nisley, eds. Woody plant Syrett, P., S.V. Fowler, E.M. Coombs, J.R.
seed manual. U.S. Department of Agriculture, Hosking, G.P. Markin, Q.E. Paynter, and A.W.
Forest Service, Washington, DC. Sheppard. 1999. The potential for biological
http://wpsm.net/Cytisus.pdf. 4 p. control of Scotch broom (Cytisus scoparius)
(Fabaceae) and related weedy species.
Michigan State University Extension. 1998. Biocontrol News and Information 20: 17N-34N.
Cytisus scoparius—Scotch broom. http://www.
msue.msu.edu/msue/imp/modop/00000485.html. University of Connecticut. 2003. University of
2 p. Connecticut plant database: Cytisus scoparius.
http://www.hort.uconn.edu/plants/c/cytsco/
National Heritage Trust. 2003. English broom cytsco3.html. 4 p.
(Cytisus scoparius). http://www.bushcare.
tas.gov.au/ToAdd/Data/weeds/English%20%20 Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
broom.pdf. 2 p. Higgins. 1987. A flora of Utah. Great Basin
Naturalist Memoirs 9. Brigham Young
Natural Resources Conservation Service. 2003. University, Provo, UT. 894 p.
Plant profile: Cytisus scoparius (L.) Link.
http://plants.usda.gov/chi_bin/plant_profile.cgi?
symbol=CYSC4. 7 p.
277
Dalbergia ecastaphyllum (L.) Taubert coin vine
FABACEAE
Synonyms: Hedysarum ecastaphyllum L.

Pterocarpus ecastaphyllum L.
Ecastophyllum ecastophyllum (L.) Britton
Amerimnon ecastophyllum (L.) Standl.
John K. Francis
pubescent on the under side. The small white

flowers are grouped in panicles in the leaf axils.
These develop into small groups of copper-colored
to gray-brown, elongated lenticular fruits about 1.5
to 3 cm long that contains one flattened, brown
seed (Holdridge and Poveda 1975, Howard 1988,
Liogier 1988, Nelson 1996).
Range.—Coin vine is native to Florida, the

Bahamas, the Greater and Lesser Antilles,
Trinidad, the east coasts of Mexico, Central
America, and South America to southern Brazil,
and the coast of tropical West Africa (Liogier
1988, Burkill 1995).
Ecology.—Coin vine inhabits coastal sand dunes,

open coastal forests and brush fields, lagoon sides,
landward sides of mangrove swamps, shell mounds
within mangroves, coastal hammocks, and
estuaries (Burkill 1995, Nelson 1996). It also
encroaches on dense grass swards. The species is
most common on sands or sandy soils, but also
grows on heavier soils. Coin vine inhabits areas in
Puerto Rico that receive from about 1200 to 2200
mm of annual precipitation. It also survives
droughts well (Workman 1980). The species
General Description.—Coin vine is a scrambling frequently grows on sites where there is constant
and climbing shrub that sometimes becomes a salt spray and sometimes in moderately salty soils.
small tree with stems that extend as much as 10 m Old plants tend to form dense thickets.
(Howard 1988). It is also known by the common
names maraimaray, palo de pollo, mangle médaille, Reproduction.—A collection of mature fruits
popian, and a host of other common names (Burkill from Puerto Rico weighed an average of 0.290 +
1995, Liogier 1988). Coin vine develops a tap and 0.010 g/fruit. Seeds extracted from 24 of the pods
lateral root system. The tan colored roots support constituted 69 percent of the pod weight. The pods
many nodules that attach directly to the tap and are tough and the seeds are fragile. Extraction is
lateral roots. The older stems, which may reach 7 difficult and unnecessary because seeds germinate
cm or more in diameter, are gray and extend into through the exocarp without difficulty. Seventy-
the crowns of low trees and scramble over low three percent of 100 fruits sown in commercial
obstacles. There are many long, vine-like branches potting mix germinated between 31 and 181 days
that bear leaves only on the current year’s growth. after sowing. Germination is hypogeal. Seeds
The simple leaves have petioles 5 to 12 mm long disburse by lateral branch extension and by water.
and elliptical or ovate blades 2.5 to 14 cm long and Seed may be collected after they wash up on the
2 to 8 cm broad. They are leathery, glossy green, shores (Workman 1980), directly from the shrubs,
rounded at the base, pointed at the tip, and and from the ground under them. Natural seedlings
278
are abundant near coin vine plants. However, few Holdridge, L.R. and L.J. Poveda A. 1975. Arboles
survive more than a few weeks. Branches layer de Costa Rica. Vol. 1. Centro Cientifico
whenever they come in contact with the ground. Tropical, San Jose, Costa Rica. 546 p.
Growth and Management.—Early height growth Howard R.A. 1988. Flora of the Lesser Antilles,
is moderate but steady. A small group of nursery Leeward and Windward Islands.
seedlings reached about 15 cm in 5 months. Older Dicotyledoneae, Part 1. Vol. 4. Arnold
plants grow about 2 m per year. By resprouting and Arboretum, Harvard University. Jamaica Plain,
layering, plants (clones) can survive almost MA. 673 p.
indefinitely.
Benefits.—The extensive horizontal branches and Rico and adjacent islands, Spermatophyta. Vol.
their roots cover, bind, and stabilize coastal sand 2. Editorial de la Universidad de Puerto Rico,
dunes (Burkill 1995). The stems are brittle and not Río Piedras, PR 481 p.
suitable for wooden implements or basketry. The
crushed roots and bark contain a chemical that was Liogier, H.A. 1990. Plantas medicinales de Puerto
used by Native Americans to stupefy and catch fish Rico y del Caribe. Iberoamericana de Ediciones,
(Workman 1980). Coin vine is listed among the Inc. San Juan, Puerto Rico. 566 p.
honey plants of the Dominican Republic (Marcano
Fondeur 2001). In Senegal, the leaves are put into Marcano Fondeur, E. de J. 2001. La flora apícola
inhalations and baths to treat various debilities de la Republica Dominicana. http://marcano.
(Burkill 1995). Various extracts are used in herbal Freeservers.com/nature/estudios/apicola/dicotsp.
medicine as a diuretic, an emetic, and a vermicide. html. 11 p.
Care must be taken, because some of the tissues are
toxic (Liogier 1990). Nelson, G. 1996. The shrubs and woody vines of
References 391 p.
Burkill, H.M. 1995. The useful plants of West Workman, R.W. 1980. Growing native. The
Tropical Africa. Vol. 3. Royal Botanci Gardens, Sanibal-Captiva Conservation Foundation, Inc.,
Kew, United Kingdom. 857 p. Sanibal, FL. 137 p.
279
Dasiphora floribunda (Pursh) Kartesz, comb. nov. ined. shrubby cinquefoil
ROSACEAE
Synonyms: Potentilla fruticosa L.

Dasiphora fruticosa auct. non. (L.) Rydh.
Potentilla floribunda Pursh.
Potentilla fruticosa ssp. floribunda (Pursh) Elkington
Potentilla fruticosa var. tenuifolia Lehm.
Juanita A.R. Ladyman

other Potentilla species that are all herbaceous
(Cronquist and others 1997). Recently it has been
reported as Dasiphora floribunda (Pursh) Kartesz
comb. nov. ined (ITIS 2002). The genus
Dasiphora includes approximately 8 or 9 species
that are all exclusively in Asia except for shrubby
cinquefoil. The genus name “Pentaphylloides” is
derived from the Greek and refers to the pinnate
five-foliate leaves. “Penta-” means “five-“and
“phylloides” means “leaved.” “Fruticosa” is a
Latin adjective that means “shrubby” or “bushy.”
This shrub is usually from 10 cm to 1 m tall but
may grow to a little over 1.5 m. The herbage of the
current year is covered by soft hairs, but the stem
becomes smooth on aging with, eventually, a red-
brown shreddy bark. The leaves comprise usually
five, but sometimes three to seven, elliptic-shaped
leaflets. The flowers are solitary in the leaf axils
along the stem but form three- to seven-flowered
clusters (cymes) at the stem tips. Petal color is
usually yellow, although it may be white, or even
pink and red, in cultivated varieties, and there are
15 to 25 stamens per flower (Carter 1997,
Cronquist and others 1997, Hitchcock and
Cronquist 1973, Vines 1986). This is a wide-
ranging species, but apparently chromosome
number differs between new and old world
material. North American plants have a
chromosome number of 2n = 14 while Asian and
European plants have 2n = 28 (Cronquist and
others 1997).
Range.—Shrubby cinquefoil is a common, wide-

ranging circumboreal shrub ranging from
Illustration source: USDA—Forest Service collection, Greenland and Labrador, to Alaska and south to
Hunt Institute the mountains of California, Nevada, Arizona,
Colorado, New Mexico, and through the northern
General Description.—Pentaphylloides states into Iowa, Indiana, Illinois, Ohio,
floribunda is commonly known as shrubby or bush Pennsylvania and New Jersey in North America. It
cinquefoil, but local names also include golden also occurs in Europe, for example England,
hardback, prairie-weed, fivefinger, buckbrush, Ireland, Scandinavia and Bulgaria, and in Asia
ninebark, widdy, and yellow rose. It is also (Stace 1997, Gibbons and Davies 1994, Polunin
frequently referred to by the most common 1997).
synonym, Potentilla fruticosa (Kartesz 1994). This
is a distinctive shrub and easily separable from the
280
Ecology.—Shrubby cinquefoil grows in moist germination (Baskin and Baskin 2001). A cold,
soils in mountain regions from lower foothills to moist stratification period of 2 months has been
sub alpine. The elevation tends to depend generally recommended for Potentilla species,
somewhat upon latitude, but generally plants are including Potentilla fructicosa. (Phillips 1987).
found between approximately 1,800 m and 3,400 Shrubby cinquefoil can also be propagated from
m. It grows in open woods, wet mountain softwood cuttings in the spring or from suckers
meadows, calcareous bogs, limestone pavements, (Tykač 1990). Container grown plants, that are
and along streams (Cronquist and other 1997, very popular within the landscape business, can be
Gibbons and Davies 1994, Vines 1986, Welsh and planted in spring and fall. Even old plants are
others 1993). It has also been reported from reported to tolerate transplanting (Tykač 1990).
relatively xeric habitats such as mountain shrub
lands and rocky mountain slopes (McGregor and Benefits.—Shrubby cinquefoil leaves are bitter
others 1986, Stace 1997, Thilenius 1972). The tasting, and the forage and browse value varies
degree of permanent soil moisture required is depending upon the part of the world in which it
likely to depend on average precipitation, grows. It is generally on summer range, although at
temperature, and localized water collection sites or lower elevations it may be valuable for fall
drainage channels. It is a frequent member of browsing (Dayton 1931). It is likely that just as the
willow (Salix)-dominated communities, and in shrubs’ morphology varies, the levels of secondary
some parts of its range, for example New Mexico, plant products that impart the bitter taste differ
it is a member of the Krummholz communities between habitats and geographic range. Generally,
found at the lower tundra edge (Dick-Peddie it is considered inferior forage for livestock, but it
1993). Shrubby cinquefoil may be the dominant is used by cattle in Arizona and Montana (Epple
shrub in many situations and can also grow in 1995, Vines 1986). It is also often an important
fairly pure stands. For example, it may be the browse for sheep and goats especially in the
dominant shrub in aspen stands or other forest Southwestern United States and in southwestern
clearings where snow accumulation prevents tree Montana and contiguous portions of southeastern
establishment (Knight 1994, Thilenius 1972). In Idaho (Dayton 1931). In Montana it is rated 18th
addition, it frequently attains dominance in among the most important browse species.
limestone-derived soils (Thilenius 1972). Similarly, it is reported unpalatable to deer
(Thilenius 1972) but does provide browse for deer
Reproduction.—Shrubby cinquefoil generally and elk in some regions (Epple 1995, Vines 1986).
flowers in June through August, although flowers On some overgrazed range it is often grazed quite
may be found until the first frost (Epple 1995, closely, and the shrubs assume a hedge-like
McGregor and others 1986, USDA FS 1988). The appearance (USDA FS 1988). Such use indicates
achenes mature in summer and fall. The bright overstocking and a critical reduction, if not
yellow flowers are insect pollinated. The fruit is a elimination, of other palatable plant species. It
densely hairy achene. provides effective protection against soil erosion
(Epple 1995, Vines 1986). Native Americans have
Growth and Management.—Shrubby cinquefoil used various parts of the plant for an assortment of
increases in response to overgrazing and other purposes including as a medicine and in
disturbance. In the Western United States it does ceremonials (Moerman 1998). The Blackfoot
not appear to be an invasive species, but in New peoples have used the leaves to fill pillows and the
England it can be an aggressive invader of bark as tinder, while the Eskimos of Alaska and the
agricultural land and may be difficult to eradicate Arctic have used dried leaves for tea (Moerman
once established (USDA FS 1988). It has been 1998). Leaves were also used for tea in Russia. The
used as an indicator species of past disturbance name “potentilla” is derived from the Latin
(Dick-Peddie 1993). Shrubby cinquefoil is widely “potens,” which means powerful and may refer to
used as an ornamental and in wild land seeding. It the medicinal value of some species. A preparation
does best in full sun as it tends to flower poorly in made from the leaves was used by the Cheyenne
shade conditions. Seed apparently does not require for protection and was also considered a deadly
cold stratification for germination and will arrow poison, only to be administered by holy
germinate immediately after harvesting (Bonde people (Carter 1997, Moerman 1998). Shrubby
1965, Tykač 1990). The highest germination rate cinquefoil has commercial value in the landscape
was achieved at 18 oC (Bonde 1965). However, it and horticultural trade. It is a popular ornamental
has been proposed that cold stratification may shrub, and many cultivars exist that have been
reduce the temperature requirements for seed chosen for the degree of leaf pubescence
281
(hairiness), flower color, and growth form. A list of ITIS, the Integrated Taxonomic Information
some of the recognized cultivars available can be System. 1996. Dasiphora floribunda (Pursh)
found in Bailey and others (1976) and Tykač Kartesz comb. nov. ined. Taxonomic Serial No.:
(1990). 565123 and Plants, database (version 5.1.1)
http://www.itis.usda.gov/ [not paged].
References.
Kartesz, J.T. 1994. A synonymized checklist of the
Bailey, L.H., E.Z. Bailey, and the staff of the vascular flora of the United States, Canada and
Liberty Hyde Bailey Hortorium. 1976. Hortus Greenland. Vol. 1 - Checklist. 2nd Ed. Timber
Third – a concise dictionary of plants cultivated Press, Portland, OR. 622 p.
in the United States and Canada. Macmillian
Publishing Company, New York. 1,290 p. Knight, D.H. 1994. Mountains and plains–the
ecology of Wyoming landscapes. Yale
Baskin, C.C. and J.M. Baskin. 2001. Seeds, University Press, New Haven, CN. 338 p.
ecology, biogeography, and evolution of
dormancy and germination. Academic Press, McGregor, R.L., T.M. Barkley, R.E. Brooks, E.K.
New York. 666 p. Schofield. 1986. Flora of the Great Plains.
University Press of Kansas, Lawrence, KN.
Bonde, E.K. 1965. Further studies on the 1,402p.
germination of seeds of Colorado alpine plants.
University of Colorado Studies 18. Moerman, D.E. 1998. Native American ethno-
botany. Timber Press, Portland, OR. 927 p.
Carter, J.L. 1997. Trees and shrubs of New
Mexico. Johnson Books, Boulder, CO. 534 p. Phillips, J. 1987. Southwestern landscaping with
native plants. University of New Mexico Press,
Cronquist, A., N.H. Holmgren, and P.K. Albuquerque, NM. 140p.
Holmgren. 1997. Intermountain flora, vascular
plants of the Intermountain West, U.S.A. Vol. 3, Polunin, O. 1997. Flowers of Greece and the
Part A. New York Botanical Garden, Bronx, Balkans–a field guide. Oxford University Press,
NY. 446 p. New York. 567p.
Dayton, W.A. 1931. Important western browse Stace, C. 1997. New Flora of the British Isles. 2nd
plants. Miscellaneous Publication 101. U.S. edition. Cambridge University Press, New York.
Department of Agriculture, Washington DC. 1,130 p.
214 p
Thilenius, J.F. 1972. Classification of deer habitat
Dick-Peddie, W.A. 1993. New Mexico vegetation in the ponderosa pine forest of the Black Hills,
– past, present, and future. University of New South Dakota. Research Paper RM-91. USDA
Mexico Press, Albuquerque, NM. 244 p. Forest Service Rocky Mountain Forest and
Range Experiment Station, Fort Collins, CO. 28
Epple, A.O. 1995. A field guide to the plants of p.
MT. 347 p. United States Department of Agriculture Forest
Service. 1988. Range Plant Handbook. Dover
Gibbons, B. and P. Davies. 1994. Field guide to Publications, Inc. New York. 838 p.
wild flowers of Britain and Northern Europe.
The Crowood Press, Marlborough, Wiltshire, Vines, R.A. 1986. Trees, shrubs, and woody vines
UK. 320 p. of the Southwest. University of Texas Press.
Austin, TX. 1,104 p.
Hitchcock, C.L. and A. Cronquist. 1973. Flora of
the Pacific Northwest – an illustrated manual. Welsh, S.L., N.D. Atwood, S. Goodrich, L.C.
University of Washington Press, Seattle, WA. Higgins, eds. 1993. A Utah flora. Second
730 p. Edition, revised. Brigham Young University,
Provo, UT. 986 p.
282
Dasylirion wheeleri S. Wats. Wheeler sotol
AGAVACEAE
Synonyms: None.
northern Mexico (Kearney and others 1960,

Powell 1998, Tesky 1993, Vines 1960).
Ecology.—Wheeler sotol occurs in chaparral,
southwestern oak (Quercus) woodland, desert and
semi-desert grasslands communities at 914 to
1,830 m in elevation (Carter 1997). It occurs in
seral, climax, and postclimax communities. In the
desert plains grasslands it is often sub-dominate in
the beargrass-scrub oak postclimax community
and the curly mesquite (Hilaria belangeri)-blue
grama climax community. It has been described
as a diagnostic climax member of desert
grasslands (Dick-Peddie 1993). In Trans-Pecos,
Texas, it is commonly found growing on limestone
and granite (Powell 1998, Schmutz and others
1992). Wheeler sotol usually grows on rocky soils
in desert grasslands that are often described as too
moist for open desert scrub but too dry for oak or
juniper woodland (Burgess 1995, Dick-Peddie
1993, Schmutz and others 1992). These are areas
Illustration credit: Robert De Witt Ivey where subsurface water is generally not available.
Wheeler sotol can store relatively large amounts of
General Description.—The common names for water internally that act as a buffer against the
Dasylirion wheeleri are Wheeler sotol, sotol, rapid onset of drought stress. It is classed as a
desert spoon, spoonplant and spoonleaf. The latter water storer, or “succulent” plant (Burgess 1995,
names refer to the enlarged, spoon-like bases of Thomas 1991). The species can also tolerate
the long serrated leaves. Dasylirion wheeleri several degrees of frost (Vines 1960).
grows from a thick, woody, mostly subterranean
caudex with a large rosette of rigid leaves that are Reproduction.—The flowering stalks elongate to
armed with sharp curved spines, or teeth, around up to 5 m in late spring and bloom from May to
the margins (Kearney and others 1960). The long August (Bowers 1993). Wheeler sotol is dioecious,
flowering stalk arises out of the center of the the male and female flowers being borne on
rosette and is topped by a slender compound separate plants. Female flowers frequently flower
panicle. The genus name, Dasylirion, means thick, later than the male plants (Warnock 1974). The
or hairy, white lily from “dasy-“ and “lirio” female flowers are wind pollinated, and wind also
(Gledhill 1992). Dasylirion wheeleri is disperses the seed that are contained within a
distinguished from Dasylirion leiophyllum papery, one-celled, three-winged capsule. Wheeler
Engelm. ex Trel. by having the leaf margin spines sotol grows readily from seed.
directed apically rather than basally (Carter 1997).
There are two recognized varieties of Dasylirion Growth and Management.—Wheeler sotol
wheeleri, var. wheeleri and var. wislizenii (Vines occurs in desert grasslands that are being
1960). increasingly managed with fire (Thomas and
Goodson 1992). In these areas, fire is primarily
Range.—Wheeler sotol grows on well-drained used to stimulate grass production and reverse
rocky and gravelly hillsides or slopes in Western dense scrub invasion. The apical meristem of sotol
Texas, southern to central New Mexico, southern is usually protected from fire by a cluster of green
Arizona in the U.S.A. and in Chihuahua in leaves, and this species can also resprout from the
283
caudex that is the region at the base of the stem by Native Americans (Moerman 1998). Wheeler
(McPherson 1995, Thomas 1991). However, sotol plants are commercially used as ornamentals
frequent fires and stable soil moisture regimes tend and described as “one of the most striking accent
to exclude wheeler sotol (Burgess 1995). The plants available” (Starr 1999). The leaves were
response of sotol to fire tends to vary depending once sold for use in floral arrangements, but this
upon conditions. Kittams (1973) observed that practice has been discouraged due to conservation
mature sotol (in this case Dasylirion leiophyllum) considerations (Epple 1995).
were usually killed by fire. White (1969) randomly
selected 50 individuals after a fire in desert References
grassland in Arizona and classified them with
respect to the level of the damage (severe, Ahlstrand, G.M. 1982. Response of Chihuahuan
moderate, or light) experienced. All of the lightly Desert Mountain Shrub Vegetation to Burning.
or moderately damaged plants survived, but only 3 Journal of Range Management 35 (1): 62-65.
percent of the severely damaged plants survived.
Sprouting did not occur except as refoliation from Bowers, J.E. 1993. Shrubs and trees of the
the terminal bud (White 1969). Ahlstrand (1982) southwest deserts. Southwest Parks and
investigated the response of Dasylirion Monumens Association, Tucson, AZ. 140 p.
leiophyllum to fire at sites in New Mexico and
western Texas. He reported losses of 50 percent or Burgess, T.L. 1995. The dilemma of coexisting
more on burned sites, but surviving top-killed growth forms. In: The desert grassland. M.P.
plants regained much of their pre-burn coverage McClaran and T.R. Van Devender, eds. The
after three growing seasons. This was similar to University of Arizona Press, Tucson, AZ. p. 31-
the observations after a semidesert grassland fire 67.
in southern Arizona where 47 percent of the
Wheeler sotol plants burned to death (Thomas and Carter, J.L. 1997. Trees and shrubs of New
Goodson 1992). Of the 71 Wheeler sotol plants Mexico. Johnson Books, Boulder, CO. 533 p.
that survived, all had apical regrowth, but despite
the fact that sotol may occasionally produce new Dayton, W. A. 1931. Important western browse
shoots from below ground, no offsets or seedlings plants. Miscellaneous Publication 101. U.S.
were found 11 months after the fire (Thomas and Department of Agriculture, Washington, DC.
Goodson 1992). In this study no individuals 214 p.
evaded the fire in refugia.
Dick-Peddie, W.A. 1993. New Mexico vegetation:
Benefits.—The head-like trunks and leaf bases of past, present, and future. University of New
Wheeler sotol are edible and American Indians Mexico press, Albuquerque NM. 244 p.
roasted and ate the young flower stalks (Moerman
1998, Powell 1998). The basal trunks contain Epple, A.O. 1995. A field guide to the plants of
starch and are also roasted, fermented and distilled Arizona. Falcon Press Publishing Co., Helena,
to make alcohol (Powell 1998). Currently an Montana. 347 p.
alcoholic beverage, generally known as sotol, is
made in parts of Mexico (Bowers 1993). Bighorn Gledhill, D. 1992. The names of plants. 2nd ed.
sheep and javelinas browse sotols (Epple 1995, Cambridge University Press, Cambridge,
Kearney and others 1960, Starr 1999). In times of England. 202 p.
stress Wheeler sotol has been used as feed for
livestock in a variety of ways. The trunks and leaf Ivey, R.D. 1995. Flowering plants of New Mexico.
bases have been used during drought, and the 3rd ed. Rio Rancho Printing, Albuquerque, NM.
plants have also been machine-cut and shredded 504 p.
for emergency feed (Dayton 1931, Powell 1998).
In addition, ranchers have burned off the leaves Kearney, T.H., R.H. Peebles, J.T. Howell, and E.
and split the heads (trunks) for livestock (Vines McClintock. 1960. Arizona flora. 2d ed.
1960). The leaves have been used to make mats, University of California Press. Berkeley, CA.
baskets, thatch, and paper (Moerman 1998, Powell 1,085 p.
1998). The long flowering stalks have been used
extensively within its range for corral, porch and Kittams, W.H. 1972. Effect of fire on vegetation of
house roofs, and other structures (Powell 1998). the Chihuahuan Desert region. Proc. Tall
They have also been used for ceremonial purposes Timbers Fire Ecology Conference 12: 427-444
284
McPherson, G.R. 1995. The role of fire in desert Sciences Laboratory. Fire Effects Information
grasslands. In: The desert grassland. M.P. System. http://www.fs.fed.us/database/feis/. 8
McClaran and T.R. Van Devender, eds. The p.
University of Arizona Press, Tucson, AZ. p. 130-
151. Thomas, P.A. 1991. Response of succulents to
fire: A review. International Journal of Wildland
Moerman, D.E. 1998. Native American ethno- Fire 1(1): 11-22.
Thomas, P.A. and P. Goodson. 1992. Conservation
Powell, A. M. 1998. Trees & shrubs of Trans- of succulents in desert grasslands managed by
Pecos and adjacent areas. University of Texas fire. Biological Conservation. 60(2): 91-100.
University Press, Austin, TX . 498 p.
Vines, R.A. 1960. Trees, shrubs, and woody vines
Schmutz, E.M., E.L. Smith, P.R. Ogden, M.L. of the Southwest. Sixth printing 1986.
Cox, J.O. Klemmedson, J.J. Norris, and L.C. University of Texas Press. Austin, TX.
Fierro. 1992. Desert grassland. In: R.T. 1,104 p.
Coupland, ed. Natural grasslands: Introduction
and Western Hemisphere. Elsevier, New York. Warnock, B.H. 1974. Wildflowers of the
p. 337-362. Guadalupe Mountains and the Sand Dune
Country, Texas. Sul Ross State University,
Starr, G. 1999. Dasylirion – The shaggy lilies. Alpine, TX. 176 p.
Desert Plants 15(1): 29-31.
White, L.D. 1969. Effects of a wildfire on several
Tesky, J.L. 1993. Dasylirion wheeleri. In: U.S. desert grassland shrub species. Journal of Range
Department of Agriculture, Forest Service, Management 22: 284-285.
Rocky Mountain Research Station, Fire
285
Datura inoxia P. Mill. devil’s trumpet
SOLANACEAE
Synonyms: Datura metel auct. non L.

Datura fastuosa sensu Griseb. non L.
John K. Francis
The ovoid capsule is nodding, about 3 cm in
diameter and covered with stout, soft prickles 2 to
4 mm long. The capsules remain on the plant for a
long period. The yellowish-brown seeds are flat,
kidney-shaped, about 5 mm long, and have a small
fleshy aril. Datura species normally have 12 pairs
of chromosomes (Bonde 2001, Burkill 2000,
Howard 1989, Liogier 1995, Stevens and others
2001).
Range.—The native range of devil’s trumpet

appears to have been Mexico and the U.S.
Southwest (Schultes and Hofmann 1992), the
Caribbean Islands (Howard 1989), India, and
China (Schultes and Hofmann 1992). There are
related species in all these areas and ancient
traditions of its use (Bonde 2001, Schultes and
Hofmann 1992). The species apparently
naturalized in Africa hundreds of years ago
(Burkill 2000). Today, devil’s trumpet grows
throughout the Tropics.
Ecology.—Devil’s trumpet grows naturally in

disturbed areas such as eroded sites, old fields,
vacant lots, overgrazed pastures and rangeland,
roadsides and abandoned roadbeds, and fencerows.
Apparently, disturbance and reduced competition
are required for the plant to become established
Drawing source: Missouri Botanic Garden 2003
and grow. A wide variety of well-drained soils on
both igneous and sedimentary parent materials are
General Description.—Devil’s trumpet is also
suitable. In Puerto Rico, the species grows
known as angel’s trumpet, thorn apple, Indian
naturally in areas that receive from 750 to about
apple, purple datura, garden datura, horn-of-plenty,
1000 mm of mean annual precipitation from near
David bush, chaico blanco, buenas tardes,
sea level to about 400 m. Devil’s trumpet grows on
concombre zombi, cornicopio, trompette du
sites up to 900 m in elevation in Nicaragua
jugement, and pomme épineuse. It is a shrub or
(Stevens and others 2001)
woody herb to 2 m in height that is often grown as
an annual in temperate zones. The stems are
Reproduction.—Devil’s trumpet flowers and
semiwoody and suffruticose in the moist tropics. In
fruits throughout the year in some environments,
drier environments, it dies to the ground annually.
but in India principally from July to September
The alternate leaves have petioles 3 to 7 cm long
(Parrotta 2001), and from September through
and ovate to elliptic blades 6 to 15 cm long with
November in Nicaragua (Stevens and others 2001).
sinuate to irregularly toothed edges. The tubular
Hummingbirds sometimes visit the flowers, but are
flowers are axillary and usually solitary. They are
affected by the alkaloids in the nectar and must
erect or nodding, have a five-toothed calyx 5 to 7
limit their consumption. Honeybees are apparently
cm long and a white, purple, or yellow corolla 8 to
unaffected. The flowers have an intense night
20 cm long, often double in horticultural varieties.
286
fragrance (Annie’s Annuals 2002), which perhaps D%20-%20G/Datura_metel_’Belle_Blanche’.
helps attract night-flying moths. The seeds are htm. 1 p.
distributed by ants and some species of birds that
are resistant to the chemicals they contain (Bonde Bonde, K. 2001. The genus Datura: From research
2001). Seeds germinate in less than 2 weeks at soil subject to powerful hallucinogen.
temperatures of 27 °C (Hardy Plants 2002). Devil’s http://leda.lycaeum.org/Documents/The_Genus_
trumpet is normally propagated with seeds. Datura:_From_Research_Subject_to_Power.
9 p.
Growth and Management.—In dry environments,
devil’s trumpet reaches a maximum height of about Burkill, H.M. 2000. The useful plants of West
0.5 m in one growing season. In moist areas, it can Tropical Africa. Vol. 5. 2nd ed. Royal Botanical
reach two to four times this height although it is Gardens, Kew, UK. 686 p.
not clear whether it does so in a single season. New
stands can be established by cultivation and Erowid. 2002. Erowid experience vaults.
planting. Although control information has not http://www.erowid.org/experiences/subs/ex_Dat
been published, weedy plants can probably be ura.shtml#retrospective_/_Summary. [not
reduced or eliminated without difficulty by paged].
cultivation or spraying with broadleaf herbicides.
Gupta, M., A. Bagchi, and A.B. Ray. 1991.
Benefits and Detriments.—Devil’s trumpet is Additional withanolides of Datura metel.
grown in all but the coldest climates as a flowering Journal of Natural Products. 54(2): 599-602.
ornamental. Beautiful white, purple, and yellow
varieties with large, single and double blossoms are Hardy Plants. 2002. Specialty perennials.
available. From ancient times continuing to the http://hardyplants.com/D.htm. [not paged].
present, the taking of Datura spp. tissues,
particularly the seeds, was used in shamanistic Howard, R. A. 1989. Flora of the Lesser Antilles,
rituals as a path to enlightenment (Schultes and Leeward and Windward Islands. Vol. 6. Arnold
Hofmann 1992). Today, people frequently Arboretum, Harvard University, Jamaica Plain,
experiment with it for the hallucinogenic effect, but MA. 658 p.
the results are so unpleasant (dark visions,
disorientation, amnesia, blurred vision, dry mouth, Janthangvith, J., P. Chumsri, K. Kraisintu, and A.
and incontinence) that they seldom recommend the Pongpan. 2000. Comparison of tropane alkaloid
experience (Erowid 2002). Overdoses can result in production by Datura innoxia and Datura metel
death. The plant has been used to treat impotence, varieties of white, soil, hydroponic, in vitro plant
asthma, diarrhea, as an analgesic, to control fever, culture and tranformed root culture. Mahidol
kill parasites, and as a drug for criminal purposes University annual Research Abstracts.
(Bonde 2001, Parrotta 2001). Devil’s trumpet http://www.hahidol.ac.th/abstracts/annual1999/0
contains a host of phytoactive chemicals including 355.htm. 2 p.
atropine, hyoscyamine, hyoscine, scopolamine,
norscopolamine, meteloidine, hydroxy-6- Liogier, H.A. 1995. Descriptive flora of Puerto
hyoscyamine, tiglic esters of dihydroxytropine, and Rico and adjacent islands. Vol. 4. Editorial de la
a number of withanolides (Gupta and others 1991, Universidad de Puerto Rico, San Juan, PR.
Janthangvith and others 2000). Alkaloid 617 p.
concentration in tissues is around 0.5 percent
(Niber and others 1992) but varies according to Missouri Botanic Garden. 2003. Missouri Botanic
environmental circumstances (Burkill 2000). Garden. Missouri Botanic Garden, St. Louis,
Devil’s trumpet is a minor weed in tropical and MO. http://www.mobot.org/. [not paged].
some subtropical countries. It causes erratic
behavior and even death of livestock that have Niber, B.T., J. Helenius, and A.L. Varis. 1992.
eaten it, but it is seldom a problem for pastured Toxicity of plant extracts to three storage beetles
animals because they carefully avoid consuming it. (Coleoptera). Journal of Applied Entomology
113(2): 202-208.
References
Parrotta, J.A. 2001. Healing plants of India. CABI
Annie’s Annuals. 2002. Datura metel, ‘Belle Publishing, Wallingford, UK and New York.
Blanche.’ http://www.anniesannuals.com/signs/ 917 p.
287
Schultes, R.E. and A. Hofmann. 1992. Plants of Stevens, R.D., C. Ulloa-U., A. Pool, and O.M.
the gods. Healing Plants Press, Rochester, NY. Montiel, eds. 2001. Flora de Nicaragua.
192 p. Monographs in Systematic Botany Vol. 85, No.
3. Missouri Botanic Garden, St. Louis, MO. pp.
911-2,666.
288
Desmanthus virgatus (L.) Willd. wild-tantan
FABACEAE
Synonyms: Mimosa virgata L.

Acuan virgatum (L.) Medikus
Acuan guadeloupense Britton & Rose
Desmanthus depressus Humb. & Willd.
John K. Francis
short petiole, less than seven pinnae, 10 to 25 pairs
of leaflets per pinnae that are linear-oblong and 4
to 9 mm long. The flowers occur in axillary heads
with peduncles 2 to 7.5 cm long. The petals are
white or creamy yellow. The pod, a legume, is
linear, 50 to 90 mm long, 3 to 4 mm broad,
reddish-brown, and contains 20 to 30 seeds
(Howard 1988, Liogier 1988, Skerman and others
1988). There are 2n = 28 chromosomes (Long and
Lakela 1976).
Range.—Wild-tantan is native to the Greater and

Lesser Antilles, Trinidad and Tobago, from Texas
to Argentina, and the Galapagos Islands (Liogier
1988, Howard 1988, Skerman and others 1988). It
has naturalized in Florida and Hawaii (Institute of
Systematic Botany 2002, Neal 1965). The species
has been planted throughout the tropics and
subtropics and has probably naturalized in many
other areas.
Ecology.—Wild-tantan grows well in deep, well-

to moderately well-drained silts, clays, loams, and
sands with pH’s from about 5 to about 8, formed
over both igneous and sedimentary rocks. It is
seldom seen in shallow, rocky sites. The species is
recommended for planting in Australia in areas
receiving from 550 to 750 mm of mean annual
precipitation (Department of Primary Industries
2002) and grows naturally in Puerto Rico in areas
General Description.—Wild-tantan, also called
that receive from 750 to about 1600 mm of mean
prostrate bundleflower, dwarf koa, desmanthus,
annual precipitation. Wild tantan tolerates seasonal
desmanto, acacia courant, acacia savane, and
drought without difficulty. The species grows from
pompon blank, is a suffruticose shrub or perennial
near sea level up to 300 m in elevation (Skerman
woody herb that reaches 0.5 to 2 m in height and 3
and others 1988). Wild-tantan is intolerant of
to 10 mm in basal diameter. The plant is supported
shade. It tolerates severe competition from grass
by a deep woody taproot and descending laterals.
and herbs but disappears when overtopped by trees.
Each dry season, wild-tantan dies back to the root
The species is usually more or less evenly
crown or lower stems and grows anew when the
dispersed in stands of grass and herbs, often at 1 or
rains return. The stems and branches are semi-
2 per square meter. Although plants are killed to
woody to woody, pithy in the center, angular,
the ground by heavy frost, they regrow from the
slender, green turning brown, and erect or
root crown (Forlin and others 2000). It withstands
semiprostrate. There may be as many as 50 stems,
heavy grazing pressure and recovers well from fire.
depending on the age, vigor, and disturbance
history of the plant. The 2- to 8-cm leaves have a
289
Reproduction.—Wild-tantan blooms at the end of Government, Brisbane, Australia. http://dpi.qld.
the wet season, and fruits mature about 1 month gov.au/pastures/4490.html. 2 p.
later. Seeds collected in Puerto Rico weighed an
average of 0.0041 + 0.0005 g/seed or 244,000 Forlin, S.M., H.Y. Rey, and L.A. Mroginski. 2000.
seeds/kg. Placed to germinate on moist filter paper, Cultivo de tejidos de Desmanthus virgatus:
only 28 percent germinated in 1 year. The atención de plantas a partir de hojas.
remaining seeds were scarified (by nicking) and Universidad Nacional del Nordeste, Corrientes,
100 percent germinated starting in 2 days (author’s Argentina. Comunicaciones Científicas y
observation). In nature, scarification occurs by Tecnológicas 2000. 3 p.
weathering of the seed coat in the soil or while
passing through an herbivore. Artificial Fox, J.E.D., A.I. Doronila, D.R. Barrett, and I.K.
scarification by treatment with concentrated Surata. 1996. Desmanthus virgatus (L.) Willd.
sulfuric acid for 8 minutes is recommended An efficient intermediate host for the parasitic
(Skerman and others 1988). The seeds are species Santalum album L. in Timor, Indonesia.
dispersed by ruminant animals. Many seedlings Journal of Sustainable Forestry 3(4): 13-23.
spring up under favorable conditions in natural
stand, but few survive. The species can be Howard R.A. 1988. Flora of the Lesser Antilles,
propagated by rooting hormone-treated leaves Leeward and Windward Islands.
(Forlin and others 2000). Dicotyledoneae, Part 1. Vol. 4. Arnold
Growth and Management.—Wild-tantan MA. 673 p.
normally grows from 0.5 to 1.5 m/year in height.
After establishment, the root crown gradually Institute of Systematic Botany. 2002. Atlas of
increases in size until it reaches 15 cm across in 3 Florida plants: Desmanthus virgatus. University
years. Plantations are established by sowing 2 kg of of Florida, Gainesville, FL. http://www.
scarified seed per hectare into well-prepared plantatlas.usf.edu/maps.asp?plantID=1474. 1 p.
seedbeds at a depth of 1 to 1.5 cm. Plantations can
be cut for feed 4 times/year and yield 35 Labios, R.V., J.G. Montesur, and R.O. Retales.
tons/ha/year of dry matter. There is little mortality 1994. Alley cropping in sloping upland rice
up to at least the fourth year (Skerman and others areas of Cavite, Philippines. Philippine Journal
1988). of Crop Science 19(1): 33-37.
Benefits.—Wild-tantan is planted throughout the Liogier H.A. 1988. Descriptive flora of Puerto
tropics as a forage and hay crop and is appreciated Rico and adjacent islands, Spermatophyta. Vol.
as a forage in range land. The forage is nontoxic 2. Río Piedras, PR: Editorial de la Universidad
and palatable to cattle and other ruminants. It is a de Puerto Rico. 481 p.
favorite of white-tailed deer in southern Texas
(Soltero-Gardea and others 1994). Crude protein Long, R.W. and O. Lakela. 1976. A flora of
content of whole plants from plantations ranges Tropical Florida. Banyon Books, Miami, FL.
from 10.5 to 15.5 percent, the leaves having 22.4 962 p.
percent and the stems 7.1 percent (Skerman and
others 1988). Wild-tantan is planted as a hedge- Neal, M.C. 1965. In gardens of Hawaii. Special
row plant in the alley cropping system on steep Publication 50. Bernice P. Bishop Museum
land in the Philippines (Labios and others 1994). It Press, Honolulu, HI. 924 p.
has been shown to be an efficient nursery-stage
host for sandalwood (Santalum album L.) (Fox and Skerman, P.J., D.G. Cameron, and F. Riveros.
others 1996). The species can become a serious 1988. Tropical forage legumes. 2nd. Ed. Food
weed in sugar cane fields (Neal 1965). and Agriculture Organization of the United
Nations, Rome. 692 p.
References
Soltero-Gardea, S., I.M. Ortega, and F.C. Bryant.
Department of Primary Industries. 2002. Legumes 1994. Nutrient content of important deer forage
for the tropics and subtropics: Desmanthus plants in the Texas costal bend. Texas Journal of
(Desmanthus virgatus). Queensland Science 46(2): 133-142.
290
Distictis lactiflora (Vahl) DC. pega palo
BIGNONIACEAE
Synonyms: Bignonia lactiflora Vahl

Bignonia rigescens Jacq.
Bignonia odorata Bello
Distictis rigescens (Jacq.) DC.
Macrodiscus lactiflorus (Vahl) Bureau ex Baill.
John K. Francis
in the same areas and is similar in appearance to

Macfadyena unguis-cati (L.) A. Gentry. They are
distinguished by the tripartite tendril, yellow
flower, and long pod of M. unguis-cati.
Range.—Pega palo is native to Cuba, Hispaniola,

Puerto Rico, and the Virgin Islands. It has been
introduced into Guadeloupe (Liogier 1995).
Ecology.—Pega palo grows in areas that receive

from about 750 to 2000 mm of annual
precipitation. It grows on most types of well-
drained to excessively drained soils derived from
metamorphic (including ultramafic) rocks. Pega
General Description.—Pega palo, also known as palo grows from near sea level to over 600 m in
liana fragante and viuda, is a climbing woody vine elevation. The species is moderately intolerant,
that clings to objects by means of sticky disks and growing in low basal-area forests, forest edges and
may extend 6 m or more into the crowns of trees or openings, brushy pastures, fencerows, and
laterally along rocks and fences (Acevedo- disturbed forests. Pega palo is not eaten by cattle
Rodríguez 1985). Older plants may have multiple and is often common in overgrazed range.
stems arising from ground level. Like many other Although the usual habit of the species is to ascend
vines in the Bignoniaceae, pega palo has a into trees or over rocks, it forms mounds and
characteristic pattern in the stem cross-section, scrambles over the ground when trees are absent.
shamrock-shaped in this case. The brown stems are Pega palo resprouts after fires. An unknown insect
slender, furrowed, and fissured. Older plants have in Puerto Rico often destroys all the seeds in
no taproot, only lateral roots with sinkers and a immature capsules.
moderate amount of fine roots. The larger roots
have the same general appearance as the stems. Reproduction.—Pega palo flowers from
There are few branches until the crown of the plant December to July (Acevedo-Rodríguez 1985).
is reached. Branches usually arise in three’s at the Because the species requires nearly full sunlight to
nodes (author’s observation). The opposite, bloom and fruit and because it does not climb to
compound leaves have two leaflets with the tendril the tops of most of the trees it invades, fruiting
in place of a third leaflet. The leaf blades are ovate, plants are not abundant. A collection of fruits from
3 to 8 cm long, and obtuse, rounded, or acute at the Puerto Rico contained seeds of which only about
tip. Terminal panicles support a few white tubular one in four contained endosperm. Visually culled,
flowers with a yellow throat, 3 to 7 cm long. A few filled seed weighed an average of 0.0153 g or
of the flowers develop into narrowly ellipsoid, 65,000 seeds/kg. Sown without any pretreatment
somewhat flattened capsules 7 to 12 cm long and on peat, 55 percent germinated within 44 days.
brown at maturity. They split open by a medial Germination is hypogenous. Seedlings, whose first
suture and release brown, winged seeds (Liogier two leaves have an unusual dark-green color tinged
1995, Acevedo-Rodríguez 1985). Pega palo occurs with purple and a whitish net-like vein pattern, are
common, but few survive the first year (author’s
291
observation). Vines layer (root) when they come in References
contact with the ground.
Growth and Management.—Pega palo grows at Puerto Rico. Vol. 1. Gen. Tech. Rep. SO-58.
least a meter per year from sprouts. Individual U.S. Department of Agriculture, Forest Service,
vines live several years, and strong plants develop Southern Forest Experiment Station, New
new stems periodically. Grazing of dry and moist Orleans, LA. 331 p.
forest areas probably promotes the species.
Benefits.—Pega palo contributes to the Rico and adjacent islands. Vol. 4. Editorial de la
biodiversity of moist and dry forests and helps Universidad de Puerto Rico, San Juan, PR.
protect the soil. Its foliage and white flowers add to 617 p.
the beauty of the forest. The species has been used
as an ornamental to a limited extent.
292
Dodonaea viscosa (L.) Jacq. ‘a‘ali‘i
SAPINDACEAE
Synonymns: Dodonaea angustifolia L.

Dodonaea eriocarpa Sm.
Dodonaea sandwicensis Sherff
Dodonaea scottsbergii (Sherff) Degener & I. Degener
Dodonaea spathulata DC.
Amanda J. Stevens, Sarah A. Taylor, and Randy S. Senock
Ecology.―‘A’ali’i grows on most types of soils

and substrates in Hawaii, from old deeply leached
soils, to fertile soils high in organic materials, to
barren ‘a‘a and pahoehoe lava, and ash and cinder.
A’ali’i seedlings are considered fairly shade
intolerant and usually grow in open sites (Williams
and Buxton 1989). The ‘a‘ali‘i has a broad
ecological amplitude and can occur from near sea
level up into the sub-alpine zone, and from wet
forests to arid shrub and grasslands. It resists
droughts well and is generally one of the heartiest
plants in the arid shrublands of leeward Hawaii. In
response to wildfire ‘a’ali’i resprout from both the
root crowns and from seed. Following wildfire in
diverse ‘a‘ali‘i-dominated montane shrublands, it
General Description.—‘A’ali’i is also known as
doubled its preburn density within 1 year. Its
a’ali’i-ku ma kua, a’a’li’i ku makani, lampuaye,
density, however, declined over subsequent years
hop bush, hopseed bush, togovao, and akeake. It is
but did regain preburn cover values 4 years
usually a many-branched shrub or small tree
following the burns (Sherry and others 1999). In
reaching a height of 4.5 to 9.0 m, and 7.5 cm in
Hawaii, a’ali’i can be affected by a yellowing
diameter (Little and Skolmen 1989). The bark is
disease distinguished by chlorotic witches’-
reddish brown to gray or blackish gray, finely
brooms. Infected areas of the plants fail to produce
furrowed, black heartwood. Leaves are simple,
flowers, while flowering is normal on the
alternate, and membranous to chartaceous, 3 to 10
nonafflicted parts of the same plants. The terminal
cm long and 1 to 3 cm wide and vary in shape
leaves of the portion of the afflicted branches
from elliptic to spatulate, oblanceolate or
become chlorotic, crinkled, and reduced in size.
lanceolate, entire margins, finely hairy when
This disease can affect all stages of maturity, and
young, yellow green, sticky coating, slightly shiny
plants will eventually die. A relatively small
above, and paler below. Petioles are 0.1 to 1.1 cm
portion of these plants are infected in Hawaii, but
long and slightly winged (Wagner and others
this disease is spreading and can be found on all
1990, Little and Skolemen 1989, Lamb 1981).
the major islands (Borth and others 1995).
Range.—‘A’ali’i has a pantropical distribution
Reproduction.—The flowers are unisexual,
occuring in temperate regions of Australia, Africa,
sometimes perfect, in paniculate inflorescences
Mexico, New Zealand, India, Samoa, Guam,
usually 1 to 5 cm long, clustered at the ends of the
Northern Mariana Islands, Virgin Islands, Puerto
twigs (Wagner and others 1990). The flowers are
Rico, Florida, Arizona, South America, Hawaii
sticky with fine hairs, three to five sepals, and
and elsewhere (West and Noble 1984). ‘A‘ali‘i
without petals. Female flowers have four stigmas
occurs on all the main Hawaiian islands except
and male flowers have ten stamens (Little and
Kaholawe.
Skolmen 1989). Male and female flowers are born
on different trees. The fruit is a capsule straw-
293
colored to dark reddish purple, exceedingly phytoplasmas in Hawaii. Plant Disease. 79(11):
variable, turgid to inflated bladder or compressed. 1,094-1,097.
Seeds are compressed-ovoid, about 3 mm long.
The two to four-winged, papery capsules are either Burrows, C.J. 1995. Germination behavior of the
dispersed by wind or adhere to plumage of birds seeds of six New Zealand woody plant species.
(Burrows 1995). The seed coats of most of the New Zealand Journal of Botany 33: 365-377.
‘a’ali’i seeds are impermeable to water and there
also appears to be a micropylar plug present. The Cribb, A.B. and Cribb, J.W. 1981. Wild Medicine
hard seed coat and the micropylar plug may play a in Australia. Collins, Sydney. 228 p.
role in delayed germination. Some evidence
suggests that higher temperatures, which may Jain, R.K. 1998. Fuelwood characteristics of
enhance the breakdown of the plug, as well as medium tree and shrub species of India. In:
scarification of the seed coat, may be necessary for Bioresource technology. Volume 68, Issue 3.
germination (Burrows 1995). ‘A‘ali‘i is able to Elsevier Science Limited, Essex, UK. p. 305-
reproduce through sprouting following 308.
disturbance.
Lamb, S.H. 1981. Native trees and shrubs of the
Growth and management.—‘A‘ali‘i is popular as Hawaiian Islands. The Sunstone press, Santa
an ornamental in xeriscape gardens. Seedlings Fe, NM. 160 p.
should be planted from containers after last frost in
any kind of soil. Water infrequently once Little, E.L. and R.G. Skolmen. 1989. Common
established and prune if a more formal appearance forest trees of Hawaii (native and introduced).
is desired. Plant emits strong odor during pollen Agriculture Handbook. 679. U.S. Department of
season. Plant is frost damaged at approximately – Agriculture, Washington, DC. 321 p.
6.6° C (Anonymous 1999).
Pengelly, A. Dodonaea viscosa. www.nhaa.org.au/
Benefits.—‘A‘ali‘i has enormous potential as a forum/text/dodvis2.html.
landscape species. ‘A’ali’i also possesses many
medicinal properties and has been used by native Rojas, A., S. Cruz, H. Ponce-Monter, and R. Mata.
peoples from all regions where it is found. Its 1996. Smooth muscle relaxing compounds from
leaves can be used to combat rheumatism, skin Dodonaea viscosa. Planta Medica. 62:154-159.
infections, fevers, swellings, aches, and
“gastrointestinal disorders” including diarrhea, and Sherry, K., J.M. Castillo, and R.B Shaw. 1999.
can be used as a antispasmodic agent (Rojas and Effects of wildfire on vegetation and rare plants
others 1996). The Australian aborigines used the in arid montane shrublands. Pohakaloa training
leaves and roots as a painkiller to soothe area, Hawaii. 1999 Hawaii Conservation
toothaches and headaches (Crib and Crib 1981). Conference, Honolulu, HI.
The flowers (or hops) of ‘a’ali’i are used as a
“home-brew” substitute to bestow a bitter flavor, Wagner, H., C. Ludwig, L. Grotjahn, and M.S.Y.
and also used as a tonic. A red dye is extracted Khan. 1987. Phytochemistry 26(3): 697-701.
from the fruit, and the capsules are also commonly
used in leis (Little and Skolmen 1989). In India the Wagner, W.L., D.R. Herbst, and S.H. Sohmer.
seeds are used as fish poison (Wagner and others 1990. Manual of the Flowering Plants of
1987). The wood from ‘a’ali’i is extremely hard Hawaii. University of Hawaii Press, Honolulu,
and has been used for tools, spears and weapons HI. 1,854 p.
(Lamb 1981). In India ‘a’ali’i is an important
source of fuelwood (Jain 1998). Williams, P.A. and R.P. Buxton. 1989. Response
to reduced irradiance of 15 species of native and
References adventive shrub and tree seedlings from eastern
Canterbury. New Zealand Journal of Ecology.
Anonymous. 1999. Dodonaea viscosa: Hopbush. 12: 95-101.
www.plantadviser.com/plants/dodovisc.shtml
West, J.G. and I.R. Noble. 1984. Analyses of
Borth, W.B., J.S. Hu, B.C. Kirkpatrick, D.E. digitized leaf images of the Dodonaea viscosa
Gardner, and T.L. German. 1995. Occurrence of complex in Australia. Taxon 33(4):595-613.
294
Dryas octopetala L. white mountain avens
ROSACEAE
Synonyms: Dradetum octopetalae Keiner
(Kershaw and others 1998). The stalk, midrib, and

veins have yellowish-brown glands on the lower
surface. Each flower has eight (sometimes nine or
10) egg-shaped white, rarely yellowish, petals and
lance-shaped sepals. The flowers are 2 to 3 cm
across and are solitary on hairy stalks that are 1 to
20 cm long. The seeds (achenes) are 2.5 to 4 mm
long with a feathery plume (style) at the tip
(Komarov 1941). The chromosome number is 18
(Stace 1997, Welsh and others 1993). The species
tends to be heterogeneous throughout its range.
The stature and organs of white mountain avens
plants in North America and a part of the Western
Arctic appear to have smaller dimensions than
those in Russia and alpine Western Europe
(Komarov 1941). At least four subspecies and
several varieties of Dryas octopetala are
recognized (Hitchcock and Cronquist 2001,
Kartesz 1994): Dryas octopetala L. ssp.
alaskensis (Porsild) Hultén, ssp. hookeriana (Juz.)
Breit., ssp. punctata (Juz.) Hultén, and ssp.
octopetala of which there are at least three
varieties; namely var. angustifolia Hitch., var.
kamtschatica (Juz.) Hultén, and var. luteola
Hultén. Some other varieties are described in
Drawing source: USDA—Forest Service collection, Kartesz (1994) and Komarov (1941).
Hunt Institute
Range.—White mountain avens is a circumpolar
General Description.—Dryas octopetala is species occurring in arctic and alpine zones in
commonly known as alpine dryad, mountain Europe, Asia, and North America. In North
avens, or white mountain avens in areas where America, it grows in sub-alpine and alpine tundra
more than one Dryas species occur (Stace 1997, zones from Greenland, Labrador, and Alaska to
Kershaw and others 1998, Carter 1988). The the Cascade Mountains in Washington and Oregon
epithet “dryad” and “dryas” is derived from the and through the Rocky Mountains to Montana,
mythical oak-nymph, Dryas, because the shape of Wyoming, and Colorado and west to high
its leaves resembles that of an oak species elevation sites in Utah and Idaho (Britton 1901,
(Gledhill 1992, Zwinger and Willard 1996). This Hitchcock and Cronquist 2001, Welsh and others
prostrate, trailing perennial shrub is usually less 1993, Kershaw and others 1998). It is found in
than 30 cm tall, but can grow up to 50 cm, and Scandinavia, the British Isles, Ireland, and the
commonly forms mats. The oval to oblong leaves Alps, Pyrenees, Carpathians and Dinarid
are 5 to 25 cm long and the margins are coarsely Mountains of Europe (Praprotnik 1997, Ellenberg
round-toothed (crenate) and somewhat rolled 1988). It also dominates the tundras of Novaya
under (Carter 1988, Kershaw and others 1998, Zemlya, Wrangel Island, the Urals and mountin
Stace 1997). Typically the shrub is described as slopes in Siberia (Komárková 1979, Botch and
“evergreen” but has also been observed as “semi- Masing 1983).
deciduous,” because it loses some of its foliage in
winter (Williams and others 1986). The leaves are Ecology.—Mountain avens is often the dominant
dark green, wrinkled and hairless on the upper side or co-dominant species in areas where it occurs
but densely white-hairy on the lower surface throughout its range. The elevation at which it
295
grows is associated with latitude. Plants are found when sown outdoors in the autumn while seeds did
at 3,500 m to 3,965 m in Utah whereas they may geminate the following spring, suggesting that
occur at 100 m or less in northern parts of its cold stratification lowers the temperature
range, such as Greenland (Borge Wills 1996, requirement for germination (Elkington 1971). The
Welsh and others 1993). It typically grows above stems root easily and contribute to the rapid spread
the timberline in sites with low snow cover on of plants. As part of studies on the consequences
calcareous or basic soils (Dorn 2001, Dorn 1984, of global climate change, experiments have been
Komárková 1979, Ellenberg 1988, Stace 1997, made on the response of white mountain avens to
Weber and Wittman 2001). It is not an indicator of elevated temperatures in the Colorado Rocky
calcareous sites per se as it is common on granitic Mountains (Welker and others 2001). The results
substrates in the Colorado Rocky Mountains suggested that warmer, wetter conditions would
(Weber 1960). However, on this substrate calcium initially lead to greater productivity (increased leaf
is likely available to the plants as a leachate mass and higher numbers of seed per ramet)
(Weber and Wittman 2001). Mountain avens although decomposition may be retarded, leading
grows in dry meadows, rocky tundra, fellfields, to a lower nutrient availability in subsequent years
and in relatively moist sites in fen lands. In Russia, (Welker and others 2001). Under longer term
it has been reported as a co-dominant in “frost conditions, a switch in the source-sink activity of
mound bogs” that are essentially permanently the community occurred. Under current conditions
frozen ridges, or mounds, that alternate with wet the mountain avens community is a carbon sink of
hollows (Botch and Masing 1983). In contrast, about 8 g CO2-C m-2. However, after 4 years of
mountain avens and lichens are of primary experimental warming, the community switched to
importance on the dry upland meadows in the a carbon source of almost 10 g CO2-C m-2 per
Alaskan Arctic and on the windswept fellfield summer due primarily to an increase in the rate of
slopes of the Rocky Mountains of Colorado U.S.A. respiration without a concomitant increase in
(Komárková 1979, Langenheim 1962). These photosynthesis (Welker and others 2001). This
shrubs have an trellis-like growth habit clinging switch in source sink activity has been reported for
closely to rocks and stones and anchoring its roots Dryas and other communities in the Arctic
in cracks or between stones. The radiation is so (Welker and others 2001, Oechel and others 1993).
intense at high elevations that the temperature can This finding may be significant when considering
rise to more than 65 oC where the ground is dark the repercussions of increasing industrial
colored. In these locations the prostrate seedlings “greenhouse gas” emissions
and young plants can be heat damaged (Ellenberg
1988). A mutualistic association exists between Benefits.—White mountain avens is a pioneer
mountain avens and ectomycorrhizal fungi species and is important in stabilizing the thin soils
(Monson and others 2001). The most dominant on mountain slopes (Zwinger and Willard 1996,
fungus is Cenococcum geophilum, an ascomycete. Ellenberg 1988). It spreads relatively rapidly and
is popular in landscape rock gardens (Williams
Reproduction.—White mountain avens reproduce and others 1986). Several cultivated varieties,
by seed. Flowering is from June into August. The including hybrids, are available (Kershaw and
flowers always face the sun on calm, sunny days, others 1998). One hybrid, xSuendermannii
and their parabolic shape directs the sun’s heat on Kellerer, between Dryas octopetala and D.
to the stigma, which can then become up to 3.5 oC drummondii is morphologically similar to D.
warmer than the surrounding air temperature octopetala, but the flowers are yellow in bud
(Kershaw and others 1998). The warmer becoming white and nodding upon opening
temperatures can attract insects and also speed (Bailey and Bailey 1976). White mountain avens is
pollen germination. Hybridization occurs between an important food source for ptarmigian (Nelson
sympatric Dryas species (Komarov 1941). Seed is, 1992) and pikas (Martin and others 1951).
at least in part, distributed by wind.
References.
Growth and Management.—White mountain
avens seeds have physiological dormancy but do Bailey, L.H., E.Z. Bailey, and the staff of the
not appear to require stratification for germination Liberty Hyde Bailey Hortorium. 1976. Hortus
(Baskin and Baskin 2001). Forty-eight percent of Third. Macmillian Publishing Company, New
freshly matured seeds germinated after 4 days in York, NY. 1,290 p.
the light at 25 oC (Baskin and Baskin 2001).
However, in England seeds failed to germinate
296
Baskin, C.C. and J.M. Baskin. 2001. Seeds, Komárková, V. 1979. Alpine vegetation of the
ecology, biogeography, and evolution of Indian Peaks area, Front Range, Colorado Rocky
dormancy and germination. Academic Press, Mountains. J. Cramer, Vaduz, Germany.
New York, NY. 666 p.
Komarov, V.L. 1941. Flora of the U.S.S.R.
Borge Wills, L. 1996. Sanderling review. Marine Volume 10. Izdatel’stvo Akademii Nauk SSSR.
and Coastal Species Information System, Translated by N. Landau 1970. The Smithsonian
Conservation Management Inst., Roanoke, VA. Institution, Washington, D.C. p. 272-273.
http://fwie. fw.vt.edu [not paged].
Langenheim, J.H. 1962. Vegetation and
Botch, M. and V. Masing. 1983. Mire ecosystems environmental patterns in the Crested Butte area.
in the U.S.S.R. In: A.J.P. Gore, ed. Ecosystems Ecological Monographs 32(3): 249-285.
of the World, Vol. 4B. Elsevier Company, New
York. p. 95-152. Martin, A.C., H.S. Zim, and A.L. Nelson. 1951.
American wildlife and plants. McGraw-Hill
Britton, N.L. 1901. Manual of the flora of the Book Company, New York, NY. 500 p.
northern states and Canada. Henry Holt and
Company, New York, NY. 1,080 p. Monson, R.K., R. Mullen, and W.D. Bowman.
Plant nutrient relations. 2001. In: W.D. Bowman
and TR. Seastedt, eds. Structure and function of
Carter, J.L. 1988. Trees and shrubs of Colorado.
an alpine ecosystem. Oxford University Press,
Johnson Books, Boulder, CO. 165 p.
Dorn, R. D. 1984. Vascular plants of Montana.
Nelson, R.A. 1992. Handbook of Rocky Mountain
Mountain West Publishing Cheyenne, WY. 412
plants. Robert Rinehart Publishers, Niwot, CO.
p.
444 p.
Dorn, R. D. 2001. Vascular plants of Wyoming.
Oechel, W.C., S.J. Hastings, G.L. Vourlitis, M.A.
Mountain West Publishing Cheyenne, WY. 412
Jenkins, G. Riechers, and N. Grulke. 1993.
p.
Recent changes of arctic tundra ecosystems from
a carbon sink to a source. Nature 361: 520-523.
Elkington, T.T. 1971. Biological flora of the
British Isles. Dryas octopetala. Journal of
Ecology 59: 887-905. Praprotnik, N. 1997. Alpski botanicni vrt Juliana v
Trenti. Prirodoslovni muzej Slovenije,
Ellenberg, H. 1988. Vegetation ecology of central Ljubljana, Slovenia. 126 p.
Europe. 4th edition. Cambridge University Press,
New York, NY. 731 p. Stace, C. 1997. New Flora of the British Isles. 2nd
edition. Cambridge University Press, New York,
Gledhill, D. 1992. The names of plants. NY. 1,130 p.
Cambridge University Press, Cambridge,
England. 202 p. Weber, W.A. 1960. Some features of the
distribution of Arctic relicts at their Austral
Hitchcock, C.L. and A. Cronquist. 2001. Flora of limits. Twelfth International Botanical Congress.
the Pacific Northwest. University of Washington Recent Advances in Botany. p. 912-914
Press, Seattle, WA. 730 p.
Weber, W.A., and R.C. Wittman 2001. Colorado
Kartesz, J.T. 1994. A synonymized checklist of the flora, western slope. University Press of
vascular flora of the United States, Canada and Colorado, Boulder, CO. 488 p.
Greenland. Vol. 1 - Checklist. Timber Press,
Portland, OR. 622 p. Welker, J.M., W.D. Bowman, and T.R. Seastedt.
2001. Environmental change and future
Kershaw, L., A. MacKinnon, and J. Pojar. 1998. directions in alpine research. In: W.D. Bowman
Plants of the Rocky Mountains. Lone Pine and T.R. Seastedt, eds. Structure and function of
Publishing, Edmonton, Canada. 383 p. an alpine ecosystem. Niwot Range, Colorado.
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Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. Williams, J., R. Radebaugh, D. Hall, G. Keladis, P.
Higgins, eds. 1993. A Utah flora. Second Keladis, and P. A. Pachuta. 1986. Rocky
Edition, Brigham Young University, Provo, UT. Mountain Alpines. Timber Press, Portland, OR.
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the trees, a guide to American alpine tundra.
Johnson Printing, Boulder, CO. 425 p.
298
Duranta erecta L. golden dewdrop
VERBENACEAE
Synonyms: Duranta repens L.

Duranta ellisia Jacq.
Duranta plumieri Jacq.
John K. Francis

1974).
Range.—Golden dewdrop is a native of Mexico,

Central America, South America to Argentina,
southern Florida (possibly naturalized), Bermuda,
the Bahamas, and the West Indies (Howard 1989,
Liogier 1995, Little and others 1974). The species
is widely cultivated and escaped in the tropics and
subtropics including Hawaii, American Samoa, and
Guam (Pacific Islands Ecosystems at Risk 2002).
Ecology.—Golden dewdrop grows wild mostly in

dry coastal areas (750 to 900 mm of annual
precipitation in Puerto Rico) from near sea level to
over 100 m in elevation (Little and others 1974). It
also grows in disturbed areas in moister habitat,
especially along roads (Pacific Island Ecosystems
at Risk 2002). Because it is moderately intolerant
of shade and does not compete well with taller
vegetation, golden dewdrop is usually found in
rocky or sandy areas with low shrubs and sparse
grass and herbs. Although it is more common on
limestone, the shrub also grows in areas with
General Description.—Golden dewdrop, also igneous rocks. The species tolerates light to
known as skyflower, pigeon berry, angels-whisper, moderate salt spray. In natural stands, golden
duranta, cuentas de oro, azota-caballo, fruita de dewdrop may be found in small tangled stands or
paloma, lila, lluvia, and grão de galo, is an upright as occasional plants. Plants under droughty and
to drooping shrub that sometimes takes the form of infertile natural conditions tend to be short (about 1
a scrambling shrub or rarely a small tree (Liogier m in height). Where cultivated in deep, moist,
1995, Little and others 1974). It usually matures fertile soils, golden dewdrop grows faster and to
with a height 1 to 3 m. The species develops tap larger sizes. As an ornamental, it can be grown in
and lateral root systems with abundant fine roots. moderate shade at elevations up to 1,300 m
There are normally several stems arising from (Bruggeman 1974). Plants in the northerly extent
below and just above the ground line. Crowns of of the naturalized range die to the ground after
golden dewdrop are replete with fine branches and frosts but resprout from the roots in the spring
twigs that are often thorny. The bark is light gray, (Floridata 1999). The species is subject of attack by
becoming rough and fissured when old. Light- scale, mealy bugs, caterpillars, and nematodes
green, opposite leaves are elliptic to ovate and 1.5 (Watkins and Sheehan 1975, Woman’s Club of
to 8 cm long. Light-blue, lavender, or white, Havana 1958).
tubular, five-lobed flowers are borne on terminal or
axillary racemes. The yellow or yellow-orange Reproduction.—Natural stands of golden dewdrop
fleshy fruits are ellipsoidal with five lobes and in Puerto Rico flower and fruit in spring and
grow in hanging clusters. These fruits may contain summer (Little and others 1974). However,
up to eight nutlets (Hardin and Arena 1969, ornamental plants in Puerto Rico flower and fruit
299
throughout the year (author’s observation), and in Castro, O., M. Barrios, M. Chinchilla, and O.
Florida, both flowers and fruit are present on the Guerrero. 1996. Chemical and biological
shrubs at the same time (Nelson 1996). A evaluation of the effects of plant extracts against
collection of golden dewdrop fruits from Puerto Plasmodium berghei. Revista de Biologia
Rico averaged 0.515 + .009 g/fruit. Air-dried seeds Tropical 44(2A): 361-367.
separated from them averaged 0.0346 + 0.0005
g/seed or 28,900 seeds/kg. Set to germinate on Floridata. 1999. Duranta erecta. http://
moist filter paper without any pretreatment, 29 floridata.com/ref.d.dra_ere.cfm. 3 p.
percent of the seeds germinated beginning 128
days after sowing (Francis and Rodíguez 1993). Francis, J.K. and A. Rodríguez. 1993. Seeds of
Birds disseminate the seeds (Watkins and Sheehan Puerto Rican trees and shrubs: second
1975). However, seedlings are seldom common. installment. Research Note SO-374. U.S.
After establishment, plants thicken and spread Department of Agriculture, Forest Service,
laterally a few decimeters vegetatively. Golden Southern Forest Experiment Station, New
dewdrop are produced commercially from seeds, Orleans, LA. 5 p.
cuttings, and by layering (Floridata 1999).
Hardin, J.W. and J.M. Arena. 1969. Human
Growth and Management.—Golden dewdrop poisoning from native and cultivated plants.
plants live at least 15 years. The species has a Duke University Press, Durham, NC. 167 p.
moderate growth rate, usually about 0.5 m/year for
the first few years. Ornamental plants need regular Howard, R.A. 1989. Flora of the Lesser Antilles.
thinning and pruning to keep them under control Vol. 6. Arnold Arboretum, Harvard University,
and to remove dead branches (Floridata 1999). Jamaica Plain, MA. 658 p.
Although they withstand poor soil and drought
well, plants grow best if watered and fertilized Liogier, H.A. 1995. Descriptive flora of Puerto
(Woman’s Club of Havana 1958). Rico and adjacent islands. Vol. 4. Editorial de la
Benefits and Detriments.—Golden dewdrop 617 p.
forms a part of the coastal scrub community and
contributes to soil and ecosystem stability. It is a Little, E.L., Jr., R.O. Woodbury, and F.H.
popular ornamental used for accent plants and Wadsworth. 1974. Trees of Puerto Rico and the
hedges in tropical and subtropical parts of the Virgin Islands. Vol. 2. Agriculture Handbook
world because of its profuse displays of flowers 449. U.S. Department of Agriculture,
and fruits (Floridata 1999). Golden dewdrop Washington, DC. 1,024 p.
flowers attract butterflies and hummingbirds
(Floridata 1999). It is sometimes grown in Nelson, G. 1996. The shrubs and woody vines of
greenhouses in areas too cold for natural plants. Florida. Pineapple Press, Inc., Sarasota, FL.
The wood is light brown and hard and useful for 391 p.
stakes and fuel. Golden dewdrop is a poisonous
plant and has caused deaths of children. Saponins Pacific Island Ecosystems at Risk. 2002. Duranta
in the fruits and foliage cause gastroenteric erecta L., Verbenaceae. http://www.hear.
irritation, drowsiness, fever, nausea, vomiting, and org/pier3/duere.htm. 2 p.
convulsions. Dermatitis sometimes occurs from
handling the plants (Hardin and Arena 1969, Watkins, J.V. and T.J. Sheehan. 1975. Florida
Westbrooks and Preacher 1986). However, birds landscaping plants, native and exotic. The
feed on the fruits without difficulty (Nelson 1996). University Presses of Florida, Gainesville, FL.
Ethyl acetate and aqueous extracts of leaves 420 p.
showed significant antimalarial activity when
administered to mice (Castro and others 1996). In Westbrooks, R.G. and J.W. Preacher. 1986.
small quantities, fruits are used to treat intestinal Poisonous plants of Eastern North America.
worms (Whistler 2000). University of South Carolina Press, Columbia,
SC. 226 p.
References
Whistler, W.A. 2000. Tropical ornamentals, a
Bruggeman, L. 1964. Tropical plants and their guide. Timber Press, Inc., Portland, OR. 542 p.
cultivation. Thames and Hudson, London. 228 p.
300
Woman’s Club of Havana. 1958. Flowering plants
from Cuban gardens. Criterion Books, New
York. 365 p.
301
Encelia farinosa Gray ex Torr. brittlebush
ASTERACEAE
Synonyms: none
Catherine E. Koehler and Arlee M. Montalvo
Gray, E. californica Nutt. and E. actoni Elmer

(Ehleringer and Clark 1988, Hickman 1993).
Intergeneric hybrids with the annual Gerea
canescens T. & G. have also been reported (Kyhos
1967).
Range.—Brittlebush occurs within sage scrub of

the inland valleys of southern California, eastward
and southward into arid habitats of the Sonoran
and Mojave deserts and into Arizona, northwestern
Mexico, southern Nevada, and southwestern Utah
at under 1,000 m elevation (Munz and Keck 1968,
Hickman 1993, Sandquist and Ehleringer 1997). In
California, it has been extending westward and
northward due to roadside and other plantings.
Geographic Variation.—Geographic variation
and adaptation to local environments have been
documented for many brittlebush traits. In
General Description.—Brittlebush, a dry particular, adaptation to water availability has been
environment-adapted shrub with weak branches, is well demonstrated, including genetic
also known as incienso because it has fragrant differentiation in water use efficiency (Sandquist
resin, and as desert encelia because it occurs in and Ehleringer 2003). Also, individuals with
desert and semidesert habitats. The specific epithet brown-purple disk florets (var. phenicodonta)
farinosa refers to the mealy pubescence of the occur in areas with higher levels of soil moisture
leaves, which is composed of appressed wooly and are replaced by the yellow-disked form (var.
hairs. These shrubs have dense rounded canopies farinosa) in drier sites. This pattern may involve
with alternate whitish to greenish-gray leaves natural selection in response to water availability
clustered near stem tips. Adults are 0.3 to 1.5 m (Kyhos 1971) but may also be linked to the lower
tall, shallowly rooted (40 cm), with one to several frost tolerance of var. phenicodonta (Sandquist
many-branched stems. Leaves are simple, 3 to 8 and Ehleringer 1996). Similarly, Monson and
cm long, ovate to rhombic in outline, with wavy others (1992) found localized physiological and
margins, and short petioles. Heads are arranged in genotypic differences in water use between plants
loose, naked panicles and have both disk and ray at the base and the top of a slope that coincided
flowers subtended by green, glandular bracts that with a moisture stress gradient.
embrace the flat, obovate, 4.5 mm long achenes.
Involucre bracts are imbricate in three to four The degree of leaf pubescence varies across
series. Disk florets are yellow or rich purple and regions with different mean annual rainfall, and
are surrounded by 8 to 12 mm long yellow ray variation in number of leaf hairs is both a plastic
florets (Munz and Keck 1968, Hickman 1993). response and genetically determined. Leaves of
Munz and Keck (1968) recognize two varieties: E. plants growing in arid regions are more pubescent,
f. var. radians Bdg. ex Blake from southeastern thereby having greater control over leaf
California with glabrate leaves and involucre and temperature and water loss, but they have lower
yellow disk flowers, and E. f. var. phenocodonta photosynthetic capacity due to higher reflectance
(Blake) Jtn. from the southern Colorado desert of light than plants from more mesic regions.
with purple disk flowers. Plants have n = 18 Differences are maintained when offspring are
chromosomes (Hickman 1993). Brittlebush planted together in common gardens (Sandquist
hybridizes readily with E. fructescens (A. Gray) A. and Ehleringer 1997, Housman and others 2002).
However, an additional plastic response allows
302
plants in more mesic gardens to grow less- capacity to survive drought conditions (Ehleringer
pubescent leaves than sibling plants in desert 1993). In general, however, brittlebush is drought
gardens, a response which allows plants to take adapted and responds quickly to water addition
advantage of higher water availability and through rapid CO2 uptake, leaf production, and
increased photosynthetic ability through leaf-hair stem growth (Nobel and others 1998).
reduction (Ehleringer and Clark 1988). The
inverse relationship for leaf pubescence and Reproduction.—Brittlebush flowers from March
photosynthetic activity may involve pleiotropy or through May (Munz and Keck 1968) and
linkage to other genetically determined individuals must be cross-pollinated in order to
morphological differences between populations or produce seed (Ehleringer and Clark 1988).
from environments with different moisture Flowers are visited by various insects including
regimes (Housman and others 2002). butterflies, moths, flies, bees, wasps, and beetles
(Kyhos 1971, Moldenke 1976). In one study, a
There is also clinal variation in production of beetle (Malachiidae) was found to be 10 times
chemical compounds that provide defense against more common than all other insect species
herbivores. From north to south in Baja California combined (Kyhos 1971). These potential
and east to west from the Sonoran desert to coastal pollinators do not discriminate between plants with
regions of California, plants produce progressively different disk-flower color (Kyhos 1971). Achenes
less of a sequiterpene and more of a chromene are collected from May to July (Mirov and Kraebel
toxin, which may influence local resistance to 1939) with about 770,000 bulk seeds/kg (personal
herbivores (Wisdom 1985, Kunze and others communication with S&S seeds, Carpenteria, CA).
1995). Seasonal production of these compounds Seed viability varies among years, with as few as
may also influence herbivores (Wisdom and 35 percent of seeds viable (personal
Rodriquez 1983). There are higher concentrations communication with M. Wall, Rancho Santa Ana
of these chemicals and nitrogen in young tissues. Botanical Garden, Claremont, CA). Seed
In addition, populations differ in the relative production is influenced by water stress
amount of different compounds. The specialist heightened by competition. In a desert study in
beetle Trirhabda geminata Horn experienced which nearby neighbors were removed, shrubs
lowered larval growth rates when fed higher levels experienced lower water stress, had higher
of the secondary compounds. survival, grew to nearly twice the mass, and
Ecology.—In sage scrub, brittlebush occurs on produced 53 percent more flower heads per twig
flats and slopes, primarily on weathered sandstone, and 220 percent more achenes per head than
granite, diorite, and alluvial deposits; in the desert shrubs with brittlebush neighbors within 2 m
it occurs on rocky slopes, flats, and in washes (Ehleringer 1984).
(Munz and Keck 1968, Hickman 1993). Studies on Establishment and Growth.—Brittlebush
the adaptive significance of variation in leaf form becomes established rapidly following
and pubescence both within and among disturbance. Seedlings emerge and become
populations is reviewed by Housman and others established in open areas in the winter rainy
(2002) and by Sandquist and Ehleringer (1997, season. Plants become dormant and drop many
1998). Plants react to seasonal increase in water leaves during the dry season and then sprout new
stress at the end of the rainy season by replacing leaves with the onset of winter rains. Plants can
the larger, less hairy leaves produced earlier in the reach maturity within 2 years and often live for 10
growing season with more pubescent leaves that to 15 years. In sage scrub vegetation, resprouting
are smaller and thicker. This reduces water loss success of shrubs from the base is inversely related
and regulates leaf temperature, but it also to fire intensity (Westman and others 1981, Martin
decreases photosynthetic capacity. Prolonged 1984). In one study, 2 to 30 percent of brittlebush
drought leads to dormancy and leaf drop. resprouted on slopes previously dominated by the
Brittlebush shows variation among shrub, and resprouts and seedlings surpassed
individuals and populations in water-use efficiency prefire densities within 2 years (Martin 1984). For
(the ratio of photosynthesis to transpiration, or coastal sage scrub in general, both resprouting and
delta value). Individuals with a high delta have a seedling emergence from a soil seed bank are
higher growth response if water stress is decreased negatively correlated with fire intensity (Keeley
but perform poorly in response to drought stress, 1998).
while those with a low delta show lower growth
response under low water stress and a greater Seed germination.—Some authors report seed
germination without pretreatment (Mirov and
303
Kraebel 1939, Emery 1988), but pretreatment can pellets in spring, summer, and fall in trace amounts
increase otherwise low germination rates. Padgett (Perry and others 1987). The dominant herbivores
and others (1999) found that seed stored for 6 on brittlebush leaves are the larvae and adults of
months at room temperature had 2 to 4 percent the beetle, Trirhabda geminata (Wisdom 1985).
germination while seed stored at 5 to 10 oC in a The fly Neotephritis finalis Loew lays its eggs
refrigerator had 10 to 12 percent germination. In between the florets, and the larvae feed on the
addition, treatment with gibberelic acid or achenes (Goeden and others 1987).
Ca(NO3)2 increased germination of both warm-
and cold-stored seeds approximately two-fold, and Horticulture.—Brittlebush does best with full sun
leaching with water for several days increased and good drainage (Hickman 1993, Keator 1994).
germination by about 50 percent. The rounded form with striking yellow flowers is
attractive near the back of borders or rock gardens,
Growth and Management.—Maximum growth and it is especially suitable on dry slopes (Perry
of roots occurs in the winter and early spring 1992, Keator 1994). Plants can be established
(Drennan and Nobel 1996), so plants will establish quickly from seed or containers (Perry 1992).
best if sown in late fall. Planting brittlebush seeds Provision of occasional summer water allows
can yield vegetative cover and visual appeal plants to remain attractive throughout the year
relatively quickly. However, seed mixtures should (Keator 1994).
be balanced carefully because overuse can retard
establishment of other species (Went 1942, Gray References
and Bonner 1948). Ample evidence for local
adaptation in this species underscores the need to Bean, J.L. and K.S. Saubel. 1972. Temalpakh:
collect seed material for wildland restoration from Cahuilla Indian Knowledge and Usage of Plants.
within the same ecological zone and vegetation Malki Museum Press, Morongo Indian
type as the targeted planting site so as to maximize Reservation, CA. 225 p.
success of planting projects. Because of potential
competition and hybridization, it is also important Drennan, P.M. and P.S. Nobel. 1996. Temperature
that correct native species are specified and used. influences on root growth for Encelia farinosa
Mistaken plantings of E. californica instead of (Asteraceae), Pleuraphis rigida (Poaceae), and
brittlebush, or vice versa, abound (authors’ Agave deserti (Agavaceae) under current and
observation), and hybrids between species have doubled CO2 concentrations. American Journal of
been found in such locations (personal Botany 83: 133-139.
communication with A. Sanders, University of
California, Riverside). Improper seed choices can Ehleringer, J.R. 1984. Intraspecific competitive
compromise the success of restoration efforts and effects on water relations, growth and
the genetic integrity of wild populations. reproduction in Encelia farinosa. Oecologia 63:
153-158.
Benefits.—Brittlebush was used by native tribes
for medicinal and other purposes. The resinous Ehleringer, J.R. 1993. Variation in leaf carbon
gum, heated or made into a salve, was applied to isotope discrimination in Encelia farinosa:
the chest to relieve pain and loosen bronchial implications for growth, competition, and drought
mucous. A decoction of boiled blossoms, leaves, survival. Oecologia 95: 340-346.
and stems was held in the mouth to relieve gum
and tooth ache (Bean and Saubel 1972, Moore Ehleringer, J.R. and C. Clark. 1988. Evolution and
1989). In addition, tea made from the gum has a adaptation in Encelia (Asteraceae). In: L.
numbing effect and was used to relieve arthritic Gottlieb, L. Jain and S. Jain, eds. Plant
pain (Moore 1989). The resin was also burnt as Evolutionary Biology. Chapman and Hall,
incense or melted and used as a varnish (Moore London. p. 221-248.
1989, Hickman 1993).
Brittlebush feeds numerous pollinators and Emery, D.E. 1988. Seed propagation of native
herbivores. It is an important nectar and pollen California Plants. Santa Barbara Botanical
source of the bee, Calliopsis pugionis Cockerell, Garden, Santa Barbara, CA. 23 p.
which is the host of the rare cleptoparasitic bee,
Holcopasites ruthae Cooper in Riverside County, Goeden, R.D., T.D. Cadatal, and G.A. Cavender.
CA (Visscher and Danforth 1993). Mountain 1987. Life history of Neotephritis finalis (Loew)
sheep eat brittlebush, but it is only found in fecal on native Asteraceae in southern California
304
(Diptera: Tephritidae). Proceedings of the Monson, R.K., S.D. Smith, J.L. Gehring, W.D.
Entomological Society of Washington 89: 552- Bowman, and S.R. Szarek. 1992. Physiological
558. differentiation within an Encelia farinosa
population along a short topographic gradient in
Gray, R. and J. Bonner. 1948. An inhibitor of plant the Sonoran Desert. Functional Ecology 6: 751-
growth from the leaves of Encelia farinosa. 759.
Moore, M. 1989. Medicinal Plants of the Desert
Hickman, J.C., ed. 1993. The Jepson Manual: and Canyon West. Museum of New Mexico
Higher Plants of California. University of Press, Santa Fe, NM. 184 p.
California Press, Ltd., Los Angeles, CA. 1,400 p.
Munz, P.A. and D.D. Keck. 1968. A California
Housman, D.C., M.V. Price, and R.A. Redak. Flora with Supplement. University of California
2002. Architecture of coastal and desert Encelia Press, Berkeley, CA. 1,681 + 224 p.
farinosa (Asteraceae): consequences of plastic
and heritable variation in leaf characters. Nobel, P.S., H. Zhang, R. Sharifi, M. Castañeda,
American Journal of Botany 89: 1,303-1,310. and B. Greenhouse. 1998. Leaf expansion, net
CO2 uptake, rubisco activity, and efficiency of
Keator, G. 1994. Complete garden guide to the long-term biomass gain for the common desert
native shrubs of California. Chronicle Books, San subshrub Encelia farinosa. Photosynthesis
Francisco, CA. 314 p. Research 56: 67-73.
Keeley, J.E. 1998. Postfire ecosystem recovery and Padgett, P.E., L. Vázquez, and E.B. Allen. 1999.
management: The October 1993 large fire Seed viability and germination behavior of the
episode in California. In: J.M. Moreno, ed. Large desert shrub Encelia farinosa Torrey & A. Grey
Forest Fires. Backbuys Publishers, Leiden, (Compositae). Madroño 46: 126-133.
Netherlands. p. 69-90.
Perry, B. 1992. Landscape Plants for Western
Kunze, A., C. Muller, and P. Proksch. 1995. Regions: an Illustrated Guide to Plants for Water
Chemical variation and defense of Encelia Conservation. Land Design Publishing,
farinosa. Biochemical Systematics and Ecology Claremont, CA. 318 p.
23: 355-363.
Perry, W.M., J.W. Dole, and S.A. Holl. 1987.
Kyhos, D.W. 1967. Natural hybridization between Analysis of the diets of mountain sheep from the
Encelia and Geraea (Compositae) and some San Gabriel Mountains, California. California
related experimental investigations. Madroño 19: Fish and Game 73: 156-162.
33-43.
Sandquist, D.R. and J.R. Ehleringer. 1996. Potential
Kyhos, D.W. 1971. Evidence of different adaptability and constraints of response to the
adaptations of flower color variants of Encelia changing climates for Encelia farinosa var.
farinosa (Compositae). Madroño 21: 49-61. phenicondonta from southern Baja California,
Mexico. Madroño 43: 465-478.
Martin, B.D. 1984. Influence of slope aspect on
postfire reproduction of Encelia farinosa Sandquist, D.R. and J.R. Ehleringer. 1997.
(Asteraceae). Madroño 31: 187-189. Intraspecific variation of leaf pubescence and
drought response in Encelia farinosa associated
Mirov, N.T. and C.J. Kraebel. 1939. Collecting and with contrasting desert environments. New
handling seeds of wild plants. Forestry Phytologist 135: 635-644.
Publication 5. Civilian Conservation Corps,
United States Government Printing Office, Sandquist, D.R. and J.R. Ehleringer. 1998.
Washington, D.C. Intraspecific variation of drought adaptation in
brittlebush: Leaf pubescense and timing of leaf
Moldenke, A.R. 1976. California pollination loss vary with rainfall. Oecologia 113: 162-169.
ecology and vegetation types. Phytologia 34: 305-
361. Sandquist, D.R. and J.R. Ehleringer. 2003. Carbon
isotope discrimination differences within and
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between contrasting populations of Encelia substrate on post-fire growth of California coastal
farinosa raised under common-environmental sage scrub. In: N.S. Margaris and H.A. Mooney,
conditions. Oecologia 134: 463-470. eds. Components of Productivity of
Mediterranean-Climate Regions: Basic and
Visscher, P.K. and B.N. Danforth. 1993. Biology of Applied Aspects. W. Junk Publishers, Boston,
Calliopsis pugionis (Hymenoptera: Andrenidae): MA. p. 151-179.
nesting, foraging, and investment sex ratio.
Annals of the Entomological Society of America Wisdom, C.S. 1985. Use of chemical variation and
86: 822-832. predation as plant defenses by Encelia farinosa
against a specialist herbivore. Journal of
Went, F.W. 1942. The dependence of certain annual Chemical Ecology 11: 1,553-1,565.
plants on shrubs in southern California deserts.
Bulletin of the Torrey Botanical Club 69: 100- Wisdom, C.S. and E. Rodriguez. 1983. Seasonal
114. age-specific measurements of the sesquiterpene
lactones and chromenes of Encelia farinosa.
Westman, W.E., J.F. O'Leary, and G.P. Malanson. Biochemical Systematics and Ecology 11: 345-
1981. The effects of fire intensity, aspect and 352.
306
Ephedra nevadensis S. Wats. Nevada ephedra
EPHEDRACEAE
Synonyms: Ephedra aspera Engelm. ex S. Wats

Ephedra reedii Cory
Ephedra antisyphilitica sensu S. Watts in King
Stanley G. Kitchen
several oval-shaped male cones, 4 to 8 mm long,

are subtended by several pairs of membranous
bracts attached at the nodes (Cronquist and others
1972). Egg-shaped female cones, 5 to 11 mm long
and light-brown to yellowish-green, are attached
on short stalks (peduncles) and partially enclosed
by 8 to 10 bracts at young nodes. Fruits are smooth
brown nutlets, 4 to 9 mm long, 2 to 4 mm wide,
and occur individually or in pairs (Cronquist and
others 1972). The inner cone scales enclose the
seed mimicking the angiosperm pericarp. The
embryo has two cotyledons.
Taxonomy.—Ephedra is monotypic to
Ephedraceae with a worldwide distribution (except
for Australia) and is represented by approximately
60 species (Stevenson 1993). Two varieties, of
Nevada ephedra are recognized; E. nevadensis var.
nevadensis and var. aspera (Engelm. ex S. Wats)
L. Benson (Kartesz 1994). Chromosome number is
2n = 14 or 28 (Stevensen 1993).
Range.—Nevada ephedra is endemic to arid and

semiarid parts of the Great Basin, Colorado
Plateau, and desert Southwest of western North
America. Its range extends from Oregon and
California on the north and west, into Texas in the
east, and Mexico in the south (Tirmenstein 1990).
General Description.—Nevada ephedra, also In Utah it grows from 1,220 to 1,830 m in
known as Nevada jointfir, gray ephedra, or elevation (Dittberner and Olson 1983).
mormon tea, is an erect, 0.3 to 1.5 m tall,
coniferous shrub with opposite spreading Ecology.—Nevada ephedra grows in well drained,
branches. Roots are deep and fibrous extending sandy to rocky soils on arid hills, alluvial fans, and
plains, and on ridges and canyons partially to
from an expanded root crown. Stems are jointed
and grooved. Internodes are 1.5 to 6 cm long completely devoid of trees (Stanton 1974). It is
(Cronquist and others 1972). The combination of moderately salt-tolerant (Stanton 1974, Plummer
1977). It occurs in large pure stands and in mixed
small cones, ring porous wood, wide multiseriate
rays, and vessels in older stems separate Ephedra pinyon-juniper woodlands, salt-desert, sagebrush,
from other gymnosperms (Stevenson 1993). Bark and hot-desert transitional shrublands, and
mountain and desert grasslands (Tirmenstein
on older branches is gray and fissured. The erect,
photosynthetic branchlets (1 to 5 mm wide) are 1990). Mean annual precipitation varies from 150
to 400 mm. Plants sprout from the woody crown
pale green to gray, glaucous and smooth. Leaves
and from roots following fire (Young and Evan
are reduced to 2 to 5 mm long, deciduous scales,
positioned in opposite pairs at the nodes. A 1978, West and Hassan 1985). Fire seasonality,
intensity, and severity affect plant capacity to
thickened gray base remains after leaf-fall. One to
307
recover after burning (Stanton 1974, Wright 1980, up to 5 years (Stevens and others 1981). Although
McLaughlin and Bowers 1982). A lack of fuel seedlings tolerate drought well, they are slow
continuity in many Nevada ephedra communities growing and may take 5 to 10 years to reach 60 cm
results in fire being a rare event. Plants spread in height (Plummer and others 1968). Plants
vegetatively through underground stolons in establish well from direct seeding when more
response to burial. Natural recruitment from seeds competitive species are controlled. Successes with
is often from rodent caches (Tirmenstein 1990). transplants and cuttings to revegetate minelands
Of North American ephedra, Nevada ephedra is and other degraded shrublands have also been
considered to be the most important forage species reported (Luke and Monsen 1984, Shaw and
(Tirmenstein 1990). It is used extensively by mule Monsen 1984).
deer (Ococolius hemionus), pronghorn
(Antilocapra americana), and bighorn (Ovis Benefits.—Nevada ephedra provides forage and
canadensis) and domestic sheep during winter and cover for numerous species of wildlife, especially
in periods of drought (Stanton 1974, Smith and during winter and periods of drought (Stanton
Beale 1980). Nutritive value and palatability of 1974, Smith and Beale 1980, Dittberner and Olsen
young branchlets is considered to be fair to good 1983). It is also used as forage by domestic sheep
(Tirmenstein 1990). and cattle but is considered poor forage for horses.
The species is used effectively in restoration
Reproduction.—Nevada ephedra is dioecious planting on harsh sites (Plummer and others 1968,
(occasional monoecious) with female and male Luke and Monsen 1984, Shaw and Monsen 1984).
cones occurring on separate plants. Male plants are Indigenous humans used roasted seeds in foods
more common on dry slopes and female plants and various concoctions from seeds and stems to
dominate more favorable locations, maximizing treat a variety of symptoms including coughs,
reproductive output (Freeman and others 1976). headaches, cold, fever, and kidney ailments
Plants are wind-pollinated. The timing of cone (Steward 1938, Tirmenstein 1990). Mormon
maturation, or coning, and pollination is weather- pioneers also made teas from stems. All members
dependent and generally occurs over a period of 1 of the genus produce unique secondary compounds
to 2 weeks between early March and early June with potential medicinal applications (Caveney
(Stanton 1974, Turner and Randall 1987). Seeds and others 2001). Nevada ephedra, and other
ripen between May and August. Turner and North American members of the genus, lack the
Randall (1987) observed considerable inter-annual nueroactivating ephedrine alkaloids found in some
variation in timing of phonological development. Old World species. However, a class of
Cones are not produced every year, and good seed compounds with antimicrobial potential,
crops are typically several years apart for any kynurenates, are found in Nevada ephedra
given population (Plummer and others 1968). (Caveney and others 2001). It is occasionally used
Years of good cone production are apparently as an ornamental (Cronquist and others 1972).
synchronized among most plants of a population
by an interaction between years of above-average References
precipitation and time since last major seed crop.
There are approximately 45,000 seeds (nutlets) per Caveney, S., D.A. Charlet, H. Freitag, M. Maier-
kg (Kitchen and others 1999). Stolte, and A.N. Starratt. 2001. New
observations on the secondary chemistry of
Growth and Management.—Once ripe, seeds are world Ephedra (Ephedraceae). American
easily dislodged. Ripened seeds can be collected Journal of Botany 88: 1,199-1,208.
by gently striking or shaking seed-bearing
branches over tarps or hand-held hoppers. In good Cronquist, A., A.H. Holmgren, N.H. Holmgren,
years, large quantities of seeds can be harvested in and J.L. Reveal. 1972. Intermountain flora:
relatively short time if harvests are well timed. Vascular plants of the Intermountain West,
After drying, seeds are cleaned to remove light, U.S.A. Vol. 1. Hafner Publishing Company,
unfilled seeds, stem fragments, cone scales, and Inc., New York. 270 p.
other inert material. Purity and viability levels of
commercially available seed are generally high. Dittberner, P.L. and M.R. Olson. 1983. The plant
Seed germination is rapid across a wide range of information network (PIN) data base: Colorado,
temperatures (5 to 20 oC) with little or no Montana, North Dakota, Utah, and Wyoming.
dormancy (Young and others 1977, Kitchen and FWS/OSB-83/86. U.S. Department of the
others 1999). Seed viability has remained high for Interior, Fish and Wildlife Service, Washington
308
DC. 786 p. Smith, A.D. and D.M. Beale. 1980. Pronghorn
antelope in Utah: some research and
Freeman, D.C., L.G. Klikoff, and K.T. Harper. observations. Publication 80-13. Utah Division
1976. Differential resource utilization by the of Wildlife Resources, Salt Lake City, UT. 88 p.
sexes of dioecious plants. Science 193: 597-599.
Stanton, F. 1974. Wildlife guidelines for range fire
Kartesz, J.T. 1994. A synonymized checklist of the rehabilitation. Technical Note 6712. U.S.
vascular plants of the United States, Canada, and Department of the Interior, Bureau of Land
Greenland. Vol. II—thesaurus . 2nd ed. Portland, Management, Denver, CO. 90 p.
OR: Timber Press. 816 p.
Stevens, R., K.R. Jorgensen, and J.N. Davis. 1981.
Kitchen, S.G., S. Walker, and G.R. Wilson. 1999. Viability of seed from thirty-two shrub and forb
Addition of Ephedra nevadensis—Nevada species through fifteen years of warehouse
ephedra to the Rules. The Seed Technologist storage. Great Basin Naturalist 41: 274-277.
Newsletter 73: proposal #9.
Stevensen, D.W. 1993. Ephedraceae. In: Flora of
Luke, F. and S.B. Monsen. 1984. Methods and North America. http://flora.huh.harvard.edu/
costs for establishing shrubs on mined lands in [not paged].
southwestern Wyoming. In: A.R. Tiedemann,
E.D. McArthur, H.C. Stutz, R. Stevens, and K.L. Steward, J.H. 1938. Basin-Plateau Aboriginal
Johnson, comps. Proceedings—Symposium on Sociopolitical Groups. Bulletin 120.
the biology of Atriplex and related chenopods; Smithsonian Institution Bureau of American
1983 May 2-6; Provo, UT. General Technical Ethnology, Washington, DC. [1997 reprinted by
Repeport INT-172. U.S. Department of the University of Utah Press, Salt Lake City,
Agriculture, Forest Service, Intermountain UT]. 346 p.
Forest and Range Experiment Station, Ogden,
UT. p. 286-291. Tirmenstein, D. 1990. Ephedra nevadensis. In:
McLaughlin, S.P. and J.E. Bowers. 1982. Effects Fire Sciences Laboratory. Fire Effects
of wildfire on a Sonoran Desert plant Information System, http://www.fs.fed.
community. Ecology 63: 246-248. us/database/feis/ [not paged].
Plummer, A.P. 1977. Revegetation of disturbed Turner, F.B. and D.C. Randall. 1987. The
Intermountain area sites. In: J.C. Thames, ed. phenology of dessert shrubs in southern Nevada.
Reclamation and use of disturbed lands of the Journal of Arid Environments 13: 119-128.
Southwest. University of Arizona Press, Tucson,
AZ. p. 302-337. West, N.E. and M.A. Hassan. 1985. Recovery of
sagebrush-grass vegetation following wildfire.
Plummer, A.P., D.R. Christensen, and S.B. Journal of Range Management 38: 131-134.
Monsen. 1968. Restoring big-game range in
Utah. Publication 68-3. Utah Division of Fish Wright, H.A. 1980. The role and use of fire in the
and Game, Ephraim, UT. 183 p. semidesert grass-shrub type. General Technical
Report INT-85. U.S. Department of Agriculture,
Shaw, N. and S.B. Monsen. 1984. Nursery Forest Service, Intermountain Forest and Range
propagation and outplanting of bareroot Experiment Station, Ogden, UT: 24 p.
chenopod seedlings. In: A.R. Tiedemann, E.D.
McArthur, H.C. Stutz, R. Stevens, and K.L. Young, J.A. and R.A. Evans. 1978. Population
Johnson, comps. Proceedings—Symposium on dynamics after wildfires in sagebrush
the biology of Atriplex and related chenopods; grasslands. Journal of Range Management 31:
1983 May 2-6; Provo, UT. General Technical 283-289.
Report INT-172. U.S. Department of
Agriculture, Forest Service, Intermountain Young, J.A., R.A. Evans, and B.L. Kay. 1977.
Forest and Range Experiment Station, Ogden, Ephedra seed germination. Agronomy Journal
UT. p. 251-260. 69: 209-211.
309
Ephedra viridis Coville green ephedra
EPHEDRACEAE
Synonyms: Ephedra nevadensis subvar. pluribracteata Palmer ex Stapf.

Ephedra nevadensis var. viridis (Colville) M.E. Jones
Stanley G. Kitchen
mm), bright green to yellow green and broom-like,

with opposite or falsely whorled attachment
(Welsh and others 1987). Leaves are reduced to
1.5 to 4 mm, deciduous scales, positioned in
opposite pairs at nodes. A thickened brown base
remains after leaf-shed. Egg-shaped cones are
borne at nodes, singly or whirled. Two or more
male cones, 5 to 7 mm long, are subtended by
opposite membranous bracts, 2 to 4 mm long,
while female cones, 6 to 10 mm long, are partially
enclosed by thickened bracts, 4 to 7 mm long
(Cronquist and others 1972). Brown, three-sided
seeds are smooth and 5 to 8 mm long and 2 to 4
mm wide. The combination of small cones, ring
porous wood, wide multiseriate rays, and vessels
in older stems separate Ephedra from other
gymnosperms (Stevenson 1993).
Taxonomy.—Two varieties of green ephedra are

recognized. Variety viscida (Cutler) L. Benson
differs from the more widespread var. viridis by
the presence of short stalks (peduncles) on female
cones and the often sticky nature of stems (Welsh
and others 1987). Chromosome number is 2n = 28
(Stevensen 1993).
Range.—Green ephedra is closely associated with

the Great Basin and Colorado Plateau provinces of
western North America (Anderson 2001). Its range
extends from California and Oregon east and south
to western Texas and Chihuahua, Mexico. In Utah
and California its elevational range varies from
900 to 3,000 m (Welsh and others 1987, Anderson
2001).
General Description.—Green ephedra, also
known as Mormon tea, or jointfir, is a spreading to Ecology.—Green ephedra is found on
erect, coniferous shrub, 0.5 to 1.5 m tall and with a sedimentary, volcanic, and metamorphic substrates
crown 1 to 3 m wide at maturity (Cronquist and in sandy, gravelly, or rocky, well-drained,
others 1972, USDA Forest Service 1976). A underdeveloped soils (Anderson 2001).
fibrous, somewhat fragile root system originating Populations are found on a variety of landforms
from several, almost parallel, taproots descends including: plains, alluvial fans, mesa tops, washes,
deep into the soil profile. Stems are jointed and and canyon walls on all aspects. Salt tolerance is
grooved. Internodes are 1 to 4.5 cm long limited. Plants tolerate partial shade but do best in
(Cronquist and others 1972). Lower bark is gray full sun. Although rarely considered a dominant,
and irregularly fissured (Anderson 2001). green ephedra is associated with a wide variety of
Photosynthetic upper branches are thin (1 to 3 plant community types including: big sagebrush-
310
bunchgrass, black sagebrush, blackbrush, desert shattering after seeds ripen. After drying, seeds are
grassland, pinyon-juniper, mountain mahogany, cleaned to high purity and viability levels using
oak woodland, mountain shrub, mixed conifer, and conventional milling equipment. Freshly-harvested
ponderosa pine (Anderson 2001). It is present in seed may be partially dormant (Meyer and others
early, mid, and late successional stages (Young 1988). Dormancy can be broken with a 4-week
and Evans 1973, Koniak 1985). Mean annual moist prechill or stratification (Meyer and others
precipitation varies from 200 to 430 mm 1988). Germination for after-ripened or prechilled
(Anderson 2001). Plants sprout from crown or seed is relatively rapid. Germination of non-
roots following top-kill by fire (USDA Forest dormant seeds occurs over a range of cool
Service 1976, Young and Evans 1978, Koniak temperatures (Young and others 1977). Seed
1985, Everett 1987). Reported survival following viability remains high for at least 5, and perhaps as
fire is variable (Anderson 2001). Sprouting long as 15 years (Stevens and others 1981).
response following fire may vary by season of Seedling growth rate is slow. Seedlings may
burn, fire severity, or among population variation require 5 to 10 years to reach 60 cm in height
in fire tolerance. Green ephedra is found in (Anderson 2001). In restoration plantings,
communities for which a wide range of fire seedlings are susceptible to competition from
frequencies is known. It also establishes on weeds and faster-growing grasses and forbs in the
disturbed sites from seed (Koniak 1985). Green seed mix (Ferguson and Frischknecht 1981).
ephedra is used as winter forage by big game and Plants establish well from direct seeding and
livestock but receives only limited use during transplants (USDA Forest Service 1976, Plummer
other seasons (Jameson and others 1962, Kufeld 1977, Everett 1980, Luke and Monsen 1984). Stem
and others 1973). Palatability and nutritive quality cutting are capable of root development in
of young stems are generally ranked as fair to greenhouse conditions (Wieland and others 1971).
good for most wildlife species and classes of Green ephedra was toxic to domestic cattle and
domestic livestock (Dittberner and Olson 1983, sheep during pregnancy in a controlled diet study
Anderson 2001). Seeds are eaten by birds and (Keeler 1989). No detrimental effects were
rodents and appear to be moderately prefered observed for developing calves and lambs.
(Everett and others 1978). Rodents probably play Conflicting reports have been published on the
an important role in seed dispersal. possible presence of ephedrine alkaloids in green
ephedra. While Anderson (2001) suggests the
Reproduction.—Green ephedra is dioecious. presence of this group of compounds in this
Spatial segregation of the sexes results from high species, Caveney and others (2001) failed to find
male plant representation on dry ridges and slopes these compounds common to Old World ephedras
and high female plant representation on microsites in any New World species. Tannin content is high.
with relatively favorable soil moisture conditions
(Freeman and others 1976, Brunt and others 1988) Benefits.—Green ephedra is valuable for restoring
providing for greater reproductive efficiency. vegetation structure and diversity and for erosion
Plants are wind-pollinated. Timing of cone control on disturbed lands, including roadsides,
maturation, or coning, and seed maturation is minelands, and degraded rangelands (USDA
dependent upon variation in weather, elevation, Forest Service 1976, Everett 1980, Ferguson and
aspect, and geography. Coning and pollination Frischknecht 1981, Ferguson and Frischknecht
occur for approximately 2 weeks, usually between 1985, Herbel 1986), in areas in which it is adapted.
April and June. Seeds mature between June and When competition is controlled, it establishes well
September. Cones are not produced each year, and from seed and persists after wildfire. It provides
good seed crops may be several years apart important winter forage for wildlife and domestic
(Plummer and others 1968). There are livestock (USDA Forest Service 1976). Green
approximately 45,000 seeds per kg (Meyer and ephedra has had limited use as an ornamental
others 1988). (Anderson 2001). Native Americans made both
medicinal and nonmedicinal teas brewed from
Growth and Management.—In years of good stems and used seeds and roasted seeds (nutlets)
seed crops, significant quantities are easily for food (Steward 1938, USDA Forest Service
harvested by hand (Plummer and others 1968). 1976). Mormon settlers were also known to use it
Ripened cones are easily dislodged by gently in the production of a beverage, a cause for the
shaking or striking cone-bearing branches over frequently used moniker, ‘mormon tea’.
tarps or hand-held hoppers. Timing of harvest is Undetermined medicinal applications for a group
critical as thunder storms can cause considerable of unique secondary compounds associated with
311
members of the genus have not been fully establishment on reconstructed soils in semiarid
determined, including possible antimicrobial areas. In: Shrub establishment on disturbed arid
applications for kynurenates (Caveney and others and semi-arid lands: Proceedings of the
2001). symposium; 1980 December 2-3; Laramie, WY.
Wyoming Game and Fish Department. p. 57-63.
References
Ferguson, R.B. and N. Frischknecht. 1985.
Anderson, M.D. 2001. Ephedra viridis. In: U.S. Reclamation on Utah’s Emery and Alton coal
Department of Agriculture, Forest Service, Fire fields: techniques and plant materials. Research
Sciences Laboratory, Fire Effects Information Paper INT-335. U.S. Department of Agriculture,
System, http://www.fs.fed.us/database/feis/ Forest Service, Intermountain Research Station,
26 p. Ogden, UT. p. 78.
Brunt, J.W., M.R. Conley, and G.L. Cunningham. Freeman, D.C., L.G. Klikoff, and K.T. Harper.
1988. Sex in Ephedra trifurca (Ephedraceae) 1976. Differential resource utilization by the
with relation to Chihuahuan Desert habitats. The sexes of dioecious plants. Science 193: 597-599.
American Midland Naturalist. 119: 137-142.
Herbel, C.H. 1986. Seeding shrubs in the field.
Caveney, S., D.A. Charlet, H. Freitag, M. Maier- Reclamation and Revegetation Research 5: 377-
Stolte, and A.N. Starratt. 2001. New 385.
observations on the secondary chemistry of
world Ephedra (Ephedraceae). American Jameson, D.A., J.A. Williams, and E.W. Wilton.
Journal of Botany 88: 1199-1208. 1962. Vegetation and soils of Fishtail Mesa,
Arizona. Ecology 43: 403-410.
Cronquist, A., A.H. Holmgren, N.H. Holmgren,
and J.L. Reveal. 1972. Intermountain flora: Keeler, R.F. 1989. Investigation of maternal and
Vascular plants of the Intermountain West, embryo/fetal toxicity of Ephedra viridis and
U.S.A. Vol. 1. Hafner Publishing Company, Ephedra nevadensis in sheep and cattle. Journal
Inc., New York. 270 p. of Range Management 42: 31-35.
Dittberner, P.L. and M.R. Olson. 1983. The plant Koniak, S. 1985. Succession in pinyon-juniper
information network (PIN) data base: Colorado, woodlands following wildfire in the Great
Montana, North Dakota, Utah, and Wyoming. Basin. The Great Basin Naturalist 45: 556-566.
FWS/OSB-83/86. U.S. Department of the
Interior, Fish and Wildlife Service, Washington Kufeld, R.C., O.C. Wallmo, and C. Feddema.
DC. 786 p. 1973. Foods of the Rocky Mountain mule deer.
Research Paper RM-111. U.S. Department of
Everett, R.L. 1980. Use of containerized shrubs for Agriculture, Forest Service, Rocky Mountain
revegetating arid roadcuts. Reclamation Review Forest and Range Experiment Station, Fort
3: 33-40. Collins, CO. 31 p.
Everett, R.L. 1987. Plant response to fire in the Luke, F. and S.B. Monsen. 1984. Methods and
pinyon-juniper zone. In: Everett, R.L., comp. costs for establishing shrubs on mined lands in
Proceedings—pinyon-juniper conference; 1986 southwestern Wyoming. In: A.R. Tiedemann,
January 13-16; Reno, NV. General Technical E.D. McArthur, H.C. Stutz, R. Stevens, and K.L.
Report INT-215. U.S. Department of Johnson, comps. Proceedings—Symposium on
Agriculture, Forest Service, Intermountain the biology of Atriplex and related chenopods;
Research Station, Ogden, UT. p. 152-157. 1983 May 2-6; Provo, UT. Gen. Tech. Rep.
INT-172. U.S. Department of Agriculture,
Everett, R.L., R.O. Meeuwig, and R Stevens. Forest Service, Intermountain Forest and Range
1978. Deer mouse preference for seed of Experiment Station, Ogden, UT. p. 286-291.
commonly planted species, indigenous weed
seed, and sacrifice foods. Journal of Range Meyer, S.E., S.G. Kitchen, G.R. Wilson, and R.
Management. 31: 70-73. Stevens. 1988. Addition of Ephedra viridis
green mormon tea to the Rules. AOSA
Ferguson, R.B. and N. Frischknecht. 1981. Shrub Newsletter 62: 18-19.
312
Plummer, A.P. 1977. Revegetation of disturbed USDA Forest Service. 1976. Some important
Intermountain area sites. In: J.C. Thames, (ed). native shrubs of the west. U.S. Department of
Reclamation and use of disturbed lands of the Agriculture, Forest Service, Intermountain
Southwest. University of Arizona Press, Tucson, Forest and Range Experiment Station. Ogden,
AZ. p. 302-337. UT. 16 p.
Plummer, A.P., D.R. Christensen, and S.B. Welsh, S.L., N.D. Atwood, L.C. Higgins, and S.
Monsen. 1968. Restoring big-game range in Goodrich. 1987. A Utah Flora. Brigham Young
Utah. Publ. No. 68-3. Utah Division of Fish and University Print Services, Provo, UT. 894 p.
Game, Ephraim, UT. p. 183.
Wieland, P.A.T., E.F. Frolich, and A. Wallace.
Stevens, R., K.R. Jorgensen, and J.N. Davis. 1981. 1971. Vegetative propagation of woody shrub
Viability of seed from thirty-two shrub and forb species from the northern Mojave and southern
species through fifteen years of warehouse Great Basin deserts. Madrono 21: 149-152.
storage. Great Basin Naturalist 41: 274-277.
Young, J.A. and R.A. Evans. 1973. Downy brome-
Stevensen, D.W. 1993. Ephedraceae. In: Flora of intruder in the plant succession of big sagebrush
North America. http://flora.huh.harvard.edu/ communities in the Great Basin. Journal of
[not paged]. Range Management 26: 410-415.
Steward, J.H. 1938. Basin-Plateau Aboriginal Young, J.A. and R.A. Evans. 1978. Population
Sociopolitical Groups. Bulletin 120. dynamics after wildfires in sagebrush
Smithsonian Institution Bureau of American grasslands. Journal of Range Management 31:
Ethnology, U.S. Government Printing Office, 283-289.
Washington, DC. [1997 reprinted by the
University of Utah Press, Salt Lake City, UT]. Young, J.A., R.A. Evans, and B.L. Kay. 1977.
346 p. Ephedra seed germination. Agronomy Journal
69: 209-211.
313
Eriogonum fasciculatum Benth. California buckwheat
POLYGONACEAE
Synonyms: Eriogonum rosmarinifolium Nutt.

Eriogonum fasciculatum var. maritimum Parish
Eriogonum fasciculatum var. oleifolium Gand.
Eriogonum aspalathoides Gand.
Eriogonum fasciculatum ssp. aspalathoides S. Stokes
Arlee M. Montalvo
born in heads of tightly to loosely packed

involucres with few flowers per involucre. The
heads (or dense cymes) are born singly or in
umbels subtended by long, naked peduncles (often
> 5 cm). Branching of the umbels varies from one
to four levels. Umbels are rounded to flat-topped
depending on the extent of branching. Flowers
produce small (about 1.5 to 2 mm long), single
seeded achenes that are partially enclosed in a dry,
persistent calyx. Below is a summary of diagnostic
differences among varieties following Munz and
Keck (1968), Reveal (1989), and Hickman (1993).
Range and Taxonomy.—E. f. var. fasciculatum
(see insert) has 2n = 40 chromosomes (Munz and
Keck 1968). Shrubs are often low and spreading
and 0.6 to 1.2 m tall. Leaves are linear to linear-
oblanceolate, green and glabrate above, sparsely
pubescent beneath, with leaf margins strongly
rolled under (revolute). Involucres and flowers are
usually glabrous. Plants occur from nearly sea
level to just over 300 m in coastal scrub and sage
General Description.—California buckwheat is a scrub along coastal bluffs and cliffs, and in
widespread and morphologically variable shrub canyons and dry slopes of the foothills near the
and is one of the most important components of coast from San Luis Obispo Co. south to central
sagebrush scrub, desert scrub, and especially Baja California.
coastal sage scrub (Kirkpatrick and Hutchinson E. f. var. foliolosum (Nutt.) S. Stokes ex
1977, Westman 1981). Although the four Jones (Munz and Keck 1968), the interior flat-
taxonomic varieties partly overlap in geographic topped buckwheat, is tetraploid with 2n = 80
distribution each is associated with a different chromosomes (Stebbins 1942). Shrubs are erect to
climatic zone and combination of traits, especially spreading, 0.5-1.5 m tall and occur on interior
color, chromosome number, leaf shape, degree of slopes and mesas below 1000 m (seldom on coast)
revolute leaf margins, and pubescence (Munz and in chaparral and sage scrub from Monterey Co.,
Keck 1968, Hickman 1993). All forms have California south to northern Baja California.
alternate cauline, linear to oblanceolate leaves that Leaves are often linear, dark to gray-green and
are white wooley with appressed hairs beneath, pubescent above, and densely pubescent below.
and darker, generally less hairy to glabrous above. Involucres and perianth are usually pubescent.
Leaves are 1 to 2 cm long and cluster tightly at Synonyms are E. rosmarinifolium var. foliolosum
nodes on short axillary shoots, giving the leaves a Nutt., E. f. ssp. foliolosum S. Stokes, and E. f. var.
fascicled appearance. Flowers are small (about 3 obtusiflorum S. Stokes.
mm long) with six white to pinkish sepals and are
314
Eriogonum f. var. polifolium (Benth.) Torrey well on rocky road cuts and in shallow soils that
& A. Gray (flowering shoot in photograph) has 2n are inhospitable to annual grasses.
= 40 chromosomes (Stebbins 1942). These
greyish, pubescent, rounded shrubs occur at 600 to Reproduction.—Plants flower from early spring
2300 m on dry slopes and washes. They are through summer. The four taxa have slightly
widespread across inland mountains and deserts of shifted flowering times, with var. flavoviridae
California, from the inner coast ranges of San Luis having the earliest season (Munz and Keck 1968)
Obispo and Fresno Cos., south to San Diego Co., and var. polifolium flowering before the other two.
eastward into the Mojave Desert to southern In all taxa, stamens and stigmas are exerted and
Nevada, Utah, and Arizona, and south into open to generalist pollinators, including honey
northwestern Senora, Mexico and central Baja bees and many species of small native bees, flies,
California. In inland sage scrub, plants overlap in wasps, and beetles. Achenes mature from June to
distribution with var. foliolosum, especially in September and disperse when dried inflorescences
Riverside Co., but occupy drier sites. Leaves are shatter in autumn. The dry calyx provides
oblanceolate to obovate, densely pubescent buoyancy to detached achenes and assists dispersal
beneath, and usually tomentose above with leaf by wind and water. Harvester ants also disperse
margins sometimes barely revolute. Involucres and achenes (DeSimone and Zedler 1999). Seedlings
sepals are pubescent. Heads tend to be born in emerge in the rainy season in mid to late winter
simple to once-compound umbels. Reveal (1989) within light gaps and open areas. Survival is
reports hybridization between var. polifolium and unusually high as shown by studies in a variety of
E. cinereum Benth in areas were E. cinereum has sites and years, often ranging from 80 to 90
been introduced within the range of var. percent (Wright and Howe 1987, Miriti and others
polifolium. Synonyms are E. f. ssp. p. S. Stokes 1998, Montalvo and others 2002). Most shoot
and E. p. Benth. elongation and leaf growth occurs before
Eriogonum f. var. flavoviride Munz & flowering from late winter to mid spring (Cole
Johnson has n = 20 chromosomes (Reveal 1989) 1967). Plants can reach maturity in 1 to 2 years.
and is 2 to 3 dm tall. The rounded shrubs occur on
dry rocky slopes and washes from 200 to 1300 m Seed Germination.—Light improves germination
in creosote bush scrub of the southern Mojave (Cole 1967, Keeley 1987). Atwater (1980)
Desert, San Bernardino Co., south through eastern suggests that light increases permeability of seed
Riverside Co. to far eastern San Diego Co., and coats, water adsorption, and possibly leaching of
south to central Baja California. Leaves are light inhibitors, allowing seeds to germinate. Light-
green to yellowish-green, linear, strongly revolute, induced seed coat permeability would allow some
and glabrous above. Peduncles are glabrous; buried seeds to remain viable and dormant in the
involucres and sepals are subglabrous. Synonym is seed bank, a bet-hedging strategy. Cole (1967)
E. f. ssp. flavoviride S. Stokes. found coastal populations germinated at 15 to 25
o
C while inland populations germinated over a
Ecology.—California buckwheat occurs on dry larger range of 5 to 35 oC. Keeley (1987)
slopes and often granitic substrate (Westman examined the effects of light, dark, heat, and
1981). The woody, branched roots penetrate to leachate through charred wood (charrate) on
under 1.5 m (Kummerow and others 1977). Plants germination. Seeds germinated equally well on
lose some leaves in the dry season and can shed soil vs. filter paper, and light controls had a two-
many leaves in severe drought. Plants are fold increase in germination compared to dark
vulnerable to hot fires so resprout success is low controls (83 vs. 45 percent germination). Charrate
and most regeneration is from seeds (Keeley did not affect germination. In light, seeds exposed
1998). Frequent fires deplete the seed bank, to 120 oC for 5 min suffered a large decrease in
making populations vulnerable to local extinction. germination under all conditions relative to seeds
In Riverside Co., this shrub has decreased heated to 70 or 100 oC, suggesting fire may kill
dramatically since the 1940’s, with the largest many seeds in the upper centimeter of the
decreases on alluvial soils (Minnich and Dezzani seedbank. Heat sensitivity and a light requirement
1998). Decreases are attributed to a combination of for germination encourage colonization of
competition with exotic annual grasses, too disturbed or open sites.
frequent a fire interval, that is exacerbated by
weeds which carry fire, and by atmospheric Horticulture.—Plants can be propagated from
deposition of nitrogen, which encourages growth seeds or rooted from cuttings. Seeds and container
of competing weeds. California buckwheat does plants should be planted in the fall to early winter
315
to take advantage of the moist cool season for Vertebrates also use California
growth. In one study, seeds planted by seed buckwheat. The rare orange-throated whiptail is
imprinting and hydroseeding, shallow methods associated with open vegetation with this shrub
that allow light to reach planted seeds, had at least (Brattstrom 2000). Rare California gnatcatchers
twice the emergence success as drilling that (Polioptila californica californica) forage and nest
covered seeds with 0.5 to 1 cm of soil (Montalvo in the shrub. California buckwheat and Artemisia
and others 2002). Seeds germinate within 2 weeks californica Less. are dominant shrubs used by
if soil is moist and over about 15 oC. Plants require gnatcatchers (Mock and Bolger 1992). The rare
full sun and good drainage. No irrigation is mountain sheep (Ovis canadensis nelsoni) browses
required if planted before seasonal rains. on California buckwheat in the San Gabriel
The grey-green var. polifolium provides a Mountains of southern California (Perry and others
nice contrast to the other green varieties in the 1987), and deer eat the inflorescences
garden, but all do well in dry landscaping. The (Schopmeyer 1974).
flowers last far into the summer and become an
attractive rusty-brown in fruit. Some prostrate Genetics, Geographic Variation, Hybridization,
cultivars of California exist, including “Theodore and Fitness.—Stebbins (1942) hypothesized that
Payne,” “Prostrata,” “Dana Point,” and tetraploid populations of var. foliolosum arose
“Wildwood” (Perry 1992). from hybridization between diploid var.
fasiculatum and var. polifolium (= E. f. subsp.
Greenhouse studies indicate that California foliolosum, E. f. subsp. typicum, and E. f. subsp.
buckwheat forms facultative associations with polifolium, respectively, in the paper). He
arbuscular mycorrhizal fungi (Egerton-Warburton, examined the distribution of taxa in relation to
Montalvo, and Allen, unpublished report). The climatic factors and concluded that the
plant’s facultative dependence on mycorrhizal morphology and ecological tolerances of var.
fungi may be instrumental to successful foliolosum combined the morphology and
colonization of barren sites. Montalvo and others tolerances of the two diploid taxa. The variety
(2002) found that plants grew quickly to maturity polifolium was in areas with some snow, frost, and
on a graded site with low organic matter and only 10 to 13 cm of rainfall, fasciculatum was in
nutrients, but establishment, growth, and flowering the areas with no frost, often frequent fog, and 22
all decreased significantly with decreasing soil to 37 cm of rainfall, and foliolosum often occurred
nitrogen (NO3-N mean = 7.89 mg/g, range = 0.8- in habitats intermediate to the two diploids. Cole
69 mg/g in the study plot). (1967) tested assumptions about the distribution of
morphology and ecological tolerances of vars.
Benefits.—California buckwheat provides habitat fasciculatum and foliolosum. Cole compared their
and food for numerous animals. Osborne (1998) physiologies over an ecological gradient from the
found high abundance and diversity of arthropod cool coast across the mountains to the hot inland
species on California buckwheat. Some are valleys and found that coastal var. fasciculatum
species-specific feeders such as the larvae of the were the least hairy, and populations of var.
moth, Hemileuca electra (Wright) (Rubinoff foliolosum became more hairy inland. There was a
1998). Numerous species of butterfly larvae feed clear morphological cline from the coast to the
on California buckwheat and specialize on interior among and within ecotypes that correlated
different plant parts and taxa (Howe 1975), with habitat. Photosynthesis vs respiration ratios
sometimes according to timing of growth. For and photosynthetic rates and respiration rates all
example, the larvae of the butterfly, Apodemia varied in a clinal manner that correlated with
mormo (Felder & Felder), has three biotypes in the morphological characters. The variety
Mojave Desert. The spring morph feeds on fasciculatum forms a distinctly adaptive coastal
California buckwheat while the others feed on ecotype, while var. foliolosum forms an inland
species with different flowering times. E. ecotype.
parvifolium Sm. in Rees. is the host of the rare El
Segundo blue butterfly (Euphilotes bernardino Analysis of eight allozyme loci for eight
allyni Shields). When California buckwheat was populations of var. foliolosum and four
planted instead of later flowering E. parvifolium, fasciculatum revealed ample genetic variation
competing insect species used California (Montalvo, Ellstrand, and Clegg, unpublished
buckwheat, built up populations, and then data). All loci were polymorphic and there was an
competed with the rare butterfly for E. parvifolium average of 5.7 alleles/locus. Expected
flowers (Longcore and others 2000). heterozygosity was also high at 0.42. Nei’s genetic
distances among populations ranged from 0 to
316
0.074, with the largest distance between varieties. DeSimone, S.A., and P.H. Zedler. 1999. Shrub
Overall, the proportion of total variation explained seedling recruitment in unburned Californian
by differences among populations (GST) was coastal sage scrub and adjacent grassland.
0.025. By itself, the tetraploid var. foliolosum had Ecology 80: 2018-2032.
no significant structure (GST = 0.005), while GST =
0.028 for var. fasciculatum. These values suggest Hickman, J.C., ed. 1993. The Jepson Manual:
high levels of gene flow and outbreeding. Higher Plants of California. University of
California Press, Ltd., Los Angeles, CA. 1,400 p.
Seed Collection and Processing.—Seeds are
collected after heads have turned rusty brown. The Howe, W.H., ed. 1975. The butterflies of North
chaff can be separated from seeds by pushing America. Doubleday and Company, Inc., Garden
seeds through a screen and then separating seeds City, NY. 633 p.
from chaff with an air separator or fan (author’s
observation). Commercial seed companies usually Keeley, J.E. 1987. Role of fire in seed germination
skip the screening and just break up the chaff, air of woody taxa in California chaparral. Ecology
separate the seeds from crude chaff, and sell seeds 68: 434-443.
with the persistent calyx intact. A recommended
target for minimum quality of purchased seeds is Keeley, J.E. 1998. Postfire ecosystem recovery and
15 percent purity and 65 percent germination. management: The October 1993 large fire
Here, a bulk seed pound with about 990,000 episode in California. In: J. M. Moreno (ed.).
seeds/kg, would have about 96,800 pure live seeds Large Forest Fires. Backbuys Publishers, Leiden,
(personal communication, S & S Seeds, Netherlands. p. 69-90.
Carpenteria, CA).
Kirkpatrick, J.B., and C.F. Hutchinson. 1977. The
Growth and Management.—Seeds have been community composition of California coastal
widely planted for restoration, roadside erosion sage scrub. Vegetatio 35: 21-33.
control, slope stabilization, landscaping, and
apiary (Schopmeyer 1974, Perry 1992). Despite Kummerow, J., D. Krause, and W. Jow. 1977. Root
the clear geographic pattern and ecological systems of chaparral shrubs. Oecologia 29: 163-
affinities of the four taxonomic varieties, most 177.
researchers and consultants fail to designate
taxonomic variety on publications, plant lists, or Longcore, T., R. Mattoni, G. Pratt, and C. Rich.
plant palettes. They also commonly fail to specify 2000. On the perils of ecological restoration:
appropriate ecological zone for seeds used in lessons from the El Segundo blue butterfly. In: J.
wildland plantings. This has resulted in extensive E. Keeley, M. Baer-Keeley and C. J.
plantings outside natural ranges and habitats Fotheringham, eds. 2nd Interface Between
(Reveal 1989). Projects should use seeds of Ecology and Land Development in California:
appropriate taxa to maximize project success. U.S. Geological Survey Open-File Report 00-62.
Vol. 62. Sacramento, CA. p. 281-286.
References
Minnich, R.A., and R.J. Dezzani. 1998. Historical
Atwater, B.R. 1980. Germination, dormancy and decline of coastal sage scrub in the Riverside-
morphology of the seeds of herbaceous Perris plane, California. Western Birds 29: 366-
ornamental plants. Seed Science and Technology 391.
8: 523-573.
Miriti, M.N., H.F. Howe, and S.J. Wright. 1998.
Brattstrom, B.H. 2000. The range, habitat Spatial patterns of mortality in a Colorado desert
requirements, and abundance of the orange- plant community. Plant Ecology 136: 41-51.
throated whiptail, Cnemidophorus hyperythrus
beldingi. Bulletin of the Southern California Mock, P.J., and D. Bolger. 1992. Ecology of the
Academy of Sciences 99: 1-24. California gnatcatcher at Rancho San Diego.
Ogden Environmental and Energy Services Co.,
Cole, N.H. A. 1967. Comparative physiological Inc.; Project No. 110970000. 50 p.
ecology of the genus Eriogonum in the Santa
Monica Mountains, southern California. Montalvo, A.M., P.A. McMillan, and E.B. Allen.
Ecological Monographs 37: 1-24. 2002. The relative importance of seeding method,
317
soil ripping, and soil variables on seeding success. Rubinoff, D. 1998. Field observations on mating
Restoration Ecology 10: 52-67. behavior and predation of Hemileuca electra
(Saturniidae). Journal of the Lepidopterists'
Munz, P.A., and D.D. Keck. 1968. A California Society 52: 212-214.
Flora with Supplement. University of California
Press, Berkeley, CA. 1681 + 224 p. Schopmeyer, C.S., ed. 1974. Seeds of Woody Plants
in the United States. Vol. 450 Agricultural
Osborne, K.H. 1998. A description of arthropod Handbook. USDA Forest Service, Washington,
community structure in southern California D.C. 883 p.
coastal sage scrub. M.S. Thesis. University of
California, Riverside, CA. 133 p. Stebbins, G.L. 1942. Polyploid complexes in
relation to ecology and the history of floras. The
Perry, B. 1992. Landscape plants for western American Naturalist 76: 36-45.
regions: an illustrated guide to plants for water
conservation. Land Design Publishing, Westman, W.E. 1981. Factors influencing the
Claremont, CA. 318 p. distribution of species of California coastal sage
scrub. Ecology 62: 439-455.
Perry, W.M., J.W. Dole, and S.A. Holl. 1987.
Analysis of the diets of mountain sheep from the Wright, S.J., and H.F. Howe. 1987. Pattern and
San Gabriel Mountains, California. California mortality on Colorado Desert plants. Oecologia
Fish and Game 73: 156-162. 73:543-552.
Reveal, J.L. 1989. The eriogonoid flora of

California (Polygonaceae: Eriogonoideae).
Phytologia 66: 295-414.
318
Erithalis fruticosa L. black torch
RUBIACEAE
Synonyms: Erithalis revoluta Urban

Erithalis angustifolia DC.
Erithalis elliptica Raf.
Erithalis inodora Jacq.
Erithalis odorata Pers.
Erithalis parviflora Griseb.
John K. Francis
black fruits are globose or flattened drupes 3 to 4

mm in diameter that have a bitter-sweet flavor and
contain five to 10 nutlets (Howard 1989, Liogier
1997, Little and others 1974, Nelson 1996). It is
morphologically a highly variable species (Long
and Lakela 1976).
Range.—Black torch is native to Florida, the

Bahamas, the Greater and Lesser Antilles,
Trinidad, Margarita and other Venezuelan islands,
Quintana Roo (Mexico), and Belize (Howard 1989,
Little and others 1974).
Ecology.—Black torch grows from near sea level

to 120 m in elevation in Puerto Rico (Little and
others 1974) in areas that receive from 750 mm to
about 1800 mm of annual precipitation. The
species is drought-hardy and moderately intolerant
of shade and can grow under the canopy of low-
density forest. Because of slow growth, it is
restricted to areas with reduced competition. Black
General Description.—Black torch, also known as torch is most common near beaches and on rock
candlewood, tea, parrot apple, cubra prieta, outcroppings and bluffs near the shoreline,
jayajabico, manglillo, tarro de chivo, bois especially in moist limestone areas, and on
flambeau, and bois chandelle, is an evergreen shrub limestone hills in the interior. In Florida, black
that typically is 1 or 2 m in height but occasionally torch is found in beach strand vegetation, on sand
reaches 8 m in height. Usually the species has dunes, and coastal hammocks (Nelson 1996).
multiple stems arising near the ground line and
many branches that are stiff and sometimes Reproduction.—Black torch flowers and fruits
gnarled. The bark is gray and smoothish to warty. throughout the year (Little and others 1974, Nelson
The resinous wood is brown, hard, fine-textured, 1996). Open-grown individuals produce fruits and
heavy, and has visible growth rings. Black torch seeds in abundance. Fruits collected in Puerto Rico
plants are supported by brown roots that are often averaged 0.1087 + 0.0071 g/fruit. Air-dried seeds
contorted as they grow over rocks and into cracks. separated from them averaged 0.00092 g/seed or
The foliage may be dense or diffuse depending on 1,080,000 seeds/kg. Presumably, the seeds are
whether the shrubs grow in full sun or partial dispersed by birds that eat the fruits. Seedlings and
shade. The opposite leaves have 3 to 20 mm saplings are relatively common in Puerto Rico. The
petioles, orbicular to oblanciolate blades 2 to 12 cm stems layer (root) when they come in contact with
long with entire edges, and dark-green upper the ground.
surfaces. The inflorescences are lateral or terminal
panicles containing many small, white flowers. The Growth and Management.—Black torch is slow
319
growing, especially when situated on rock Liogier, H.A. 1990. Plantas medicinales de
outcrops. Management should mainly consist of Puerto Rico y del Caribe. Iberoamericana de
protection from fires and clearing of sites for Edicones, Inc., San Juan, PR. 566 p.
development.
Benefits.—Black torch contributes to biodiversity Rico and adjacent islands. Vol. 5. Editorial de la
in the forests where it grows, helps protect the soil, Universidad de Puerto Rico, San Juan, PR.
and furnishes food and cover for wildlife. It would 436 p.
probably make an acceptable ornamental for green
areas and nonirrigated gardens. The wood is good Little, E.L., Jr., R.O. Woodbury, and F.H.
for turned articles but available only in small sizes. Wadsworth. 1974. Trees of Puerto Rico and the
It is resistant to rot and has been used for piles and Virgin Islands. Vol. 2. Agriculture Handbook
posts. Formerly, it was split and used for torches 449. U.S. Department of Agriculture,
(Little and others 1974) but tended to be smoky Washington, DC. 1,024 p.
(University of the Virgin Islands 2002). The bark,
fruits, and the resin have diuretic and astringent Long, R.W. and O. Lakela. 1976. A flora of
properties and are used to treat inflammation of the Tropical Florida. Banyan Books, Miami, FL.
kidney and bladder, and blennorrhoea (Liogier 962 p.
1990). The leaves are parched and ground and used
to treat skin sores (University of the Virgin Islands Nelson, G. 1996. The shrubs and woody vines of
2002). Florida. Pineapple Press, Inc., Sarasota, FL.
390 p.
References
University of the Virgin Islands. 2002. Erithalis
Howard, R. A. 1989. Flora of the Lesser Antilles, fruticoas (blacktorch). http://rps.uvi.edu/
Leeward and Windward Islands. Vol. 6. Arnold VIMAS/blacktorch.htm. 1 p.
MA. 658 p.
320
Ernodea littoralis Sw. golden creeper
RUBIACEAE
Synonyms: Ernodea angusta Small
John K. Francis
Golden creeper is tolerant of salt spray and mild

salt concentration in the soil. The species is
drought-tolerant and usually grows in excessively
drained sites. However, it withstands short-term
flooding (Carney-Kulig and others 2002). In
Florida, it is common in coastal dunes (Nelson
1996). The variety angusta grows in pinelands in
Florida (Workman 1980). In Puerto Rico, golden
creeper also grows on coastal rocks (usually
limestone) and on rocky ultramafic (serpentine)
slopes. It is a sun-loving plant that usually grows
in the open. When the shrubs grow under low trees
in open forest, they do not become thick and
robust. Golden creeper benefits from mild
General Description.—Golden creeper, also disturbance that eliminates shade and taller
known as cough bush, beach creeper, wild competition.
pomegranate, ernodia de playa, wild pamgramma,
and liane-séche, is an evergreen prostrate or Reproduction.—Golden creeper blooms and
arching shrub to about 1.5 m in height. Heavy fruits all year (Gilman 1999, Nelson 1996). Fruits
lateral roots support the plants. The stems are four- collected in Puerto Rico weighed an average of
angled and orange or reddish brown. Golden 0.0467 + 0.0014 g/fruit. Air-dried seeds separated
creeper leaves have a short petiole (1 to 2 mm), from them weighed an average of 0.0078 + 0.0002
lanceolate blades, and a spiny, pointed tip. In some g/seed or 128,000 seeds/kg. Sown without
environments, leaves are somewhat fleshy; in pretreatment on commercial potting mix, 24
others they are hard or coriaceous. White-to-pink percent germinated between 35 and 146 days of
tubular flowers are solitary, axillary, and sessile. sowing (author’s observation). The fruits are
The fleshy fruits are golden orange, in the shape of presumably eaten and the seeds dispersed by birds.
tiny pomegranates, and have an apple-like flavor. Once established, plants thicken and spread by
The seeds, borne one per fruit, are brown with a root suckering and layering.
longitudinal furrow (Howard 1989, Liogier 1997,
Nelson 1996). Growth and Management.—The growth rate of
golden creeper is reported to be moderate (Gilman
Range.—Golden creeper is native to Southern 1999). Individual stems appear to live about 2 to 5
Florida, the Bahamas, the Greater Antilles, the years, but the plants, being clonal, can go on
Lesser Antilles as far south as Marie Galante, indefinitely. Golden creeper is produced and sold
Mexico, Belize, and Honduras (Howard 1989, in commercial nurseries. Plants in commercial
Liogier 1997). It is planted as an ornamental but is production are produced by ground layering and
not known to have naturalized into any new range rooted cuttings. Field planting of hormone-treated
as a result. cuttings is also recommended (Centro Ecologico
Sian Ka’an 2002). Wildlings (seedlings and plants
Ecology.—Golden creeper will grow in acidic to dug up from natural stands) transplant well
mildly alkaline soils with the whole range of (Workman 1980). Recommended spacing for
textures (Gilman 1999) and over a variety of planting is 0.9 to 1.5 m (Gilman 1999). Once
parent materials including serpentine. In Puerto planted, golden creeper needs little attention
Rico, it grows in areas receiving from 750 to about (Workman 1980); in fact, it often dies if over-
1400 mm of mean annual precipitation and at irrigated (Gilman 1999).
elevations from near sea level to about 400 m.
321
Benefits.—Golden creeper contributes to the Gilman, E.F. 1999. Ernodea littoralis. Fact Sheet
biodiversity of the sites where it grows, stabilizes FPS-196. Cooperative Extension Service,
the soils in those fragile environments, and University of Florida. http://
provides food and cover for wildlife. It is planted hort.ifas.ufl.edu/shrubs/ERNLITA.pdf. 3 p.
as an ornamental for landscaping, usually as
ground cover. Golden creeper is also used for Howard, R.A. 1989. Flora of the Lesser Antilles,
beach dune stabilization. It is particularly Leeward and Windward Islands. Dicotyledoneae.
promoted for water-restricted landscaping Part 3. Vol. 6. Arnold Arboretum, Harvard
(Workman 1980). A tea is made from the leafy University, Jamaica Plain, MA. 658 p.
shoots as a treatment for coughs (Centro Ecologico
Sian Ka’an 2002). Liogier, H.A. 1997. Descriptive flora of Puerto
References Universidad de Puerto Rico, San Juan, PR.
436 p.
Carney-Kulig, S., M. Jolly, T. Weyant, S. Buehler,
and G. Davis. 2002. Planting the right plant in Nelson, G. 1996. The shrubs and woody vines of
the right place. Monroe County Extension Florida. Pineapple Press, Inc., Sarasota, FL.
Service, University of Florida. http:// 391 p.
monroe.ifas.ufl.edu/gardeningguide_5.htm. 6 p.
Workman, R.W. 1980. Growing native. The
Centro Ecologico Sian Ka’an. 2002. Flora. Sanibel-Captiva Conservation Foundation, Inc.,
http://cesiak.org/flora1.htm. 3 p. Sanibel, FL. 137 p.
322
Erythroxylum areolatum L. false cocaine
ERYTHROXYLACEAE
Synonyms: none (genus also spelled Erythroxylon)
John K. Francis
Range.—False cocaine is native to the Bahamas,

the Greater Antilles, the Cayman Islands, southern
Mexico, and Central America (National Trust for
the Cayman Islands 2002, Stevens and others
Ecology.—False cocaine grows in areas of Puerto

Rico that receive from about 750 to 900 mm of
mean annual precipitation at elevations of a few
meters above sea level to about 300 m. It grows in
gallery forests in Nicaragua from 40 to 380 m
elevation, frequently associated with limestone
rocks (Stevens and others 2001). False cocaine is
common in limited areas but uncommon in most of
its range, growing in remnant and middle to late
secondary forests. False cocaine grows on deep,
medium-textured soil and sandy beach-strand soils
(Vásquez and Kolterman 1998). The species is
most frequent on limestone parent material, as
skeletal rock or porous solid rocks but grows in
areas with igneous and metamorphic (including
ultramaphic) rocks. It has an intermediate tolerance
to shade and will grow in openings or in the
understory of medium to low basal area forests.
General Description.—False cocaine, also known
as redwood, swamp redwood, thin-leafed Reproduction.—False cocaine has been observed
erythroxylon, indio, palo de hierro, arabo flowering from October to June in Puerto Rico
carbonero, limoncillo, huesito, cocaina falsa, and (Little and Wadsworth 1964). Fresh fruits collected
poirier, is a deciduous shrub or small tree 2 to 7 m in Puerto Rico weighed an average of 0.119 +
in height and 5 to 15 cm in stem diameter. The 0.013 g/fruit. Seeds cleaned from them weighed
plant may have a single stem or multiple stems (air-dried) an average of 0.0493 g/seed or 20,300
branching from near the ground and is supported seeds/kg. Sown on moist filter paper without
by a tap and lateral root system. The roots are dark pretreatment, they germinated at 35 percent,
brown, stiff, and woody. The bark is gray and scaly beginning 12 days after sowing (Francis and
and the inner bark is red. Moderately robust Rodríguez 1993). Apparently, birds are the
branches and many twigs form a “branchy” crown, principal dispersers of seeds. Seed production can
often with a dense display of foliage. Alternate be abundant in some years. Seedlings and saplings
leaves are tightly grouped on short side branches or vary from rare to common.
spread out on new twigs. The leaves are light green
to dark green, elliptic to oblong, 3 to 13 cm long, Growth and Management.—False cocaine grows
with entire margins and petioles 5 to 7 mm long. slowly, about 1 to 3 mm in diameter per year and is
Tiny, fragrant, five-merous white flowers are borne capable of living several decades. Published
in clusters in the leaf axils. At maturity, the thin- planting or management experience is not known
fleshed, 6- to 9-mm drupes are bright red. Each to the author. Protecting forests and stands
fruit contains one oblong, bony seed (Britton and containing the species is probably the best
Millspaugh 1962, Liogier 1988, Little and management approach until research can be
Wadsworth 1964, Stevens and others 2001).
323
performed. Rico and adjacent islands, Spermatophyta. Vol.
Benefits.—False cocaine protects the soil and Río Piedras, PR. 481 p.
furnishes food and cover for wildlife. The wood is
heavy, hard, fine-grained, durable, and strong. The Little, E.L., Jr. and F.L. Wadsworth. 1964.
sapwood is light brown and the heartwood is Common trees of Puerto Rico and the Virgin
reddish or chocolate brown (Little and Wadsworth Islands. Agriculture Handbook 249. U.S.
1964). Available only in small sizes, it is useful for Department of Agriculture, Washington, DC.
stakes, fuel, carving, and turnery. The foliage of 548 p.
this species does not contain useful amounts of
cocaine, which is extracted from a sister species (E. National Trust for the Cayman Islands. 2002.
coca Lam.) from the Andean Highlands (Little and Virtual herbarium of the National Trust for the
Wadsworth 1964). Cayman Islands: Erythroxylaceae, Erythroxylum
areolatum L. http://www.virtualherbarium.org/
References vh/CAYM/403.htm. [not paged].
Britton, N.L. and C.F. Millspaugh. 1962. The Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
Bahama flora. The New York Botanical Garden, Montiel, eds. 2001. Flora de Nicaragua.
New York. 695 p. Monographs in Systematic Botany, Vol. 85, No.
Francis, J.K. and A. Rodríguez. 1993. Seeds of 943 p.
Puerto Rican trees and shrubs: Second
installment. Research Note SO-374. U.S. Vásquez, O.J. and D.A. Kolterman. 1998. Floristic
Department of Agriculture, Forest Service, composition and vegetation types of the Punta
Southern Forest Experiment Station, New Guaniquilla Natural Reserve—Cabo Rojo,
Orleans, LA. 5 p. Puerto Rico. Caribbean Journal of Science
43(304): 265-279.
324
Erythroxylum brevipes DC. brisselet
ERYTHROXYLACEAE
Synonyms: Erythroxylum brevipedatum St. Lager

Erythroxylum rhamnoides Peyr. ex Schulz
John K. Francis
Indies (Howard 1988). It has been much confused

in the literature with E. rotundifolium Luman that
also occurs in Puerto Rico (Liogier and Martorell
2000), southern Mexico through Central America
as far south as Costa Rica, and in the Bahamas,
Cuba and Jamaica (Stevens and others 2001).
Ecology.—Brisselet grows in most types of soil,

except poorly drained or salty soils. Plants are
more frequent in eroded areas and rocky terrain
where competition is weaker. Annual rainfall in the
Puerto Rican range varies from about 750 to about
2000 mm. Elevation where it grows ranges from
near sea level to about 760 m (Little and others
1974). Brisselet is moderately intolerant of shade.
It invades abandoned pastures, fields, and disturbed
areas, usually as scattered plants. It is common in
early and mid-secondary forest and is an important
plant of the understory of remnant and late
secondary forests that are not too dense. Grazing
cattle benefits the species, probably by reducing its
competition. Cut stems will resprout. No
General Description.—Brisselet is a shrub, observations are published on sprouting after single
usually 1 or 3 m in height, that sometimes fires, but repeated fires will eliminate it from an
becomes a small tree to 8 m in height. It is also area.
known by the common names, rat-wood, jibá,
ratón, yaría de costa and rocío in Spanish and bois Reproduction.—Brisselet flowers during the
vinette and brésillette in French (Liogier 1988, summer months and fruits in late summer or early
Little and others 1974). Older plants often have fall (Little and others 1974). Fruit production can
multiple stems from below or just above the be heavy in open-grown plants but is rare in shaded
ground level. They are supported by a tap and understory plants. A collection of fruits from
lateral root system consisting of dark-brown, Puerto Rico weighed an average of 0.0936 +
tough, and brittle roots. The shrub’s crowns are 0.0035 g/fruit. Air-dried seeds separated from them
cluttered with slender branches and twigs. The weighed an average of 0.0401 + 0.0007 g/seed or
bark of brisselet is gray or light brown. Foliage on 24,900 seeds/kg. Sown in commercial potting mix
older branches is borne on short spurs. The 1- to 3- without any pretreatment, these seeds began
cm obovate leaves have prominent light-colored germinating in 19 days and completed 99 percent
central nerves and are sometimes notched at the germination in 47 days. Germination is epigenous.
tips. The small, white, five-petaled flowers are A thin, deep taproot is quickly produced. The
clustered one to four at the leaf axils. Brisselet seedlings are woody, stiff, and easy to lift and
fruits are red, ellipsoidal, fleshy drupes 5 to 9 mm transplant. Birds disperse the seeds. Seedlings are
long (Howard 1988, Little 1988, Little and others relatively common and widespread.
1974).
Growth and Management.—Both height and
Range.—Brisselet is native to Hispaniola, Puerto diameter growth of brisselet is slow. Weaver
Rico, the Virgin Islands, and St. Barts in the West (1990) measured the mean annual diameter growth
325
of 33 brisselet stems 4 to 14 cm in diameter in St. Liogier, H.A. 1988. Descriptive flora of Puerto
John, U.S. Virgin Islands at 0.5 mm/year. Rico and adjacent islands, Spermatophyta. Vol.
Apparently, plants live for several decades. No 2. Editorial de la Universidad de Puerto Rico,
management experience has been published. Río Piedras, PR. 481 p.
Benefits.—Brisselet is an important component of Liogier, H.A. and L.F. Martorell. 2000. Flora of
dry and moist forests, protects the soil, and Puerto Rico and adjacent islands, a systematic
furnishes cover for wildlife. Although seasonal, the synopsis. 2nd Ed. Editorial de la Universidad de
fruits are a consistent source of food for birds. Bees Puerto Rico, Río Piedras, PR. 382 p.
are attracted to the fragrant flowers (Little and
others 1974). The sapwood is light tan while the Little, E.L., Jr., R.O. Woodbury, and F.H.
heartwood is dark brown, hard, and flexible. Wadsworth. 1974. Trees of Puerto Rico and the
Because of its small diameters and lengths, the Virgin Islands. Vol. 2. Agriculture Handbook
wood is useful for little other than fuel. Brisselet 449. U.S. Department of Agriculture,
tissues do not contain cocaine, as obtained from its Washington, DC. 1,024 p.
more famous relative Erythroxylum coca Lam.
Stevens, W.D., C. Ulloa-U., A. Pool, and O.H.
References Montiel, eds. 2001. Flora de Nicaragua.
Howard, R.A. 1988. Flora of the Lesser Antilles, 1. Missouri Botanical Garden Press. p. 1-943.
Dicotyledoneae, Part 1. Vol. 4. Arnold Weaver, P.L. 1990. Tree diameter growth rates in
Arboretum, Harvard University, Jamaica Plain, Cinnamon Bay Watershed, St. John, U.S. Virgin
MA. 673 p. Islands. Caribbean Journal of Science 26(1-2):
1-6.
326
Eugenia foetida Pers. boxleaf stopper
MYRTACEAE
Synonyms: Eugenia buxifolia (Sw.) Willd.

Eugenia myrtoides Poir.
Myrtus axillaries Poiret in Lam.
Myrtus poiretii Sprengl
Eugenia triplinervia O. Berg.
Eugenia lateriflora sensu Griseb.
John K. Francis
obovate or oblanciolate 2 to 5 cm long and have a

1.5 to 3 mm petiole and usually have a rounded
tip. The foliage, especially when crushed, emits a
mild, offensive, skunk-like odor. Small, four-
petaled, white flowers are grouped in few-
flowered, axillary racemes. Fruits are black
globose berries 4 to 7 mm in diameter with one or
two small seeds (Howard 1989, Liogier 1994,
Range.—Boxleaf stopper is native to Florida,

Puerto Rico, the U.S. Virgin Islands, Jamaica,
Hispaniola, Cuba, Anguilla, St. Martin, St. Barts,
Netherlands Antillies, Yucatan in Mexico, Belize,
and Guatemala (Howard 1989, Liogier 1994, Little
and others 1974). The species has not been
reported to have naturalized outside its native
range.
Ecology.—Boxleaf stopper most frequently grows

in areas underlain by limestone rock, usually on
ridges and hill sides. It grows on clay and courser-
textured soils, rocky rubble, and often directly into
cracks in fractured rock. The species also occurs
General Description.—Boxleaf stopper is also on sandy flats behind beaches. In Florida, it grows
known as Spanish stopper, gurgeon stopper, on hammocks near the coast and on pinelands in
stopper bush, white watting, anguila, hoja menuda, the Lower Keys (Nelson 1996). Elevations vary
escobón, pico de paloma, guairaje, bois petites from near sea level to about 100 m. Mean annual
feuilles. It is an upright, evergreen shrub or small rainfall of the native range in Puerto Rico ranges
tree sometimes reaching 10 m in height and 12.5 from about 750 and 1700 mm (author’s
cm in diameter at breast height. There is usually a observation, Little and others 1974). The species is
single stem from the ground, but it may branch drought tolerant (Gilman 1999). Although boxleaf
near the ground. The stems and branches are stopper eventually reaches the stature of a small
slender with gray, smooth to lightly fissured or tree on favorable sites, much of its habitat is so
platy bark. The inner bark is pink and slightly difficult that the plant completes its life cycle in
bitter. The sapwood is light brown, and the sizes (2 to 5 m in height) usually associated with
heartwood is dark reddish brown. The wood is shrubs. The species is moderately intolerant of
hard, heavy, and fine-grained. This deep-rooted shade, growing in the understory, as an
species has a tap and lateral root system of slender, intermediate or codominant in low-basal area,
flexible to moderately stiff brown roots. The short-statured, dry forests.
paired, leathery, green to dark green leaves are
327
Reproduction.—Flowering of boxleaf stopper in ruminant herbivores. Although small, the fruits are
Puerto Rico is reported to occur in spring and edible to humans, and birds consume them when
summer (Little and others 1974). It flowers in the available. Boxleaf stopper is recommended and
summer in Florida (Gilman 1999). Fresh fruits used for screens, hedges, parking lot, yard, and
collected in Puerto Rico averaged 0.181 + 0.032. street trees in Florida (Gilman 1999).
Air-dried seeds separated from them averaged
14,900 seeds/kg. Placed in moist potting mix, 51 References
percent germinated beginning 69 days after
sowing. The new seedlings grow slowly. In the Gilman, E.F. 1999. Eugenia foetida. Fact Sheet
wild, seeds are dispersed by birds. Seedlings and FPS-200. Cooperative Extension Service,
saplings are widely scattered to relatively University of Florida, Gainesville, FL. 3 p.
common.
Howard, R.A. 1989. Flora of the Lesser Antilles,
Growth and Management.—The growth rate of Leeward and Windward Islands. Vol. 5. Arnold
boxleaf stopper is moderate to slow, depending on Arboretum, Harvard University, Jamaica Plain,
habitat. Apparently, it is relatively long lived (a MA. 604 p.
few decades). Potted stock is available from
nurseries for planting. Although expensive for Liogier, H.A. 1994. Descriptive flora of Puerto
wildland plantings, this may be the best way to Rico and adjacent Islands. Vol. 3. Editorial de la
establish them in order to get them above Universidad de Puerto Rico. Río Piedras, PR.
competing weeds and grass. Ornamental plants 461 p.
may be pruned into hedges or forced to single-
stemmed trees (Gilman 1999). Because they are Little, E.L., Jr., R.O. Woodbury, and F.H.
relatively slow growing and not aggressive Wadsworth. 1974. Trees of Puerto Rico and the
invaders, control of the species in natural stands Virgin Islands. Vol. 2. Agriculture Handbook
should not be necessary. 449. U.S. Department of Agriculture,
Benefits.—Boxleaf stopper protects the soil,
furnishes cover for wildlife, and contributes to the Nelson, G. 1996. The shrubs and woody vines of
aesthetics of the forest. Aromatic compounds Florida. Pineapple Press, Inc., Sarasota, FL.
released from the leaves of this species apparently 391 p.
are responsible for the slight musky smell present
in forests in Puerto Rico where it is common. Schaus, C., S. Wade and J. Dunan. 2003. Key deer
Foliage of the species is not eaten by Key deer and plants they won’t eat. http://monroe.
(Odocolleus virginanus clavium) (Schaus and ifas.ufl.edu/key_deer_plants.htm. 4 p.
others 2003) and probably is avoided by other
328
Eugenia pseudopsidium Jacq. wild guava
MYRTACEAE
Synonyms: Eugenia megalocarpa Urban

Eugenia portoricensis DC.
Eugenia thomasiana Berg
Myrtus willdenovii Spreng.
Stenocalyx portoricensis Berg
John K. Francis
axils, one to three per axil. The globose fruits

retain the calyx of the flower. They are orange,
turning to bright red, and astringent with little
flavor. Each fruit contains one rounded seed.
Range.—The native range of wild guava includes

Hispaniola, Puerto Rico, the Virgin Islands,
Guadalupe, Dominica, Martinique, and St. Lucia.
The same plant or a very similar one also occurs in
Venezuela, the Guyanas, and Amapá, Brazil
(Howard 1989). That it is wild guava is supported
by its listing in the Smithsonian (2001) checklist
of plants for Guyana.
Ecology.—Wild guava grows inland a few meters

above sea level to elevations of 670 m in Puerto
Rico (Little and others 1974). It occurs in areas
that receive from about 1400 to 2200 mm of
annual precipitation. Medium- to heavy-textured
soils derived from sedimentary, igneous, and
metamorphic (including ultramafic) rocks are
colonized. It does not appear sensitive to
topographic position but rarely if ever grows on
General Description.—Wild guava, which is also
excessively or very poorly drained soils. Wild
known as bastard guava, Christmas cherry,
guava is shade tolerant and normally is confined to
guayaba silvestre, quiebrahacha, goyavier de
the understories of remnant old growth and
montague, bois plié, and goyavier bâtard, is a
advanced secondary forests.
shrub or a small tree reaching a maximum of 20 m
in height and 30 cm in trunk diameter (Howard
Reproduction.—Wild guava flowers and fruits
1989, Liogier 1994, Little and others 1974).
irregularly throughout the year (Little and others
Mature plants are usually 3 to 5 m in height and 4
1974). Understory plants produce fruits in small
to 8 cm in stem diameter. Wild guava has a tough,
numbers; plants receiving increased sunlight in
woody root system with tap and lateral roots. The
gaps or thinned canopies produce several times
stems are light gray, slightly fissured, and scaly.
more fruits. Wild guava fruits collected in Puerto
The wood is light to reddish brown, hard and very
Rico weighed an average of 1.284 + 0.043 g.
heavy (specific gravity = 1.3). Twigs are green,
Seeds extracted from these fruits weighed an
turning to brown (Little and others 1974). The
average of 0.565 + 0.010 g (air dry) or 1,800
crowns branch to the third order and branches tend
seeds/kg. Eighty-one percent of the seeds from
to be slender. The opposite leaves have short
this collection germinated between 59 and 143
petioles, are generally elliptic, rounded or short
days after sowing in commercial potting mix.
pointed at the base and long pointed at the apex.
When the new seedlings are damaged above
They are dark green above and light green
ground by fungi or insects, they sprout from the
beneath. Small white flowers arise in the leaf
329
ground level, usually with more than one shoot References
(author’s observation). Birds disperse the seeds.
Seedlings and saplings are well-scattered and Howard, R.A. 1989. Flora of the Lesser Antilles,
relatively common. Wild guava sprouts readily Leeward and Windward Islands. Vol. 5. Arnold
after cutting or damage. Arboretum, Harvard University, Jamaica Plain,
MA. 604 p.
Growth and Management.—The growth rate of
wild guava appears to be relatively good for an Liogier, H.A. 1994. Descriptive flora of Puerto
understory shrub. A 2.5-m tall sapling, 2.5 cm Rico and adjacent Islands. Vol. 3. Editorial de la
d.b.h., under partial shade in Puerto Rico, had five Universidad de Puerto Rico. Río Piedras, PR.
growth rings in its trunk near the ground. Wild 461 p.
guava germinates, survives, and transplants well in
the nursery, but growth is somewhat slower than Little, E.L., Jr., R.O. Woodbury, and F.H.
most commercial nursery plants. Unfortunately, Wadsworth. 1974. Trees of Puerto Rico and the
no plantation or ornamental use has been reported. Virgin Islands. Vol. 2. U.S. Department of
Agriculture, Agriculture Handbook 449.
Benefits.—Wild guava is used to a limited extent Washington, DC. 1,024 p.
for firewood and fence posts. With its dark green
leaves, bright red fruits, and clean form, the Smithsonian. 2001. Regional plant species lists for
species has a pleasing appearance and probably Guyana. Mason.gmu.edu/~ckelloff/region2.pdf.
would make a fine ornamental, especially in shady 6 p.
locations. It provides food and cover for wildlife
and contributes to the scenic beauty and
biodiversity of the forests where it grows.
330
Euonymus alatus (Thunb.) Sieb. winged burning bush
CELASTRACEAE
Synonyms: Celastrus alatus Thunb.

Celastrus striatus Thunb.
Euonymus arakianus Koidz.
Euonymus sacrosanctus Koidz.
Euonymus striatus (Thunb.) Loes.
Euonymus subtriflorus Blume
Euonymus thunbergianus Blume
than 2 mm., the leaves then appearing subsessile).

The leaf blades are glabrous, lanceolate to obovate,
4.5 to 7.3 (rarely to 10) by 1.2 to 2.3 (rarely to 4.5)
cm, leathery to papery, with cuneate or attenuate
bases, crenate to serrate margins and acuminate to
cuspidate apices. The five to seven pairs of
ascending lateral veins disappear before reaching
the leaf margin. The flowers develop on the
current season’s growth and are are bisexual
(rarely functionally unisexual). They are in small
axillary clusters (cymes), the clusters on slender
peduncles, 2.0-2.5 cm long, usually with one
dichotomous branch and three or fewer flowers.
Each flower is on a pedicel 4 to 7 mm long. The
flowers are about 9 mm in diameter with the parts
in fours, and the sepals and petals readily
distinguishable from one another. The petals are
yellowish green. The fruits are ovoid, four-lobed
capsules, 8 to 14 by 10 to 16 mm. At maturity the
capsules are red to brown-purple and they split
open along the lobes (often one or more lobe is
Illustration soruce: Ma, J.S. 2001 abortive), exposing the one to four seeds, each
enclosed in a fleshy orange aril. The seeds (sans
General Description.—This species is commonly aril) are glabrous, light brown to black, ellipsoid, 6
known as winged burning bush or winged spindle to 12 by 4.0 to 5.5 mm. Some spell the specific
tree because of its stems, which have two to four epithet “alata” instead of “alatus” used here. This
conspicuous brown, corky ridges or wings, 2 to 3.5 is because botanists disagree as to whether the
mm wide and 1.0 to 1.2 mm thick. It is the only genus name Euonymus should be treated as
species of Euonymus L. in North America to have feminine (“alata”) or masculine (“alatus”) (Paclt
such prominent wings on its twigs. It is also 1998). In classical Greek, the generic name was
occasionally called Asian spindle tree because it is treated as feminine; however, Linnaeus treated the
native to Asia. Winged burning bush is a low- genus as masculine. The spelling “Euonymus” has
growing shrub, usually up to 2 m tall, never more formally been conserved over Linnaeus’s
than 4 m. The plant does spread well horizontally, originally spelling “Evonymus,” which has also
sending out stems in many directions. Its buds are occasionally been used (Gilstra 1991).
axillary (two per axil), ovoid or obovate, scattered
along the stem. The leaves are opposite, two per Range.—Winged burning bush is native to east
node, and are spaced somewhat evenly along the Asia being distributed in far eastern Russia (Far
stem. The leaf petioles are short, round in cross East), Japan, Korea, and China. In its native range,
section, glabrous, 2 to 3.5 mm long (rarely shorter it occurs in areas with annual precipitation of 400
331
mm or more and in elevations ranging from 40 to References
2,700 m (Ma 2001). It was first introduced into the
United States about 1860 (Rehder 1940), as an Brizicky, G.K. 1964. The genera of Celastrales in
ornamental plant, and it eventually became the southeastern United States. Journal of the
naturalized in the Eastern United States (Ebinger Arnold Arboretum. 45: 210-215.
1996, Gleason and Cronquist 1991, J. Ma in
personal observation). Currently winged burning Chittenden, F.J. and P.M. Synge. 1956. Dictionary
bush is known from 21 Eastern and Mid-Western of gardening. Vol. 2, 2nd Ed. Clarendon Press,
States (Connecticut, Illinois, Indiana, Iowa, Oxford, UK. 583 p.
Kentucky, Maryland, Massachusetts, Michigan,
Minnesota, Missouri, North Carolina, New Ebinger, J.E. 1996. Euonymus alatus. In: J.M.
Hampshire, New Jersey, New York, Ohio, Randall, J.M. and J. Marinelli, eds. Invasive
Pennsylvania, Rhode Island, South Carolina, plants–weeds of the global garden. 21st Century
Virginia, Vermont, West Virginia), as well as Gardening Series. Brooklyn Botanic Garden,
Montana (USDA 2003). It can be expected to be Brooklyn, NY. p 55.
found elsewhere with the continued spread of
naturalized populations and additional material Gilstra, Z. 1991. Proposal to conserve the spelling
escaping from cultivation. of 4618 Euonymus (Celastraceae). Taxon 40:
137-139.
Ecology.—It is not as aggressive an invader as the
related Oriental bittersweet (Celastrus orbiculatus Gleason, H.A. and A. Cronquist 1991. Manual of
Thunb). It is found in disturbed habitats, such as Vascular Plants of northeastern United States
roadsides, forest margins and woodlands that are and adjacent Canada. New York Botanical
in early stages of succession (Ma, personal Garden, Bronx, NY. 910 p.
observation).
Ma, J.S. 2001. A revision of Euonymus
Reproduction.—Hymenopterous and dipterous (Celastraceae). Thaiszia Journal of Botany 11:
insects, especially bees and flies, appear to be the 1-264.
main pollinators, and the attractive arillate seeds
are widely dispersed by birds and small mammals Martin, T. 2002. Weed alert! Euonymus alata
(Brizicky 1964). The flowers bloom in the spring (Thunb.) Siebold. http://tncweeds.ucdavis.
and the fruit mature in the fall. The species can be edu/alert/alrteuon.html. [not paged].
asexually propagated through stem cuttings and
root separation. Paclt, J. 1998. Proposal to amend the gender of
Euonymus, nom. cons. (Celastraceae) to
Growth and Management.—Winged burning feminine. Taxon 47: 473-474.
bush grows in a variety of soil conditions. It is not
a fast growing plant, reaching perhaps up to 1 m of Rehder, A. 1940. Manual of cultivated trees and
growth in a single growing season. Propagation shrubs. Macmillan Publishing Co., New York.
can be effected by layering or seed (Chittenden 996 p.
and Synge 1956). However, there is concern about
the continued use of this plant in ornamental Smith, B. 2003. The bonsai guide
plantings, since it can be invasive in some areas http://www.thebonsaiguide.com/BPG/BS_BPBT
(Martin 2002) E.HTM. [not paged].
Benefit.—The foliage of winged burning bush USDA, NRCS. 2003. The PLANTS Database.
turns bright scarlet in autumn, giving the plant its Baton Rouge, LA. http:plants.usda.gov. [not
common name burning bush. The purplish paged].
capsules and seeds with orange arils add to the
“burning” effect of the plant late in the year. This
effect of the plant has made it a popular
ornamental and horticultural species. Some
specimens can be shaped into bonsai (Smith 2003).
332
Euonymus americanus L. strawberry bush
CELASTRACEAE
Synonyms: Euonymus alternifolius Moench.

Euonymus americanus var. augustifolius (Pursh) Wood
Euonymus americanus var. sarmentosus Nutt.
Euonymus augustifolius Pursh
Euonymus heterophylus Raf.
Euonymus muricatus Raf.
Euonymus sarmentosus G. Don
Euonymus sempervirens Marsh.

subsessile. The leaf blades are slightly leathery to
papery, oblong to elliptic, 4.0 to 10 by 2.0 to 3.5
cm wide, with cuneate or attenuate bases, crenulate
margins, and acuminate apices. The five to seven
pairs of ascending lateral leaf veins disappear
before reaching the margin. The flowers occur on
the current season’s growth in small axillary
clusters (cymes), each cluster usually consisting of
two peduncles, each with one to three flowers
(often one or two of the flowers will aborte,
leaving only one functional flower per peduncle).
The peduncles are slender, 1.5 to 2.2 cm long.
When there is more than one flower per peduncle
each flower is subtended by a short pedicel, 1 to
3.5 mm long. The flowers are bisexual, about 10
mm in diameter, with the sepals and petals readily
distinguishable from one another. The calyx
comprises five yellow to green sepals, 1.0 to 1.5
by 1.6 to 1.8 mm. The corolla comprises of five
greenish to reddish petals up to 4 mm by 4 mm
wide. There are five stamens, 1.5 to 2.0 mm long.
The five carpels are fused. A sticky nectarous disk
is present, 3 to 4 mm in diameter. The color and
shape of the fruit gives it its common names. The
fruit is a plump, red globose dry, five-lobed
capsule up to 1.5 cm in diameter prior to opening.
Illustration source: Britton and Brown 1913 The surface of the capsule is densely covered with
minute spines or warts on the surface. This species
General Description.—Strawberry bush is also is the only species of Euonymus L. in North
known as bursting heart, hearts–a-bursting, or America to have a capsule with a spiny or warty
American burning bush. It is a low-growing, surface. The capsule opens along the lobes,
deciduous shrub, up to 2 m high. Its stems are stiff, exposing the seeds that are covered by a fleshy,
green, glabrous, straight, round, 2 to 5 mm in scarlet aril. The seeds (sans aril) are brown,
diameter, and are occasionally four-angled. The glabrous, ellipsoid, 7 to 9 by 4.0 to 5.5 mm. Some
pith is white, round, and continuous. The leaves of spell the specific epithet “americana” instead of
the strawberry bush are opposite (two per node), “americanus” used here. This is because botanists
spaced somewhat evenly along stem. The petioles disagree as to whether the genus name Euonymus
are short, less than 3 mm and sometimes are nearly should be treated as feminine (“americana”) or
absent, the leaves then appearing sessile or masculine (“americanus”) (Paclt 1998). In
333
classical Greek, the generic name was treated as hundreds of the “bursting” red capsules. In the
feminine; however, Linnaeus treated the genus as winter, the bright green twigs are also handsome.
masculine. The spelling “Euonymus” has formally
been conserved over Linnaeus’s originally spelling References
“Evonymus,” which has also occasionally been
used (Gilstra 1991) Britton, N.L. and A. Brown. 1913. Illustrated flora
of the northern states, Canada, and the British
Range.—Strawberry bush occurs throughout possessions. Vol. 2, 2nd Ed. Scribner, New York.
much of the Eastern United States (Alabama, 735 p.
Arkansas, Delaware, District of Columbia, Florida,
Georgia, Illinois, Indiana, Kentucky, Louisiana, Brizicky, G.K. 1964. The genera of Celastralesin
Maryland, Mississippi, Missouri, New Jersey, the Southeastern United States. Journal of the
New York, North Carolina, Ohio, Oklahoma, Arnold Arboretum. 45: 206-235.
Pennsylvania, South Carolina, Tennessee, Texas,
Virginia and West Virginia), except New England. Chittenden, F.J. and P.M. Synge. 1956. Dictionary
(Shetler and Orli 2000, USDA 2003) It is also of gardening. Vol. 2, 2nd ed. Clarendon Press,
known from Ontario (Gleason and Cronquist 1991, Oxford, UK. 583 p.
Ma 2001, Scoggan 1978).
Gilstra, Z. 1991. Proposal to conserve the spelling
Ecology.—Strawberry bush grows well in many of 4618 Euonymus (Celastraceae). Taxon 40:
soils types, up to 1 m in a growing season. It is 137-139.
usually found in forests and thickets, including
lowland areas and edges of streams. It occurs from Gleason, H.A., and A. Cronquist 1991. Manual of
sea level up to 400 m in elevation (Ma 2001). vascular plants of northeastern United States and
adjacent Canada. 2nd Ed. New York Botanical
Reproduction.—Hymenopterous insects, Garden, Bronx, NY. 910 p.
especially bees, appear to be the main pollinators,
although wind may also be involved (Brizicky Ma, J.S. 2001. A Revision of Euonymus
1964). The seeds of strawberry bush are dispersed (Celastraceae). Thaiszia, Journal of Botany 11:
by birds and other small mammals (Brizicky 1-264.
1964). Strawberry bush blooms in late spring and
early summer, and the fruits mature in autumn. Paclt, J. 1998. Proposal to amend the gender of
Euonymus, nom. cons. (Celastraceae) to
Growth and Management.—Strawberry bush feminine. Taxon 47: 473-474.
prefers a humus-rich, slightly acidic soil. It does
well in shady situations, tolerating even full shade. Scoggan, H.J. 1978. The Flora of Canada Part 3 –
It can be planted in USDA Zones 5 to 9. Dicotyledoneae (Saururaceae to Violaceae).
Propagation can be effected through stem cuttings, National Museum of Natural Sciences, National
layering and seed (Chittenden and Synge 1956) Museums of Canada, Ottawa. 568 p.
Seeds require approximately 3 months of
prechilling and stratification. Shetler, S.G. and S.S. Orli. 2000. Annotated
Benefits.—Strawberry bush is an excellent Washington-Baltimore area. Part. 1. Ferns, fern
ornamental species. The leaves turn a dark red in allies, gymnosperms, dicotyledons. Smithsonian
autumn. The red capsules and scarlet arillate seeds Institution, Washington, D.C. 186 p.
add to the “burning effect” and contributes to some
of its common names. Strawberry bush is best USDA, NRCS. 2003. The PLANTS Database.
used in naturalistic settings, in the shade of larger Baton Rouge, LA. http:plants.usda.gov. [not
shrubs and trees. Large specimens can have paged].
334
Euonymus atropurpureus Jacq. eastern wahoo
CELASTRACEAE
Synonyms: Euonymus carolinensis Marsh.

Euonymus latifolius Marsh.

diameter. The capsule opens along the lobes,
exposing the seeds that are covered by a bright red
fleshy aril. The seeds (sans aril) are orange,
glabrous, ellipsoid, 7.0 to 11.0 by 4.0 to 5.5 mm,
glabrous. Some botanists spell the specific epithet
“atropurpurea” instead of “atropurpureus” used
here. This is because they disagree as to whether
the genus name Euonymus L. should be treated as
feminine (“atropurpurea”) or masculine
(“atropurpureus”) (Paclt 1998). In classical Greek,
the generic name was treated as feminine;
however, Linnaeus treated the genus as masculine.
The spelling “Euonymus” has formally been
conserved over Linnaeus’s originally spelling
“Evonymus,” which has also occasionally been
used (Gilstra 1991).
Range.—Eastern wahoo is widely distributed

throughout the Eastern United States (no records
Illustration source: Britton and Brown 1913 are known for Vermont) westward to the states of
Kansas, Montana, North Dakota, Oklahoma, South
General Description.—Eastern wahoo, also Dakota, and Texas (Ma 2001, Shetler and Orli
known as burning bush, is a shrub or small tree, up 2000, USDA 2003). It also is known from Ontario
to 8 m high, with the main stem up to 10 cm in (Gleason and Cronquist 1991, Scoggan 1978).
diameter at breast height. The stems are round,
smooth, and green. The pith is white, round, Ecology.—Eastern wahoo can be found in
continuous, and somewhat spongy. Its leaves are thickets, forests, often in lowlands or near creek
opposite (two per node) and spaced somewhat bottoms, riversides and along the margins of
evenly along stem. The petioles can be up to 14 marshes. It can also be found along mountain
mm long. The leaf blades are elliptic to oblong slopes. It usually occurs in elevations from sea
elliptic, 8.5 to 11.3 by 3.2.to 5.5 cm papery, with level to 360 m.
cuneate to attenuate bases, crenulate margins, and
acuminate apices. The blades have nine to 10 pairs Reproduction.—Hymenopterous insects,
of ascending lateral veins that disappear before especially bees, seem to be the main pollinators,
reaching the margin. The flowers are formed on although wind may also be involved (Brizicky
the current season’s growth in small axillary 1964). The fruits are dispersed by birds and small
clusters (cymes). Each peduncle is 2 to 3 cm long mammals (Brizicky 1964). Eastern wahoo blooms
and can have several flowers, each one on a in late spring and early summer and the fruits ripen
pedicel 5 to 7 mm long. The flowers are bisexual, in autumn. The plant reproduces asexually through
10 to 12 mm in diameter, with four greenish sepals rhizomes, which allows it to form large colonies.
and four brown-purple petals. There are four
stamens, each 1.5 to 2.0 mm long. The four carpels Growth and Management.—Eastern wahoo
are fused and a nectarous disk is present. The fruit refers well-drained loamy soils. It is a moderately
of the eastern wahoo is a smooth reddish to pink fast growing but short-lived shrub or tree. It can be
four-lobed (sometimes one or more of the lobes planted in Hardiness Zones 3 to 7. Seed requires 8
aborte) obrhombic capsule, up to 1.7 cm in
335
to 16 weeks cold stratification prior to sowing. The adjacent Canada. 2nd Ed. New York Botanical
plant can also be propagated with cuttings. Garden, Bronx, NY. 910 p.
Benefits.—The common name “burning bush” Ma, J.S. 2001. A Revision of Euonymus
comes from the plant’s bright red fall foliage. The (Celastraceae). Thaiszia, Journal of Botany 11:
red fruit and arillate seeds add to the “burning” 1-264.
effect. This makes the eastern wahoo an excellent
shrub for horticultural use (Rehder 1940). The Paclt, J. 1998. Proposal to amend the gender of
seeds and the vegetative organs of eastern wahoo Euonymus, nom. cons. (Celastraceae) to
are said to be poisonous (Brizicky 1964). feminine. Taxon 47: 473-474.
References Rehder, A. 1940. Manual of cultivated trees and

shrubs, Macmillan Publishing Co, New York.
Britton, N.L. and A. Brown. 1913. Illustrated flora 996 p.
possessions. Vol. 2, 2nd Ed. Scribner, New York. Scoggan, H.J. 1978. The Flora of Canada Part 3 –
735 p. Dicotyledoneae (Saururaceae to Violaceae).
National Museum of Natural Sciences, National
Brizicky, G.K. 1964. The genera of Celastrales in Museums of Canada, Ottawa. 568 p.
the Southeastern United States. Journal of the
Arnold Arboretum. 45: 206-235. Shetler, S.G. and S.S. Orli. 2000. Annotated
Gilstra, Z. 1991. Proposal to conserve the spelling Washington-Baltimore area. Part. 1. Ferns, fern
of 4618 Euonymus (Celastraceae). Taxon 40: allies, gymnosperms, dicotyledons. Smithsonian
137-139. Institution, Washington, D.C. 186 p.
Gleason, H.A. and A. Cronquist 1991. Manual of USDA, NRCS. 2003. The PLANTS Database.
vascular plants of northeastern United States and Baton Rouge, LA. http:plants.usda.gov. [not
paged].
336
Fallugia paradoxa (D. Don) Endl. ex Torr. Apache-plume
ROSACEAE
Synonyms: Sieversia paradoxa D. Don.

Geum cercocarpoides DC. ex Ser.
end of each stalk during the fruiting period. The
length of the flowering period varies depending
upon geographic location and elevation. It flowers
in June through August in all parts of its range but
flowering is extended from May to December in
Texas (Powell 1998) and from April to October in
Arizona and New Mexico (Epple 1995, Martin and
Hutchins 1980). The common name for Fallugia
paradoxa, Apache-plume, was coined because the
feathery seed clusters look like Indian feather
headdresses (Elmore 1976). Other common names
include poñil, fallugie (Dayton 1931), feather rose,
and feather duster bush (Epple 1995). Fallugia is a
monotypic genus, that is, a genus of a single
species (Cronquist and others 1997). In addition,
Apache-plume shows minimal genetic variation
throughout its wide range (Jones and Johnson,
1998). Hybrids between Apache-plume and
cliffrose (Purshia mexicana (D. Don) Henrickson)
have been reported (Blauer and others 1975,
Range.—Apache-plume occurs in western Texas,

Oklahoma, New Mexico, Arizona, southern
Colorado, south and central Utah, Nevada, and
Drawing source: USDA-Forest Service collection, Hunt southern California in the U.S.A. (USDA
Institute
PLANTS database 2002, Baldwin and others
2002, McGregor and other 1986, Vines 1986). In
General Description.—Fallugia paradoxa, Mexico it occurs in Coahuila, Chihuahua, and
commonly called “Apache-plume,” is an evergreen Durango (Powell 1998).
shrub that grows up to 2.5 m tall (Stubbendieck
and others 1993, Cronquist and others 1997). It is Ecology.—Apache-plume generally grows at
often referred to as “semi-evergreen.” In the north elevations between 915 and 2,430 m, but in
part of its range, seedlings are deciduous (Meyer northern New Mexico it has been found as high as
1974). However, the term “semi-evergreen” may 2,700 m (Dick-Peddie 1993). It grows on mesas
be applied because the leaves take on a bronze and rocky hillsides in montane shrubland,
color during some times of the year. The leaves are grassland, pinyon-juniper woodland, and yellow
alternate, somewhat fascicled, 0.8 to 1.5 cm long pine in a variety of alluvial and gravel soils
and have three to seven deep, narrow lobes. The (Tidestrom and Kittell 1941, Dick-Peddie 1993,
bark on young twigs is white, or light-colored, and Powell 1998, Baldwin and others 2002). It is
hairy (villous). The bark becomes darker, hairless especially abundant along dry margins of water-
(glabrous), and shreddy, exfoliating in flakes, as it courses and arroyos (author personal observation,
gets older. The showy, flowers have five white Powell 1998). Dick-Peddie (1993) described it as a
petals. There are numerous fruits (achenes) in each “semi-riparian” species because it tends to
head. Each achene is tipped by a 2.5 to 5 cm long colonize microhabitats, such as run-off catchments
feathery, densely hairy, tail-like style that makes and seep areas, which contain more available
the plants appear as if they have pom-poms on the
337
moisture than the surrounding area. Apache-plume year, environmental conditions, and availability of
is relatively drought tolerant and requires 20 to 50 alternate browse appears to influence palatability
cm of annual precipitation (Hayward 1990). It is and use. It provides cover for a variety of small
tolerant of weakly saline and neutral to moderately mammals and birds (Haywood 1990, McWilliams
basic soils (McWilliams 2000). Endomycorrhizae 2000). It was used for arrow shafts and brooms by
were associated with Apache-plume in New many Native American peoples (Dunmire and
Mexico (Williams and Aldon 1976). Tierney 1995, Moerman 1998). Apache-plume
brooms are specifically kept inside some Sandia
Reproduction.—Apache-plume reproduces Pueblo houses because there is a belief that the
sexually by seed and vegetatively by sprouts brooms exert a positive spiritual effect on the
(suckers) from the roots. The feather-tailed seeds household (Dunmire and Tierney 1995). The long
are probably primarily wind dispersed. Apache- roots were used as cord to tie fencing and make
plume seeds apparently have no dormant period ramadas (Dunmire and Tierney 1995, Moerman
(Baskin and Baskin 2001). They are small and 1998). An infusion of leaves was used as a
average approximately 925,932 per kg (Vines shampoo and hair growth stimulant by some
1986). Native American tribes, and petals are reported to
prevent stomach gas (Dunmire and Tierney 1995,
Growth and Management.—Seed can be sown Moerman 1998). One tribe used it in witchcraft to
by broadcasting it on a prepared bed and covered cause “insanity” (Moerman 1998). Shrubs are
by approximately 5 mm of fine loam or sand. attractive as ornamentals and are popularly used
Germination rate is from 19 to 65 percent (Vines for landscaping (Hayward 1990, Phillips 1987,
1986). In pots, the germination rate is 30 to 40 Morrow 1995).
percent, but seedlings are reported to damp off
easily and require adequate air circulation References
(Hayward 1990). A 4-litre plant is fully established
and ready for sale in 18 months for the landscape Aro, R.S. 1971. Evaluation of pinyon-juniper
market (Hayward 1990). Plants can also be grown conversion to grassland. Journal of Range
fairly easily from root cuttings. It is closely grazed Management. 24(3): 188-197
on overstocked range and, although tolerant of
grazing with “excellent recuperative powers,” it is Baldwin B.G., S. Boyd, B.J. Ertter, R.W.
also reported to decrease under grazing pressure Patterson, T.J. Rosatti, and D.H. Wilken. 2002.
(Dayton 1931, McGregor and others 1986). The Jepson Desert Manual: Vascular plants of
Availability of adequate moisture may be an southeastern California. University of California
important factor in the degree of recovery. Press, Berkeley, CA. 624 p.
Apache-plume sprouts after fire (Shaw and
Monson 1983, McWilliams 2000). In a pinyon Baskin, C.C. and J.M. Baskin. 2001. Seeds.
juniper community it was reported to tolerate fire Ecology, biogeography, and evolution of
well (Aro 1971). dormancy and germination. Academic Press,
Benefits.—Apache-plume is valuable for erosion
control (Dayton 1931, Stubbendieck and others Blauer, A. C., A.P. Plummer, and E.D. McArthur.
1993, Vines 1986). Its long roots provide effective 1975. Characteristics and hybridization of
soil stabilization especially in arid situations important Intermountain shrubs. I. Rose family.
(McWilliams 2000). Apache-plume is important Research Paper INT-169. U.S. Department of
winter range for livestock (Dayton 1931, Agriculture, Forest Service, Intermountain
McGregor and others 1986, Stubbendieck and Forest and Range Experiment Station, Ogden,
others 1993, USDAFS 1988). It is reported to have UT. 36 p.
only a fair palatability for sheep, goats and cattle
but closely cropped, stunted plants have been cited Cronquist, A., N.H. Holmgren, and P.K.
as evidence of high palatability (Dayton 1931, Holmgren. 1997. Intermountain Flora – vascular
McGregor and others 1986, Stubbendieck and plants of the Intermountain west, U.S.A. Vol. 3
others 1993, Warnock 1974). It also furnishes part A. Subclass Rosidae. The New York
important browse for some species of wildlife Botanical Garden, Bronx, NY. 446 p.
(Dayton 1931, Stubbendieck and others 1993) and
makes up 1 percent to 5 percent of desert mule Dayton, W.A. 1931. Important western browse
deer diet (Krausman and others 1997). Time of plants. Miscellaneous Publication 101. U.S.
338
Department of Agriculture, Washington, DC. Moerman, D.E. 1998. Native American ethno-
214 p. botany. Timber Press, Portland, OR. 927 p.
Dick-Peddie, W.A. 1993. New Mexico Vegetation Morrow, B.H. 1995. Best Plants for New Mexico
– past, present, and future. University of New Gardens and Landscapes. University of New
Mexico Press, Albuquerque, NM. 244 p. Mexico Press, Albuquerque, NM. 267 p.
Dunmire, W.W. and G.D. Tierney. 1995. Wild Phillips, J. 1987. Southwestern landscaping with
plants of the Pueblo Province. Museum of New native plants. Museum of New Mexico Press,
Mexico Press, Santa Fe, NM. 290 p. Santa Fe, NM. 141 p.
Elmore. F.H. 1987. Shrubs and Trees of the Powell, A.M. 1998. Trees and shrubs of Trans-
Southwest Uplands. Southwest Parks and Pecos and adjacent areas. University of Texas
Monuments Assoc. Tucson, AZ. 214 p. University Press, Austin, TX . 498 p.
Epple, A.O. 1995. A field guide to the plants of Shaw, N.L. and S.B. Monson. 1983. Phenology
Arizona. Falcon Press Publishing Co., Helena, and growth habits of nine antelope bitterbrush,
MT. 347 p. desert bitterbrush, stansbury cliffrose, and
apache-plume accessions. In: A.R. Tiedemann
Hayward, P. 1990. Fallugia paradoxa. American and K.L. Johnson, comps. Proceedings:
Nurseryman. 172(7): 126 Research and Management of bitterbrush and
cliffrose in Western North America: 1982, April
Jones, T.A. and D A. Johnson. 1998. Integrating 13-15, Salt Lake City, UT. General technical
Genetic Concepts into Planning Rangeland Report: INT-152, USDA Forest Service,
Seedings. Journal of Range Management Intermountain Forest and Range Station, Ogden,
51(6): 594-606. UT.
Krausman, P.R., A.J. Kuenzi, R.C. Etchberger, Stubbendieck, J., S.L. Hatch and C.H. Butterfield.
K.R. Rautenstrauch, L.L. Ordway, and J.J. 1993. North American range plants. 4th edition.
Hervert. 1997. Diets of mule deer. Journal of University of Nebraska Press, Lincoln, NE.
Range Management. 50(5): 513-522. 493 p.
Martin, W.C. and C.R. Hutchins. 1980. A Flora of Tidestrom, I. and T. Kittell. 1941. A flora of
New Mexico. Strauss & Cramer, Hirschberg, Arizona and New Mexico. The Catholic
Germany. 2,592 p. University of America Press, Washington, DC.
897 p.
McArthur, E.D., B.C. Giunta, and A.P. Plummer.
1977. Shrubs for restoration of depleted range USDA Forest Service. 1988. Range Plant
and disturbed areas. Utah Science. 35: 28-33. Handbook. Dover Publications, Inc., New York,
NY. 842 p.
McGregor, R.L. and the Great Plains Flora
Association. 1986. Flora of the Great Plains. USDA PLANTS database. 2002. Internet site:
University Press of Kansas, Lawrence, KS. http:// plants.usda.gov/cgi_bin/plant_profile.
1,402 p. cgi? symbol=FAPA#. [not paged].
McWilliams, J. 2000. Fallugia paradoxa. In: U.S. Vines, R.A. 1986. Trees, shrubs, and woody vines
Department of Agriculture, Forest Service, of the Southwest. Sixth printing. University of
Rocky Mountain Research Station, Fire Sciences Texas Press. Austin, TX. 1,104 p.
Laboratory. Fire Effects Information System.
http://www.fs.fed.us/database/feis/ 19 p. Warnock, B.H. 1974. Wildflowers of the
Guadalupe Mountains and the sand dune
Meyer, S. 1974. Fallugia paradoxa (D. Don) Endl. country, Texas. Sul Ross university, Alpine, TX.
ex Torr. Apache-plume. In: . F.T. Bonner and 176 p.
R.G. Nisley, eds. USDA Forest Service Woody
Plant Seed Manual Internet site: http:// wpsm.net Williams, S.E. and E.F. Aldon. 1976.
4 p. Endomycorrhizal (vesicular arbuscular)
339
associations of some arid zone shrubs. The
Southwestern Naturalist. 20(4): 437-444.
340
Faramea occidentalis (L.) A. Rich. cafeíllo
RUBIACEAE
Synonyms: Coffea occidentalis Jacq.

Faramea odoratissima DC.
Pavetta coffeoides Ham.
Tetramerium odoratissimum (Gaertn. f.) DC.
Ixora occidentalis L.
John K. Francis
(Holdridge 1967). Alluvial soils and residual soils

derived from sedimentary and volcanic rocks are
suitable habitat. Excessively drained and very
poorly drained soils are usually not colonized.
Although not forming pure or dense stands,
cafeíllo is common, at least in Puerto Rico and
Panama.
Reproduction.—Small, white, fragrant flowers

have a short tube tipped by four pointed lobes.
Cafeíllo flowers generally occur in threes. The
fruits are blue-black or dark-purple drupes borne
singly or in pairs from the leaf axils near the twig
ends. The fruits measure 8 to 10 mm in diameter
(Howard 1989). Each fruit contains a single seed.
A Puerto Rican collection averaged 0.825 + 0.123
g/fruit. The seeds from the same collection
numbered 3,980 seeds/kg. Ninety-six percent
General Description.—Cafeíllo, also known as germinated beginning 57 days after sowing
false coffee, café cimarrón, palo de toro, hueso, (Francis and Rodríguez 1993). In Central America,
jasmín de estrella, and bois flèche, is a tall shrub in the seeds are disbursed by birds and monkeys.
the Caribbean islands and a small understory tree Rodents and a number of other animal species eat
in continental tropical America. It has dark-green, and destroy the seeds after fruit-fall (Schupp
opposite lanciolate leaves and straight, usually 1990). A rust, Aecidium farameae Arthur, which
single, gray or dark-brown stems. The twigs are attacks the ovaries and pedicels of the flowers,
slender and green with rings at the nodes (Little results, in some years, in the destruction of 32 to
and Wadsworth 1964). The stems are straight and 83 percent of the fruits of individual plants
often without branches for a meter or more. (Travers 1998). A high recruitment of seedlings
(compared to other species) results in high
Range.—The native range extends from Cuba population densities in the forest understorey in
through Trinidad and from central Mexico through Panama (Schupp 1990).
Central America to Ecuador, Peru, Brazil, and the
Guyanas (Little and Wadsworth 1964). The Growth and Management.—Cafeíllo
species is not known to have naturalized occasionally exceeds 20 m in height and 20 cm in
elsewhere. diameter at breast height in Panama (O’Brien and
others 1995). However, in Puerto Rico, mature
Ecology.—Cafeíllo is tolerant and grows and shrubs usually reach 4 m in height and sometimes
reproduces in the understories and mid-stories of attain as much as 10 m in height (Liogier 1997).
secondary forests. The species is most common in Total above-ground, ovendried weight of cafeíllo
mid- to late-secondary forests. Cafeíllo is found in plants in Puerto Rico that ranged between 1.37 m
two of the Life Zones in Puerto Rico: moist in height (with diameter greater than 0) and 5 cm
subtropical forest and much of the wet subtropical in d.b.h. may be predicted by the model:
forest from about 900 to 2800 mm of precipitation
341
TDW = 30.045D2L (adjusted R-squared = 0.962), Holdridge, L.R. 1967. Life zone ecology. Tropical
where TDW is total dry weight in grams, D is Science Center, San José, Costa Rica. 206 p.
diameter in cm at 30 cm above the ground, and L
is stem length in meters. Lower stem density of Howard, R.A. 1989. Flora of the Lesser Antilles,
Puerto Rican shrubs averaged 0.62 + 0.04 g/cm3 Leeward and Windward Islands. Vol. 6. Arnold
(Francis 2000). Stem density of saplings from Arboretum, Harvard University, Jamaica Plain,
Panama was 0.57 g/cm3 (King 1990). Allometric MA. 658 p.
relationships for cafeíllo in Panama may be found
in O’Brien and others (1995) and King (1990). As King, D.A. 1990. Allometry of saplings and
an understory tree, cafeíllo is slow-growing and understory trees of a Panamanian forest.
long lived. Based on recent growth rates, the Ecology 4(1): 27-32.
largest individuals in a Panamanian stand were
estimated to exceed 200 years in age. Liogier, H.A. 1990. Plantas medicinales de Puerto
There is no record of plantations or Rico y del Caribe. Iberoamericana de Ediciones,
planted ornamental cafeíllo. The species is able to Inc., San Juan, PR. 566 p.
reproduce by sprouting after injury. Thirty-six
percent of the individuals in a Panamanian stand Liogier, H.A. 1997. Descriptive flora of Puerto
were of sprout origin (Putz and Brokaw 1989). Rico and adjacent islands. Vol. 5. Editorial de la
Benefits.—Cafeíllo wood is heavy, hard, and 436 p.
tough. However, uses are mainly limited to fence
posts, vegetable stakes, tool handles, and Little, E.L., Jr. and F.H. Wadsworth. 1964.
fuelwood. In herbal medicine, cafeíllo is Common trees of Puerto Rico and the Virgin
reportedly used as an antiseptic, astringent, and for Islands. Agriculture Handbook 249. U.S.
diarrhea and anemia (Liogier 1990). Birds Department of Agriculture, Forest Service,
consume the fruits. The use of cafeíllo as an Washington, DC. 548 p.
ornamental or in forest restoration efforts has not
been reported. Cafeíllo is a fine looking plant and O’Brien, S.T., S.P. Hubbell, P. Spiro, R. Condit,
could probably be used as a back-ground plant or and R.B. Foster. 1995. Diameter, height, crown,
screen under shade in yards, estates, and parks. and age relationships in eight neotropical tree
species. Ecology 76(6): 1926-1939.
References
Putz, F.E. and N.V.L. Brokaw. 1989. Sprouting of
Francis, J.K. 2000. Estimating biomass and carbon broken trees on Barro Colorado Island, Panama.
content of saplings in Puerto Rican secondary Ecology 70(2): 508-512.
forests. Caribbean Journal of Science 36(3-4):
346-350. Schupp, E.W. 1990. Annual variation in seedfall,
postdispersal predation, and recruitment of a
Francis, J.K. and A. Rodríguez. 1993. Seeds of neotropical tree. Ecology 71(2): 504-515.
Puerto Rican trees and shrubs: second
installment. Research Note SO-374. U.S. Travers, S.E. 1998. The effect of rust infection on
Department of Agriculture, Forest Service, reproduction in a tropical tree (Faramea
Southern Forest Experiment Station, New occidentalis). Biotropica 30(3): 438-443.
Orleans, LA. 5 p.
342
Forestiera acuminata (Michx.) Poir. swamp privet
OLEACEAE
Synonyms: Adelia acuminata Michx.

Borya acuminata (Michx.) Willd.
Kristina Connor
Ecology.—Swamp privet grows primarily in wet

areas, in swamps and bottomland forests, along
streams, in wet woods and sloughs, and around
ponds and lakes (Johnson and Hoagland 1999,
USGS 2002). It will grow in either full or partial
sun and is highly adaptable to various soils
(MacKay and Finical 1999). The species
reportedly thrives in a variety of soil types, sand,
loam or clay, and at pH’s ranging from acidic to
basic. Swamp privet does, however, require that
the soils be moist (Plants for a Future 2001). It
frequently produces new stems from the base of
senescing individuals, especially when soils are
flooded (Mackay and Finical 1999, U.S. Army
Corps of Engineers 1997).
Drawing credit: USDA—Forest Service collection, Hunt Reproduction.—Swamp privet plants are
Institute monoecious; flowers emerge in the spring before
the leaves unfold. They are yellow, fragrant and
General Description.—Swamp privet, also called small, the males sessile or almost so, female
eastern swamp privet, is a deciduous shrub or flowers in small panicles or fascicles (Bailey and
small, open-crowned tree, occasionally reaching Bailey 1976, Botanical Institute 2002, Johnson and
10 m in height but more often averaging 1.5 to 2.5 Hoagland 1999, Krüssmann 1986). They have no
m (Krüssmann 1986, Johnson and Hoagland petals and minute or absent sepals. Male flowers
1999). It is commonly multitrunked (Mackay and have one to four stamens, female flowers have
Finical 1999, U.S. Army Corps of Engineers simple or two-lobed stigmas (Johnson and
1997). The bark ranges from gray to dark brown in Hoagland 1999, Botanical Institute 2002, USGS
color and is either smooth or ridged (Johnson and 2002). Fruits are dark purple or black oblong
Hoagland 1999, USGS 2002). Twigs are light drupes, 8 to 12 mm long with one, rarely two,
brown, glabrous, and have conspicuous lenticels seeds. They appear in summer and are promptly
(Johnson and Hoagland 1999). Leaves are simple, shed (Johnson and Hoagland 1999). Although no
opposite, and dull, yellowish green. They range in data are provided to support this, the Plants for a
size from 3 to 11 cm in length, and 2.5 to 5 cm in Future website (2001) suggests that seeds be sown
width. Leaves have long, pointed tips (acuminate) as soon as they ripen and that cuttings can be taken
and petioles that are slender and slightly winged at of half-ripe wood in summer or mature wood in
the base. While leaf margins are usually smooth, it winter.
is not uncommon for the apex half to be finely
serrated. Growth and Management.—Swamp privet,
while adaptable to many soil types, must be
Range.—Swamp privet is found in the wet located in moist areas. It is moderately tolerant of
woodlands and swamps of the Southeastern United heat and can tolerate full or partial sunlight
States, from South Carolina south to Florida and (Mackay and Finical 1999). It has been reported in
westward into Texas, Oklahoma, and Kansas. It cultivation (Krüssmann 1986).
ranges as far north as Illinois, Indiana, and
Missouri (Krüssmann 1986, USGS 2002). Benefits.—The fruits of swamp privet are
considered good food for wildlife (Johnson and
343
Hoagland 1999). A decoction of the roots and bark Mackay, W. and L. Finical. 1999. A special
has been used as a health beverage, and the wood project of the Texas Agricultural Experiment
has been used for turning (Bailey and Bailey Station. Benny Simpson’s Texas Native Trees.
1976), although reports on its durability vary Texas A&M University, College Station.
(Plants for a Future 2001). http://aggie-horticulture.tamu.edu/natives. 1 p.
References Plants for a Future (PFAF). 2001. Resource and

Information Centre for Edible and Other Useful
Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third: Plants. Forestiera acuminata.
A Concise Dictionary of Plants Cultivated in the http://www.scs.leeds. ac.uk/ pfaf/index.html. 4
United States and Canada. McMillan Publishing p.
Co., Inc., New York. 1,312 p.
U.S. Army Corps of Engineers. 1997. Wetlands
Botanical Institute. 2002. Forestiera Poiret. Delineation Manual. Appendix C: Vegetation.
Goteborg University. http://www.systbot. The Wetlands Regulation Center
gu.se/staff/evawal/ oleaceae/forestiera.html. 2 p. http://www.wetlands. com/coe/87manapc.htm.
6 p.
Johnson, F.L. and B.W. Hoagland. 1999.
Forestiera acuminata (Michx.) Poir. Catalog of USGS. 2002. Swamp privet (Forestiera
the Woody Plants of Oklahoma, Oklahoma acuminata). Northern Prairie Wildlife Research
Biological Survey http://www.biosurvey.ou.edu. Center. Directory of Resource
1 p. http://www.npwrc.usgs.gov/resource/
othrdata/plntguid/species. 2 p.
Krüssmann, G. 1986. Manual of Cultivated Broad-
Leaved Trees and Shrubs. Volume II, E-Pro.
Timber Press, Beaverton, OR. 445 p.
344
Forestiera segregata (Jacq.) Krug & Urban Florida privet
OLEACEAE
Synonyms: Myrica segregata Jacq.

Adelia porulosa Michx.
Forestiera porulosa (Michx.) Poir.
Adelia segregata (Jacq.) O. Kuntze
Adelia pinetorum Small
Forestiera cassinoides Poir.
John K. Francis
Antilles, the Cayman Islands and Anguilla,

Barbuda, Antigua, La Désirade, and Marie Galante
in the Lesser Antilles (Howard 1989, Liogier 1995,
National Trust for the Cayman Islands 2002,
Long and Lakela (1976) identify two varieties: var.
segregata with glabrous stems and leaf blades
more than 3 cm long in Florida and Georgia, and
var. pinatorum, which has puberulent stems and
leaf blades 1 to 3 cm long in South Florida.
Ecology.—Florida privet grows best in partial

shade (Horticopia 2002). It is tolerant of drought,
salt, and alkaline soils (University of Florida
Cooperative Extension Service 2002). Florida
privet competes well with shrubs and herbs and
grows in hammocks and pinelands in Florida (Long
and Lakela 1976). In Puerto Rico and other
Caribbean Islands, it is most frequently seen in dry
forests in limestone areas near the coast. The
species is rare in the United States and has become
a candidate for Federal listing (Everglades
Mitigation Bank 2002).
General Description.—Florida privet, also known Reproduction.—In Florida, Florida privet flowers
as Florida swamp privet, wild olive, and ink-bush, in the spring (Nelson 1996). It is insect pollinated.
is a semideciduous shrub or small tree to 7 m in A collection of fruits from Puerto Rico weighed an
height. The plant is supported by a weak taproot average of 0.1209 + 0.0035 g/fruit. Air-dried seeds
and more robust, tan-colored lateral roots. The cleaned from them weighed an average of 0.0359 +
branches are slender. The opposite leaves have 0.0006 g/seed or 27,900 seeds/kg. Sown in moist
short petioles, 1 to 6 mm long, and narrowly potting mix, they began germinating in 7 days and
elliptic to obovate blades, 1 to 7 cm long, with completed 89 percent germination in 49 days
entire margins. Tiny, greenish-yellow flowers are (author’s observation). The seeds are dispersed by
borne in clusters at the leaf axils. The fruits are birds that eat the fruits. Seedlings do not occur in
slightly curved, ovoid, black drupes, 6 to 10 mm large numbers.
long, that have a bitter, unpleasant flavor and stain
cloth and skin. Each contains one greenish-yellow Growth and Management.—Florida privet grows
seed (Howard 1989, Liogier 1995, Long and slowly in the early seedling stage but later
Lakela 1976, Nelson 1996). accelerates so that seedlings are ready to transplant
at the end of 1 year. Nursery production is
Range.—Florida privet is native to Florida, normally from seed. Wildlings of all sizes can be
Georgia, South Carolina, the Bahamas, the Greater
345
successfully transplanted (Workman 1980). Howard, R.A. 1989. Flora of the Lesser Antilles,
Beyond the seedling stage, it has a moderate Leeward and Windward Islands.
growth rate (University of Florida Cooperative Dicotyledoneae. Vol. 6. Arnold Arboretum,
Extension Service 2002). The largest Florida privet Harvard University, Jamaica Plain, MA. 658 p.
recorded measures 20.2 cm diameter at breast
height and 5.5 m in height (American Forests Liogier, H.A. 1995. Descriptive flora of Puerto
2002). Rico and adjacent islands. Vol. 4. Editorial de la
Benefits.—Florida privet is recommended for 617 p.
xeriscape (unwatered) landscaping (South Florida
Water Management District 2001) and is available Long, R.W. and O. Lakela. 1976. A flora of
through a number of commercial nurseries. It is tropical Florida. Banyon Books, Miami, FL.
useful for hedges and foundation and specimen 962 p.
plantings. Warblers, bluebirds, mockingbirds, and
vireos feed upon the fruits in South Florida National Trust for the Cayman Islands. 2002.
(Broschat and Verkade 2002, Cowley 2002). Virtual herberium: Oleaceae, Forestiera
Honeybees, butterflies, and other insects are segregata (Jacq.) Krug & Urban. http://www.
attracted to the flowers for the nectar (Workman virtualherbarium.org/vh/CAYM/1002.html. 2 p.
1980).
References Plants profile: Forestiera Poir. swampprivet.
http://plants.usda.gov/cgi_bin/plant_profile.cgi?
American Forests. 2002. National register of big symbol=FORES. 3 p.
trees: Florida-privet, Forestiera segregata.
http://www.americanforests.org/resources/bigtre Nelson, G. 1996. The shrubs and woody vines of
es/register.php?details=2000. 1 p. Florida. Pineapple Press, Inc., Sarasota, FL.
391. p.
Broschat, T.K. and S.V. Verkade. 2002.
Landscaping to attract birds in South Florida. South Florida Water Management District. 2001.
http://www.floridaplants.com/landscape/birds. Quick facts on xeriscape: how to save water
htm. 3 p. through water-smart landscaping. http://www.
sfwmd.gov/images/pdfs/splash/splxeris.pdf. 2 p.
Cowley, M. 2002. Florida native plants: olive
family. Native Florida Web Ring. University of Florida Cooperative Extension
http://nsis.org/garden/family/olive.html. 2 p. Service. 2002. Native shrubs of South Florida.
http://edis.ifas.ufl.edu/BODY_EH159. 9 p.
Everglades Mitigation Bank. 2002. Endangered
and threatened plants at the EMB. Workman, R.W. 1980. Growing native. The
http://www.fpl.com/environment/emb/contents/e Sanibel-Captiva Conservation Foundation, Inc.,
ndangered_plants.shtml. 1 p. Sanibel, FL. 137 p.
Hortocopia. 2002. Forestiera segregata, wild olive.

http://hortpix.com/pc1848.htm. 1 p.
346
Fouquieria splendens Engelm. ocotillo
FOUQUIERIACEAE
Synonyms: none

level to as high as 2,050 m elevation in the
Guadalupe and Del Norte mountains of the Trans-
Pecos region of Texas. Soils are generally rocky,
shallow, well-drained, and of limestone or granitic
origin and are often underlain by caliche (Mathews
1994). Although not restricted to calcareous soils,
ocotillo tends to be characteristic of a calcareous
community (Schmutz and others 1992). It has a
relatively shallow root system (Burgess 1995). It is
often found on southern facing slopes. In Mexico
it also occurs in both the Sonoran and Chihuahuan
deserts. Communities in which it commonly
occurs are mesquite-grasslands (Mason and Mason
1987), high desert bunch-grass (Schmutz and
others 1992), desert scrub (Van Devender 1995)
and mesquital (mesquite), motorral desértico
microfilo (fine-leaf desert shrubland), and pastizal
(grassland) defined by Flores-Mata and others
Illustration credit: Tonia Masaood (1971).
General Description.—The common name for Reproduction.—Ocotillo may self-pollinate to a

Fouquieria splendens is Ocotillo. This is a relative limited extent, but flowers are generally cross-
of the renowned boojum tree (Fouquieria pollinated by hummingbirds and bees. In southern
columnaris) of Baja California. Ocotillo is an erect Arizona flowering coincides with the northward
shrub with several spiny ascending stems that migration of hummingbirds (Mathews 1994). Seed
grow up to 3 m tall. A mature individual may have set per flower is greater for plants flowering during
as many as 75 stems (Epple 1995). The leaves are migration than for plants flowering after
oblanceolate to oblong-obovate and only 10 to 25 hummingbirds have migrated (Rathcke and Lacey
mm long. The bright scarlet flowers are showy, 1985, Waser 1979). The winged seeds of ocotillo
and an individual may flower several times during are probably wind dispersed.
the year. The flowers are arranged in an 8 to 25-cm
-long panicle. Each flower has a tubular corolla, Growth and Management.—Ocotillo is a long-
five petals, and 10 to 15 exerted stamens. The seed lived stable element of desert vegetation but with
capsule is ovoid and incompletely three-celled. apparently specialized requirements for seed
The flat seeds are white with a fringe of hair-like germination that limit its capacity to exploit
filaments. Flowering is from March to June opportunities for population expansion (Zedler
depending upon latitude (Martin and Hutchins 1981). There seems to be no significant change in
1980). the distribution and abundance of this species
within the recent past. Evidence suggests that
Range.—Ocotillo occurs in Arizona, southern ocotillo habitat has increased as grasslands have
California, New Mexico, Nevada, and Texas in the declined (Van Devender 1995). However,
USA. In Mexico it occurs in Sonora, Coahuila, considering the amount of urban development
Chihuahua, Durango, Zacatecas, San Luis Potosí, within octotillo habitat, the actual area occupied by
Hildalgo, and Baja California (Mason and Mason the species may not have changed significantly. It
1987). may be significant to overall species sustainability
that seedlings are rarely found on disturbed soils
Ecology.—Ocotillo grows in shallow soils on (Yeaton and others 1977) but are found under the
mesas, outwash plains, and rocky slopes from sea canopy of mature plants (McAuliffe 1988, Young
347
and Young 1986). It is a slow-growing species, cultivated genotypes originally adapted to another
and estimates of its life span range from less than region. This may pose a threat although, because
60 to greater than 72 years (Mathews 1994). It is of the slow growth and low turnover of
likely that life span is related to latitude and individuals, the effects would be a long time in
substrate. The leaves are winter- and drought- becoming apparent. For example, the cultivated
deciduous (Schmutz and others 1992). The leaves genotypes may have particularly high fecundity
appear after rains and are a reliable indicator of and may be physiologically robust but do not have
recent rainfall (Powell 1998). They tend to mature the genetic composition to be resistant to
and fall within a few weeks indicating that environmental or biological (e.g. disease or insect
available photosynthate is limited and that growth infestation) events that are periodic (perhaps
is dependent upon precipitation. Ocotillo is a decades apart) in the local area. Hedges or fences
“water storer” in that the stems store relatively of living stems are common, especially in Mexico.
large amounts of water that buffer against drought It does not appear to be grown for its medicinal
stress (Burgess 1995). Wildfires may be value. Historically the Mahuna Native Americans
detrimental to ocotillo populations. It rarely used it as a “blood medicine” (Moerman 1998).
sprouts from the root crown following damage The Apache Indians used powdered roots to treat
from fire, and its sprouting ability is probably wounds and painful swellings and they also bathed
dependent on fire severity (Ahlstrand 1982, in an ocotillo root mixture to relieve fatigue
McPherson 1995, White 1969). Mortality rates of (Krochmal and others 1954, Powell 1998, Vines
40 percent to 50 percent were reported after light 1960). A beverage made from ocotillo flowers was
to moderate fire in Arizona (Thomas 1991). used for cough medicine (Vines 1960). The
Seedlings are not known to establish in recently Cahuilla, Papago and Yavapai tribes used the
burned areas (Mathews 1994), and its seeds are flowers in beverages, the fruits for a “candy,” and
unlikely to survive for long in the soil (Zedler the stalks as building material (Moerman 1998).
1981). In fact, burning has been suggested as a The thorns were used for ear piercing by the
control method for woody species, such as Papago. The resin and wax from ocotillo bark was
Fouquieria splendens, in desert grasslands used for conditioning leather (Krochmal and others
(Mathews 1994). 1954, Powell 1988, Vines 1960). It seems that all
uses are localized, personal, or historic in the
Benefits.—The endangered Lucifer humming- Southwestern United States. It is not known if
bird's habitat in New Mexico centers on slopes and ocotillo is more widely used today in Mexico.
adjacent canyons in arid montane areas dominated
by ocotillo and agave species (New Mexico References
Department of Game and Fish 1991). Ocotillo has
no recorded forage value, presumably because its Ahlstrand, G.M. 1982. Response of Chihuahuan
leaves are so short-lived (Dayton 1931). Ocotillo Desert Mountain Shrub Vegetation to Burning.
plants are popularly used for landscaping. Plants Journal of Range Management 35 (1): 62-65.
sold in reputable nurseries are obtained from
private land under permit. There are typically two Burgess, T.L. 1995. The dilemma of coexisting
“types” of product. Smaller plants are sold in 1 growth forms. In: The desert grassland. M.P.
gallon pots or larger specimens are sold “balled McClaran and T.R. Van Devender, eds. The
and burlapped.” Although the plants transplant University of Arizona Press, Tucson, AZ. p. 31-
well if there are sufficient roots attached, many 67.
that are dug are not treated well and two thirds or
more of those dug up are likely not to survive. Dayton, W.A. 1931. Important western browse
Propagation has been reported using cuttings plants. Miscellaneous Publication 101. U.S.
although conversations with horticulturalists Department of Agriculture, Washington, DC.
indicate that this may be less easy than the 214 p.
literature indicates. The plant can also be grown
from seed but is a relatively slow grower. Seeds Epple, A.O. 1995. A field guide to the plants of
apparently collected from Mexico can be Arizona. Falcon Press Publishing Co., Helena,
purchased in the United States (Tradewinds MT. 347 p.
Import-Export Company 2002). This trade
between regions where the plant is native does Flores-Mata, G.F. J.J. Lopez, X.M. Sánchez, F.M.
raise the potential for genetic modification, or Ruiz, and F.T. Takaki. 1971. Memoria del Mapa
“dilution,” of locally adapted genotypes by Tipos de Vegetación de la República Mexicana.
348
Secretaria de Recursos Hidráulicos, Dirección Schmutz, E.M., E.L. Smith, P.R. Ogden, M.L.
de Agrología, Mexico, DF. 59 p. Cox, J.O. Klemmedson, J.J. Norris, and L.C.
Fierro. 1992. Desert grassland. In: Natural
Krochmal, A., S. Paur, and P. Duisberg. 1954. grasslands: Introduction and Western
Useful native plants in the American Hemisphere. R.T. Coupland, ed. Elsevier, New
Southwestern deserts. Economic Botany. 8: 3- York. p. 337-362.
20.
Thomas, P.A. 1991. Response of succulents to
Martin, W.C. and C.R. Hutchins. 1980. A Flora of fire: A review. International Journal of Wildland
New Mexico. Strauss & Cramer, Hirschberg, Fire 1(1): 11-22.
Germany. 2,592 p.
Tradewinds Import-Export Company. 2002. Seeds
Mason, C.T. and P.B. Mason. 1987. A handbook Department – USA. P.O. Box 218650, Houston,
of Mexican roadside flora. The University of Texas 77218 USA. http://www.abonline.
Arizona Press, Tucson, AZ. 380 p. com/products/seeds_f.htm#f. [not paged].
Matthews, R.F. 1994. Fouquieria splendens. The Van Devender, T.R. 1995. Desert grassland
Fire Effects Information System Data base. U.S. history: changing climates, evolution,
Department of Agriculture, Forest Service, biogeography, and community dynamics. In:
Intermountain Research Station, Intermountain The desert grassland. M.P. McClaran and T.R.
Fire Sciences Laboratory, Missoula, MT. http:// Van Devender, eds. The University of Arizona
www.fs.fed.us/database/feis/plants/shrub/fouspl/ Press, Tucson, AZ. p. 68-99.
all.html. 15 p.
McAuliffe, J.R. 1988. Markovian dynamics of of the Southwest. Sixth printing. University of
simple and complex desert plant communities. Texas Press. Austin, TX. 1,104 p.
American Naturalist. 131(4): 459-490.
Waser, N.M. 1979. Pollinator availability as a
McPherson, G.R. 1995. The role of fire in desert determinant of flowering time in ocotillo
grasslands. In: The desert grassland. M.P. (Fouquieria splendens). Oecologia. 39(1): 107-
McClaran and T.R. Van Devender, eds. The 121.
University of Arizona Press, Tucson, AZ. p.
130-151. White, L.D. 1969. Effects of a wildfire on several
desert grassland shrub species. Journal of Range
Moerman, D.E. 1998. Native American ethno- Management 22: 284-285.
Yeaton, R.I., J. Travis, and E. Gilinsky. 1977.
New Mexico Department of Game and Fish. 1991. Competition and spacing in plant communities: the
Handbook of species endangered in New Arizona upland association. Journal of Ecology. 65:
Mexico. Dept. of Game and Fish., Santa Fe, 587-595
NM. 185 p.
Young, J.A. and C.G.Young. 1986. Collecting,
Powell, A.M. 1988. Trees & shrubs of Trans- processing, and germinating seeds of wildland
Pecos Texas including Big Bend and Guadalupe plants. Timber Press. Portland, OR. 236 p.
Mountains National Parks. Big Bend Natural
History Association, Big Bend National Park, Zedler, P.H. 1981. Vegetation change in chaparral
TX. 536 p. and desert communities in San Diego County,
California. In: D.C. West, H.H. Shugart, D. B.
Powell, A.M. 1998. Trees & shrubs of Trans- Botkin, eds. Forest succession: Concepts and
Pecos and adjacent areas. University of Texas application. Springer-Verlag, New York. p.
University Press, Austin, TX . 498 p. 406-430
Rathcke, B. and E.P. Lacey, 1985. Phenological

patterns of terrestrial plants. Annual Reviews of
Ecological Systems. 16: 179-214.
349
Furcraea foetida (L.) Haw. Mauritius hemp
AGAVACEAE
Synonyms: Furcraea gigantea Vent.

Agave gigantea D. Dietr.
John K. Francis
Range.—Mauritius hemp is native to the Greater
Antilles, and from Guadaloupe south through
northern South America to Brazil (Grisebach
1963, Howard 1979). The species has been widely
planted. It has naturalized in Florida, Hawaii,
Marquesas Islands, French Polynesia, Tonga, and
is present and probably naturalized in many other
places (Pacific Island Ecosystems at Risk 2003).
Taxonomy.—Mauritius hemp in Puerto Rico was

known previously as Furcraea tuberose (Miller)
Ait. f. (Liogier and Martorell 2000). The current
assignment of the species to F. foetida is based on
the statement by Howard (1979) that F. foetida
occurred throughout the Antilles and that F.
General Description.—Mauritius hemp, also tuberosea is endemic to the Lesser Antilles,
known as green aloe, female karata, maguey, specifically from St. Barts southward.
mayuey criollo, cocuisa, giant cabuya, and aloes Characteristics given to separate the two species
vert, is a robust shrub with a basal rosette about are unreliable by Howard’s own admission.
2.5 to 3.5 m in diameter and flowering stalks 5 to Mature plants examined by the author had
10 m in height. The plant can be called a shrub maximum leaf widths of 18 to 20 cm, leaf lengths
because it is perennial, because it is shrub size, and of 1.5 to 2 m, oblanceolate leaves, weak fetid
because it has a fibrous-woody sheath surrounding smell of leaf juices, and bulbiferous reproduction.
the core of the short stem (20 to 30 cm long) Reliable identification of the Puerto Rican and
within the rosette, and a woody flowering stalk. Caribbean Furcrea species will have to wait for
Mauritius hemp has no taproot, relatively fine further study.
lateral roots, and many fine roots. Its green to
yellow-green leaves are linear-lanceolate to Ecology.—In Puerto Rico, Mauritius hemp grows
oblanceolate, pointed at the tip, and are fleshy with in moist areas that receive from about 1200 to
thread-like parallel fibers. They are up to 25 cm 2500 mm of mean annual precipitation from near
wide, have marginal spines but lack a terminal sea level to 1,000 m in elevation. The species
spine. The inflorescences (panicles) are terminal grows in all types of well-drained soils, including
and contain many pendulous, fragrant, white, poor and eroded soils, and frequently grows on
greenish-white, yellowish-green, or pale blue- rocks, cliffs, and rarely in crotches of trees. It is
green, 2.5 to 3.3 long by 1.0 by 1.8 cm wide resistant to short-term drought and salt spray and
flowers. Flowers open a few at a time for several can survive temperatures as low as -7 to -4 °C.
weeks. Seed development has not been observed Growth is best in full sun, which most often leads
by the author in Puerto Rico and is apparently rare to successful flowering. Mauritius hemp also
elsewhere. Bulbils 1 to 16 cm long develop grows in partial but not heavy shade. When plants
abundantly on the peduncles after flower are young, competition with tall grass, weeds,
dehiscence. There are 2n = 60 chromosomes shrubs, and trees is a critical limiting factor. Cattle
(author’s observation, Bailey 1941, Howard 1979, graze and often completely uproot small plants.
Pacific Island Ecosystems at Risk 2003, Schlegel They are ignored by cattle once they become large
2003). (author’s observation, Desert-Tropicals Nursery
2003, Pacific Island Ecosystems at Risk 2003).
This species, as others of the genus, has
350
crassulacean acid metabolism by which it fixes although not heavily, used as a landscaping plant
CO2 during the night and incorporates it into for accent and curiosity. A variegated form is
carbohydrates during the day (personal available (Desert-Tropicals Nursery 2003).
communication with Ernesto Medina, plant Mauritius hemp was once widely cultivated for
physiologist, Centro de Ecología, Instituto de fiber, hence the common name. Extracts of the
Investigaciones, Caracas, Venezuela). roots are used as ingredients in tonics for purifying
the blood, and the dried leaves are used to control
Reproduction.—Flowering, which may occur at swelling and to aid in wound healing (Núñez-
any time of the year, apparently begins when Meléndez 1982).
plants attain sufficient size and vigor to support the
large flower stalk. Plants die about 1 year after the References
onset of flowering. The flowers are visited by
honey bees (Apis mellifera L.). There is no record Bailey, L.H. 1941. The standard cyclopedia of
in the literature of seed weights. Fresh bulbils (n = horticulture. Vol. 2. The MacMillan Company,
100) collected by the author in Puerto Rico New York. p. 1,201-2,422.
weighed an average of 2.72 + 0.31 g/bulbil with a
range of 0.18 to 16.27 g. Placed in moist potting Desert-Tropicals Nursery. 2003. Mauritius hemp,
mix, 100 percent of them rooted within 1 week. sisal, maguey. http://www.desert-tropicals.com/
Bulbils are formed by the thousands on single Plants/Agavaceae/Furcraea_foetida.html. 2 p.
plants and can form dense thickets. Gravity is the
only known natural means of dispersal. The Grisebach, A.H.R. 1963. Flora of the British West
presence of scattered individuals indicates other Indian Islands. J. Cramer, Weinheim, Germany.
transport, perhaps fruit bats. 789 p.
Growth and Management.—The best way to Howard, R.A. 1979. Flora of the Lesser Antilles,
artificially propagate Mauritius hemp is to place Leeward and Windward Islands. Vol. 3. Arnold
bulbils in pots of potting mix. A group of bulbils Arboretum, Harvard University, Jamaica Plain,
planted in Puerto Rico grow rapidly with no MA. 586 p.
mortality, reached an average of 17.8 + 0.4 cm in
height in 1 month, and were ready to outplant in 1 Liogier, H.A. and L.F. Martorell. 2000. Flora of
to 2 months. New plants need protection from Puerto Rico and adjacent islands: a systematic
weeds for a few months, but afterwards the plants synopsis. 2nd ed. Editorial de la Universidad de
need little care. Individual plants probably live Puerto Rico, Río Piedras, PR. 382 p.
from 5 to 20 years depending on growing
conditions. One owner reported that an ornamental Núñez-Meléndez, E. 1982. Plantas medicinales de
plant died after flowering at 15 years old (author’s Puerto Rico. Editorial de la Universidad de
observation). Mauritius hemp is rarely common Puerto Rico, Río Piedras, PR. 498 p.
enough in its native range to need control. When
control is needed, particularly in invaded areas, Pacific Island Ecosystems at Risk. 2003. Furcraea
individual plants can be killed by grubbing them foetida (L.) Haw., Agavaceae. http;//www.hear.
out or by spray or drizzle application of 2,4-D or org/pier_v3.3/fufoe.htm. 3 p.
triclopyr in oil (Pacific Island Ecosystems at Risk
2003). Schlegel, R. 2003. Plant breeding update: crop
plants II. http://www.desicca.de/plant_breeding/
Benefits.—Mauritius hemp helps hold the soil, Crop_plants/Crops_II/hauptteil_crops_ii.html.
furnishes cover for wildlife, and adds to the [not paged.].
aesthetics of wildlands. The species is widely,
351
Gesneria pedunculosa (DC.) Fritsch arbol de navidad
GESNERIACEAE
Synonyms: Conradia pedunculosa A.P. DC.

Gesneria exogenia Sassé & Moc. ex A.P. DC.
Pentarhaphia pedunculosa (A.P. DC.) Decne.
Codonoraphia albiflora (Decne.) Oersted
John K. Francis
pinkish-white and 1.1 to 1.9 cm long. The fruit that

forms is a four-valved, 6 to 9 mm capsule with five
long, narrow calyx lobes that release numerous tiny
reddish-brown or brown seeds (Liogier 1995, Little
and others 1974).
Range.—Arbol de navidad is endemic to Puerto

Rico (Liogier 1995, Little and others 1974). It has
been planted elsewhere in botanic gardens and
arboretums but is not reported to have naturalized.
A hybrid between arbol de navidad and G.
pedicellaris Alain, dubbed “golden oldie,” has
been developed for ornamental purposes
(American Gloxinia and Gesneriad Society 2002).
Ecology.—Arbol de navidad is locally common

and usually grows on steep slopes and shallow,
rocky soils, sometimes in fractured rock. The
parent materials of the soils may be sedimentary
(including ultramafic) rocks. The species is found
as scattered individuals or in nearly pure thickets.
Plants are less common on south slopes than on
north-, east-, or west-facing slopes in areas that
General Description.—Arbol de navidad is an
receive from about 1000 to 3000 mm of
upright, evergreen shrub or occasionally a small
precipitation from about 50 to 800 m above sea
tree 3 to 5 m in height and 5 to 10 cm in basal
level. Arbol de navidad is moderately shade
diameter. On rocks, the shrub sometimes takes a
tolerant. It grows and flowers in openings and
natural bonsai form. The root system, which may
below forest canopies, if they are not too dense.
or may not have a taproot, takes whatever form
The species does not appear to compete in dense
necessary to anchor itself in the rocky ground
stands of grass. It grows in remnant forest stands
where it grows. The numerous fine roots have
and advanced secondary forests, some of which are
clearly visible, white mycorrhizal growth. There
old road cuts, abandoned pastures, and fields.
are usually multiple stems from the base of the
trunk. Arbol de navidad bark is gray or brown,
Reproduction.—Arbol de navidad flowers and
furrowed and rough. The inner bark is light brown
and fibrous. Twigs are slender, greenish gray, and fruits nearly throughout the year. The flowers are
probably pollinated by bats (Little and others
form a relatively dense crown. The leaves, which
1974). Seed production is high and consistent from
are crowded near the ends of branches, have
elliptic to obovate blades, 4 to 11 cm long, with year to year. Samples of seed collected in Cayey
and Susúa, Puerto Rico, in different years weighed
smooth to fine serrate margins. The inflorescences
an average of 1.78 x 10-5g and 1.75 x 10-5g,
usually have 2 to 4 flowers borne at the end of
respectively, or 56 million seeds/kg. Both
glandular-resinous peduncles 8 to 16.5 cm long.
collections failed to germinate when placed on
The five-lobbed tubular corolla is yellow-green to
moist filter paper. The seeds are dispersed by the
352
wind. Seedlings are relatively common on bare References
ground near fruiting adult plants. Adult and sapling
plants sprout when cut or damaged. American Gloxinia and Gesneriad Society. 2002.
Registered gesneriads. American Gloxinia and
Growth and Management.—Arbol de navidad is Gesneriad Society, Salsbury, CT. http://
slow growing at all stages. It probably lives several www.aggs.org/ir_ges/. 7 p.
decades. No plantations are known, and
recommended management practices have not been Liogier, H.A. 1995. Descriptive flora of Puerto
published. Disturbance of the soil surface in steep, Rico and adjacent islands. Vol. 4. Editorial de la
rocky terrain in moist areas near seed sources Universidad de Puerto Rico, San Juan, PR.
would likely result in seedling establishment. 617 p.
Benefits.—Arbol de navidad is a pretty plant that Little, E.L., Jr., R.O. Woodbury, and F.H.
adds to the aesthetics of the forest. It helps protect Wadsworth. 1974. Trees of Puerto Rico and the
the soil in fragile areas and furnishes cover for Virgin Islands. Vol. 2. Agriculture Handbook
wildlife. Because of the small diameters and 449. U.S. Department of Agriculture,
quantities in which it is available, the wood, which Washington, DC. 1,024 p.
is light brown and hard, is useful mainly for fuel.
353
Gonzalagunia hirsuta (Jacq.) K. Schum. rabo de ratón
RUBIACEAE
Synonyms: Barleria hirsuta (Jacq.) Jacq.

Coccocypselum spicatum (Lam.) HBK
Duggena hirsuta (Jacq.) Britton & Wilson
Duggena richardii West
Duggena spicata (Lam.) Gómez
Gonzalagunia spicata (Lam.) DC.
Gonzalagunia coccocypselum C. & S.
Gonzalia spicata (Lam.) Standl.
Heydyotis secunda Spreng. ex Schultes
Justicia hirsuta Jacq.
Lygistum spicatum Lam.
John K. Francis
areas in advanced secondary forests, and along
roads. In a Puerto Rican study (Myster and Walker
1997), rabo de ratón was the fifth most common
species colonizing recent landslides. Dense
thickets sometimes form. The species grows in
areas receiving from about 1100 to 3400 mm of
annual rainfall and at elevations ranging from near
sea level to more than 600 m. Apparently, all types
of soils are colonized, if not excessively well
drained or very poorly drained. The root system is
shallow, much branched, and extensive.
Reproduction.—Rabo de rató begins flowering

General Description.—Rabo de ratón, also near the end of its first year. The inflorescences,
known as mata de mariposa, palo pelado, yerba which arise from leaf axils, are spiciform-
pelada, bios-foufou, and bois colibri (Howard paniculate and up 56 cm in length. The flowers are
1989, Liogier 1997), is a suberect, medium-sized white with four or five lobes (Liogier 1997).
shrub. It has membranous, opposite, ovate to Flowering and fruiting begin at the base and
lanceolate leaves, 6.5 to 19.0 cm in length and 2.0 progress toward the tip. The flowers are pollinated
to 8.5 cm in width (Howard 1989) with by butterflies and small insects. When the
pronounced, sunken veins. Plants usually have inflorescence is laden with fruit progressing in size
single straight stems but begin branching near the from base to tip, it resembles a mouse’s tail—
ground. The branches and twigs are slender and hence the name, rabo de ratón (Vélez and von
sometimes extend a meter horizontally. Overbeek 1950). An inflorescence may contain
100 or more fruits. The fruits are two-seeded
Range.—The natural range of rabo de ratón fleshy drupes that are white, turning pink, violet,
includes Hispaniola, Puerto Rico, the Lesser or blue (Howard 1989) and are available most of
Antilles, Trinidad, Venezuela, the Guayanas, and the year. The fruits in a Puerto Rican collection
northern Brazil (Howard 1989). The species is not averaged 0.0885 + 0.0419 g and contained seeds
known to have naturalized elsewhere. numbering 182,000/kg. Seeds in this collection
failed to germinate (Francis and Rodríguez 1993).
Ecology.—Rabo de ratón is intolerant and requires Another collection from Puerto Rico had between
disturbance to grow and reproduce. It is most 55 and 68 percent germination in a field test. Of
frequently found in brushy pastures, in early those failing to germinate, about 5 percent were
secondary forests after abandonment of lost to pathogens. There was no significant
cultivation, in tree-fall gaps and other disturbed
354
predation (Myster 1997). The seeds are dispersed Howard, R.A. 1989. Flora of the Lesser Antilles,
by birds (Parrotta 1995). Leeward and Windward Islands. Dicotyledoneae.
Part 3. Vol. 6. Arnold Arboretum, Harvard
Growth and Management.—Rabo de ratón is University, Jamaica Plain, MA. 658 p.
fast growing and may reach 4 m in height and 3
cm in lower stem diameter. Individual plants Liogier, H.A. 1997. Descriptive flora of Puerto
usually live 1 to 2 years (Vélez and von Overbeek Rico and adjacent islands. Vol. 5. Editorial de la
1950). The species can probably be established by Universidad de Puerto Rico, San Juan, PR.
traditional methods (planted seedlings and direct 436 p.
seeding), but no plantations have been reported.
Natural reproduction may be encouraged by Myster, R.W. 1997. Seed predation, disease and
cultivation near seed sources. germination on landslides in neotropical lower
montane wet forest. Journal of Vegetation
Benefits.—Rabo de rató is beneficial to the Science 8: 55-64.
environment in that it readily invades and helps to
stabilize disturbed areas. Because the shrub forms Myster, R.W., and L.R. Walker. 1997. Plant
a thin canopy, it does not inhibit the invasion of successional pathways on Puerto Rican
later successional vegetation. While probably not landslides. Journal of Tropical Ecology 13: 165-
suitable for formal gardens, rabo de rató would 173.
contribute to wildflower gardens, forest parks, and
greenbelts. The species is an abundant producer of Parrotta, J.A. 1995. Influence of overstory
fruits that are eaten by birds. composition on understory colonization by
native species in plantations on a degraded
References tropical site. Journal of Vegetation Science 6:
627-636.
Francis, J.K. and A. Rodríguez. 1993. Seeds of
Puerto Rican trees and shrubs: second Vélez, I., and J. von Overbeek. 1950. Plantas
installment. Research Note SO-374. U.S. indeseables en los cultivos tropicales. Editorial
Department of Agriculture, Forest Service, Universitaria, Río Piedras, PR. 497 p.
Orleans, LA. 5 p.
355
Gossypium hirsutum L. wild cotton
MALVACEAE
Synonyms: Gossypium religiosum L.

Gossypium latifolium Murry
Gossypium punctatum Schum.
Gossypium taitense Parl.
Gossypium mexicanum Tod.
Gossypium marie-galante Watt
John K. Francis
with tightly adhering fuzz (Howard 1989, Liogier

1994, Watt 1907). In certain instances (Little and
others 1974), wild cotton has been misidentified as
Gossypium barbadense L., which occurs today as a
residual from cultivation and as a true wild plant
only on the coast of Ecuador (Izuko Museums of
Cape Town 2000). Wild cotton occurs in at least
three botanical varieties. The most commonly seen
in the West Indies is G. hirsutum var. marie-
galante (Watt) J.B. Hutchinson (Liogier 1994).
Doubtless a legacy of agricultural cultivation, wild
populations in Puerto Rico have a wide range of
leaf and bract shapes, seed characteristics, and lint
length and color (author’s observation). In fact,
wild cotton is a tetraploid. There is a theory that
wild cotton is the result of natural hybridization
between Old World diploid cottons, G. arborium
L. or G. herbacium L. and G. raimondii Ulbrich.
(Bajaj 1998).
General Description.—Wild cotton, also known
as upland cotton, in wild populations is a shrub or Range.—The native range of wild cotton includes
occasionally a small tree to 5 m in height and 15 Tropical Florida, the West Indies, Northern South
cm in stem diameter. The plant is deeply rooted America, Mexico, and Central America (Howard
with a taproot and laterals. Wild cotton usually has 1989, Liogier 1994, Long and Lakela 1976) and
a single trunk, unless disturbed, but may have apparently several Pacific Islands including Samoa
branches arising just above the ground. The bark is (Kohel and Lewis 1984). Because of eradication
gray and smooth, but tough and stringy. The stem intended to control the pink bole worm, the species
wood of older plants is of medium density and has become very rare and has been listed as
brittle. The twigs are relatively stout. The leaves on endangered in Florida (Nelson 1996). Gossypium
young plants are usually without lobes; older plants hirsutum is the principal cultivated cotton
usually have three lobes or occasionally are deeply throughout the world (Vijayakumar 2002). It has
incised with five lobes. The light green leaves have naturalized in many locations, especially in island
petioles 3 to 8 cm long and blades 10 to 15 by 12 to habitats, including Hawaii (Neal 1965).
15 cm. Bracteoles are pointed and may have one to
seven or more teeth. The flowers of wild cotton are Ecology.—Wild cotton is mainly a plant of the
large and showy with cream to pale yellow petals, coastal strand and lower coastal plains. The species
sometimes with a red spot at the base of the petals. has a particular affinity for small islands. It also
The capsules have three to five valves with a grows in disturbed places, particularly along roads
smooth surface and many black gland dots and and on river overflow areas, well inland. Wild
yield white or brown lint with seeds embedded. cotton can grow in almost all types of well-drained
The seeds vary from black and smooth to green soils. Optimum pH’s are said to be from 5.2 to 7.0
356
(Center for New Crops and Plants Products 2002). It has been selected, bred, and, in some cases,
Soil disturbance is usually necessary for its hybridized to give the manageable and productive
establishment. It tolerates moderate amounts of varieties planted today. Evidence from Mexico
salts in the soil and salt spray. Wild cotton shows that wild cotton has been used by man for
competes well with weeds and grasses but does not making string and cloth since at least 200 BC
tolerate shade and cannot grow under tree cover. In (Iziko Museums of Cape Town 2000). Today,
Puerto Rico, wild cotton is most competitive at low cotton is the world’s most widely used natural
rainfalls (750 to 900 mm of annual precipitation) fiber. The seeds of cultivated cotton form the basis
but grows in well-drained areas that receive up to for an important oil and animal food industry. The
1700 mm of precipitation at from near sea level to seeds contain 7.3 g water, 23.1 g protein, 22.9 g
about 300 m elevation. Wild cotton does not fat, 43.2 g total carbohydrate (16.9 g fiber), and 3.5
tolerate frost. g ash per 100 g (Center for New Crops and Plant
Products 2002). Gossypol, a sesquiterpinoid, is
Reproduction.—Wild cotton is dormant during present in vegetative parts of wild cotton as well as
the winter dry season and becomes vegetative as the cultivated varieties. It is a natural deterrent to
the rains begin again in early summer. At the end insects but is toxic to man and nonruminant
of the summer, vegetative growth ceases and plants animals. It has been shown to have strong
begin to flower and fruit (Kohel and Lewis 1984). antifertility, antitumor, antiparasite, and anti-HIV
Three collections of wild cotton seeds of Puerto properties (Bajaj 1989). Gossypol is widely used in
Rico from three locations averaged 0.088 + 0.001, China as a male contraceptive (Center for New
0.075 + 0.001, and 0.081 + 0.001 g/seed or about Crops and Plant Products 2002). Raw cottonseed
12,000 seeds/kg. Germination is epigeal. Seeds oil (containing gossypol) used alone showed
from two of the collections were sown without promise as an insecticide (Tadas and others 1994).
pretreatment in commercial potting mix. One In herbal medicine, cottonseed and roots have been
collection germinated at 100 percent starting in 17 used to treat nasal polyps, asthma, diarrhea,
days and ending in 41 days. The second group hemorrhoids, dysentery, uterine fibroids and
germinated at 67 percent starting in 22 days and certain cancers, to induce abortions, and as a
ending in 140 days. The wind disburses the seeds diuretic. (Center for New Crops and Plant Products
over short distances by blowing the lint until it 2002). Wild cotton is occasionally planted as an
catches on vegetation. Birds also move seeds when ornamental. The species contributes to biodiversity,
they use cotton lint for nest building materials. soil stability, wildlife habitat, and scenic beauty in
Under greenhouse conditions, seedlings may grow its beach strand and lowland habitat.
more than 1 m in 6 months. In wild stands,
seedlings can be abundant but they naturally thin References
themselves quickly.
Bijaj, Y.P.S. 1998. Cotton. In: Y.P.S. Bijaj, ed.
Growth and Management.—Although wild Biotechnology in Agriculture and Forestry 42.
cotton in its cultivated form is managed as an Springer-Verlag, New York. p. 3-36.
annual, it is a true perennial, and if allowed to do
so under favorable conditions, will live for several Center for New Crops and Plant Products. 2002.
years (Kohel and Lewes 1984) and become tall and Gossypium hirsutum L. Purdue University.
woody. Wild cotton in Puerto Rico grows 1 to 1.5 http://www.hort.perdue.edu/newcrop/duke_ener
m in its first year. The wild variety requires 15 or gy/Gossypium_hirsutum.html. 6 p.
more nodes to be formed along the main axis
before it can flower. Consequently, the plant is Howard, R.A. 1989. Flora of the Lesser Antilles,
rarely able to bloom during its first year (Kohel and Leeward and Windward Islands. Vol. 5. Arnold
Lewis 1984). Cotton can be transplanted using Arboretum, Harvard University, Jamaica Plain,
containerized nursery plants. However, direct MA. 604 p.
seeding following removal of overhead
competition and cultivation of the soil is Iziko Museums of Cape Town. 2000. Gossypium
recommended. Soil temperatures must be at least (cotton). Biodiversity explorer. http://www.
18 °C before planting (Center for New Crops and museums.org.za/bio/plants/malvaceae/gossypiu
Plant Products 2002). m.htm. 2 p.
Benefits.—About 87 percent of the cotton grown Kohel, R.J. and C.F. Lewis, eds. 1984. Cotton.
commercially is G. hirsutum (Vijayakumar 2002). Agronomy 24. American Society of Agronomy,
357
Inc., Crop Science Society of America, Inc., and Neal, M.C. 1965. In gardens of Hawaii. Spec. Pub.
Soil Science Society of America, Inc., Madison, 50. Bernice P. Bishop Museum Press, Honolulu,
WI. 605 p. HI. 924 p.
Liogier, H.A. 1994. Descriptive flora of Puerto Nelson, G. 1996. The shrubs and woody vines of
Rico and adjacent islands. Vol. 3. Editorial de la Florida. Pineapple Press, Sarasota, FL. 391 p.
461 p. Tadas, P.L., H.K. Kene, and S.D. Deshmukh.
1994. Effect of raw cottonseed oil against
Little, E.L., Jr., R.O. Woodbury, and F.H. sucking pests of cotton. PKV Research Journal
Wadsworth. 1974. Trees of Puerto Rico and the 18(1): 142-143.
449. U.S. Department of Agriculture, Vijayakumar, T. 2002. Cotton. http://www.
Washington, DC. 1,024 p. geocities.com/vijayakumar777/cotton1.html.
14 p.
Tropical Florida. Banyon Books. Miami, FL. Watt, Sir G. 1907. The wild and cultivated cotton
962 p. plants of the world. Longmans, Green, and Co.,
London. 406 p.
358
Gouania lupuloides (L.) Urban chew-stick
RHAMNACEAE
Synonyms: Banisteria lupuloides L.

Rhamnus domingensis Jacq.
Gouania domingensis (Jacq.) L.
Gouania glabra Jacq.
Lupulus lupuloides Kuntze var. domingensis (Jacq.) Kuntze
John K. Francis
elsewhere.
Ecology.—In Puerto Rico, Chew-stick occurs in

forests that receive from 750 to over 2000 mm of
precipitation. It grows on soils with the entire range
of textures, both poor and rich in nutrients,
excessively well drained to somewhat poorly
drained over sedimentary (including limestone),
igneous, and metamorphic (including ultramafic)
rocks. The species is found from near sea level to
elevations of 1,300 m (Acevedo-Rodríguez 1985,
Stevens and others 2001). Chew-stick has an
intermediate tolerance of shade. It can live and
grow slowly in all but dark forest understories but
requires increased sunlight to flower and fruit. The
General Description.—Chew-stick, also known as species grows on roadsides, in brushy pastures, and
toothbrush tree, bejuco de sopla, bejuco de indio, in secondary and remnant forests. In Florida, chew-
mascapalo, bejuco de fuego, liane-savon, and stick grows in mangroves and coastal hammocks
lyenn-savon, is a scrambling and climbing woody (Nelson 1996).
vine that may reach 7 to 12 m of extension. The
plant is supported by an extensive system of Reproduction.—In Puerto Rico, chew-stick
medium and fine, white, lateral roots. The stem and blooms from August to March, and fruit matures
branches are relatively slender and support from November to March (Acevedo-Rodríguez
themselves by means of coiled tendrils. The 1985). In Panama, it flowers in the early dry season
alternate leaves have petioles 0.5 to 2 cm long and (November to March), and the seeds are dispersed
thin, ovate to elliptic blades 4 to 10 cm long with a from February to May (Croat 1978). Fruits
serrate or wavy-toothed margin and a pointed tip. collected in Puerto Rico averaged 0.0374 + 0.0005
Tiny white, yellow, or greenish flowers are borne g/fruit. Seeds separated from them averaged 0.0052
on slender axillary or terminal racemes. Few-to- + 0.0001 g/seed or 192,000 seeds/kg. Seed should
many, 6- to 13-mm, gray, three-winged fruits be collected by hand from fruiting vines. Hand
(schizocarps splitting into three mericarps) develop separation is practical for small lots; larger lots
later in the season. Each mericarp contains a 3- to could be pulverized and winnowed. Fifty-seven
4-mm dark-brown seed (Acevedo-Rodríguez 1985, percent of untreated seed from the previously
Croat 1978, Howard 1989, Liogier 1994, Stevens mentioned collection germinated between 16 and
and others 2001). 75 days after sowing on commercial potting mix
(author’s observation). Seed is dispersed by its
Range.—The native range of chew-stick includes lateral tumbling action in still air or wind and by
Florida, the West Indies, Mexico, Central America, lateral extension on the vines. Seedlings and young
and South America into Brazil and Colombia plants are relatively common in Puerto Rico. The
(Howard 1989, New York Botanical Garden 2002, plant sprouts after fire and cutting.
Pérez-Arbelaez 1978, Secritaria de Medio
Ambiente y Recursos Naturals 2002). It is not Growth and Management.—Once established,
359
chew-stick grows relatively rapidly. In the presence Kennelly, E.J., W.H. Lewis, R.E.K. Winter, S.
of seed sources, reproduction is usually adequate. Johnson, M. Elvin-Lewis, and J. Gossling. 1993.
Thinning the overstory would probably allow Triterpenoid saponins from Gouania lupuloides.
existing suppressed plants to grow to larger sizes Journal of Natural Products 56(3): 402-410.
and fruit.
Benefits.—Chew-stick contributes to biodiversity Rico and adjacent Islands. Vol. 3. Editorial de la
and biomass accumulation, helps protect the soil, Universidad de Puerto Rico. Río Piedras, PR.
and furnishes wildlife cover in forests where it 461 p.
grows. It is an important honey plant (Townsend
1984). In Jamaica and the West Indies, stem Nelson, G. 1996. The shrubs and woody vines of
cuttings are used to clean teeth and gums and Florida. Pineapple Press, Inc., Sarasota, FL.
extracts are used to manufacture a dentifrice 391 p.
(Secretaría de Medio Ambiente y Recursos
Naturales 2002). Triterpenoid saponins may New York Botanical Garden. 2002. Flora and
contribute to the beneficial activity (Kennelly and mycota of Acre, Brazil. http://nybg.org/
others 1993). In Colombia, the plant is used for a bsci/acre/rhamnaceae.html. [not paged].
bitter flavoring in alcoholic beverages and to treat
water retention and stomach problems (Pérez- Pérez-Arbelaez, E. 1978. Plantas útiles de
Arbelaez 1978). Colombia. Litografía Arco, Bogotá, Colombia.
831 p.
References
Secretaría de Medio Ambiente y Recursos
Acevedo-Rodríguez, P. 1985. Los bejucos de Naturales. 2002. Especies con usos no
Puerto Rico. Vol. 1. General Technical Report maderables en bosques tropicales y
SO-58. U.S. Department of Agriculture, Forest subtropicales. http://www.semarnat.gob.mx/
Service, Southern Forest Experiment Station, pfnm2/fichas/gouania_lupuloides.htm. 2 p.
Stevens, W.D., C. Ulloa-Ulloa, A. Pool, O.M.
Croat, T.B. 1978. Flora of Barro Colorado Island. Montiel, eds. 2001. Flora de Nicaragua.
Stanford University Press, Stanford, CA. 943 p. Monographs in Systematic Botany Vol. 85, No.
3. Missouri Botanic Garden Press, St. Louis,
Howard, R.A. 1989. Flora of the Lesser Antilles, MO. 2,666 p.
Arboretum, Harvard University, Jamaica Plain, Townsend, G.F. 1984. Multi-purpose trees for
MA. 604 p. honey production. Echo technical note.
http://www.echonet.org/tropicalag/technotes/Mu
ltiPur.pdf. 3 p.
360
Grayia spinosa (Hook.) Moq. spiny hopsage
CHENOPODIACEAE
Synonyms: Atriplex spinosa Collotzi

Atriplex grayia Collotzi
Grayia polygaloides Hook. & Arn.
Nancy L. Shaw, Marshall R. Haferkamp, and Emerenciana G. Hurd
flower consists of a pistil enclosed in two

obcompressed chordate bracteoles united along
their margins except for a minute apical opening.
Bracteoles enlarge at maturity to form a papery
sac, 9 to 15 mm in diameter with a dorsally
winged margin. Fruits are utricles with the papery
pericarp free from the seed. Seeds are 1 to 2 mm in
diameter. The testa is comprised of a thin, dark
brown outer layer and an elastic inner layer. A
well-developed embryo with an inferior radicle
encircles the perisperm (Hitchcock and others
1964, Shaw and others 1996, 2001).
Range.—Spiny hopsage is widely distributed in

the interior Western United States. It occurs east of
the Cascade and Sierra Nevada Mountains from
central Washington to southern California and
eastward from southern Montana to western
Colorado and northern Arizona (Hitchcock and
others 1964, Welsh and others 1987).
Ecology.—Spiny hopsage occurs at elevations

General Description.—Alternative common from 160 to 2,130 m in Wyoming big sagebrush
names for spiny hopsage are grayia, Gray’s (Artemisia tridentata Nutt. ssp. wyomingensis
saltbush, saltbrush, spiny-sage, horsebrush, and Beetle & A. Young), salt desert shrub, Mohave
applebush (Shaw and others 2001, Welsh and Desert, and pinyon-juniper communities receiving
others 1987). Shrubs are 0.3 to 1.2 (1.5) m tall, 125 to 300 mm of annual precipitation (Welsh and
summer deciduous, and erect to rounded with others 1987). It grows on soils that are silty to
divergent, thorn-tipped branches and gray to sandy, frequently high in calcium, and neutral to
brownish exfoliating bark. Young twigs and strongly basic. It also occurs on sand dunes. Spiny
herbage are scurfy to pubescent, becoming hopsage generally occurs in small patches or as
glabrous with age. Alternate, entire, gray-green, scattered plants. Extensive stands are rare. Plants
fleshy leaves, 5 to 30 (40) mm long and 2 to 13 initiate growth in early spring and rapidly
mm wide, develop from prominent, globose, complete their vegetative and reproductive cycles
axillary buds. Blades are linear-oblanceolate with prior to entering summer dormancy. They are most
sessile or short-petiolate bases. The species is tolerant of summer wildfires after summer leaf fall
tetraploid (4x = 36) (McArthur and Sanderson and can resprout from surviving root crowns
1984). Inflorescences develop on floral shoots that (Daubenmire 1970). Regeneration is episodic.
die back following fruit dispersal. Flowers are Germination occurs in early spring. Most seedlings
inconspicuous. Staminate flowers develop in establish beneath canopies of spiny hopsage or
glomerate spikes and consist of a four or five other shrubs. Growth of cheatgrass (Bromus
lobed perianth and four or five stamens. Pistillate tectorum L.), a highly competitive annual,
flowers are produced on dense terminal spicate however, is favored by nutrient accumulation
inflorescences. Some flowers are vestigial. Each beneath spiny hopsage canopies. This can interfere
361
with natural regeneration of native vegetation and 163. U.S. Department of Agriculture, Forest
result in increased fine fuel accumulation. Service, Intermountain Research Station, Ogden,
UT. 52 p.
Reproduction.—Most plants are dioecious, but
the proportion varies among populations. Flowers
Daubenmire, R. 1970. Steppe vegetation of
appear in February to May and are wind
Washington. Technical Bulletin 62. Washington
pollinated. Fruits ripen in March to June (Shaw
State University, Pullman, WA. 131 p.
and others 2001) and are dispersed by wind,
gravity, and insects. Herbage, flower, and fruit
Hitchcock, C.L., A. Cronquist, M. Ownbey, and
production are highly variable among years
J.W. Thompson. 1964. Vascular plants of the
(Rickard and Warren 1981); many plants fail to
Pacific Northwest. Part 2. Flora of the Pacific
produce flowers in dry years (McArthur and
Northwest. University of Washington Press,
Sanderson 1984). During a drought, spiny hopsage
Seattle, WA. 730 p.
in a southern Idaho wildland planting began
flowering 4 years after seeding. Incubating
debracted utricles at 15 and 5 oC (8 hrs, 16 hrs) or McArthur, E.D. and S.C. Sanderson. 1984.
at 15 oC for 14 days provides an estimate of Distribution, systematics, and evolution of the
germination. A 30 to 60 day prechill at 3 to 5 oC is Chenopodiaceae, In: Proceedings—symposium
required to enhance germination of utricles from on the biology of Atriplex and related
northern populations. Shaw and others (2001) chenopods. General Technical Report INT-172.
reported an average of 1,030,600 debracted U.S. Department of Agriculture, Forest Service,
utricles/kg. Germination is epigeal. Intermountain Research Station, Ogden, UT. p.
14-24.
Growth and Management.—Seedlings develop
rapidly if adequate water is available. Mature Rickard, W.H. and R.F. Keough. 1968. Soil-plant
plants are drought tolerant. Spiny hopsage is relationships of two steppe desert shrubs. Plant
considered one of the most palatable shrubs in salt and Soil. 19:205-212.
desert shrub communities in spring and early
summer. Its use by wildlife and livestock declines
rapidly as plants enter summer dormancy. Rickard, W.H. and J.L. Warren. 1981. Response of
steppe shrubs to the 1977 drought. Northwest
Although fairly tolerant of grazing, some
populations receiving heavy use have been Science. 55:108-112.
replaced by less palatable species (Blaisdell and
Holmgren 1984). Shaw, N.L., M.R. Haferkamp, and E.G. Hurd.
2001. Grayia Hook. & Arn. hopsage. In: F.T.
Bonner, and R.G. Nisley, eds. Woody Plant
Benefits.—Spiny hopsage contributes to diversity
on native sites that generally support few other Seed Manual. U.S. Department of Agriculture,
woody species. It provides cover for birds and Forest Service, Washington, D.C.
http://wpsm.net/ 12 p.
other small animals, spring and early summer
forage for big game and livestock, and soil
stabilization on moderate slopes. Litter rich in Shaw, N.L., E.G. Hurd, and M.R. Haferkamp.
1996. Spiny hopsage fruit and seed morphology.
potassium and other accumulated cations
enhances growth and nutrient content of plants Journal of Range Management. 49:551-553.
growing beneath its canopy (Rickard and Keough
1968). Native Americans ground parched seed of Stubbendieck, J., S.L. Hatch, and K. Hirsch. 1986.
spiny hopsage to prepare pinole flour North American range plants. University of
(Stubbendieck and others 1986). Nebraska Press, Lincoln. 465 p.
References Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.

Higgins, eds. 1987. A Utah flora. Great Basin
Blaisdell, J.P. and R.C. Holmgren. 1984.
Managing Intermountian rangelands—salt desert University, Provo, UT. 894 p.
shrub ranges. General Technical Report INT-
362
Guettarda elliptica Sw. velvetseed
RUBIACEAE
Synonyms: Guettarda bladgettii Shuttlew. ex Chapm.

Matthiola elliptica (Sw.) Kuntze
John K. Francis
Range.—The native range of velvetseed extends

from southern Florida through the West Indies,
from Mexico through Central America, and
Venezuela and Guyana in South America (Little
and others 1974).
Ecology.—Velvetseed grows in pinelands and

hammocks in Florida (Nelson 1996). In Puerto
Rico, it is most common in limestone areas and
occasionally in areas of igneous rock, on rocky
ridge sites too difficult for tall vegetation, and in
coastal thickets and rocky hills (Liogier 1997,
Little and others 1974). Annual rainfall varies from
750 mm to 2200 mm, and elevations range form
near sea level to about 400 m. Vegetation sites
include relatively pristine to highly disturbed
remnant forests and late secondary forests.
Velvetseed is moderately intolerant of shade. It
competes well with slow-growing vegetation but
not with tall grass or overtopping trees. Open-
grown plants assume a symmetrical shape, but in
thickets, the stems are usually crooked and the
General Description.—Velvetseed, also known as
crowns asymmetrical.
Everglade velvetseed, pickle-wood, cucubano liso,
cigüillo, cuero de sabana crucillo, guayabillo
Reproduction.—Velvetseed blooms in summer in
prieto, cabrito, and punteral, is a shrub or small tree
Florida (West and Arnold 1952) and in the spring
to 6 m in height and 10 cm in stem diameter. It
in Puerto Rico (Little and others 1974). Flowers
usually has multiple stems from below or just
require pollination to develop seeds and are
above the ground level. The stem bark is gray or
ordinarily pollinated by insects (Koptur and
gray-brown, smooth with light colored dots or
Richards 1996). A collection of fruits in Puerto
lines. The stem wood is light brown, hard, fine-
Rico averaged 0.350 + 0.008 g/fruit. Seeds
textured, and heavy with a specific gravity of 0.83
separated from them weighed (air-dried) an
(Little and others 1974). The plant is supported by
average of 0.0626 + 0.0015 g/seed or 16,000
hard, brittle, dark-brown roots. Velvetseed tends to
seeds/kg. Sown on commercial potting mix, 68
form branchy crowns of opposite, moderately fine
percent of the seeds germinated between 3 and 7
branches and twigs. The opposite leaves have
months after sowing (author’s observation).
short, hairy petioles and oval to elliptic blades 3 by
Although there is usually good fruit and seed
7.5 cm long. Axillary, few-flowered cymes bear
production, seedlings are relatively uncommon.
small tubular yellowish-white flowers. Three to 4
Birds disperse the seeds. Velvetseed sprouts readily
months later, globose, 4- to 8-mm drupes with a
when cut.
persistent calyx develop. They turn red to black at
maturity and contain two to four seeds enclosed in
Growth and Management.—The Florida
a bony shell (Liogier 1997, Little and others 1974,
champion velvetseed is 7 m in height and 29 cm in
Nelson 1996, West and Arnold 1952). The fruits
diameter at breast height (Champion Tree Project
have a slightly sweet, slightly sour, and slightly
2002). The species grows slowly, at least in the
astringent flavor (author’s observation).
363
difficult habitat where most occur. Individuals smooth- and rough-leaved velvetseed,
probably live from two to several decades. No Guettarda elliptica Sw. and G. scabra in the
management experience has been published. Everglades. American Journal of Botany
However, long-term protection of velvetseed sites 83(supplement): 70.
is recommended.
Benefits.—Velvetseed contributes to biodiversity Rico and adjacent islands. Vol. 5. Editorial de
and wildland biomass while it helps protects the la Universidad de Puerto Rico, San Juan, PR.
soil from erosion and provides food and cover for 436 p.
wildlife. The species is listed as the larval food
plant for the moths, Calidota strigosa Walker in Little, E.L., Jr., R.O. Woodbury, and F.H.
Florida and the West Indies (Barns 2002) and Wadsworth. 1974. Trees of Puerto Rico and the
Hylesia lineata Druce in Mexico (Pescador 1995). Virgin Islands. Vol. 2. Agriculture Handbook
The wood is useful for fuel and stakes. Velvetseed 449. U.S. Department of Agriculture,
is a pretty shrub and would make a fine foundation Washington, DC. 1,024 p.
plant in seminatural landscaping.
References Florida. Pineapple Press, Inc., Sarasota, FL.
390 p.
Barns, M.J.C. 2002. Moths of Jamaica, Calidota
strigosa. http://members.fortunecity.co.uk/ Pescador, A.R. 1995. Distribution and abundance
jamaicamoth/images/calstr.htm. 1 p. of Hylesia lineata egg masses in a tropical dry
forest in western Mexico. Southwestern
Champion Tree Project. 2002. National Champion Entomologist 20(3): 367-375.
Trees, Florida. http://www.championtrees.org/
database/championsFL.htm. 8 p. West, E. and L.E. Arnold. 1952. The native trees
of Florida. University of Florida Press,
Koptur, S. and J.H. Richards. 1996. Comparative Gainesville, FL. 212 p.
floral biology and breeding systems of
364
Gundlachia corymbosa (Urban) Britt. ex Boldingh yambush
ASTERACEAE
Synonyms: Baccharis dioica sensu Duss

Gundlachia domingensis var. corymbosa Urban
John K. Francis
Bonaire (Howard 1989, Liogier 1997). It is not

elsewhere.
Ecology.—Seaside and small island habitat is

normal for the species. Yambush mostly grows on
shoreline rocks and cliffs in Puerto Rico. It was
collected between rocks and gravel deposited at 40
to 50 m elevation in La Désirade (New York
Botanical Garden 2003) and on calcareous silt
with emergent limestone on Turks and Caicos
(Darwin Institute 2002), and on rocky shores on
San Salvador, Bahamas (Gerace and others 2003).
Britton and Millspaugh (1962) indicate that it
grows on sand dunes, pine lands, and borders of
saline marshes. Yambush can grow in a wide
variety of soils, gravel, and even in cracks in
rocks. It is tolerant of salty soils, heavy salt spray,
sea water overwash, and constant, sometimes high
winds. Mean annual precipitation in its natural
habitat varies from about 700 mm to 1,700 mm. It
grows from near sea level to over 50 m in
elevation. The species can grow with light
General Description.—Yambush, also known as overhead shade but generally grows where few
horsebush, broombush, soldier’s-bush, Jamaican other species can survive to avoid competition.
trash, sereno, and bois-Guillaume, is an evergreen
shrub to 1.5 m in height. Stems are typically Reproduction.—Yambush probably blooms
slender, numerous, branched near the ground, and throughout the year. The seeds are small, produced
rooted at intervals where they touch the ground. in great numbers, and dispersed by the wind.
The plant is supported by brown lateral and fine Seedlings are uncommon, but once established in
roots that are flexible and weak. The leaves are favorable environments, they layer profusely and
mostly clustered near the twig ends. The subsessile form interconnected mats.
leaves are fleshy, sticky, oblanceolate to oblong-
lanceolate, mostly rounded at the apex and 3 to 8 Growth and Management.—Yambush grows
cm long by 0.5 to 2 cm wide. The terminal slowly. In seaside environments in Puerto Rico, it
inflorescences are dense corymbs of heads on adds about 20 cm per year to its multiple stems.
short peduncles. The involucre is about 5 mm long Individual plants and clumps apparently live for
and surrounds five to nine white florets. Fruits are many years. The author knows of no reports of
oblong, angular achenes with tawny pappus planting or management of the species. Because it
(Britton and Millspaugh 1962, Howard 1989, is uncommon in much of its range, not aggressive,
Liogier 1997). and grows in areas not normally used for
agriculture or forestry, control of stands of the
Range.—Yambush is native to the Bahamas, species should rarely be necessary.
Cuba, Hispaniola, Puerto Rico, the Virgin Islands,
Barbuda, Saba, Monserrat, Guadeloupe, La Benefits.—Yambush adds to the aesthetics of sites
Désirade (Guadaloupe), Curaçao, Aruba, and where it grows, helps protect the soil, and
365
furnishes cover for wildlife. Although there is no http://www.unesco.org/csi/pub/papers/gerace.
record of it having been used for ornamental htm. 20 p.
purposes, yambush seems an excellent choice for
groundcovers and borders in seaside and xeric Howard, R.A. 1989. Flora of the Lesser Antilles,
gardens. Poultices of crushed leaves relieve and Leeward and Windward Islands. Vol. 6. Arnold
help heal allergic dermatitis (Wood 2003). Arboretum, Harvard University, Jamaica Plain,
MA. 658 p.
References
Britton, N.L. and C.F. Millspaugh. 1962. The Rico and adjacent islands. Vol. 5. Editorial de la
Bahama flora. The New York Botanical Garden, Universidad de Puerto Rico, Río Piedras, PR.
New York. 695 p. 436 p.
Darwin Institute. 2002. Plan for biodiversity New York Botanical Garden. 2003. Specimens
management and sustainable development detailed results: Gundlachia domingensis A.
around Turks and Caicos Ramsar site: Gray var. corymbosa Urb. (syntype).
introduction. http://www.ukotcf.org/pdf/ http://scisun.nybg.org:8890/searchdb/owa/www
TCIRamMP/Parts%20One%20and%Two.pdf. catalog.detail_list. 1 p.
48 p.
Wood, K.M. 2003. Medicinal plants for what ails
Gerace, D.T., G.K. Ostrander, and G.W. Smith. you in Paradise. http://www.timespub.tc/
2003. San Salvador, Bahamas. Coastal region Natural%20History/Archive/Winter%20200102/
and small island papers 3. UNESCO, Rome. medicinal.htm. 2 p.
366
Gutierrezia sarothrae (Pursh) Britt. & Rusby broom snakeweed
ASTERACEAE
Synonyms: Xanthocephalum sarothrae (Pursh) Shinners

Solidago sarothrae Pursh
Gutierrezia diversifolia Greene
Gutierrezia linearis Rydb.
snakeweed is generally 20 to 60 cm tall with a

dense crown. The young, greenish stems turn tan
and then brown with age. It has linear green leaves
up to 3.8 cm long and numerous small, bright-
yellow flowers with two to seven rays that are 2 to
3 mm long. In its northern range it is deciduous
and dies back, but within its southern range it
commonly only dies back part way in winter
(Cronquist 1994, Whitson et al. 1991). Snakeweed
contains saponin, a toxic secondary product
(Whitson et al. 1991). Details of the plant may be
viewed on the Texas A&M University “Virtual
Herbarium” Website (2000).
Range.—Broom snakeweed is native to the

Americas. It is a very common shrub throughout
Western North America, from Saskatchewan and
Alberta, Canada south through western Nebraska
and Kansas into Texas and central Mexico, and
west to Washington, Oregon, California, and Baja
California. Broom snakeweed has been described
as a diagnostic climax member of desert
grasslands (Dick-Peddie 1993). It is a member of
both Chihuahuan and Sonoran desert grasslands
(Schmutz et al. 1992). It has been estimated that it
covers more acres than any other perennial plant
on Navajo Tribal lands in the Southwestern United
States (Mayes and Lacy 1994).
Illustration source: USDA-Forest Service collection,
Hunt Institute
Ecology.—Broom snakeweed grows in dry, open
places from 732 m to 2,898 m (Tirmenstein 1999).
General Description.—Gutierrezia sarothrae is a
As its wide range suggests, it grows in a variety of
woody perennial named for Pedro Gutierrez, a
soil conditions, from sandy and well-drained loam
Spanish nobleman. The epithet “sarothrae” is a
to heavy clay soils and in various plant
derivative of a Latin word meaning “broom.” Its
communities, including Douglas-fir and ponderosa
most popular common name is broom snakeweed,
pine forests, pinyon-juniper woodlands, sagebrush
or snakeweed, but is also referred to as common
shrublands, and desert grasslands. Broom
matchbrush, broomweed, brownweed, yellow-top,
snakeweed root systems are fairly shallow, which
yellow weed, resin-weed, turpentine-weed and,
is unusual for a shrub adapted to semi arid and arid
principally in Mexico, yerba de vibora (Dayton
environments (Schmutz and others 1992). They
1931). It has also many botanical synonyms and
occupy the same soil layers as those of perennial
for a complete list see Kartesz (1994). The Navajo
grasses and sometimes appear to compete with
Native Americans call it Ch’il diilyesiitoh, “big
them (Burgess 1995). Broom snakeweed tends not
dodge weed” (Mayes and Lacy 1994). Broom
to resprout after fire and frequently dies during
367
drought (Schmutz et al. 1992). It suffers from dryness, and an absence of grazing for at least one
drought to about the same extent as black grama season prior to burning has been recommended
grass, but when insect pressure is present the (Tirmenstein 1999). In many cases, fire may
snakeweed mortality is much higher (Dick-Peddie prove an ineffectual control. For example, 5 years
1993). However, by virtue of its high seed after a wildfire snakeweed had gone from 0
production and rapid seedling establishment percent canopy cover to over 40 percent canopy
(Osman and Pieper 1988) it often increases, or cover on monitored plots in Arizona (Pase and
“comes in”, after prolonged drought and other Pond 1964). In west-central Utah broom
disturbances (Dayton 1931). snakeweed was found on eight out of the nine age-
classes of burns, being absent on the 71-year-old
Reproduction.—Snakeweed flowers from July to burns and most abundant on the 22-year-old burns
December depending upon the latitude and water (Barney and Frischknecht 1974). Broom
availability. The plant is insect pollinated snakeweed is susceptible to some herbicides
(Mayeux and Leotta 1981). It is a prodigious (Whitson et al. 1991).
producer of seed that can remain viable in the soil
for a considerable length of time (Osman and Benefits.—Broom snakeweed provides some
Pieper 1988). A single plant is capable of stability to the soil against both wind and water
producing 9,000 to 10,000 seeds annually erosion (Tirmenstein 1999, USDAFS 1988). It is
(Tirmenstein 1999). The oval fruit (seeds) are fair browse for wildlife but is poisonous to
achenes and covered with stiff hairs or nearly livestock due to the saponin content, which is
glabrous. Chaffy scales, called the pappus, are on variable and depends upon both growth phase and
one end of the achene and aid seed dispersal by environmental conditions (Hatch and Pluhar
wind (Cronquist, 1994). Seeds are dispersed by 1993). The vegetation is most toxic during leaf
animals as well as wind, although it has been formation (Whitson and others 1991). At times
reported that they also accumulate under the shrub when there has been no alternative forage,
canopy (Mayeux and Leotta 1981). Light enhances snakeweed has been used by livestock with no ill
germination, which suggests that the best position effects (USDAFS 1988). Presumably saponin
for seeds is on or near the soil surface (Mayeux levels have been low at these times. It has several
and Leotta 1981). medicinal uses among the Navajo (Mayes and
Lacy 1994). It is used to heal cuts and bites in
Growth and Management.—Snakeweed is not humans and animals. A snakeweed medicine is
palatable and can be poisonous to livestock and given in childbirth and is used to treat
increases under grazing pressure (Cable 1975; stomachaches and other internal disorders. It is
Carter 1988; Cronquist 1994; Pase and Pond reported to be a powerful diuretic (Powell 1998). It
1964). Most losses in sheep and cattle are due to is also used in various Navajo ceremonies
abortions (Whitson et al. 1991), but the plant also including as an emetic and fumigent (Mayes and
apparently affects the kidneys of both sheep and Lacy 1994). A yellow dye is made from the
horses (Dayton 1931). Data suggest that the flowers, and the stems can be used to make fire by
presence of jackrabbits may reduce cattle losses friction. Birds, for example quail, eat the seeds
from poisonous plants because, under moderate (Epple 1995). Broom snakeweed also provides
stocking rates, jackrabbits consume increased cover and resting sites for species of small
amounts of poisonous plants and shrubs such as mammals, for example jackrabbits (Daniel and
snakeweed but have minimal forage competition others 1993) and birds, for example lesser prairie
with cattle (Daniel et al. 1993). When abundant, chicken (USDAFS 1988) and Columbia basin
snakeweed is generally considered an indicator of burrowing owl (Green and Anthony 1989).
overgrazing (Dayton 1931; Hatch and Pluhar
1993; Powell, 1998). Pastures are also not References
properly utilized when broom snakeweed is
present amongst the grasses (Whitson et al. 1991). Barney, M.A. and N.C. Frischknecht. 1974.
Snakeweed is killed by certain borers and other Vegetation changes following fire in the pinyon-
insects. Species from the genera, Crossidius, juniper type of west-central Utah. Journal of
Diplotaxis, and Mecas have been proposed as Range Management 27(2): 91-96.
possible biological controls (Dayton 1931). Fire
can be used to reduce snakeweed (Humphrey Burgess, T.L. 1995. The dilemma of coexisting
1949; Tirmenstein 1999). However, the success of growth forms. In: M.P. McClaran and T.R. Van
controlled burns depends upon fuel quantity and
368
Devender, eds. The desert grassland. The Greenland. Vol. 1 – Checklist. 2nd ed. Timber
University of Arizona Press, Tucson, AZ. 346 p. Press, Portland, OR. 622 p.
Cable, D.R. 1975. Range management in the Mayes, V.O. and B.B. Lacy, 1994. Nanise’ Navajo
chaparral type and its ecological basis: the status herbal. Navajo Community College Press,
of our knowledge. Research Paper RM-155. Tsaile, AZ. 153 p.
Rocky Mountain Forest and Range Experiment Mayeux, H.S., Jr. and L. Leotta. 1981. Germination
Station. Fort Collins, CO. 30 p. of broom snakeweed (Gutierrezia sarothrae) and
threadleaf snakeweed (G. microcephalum) seed.
Carter, J. L. 1988. Trees and Shrubs of Colorado. Weed Science. 29: 530-534.
Johnson Books, Boulder, CO. 165 p.
Osman, A. and Pieper, R.D. 1988. Growth of
Cronquist, A. 1994. Intermountain Flora, Vascular Gutierrezia sarothrae seedlings in the field.
Plants of the Intermountain West, USA. Vol. 5. Journal of Range Management 41(1): 92-93.
Asterales. New York Botanical Garden, Bronx,
NY. 496 p. Pase C.P. and F.W. Pond. 1964. Vegetation
changes following the Mingus Mountain burn.
Daniel, A., J. L. Holechek, R. Valdez, A. Tem, L. Note RM-18. U.S. Department of Agriculture,
Saiwana, M Rusco, and M. Carden. 1993. Range Forest Service, Rocky Mountain Research
condition influences on Chihuahuan Desert Station, Fort Collins, CO. 8 p.
cattle and jackrabbit diets. Journal of Range
Management 46(4): 296-301. Powell, A. M. 1998. Trees & shrubs of Trans-
Pecos and adjacent areas. Big Bend Natural
Dayton, W. A. 1931. Important western browse History Association. (Reprinted by University of
plants. Miscellaneous Publication 101. U.S. Texas University Press, Austin, TX.) 498 p.
214 p. Schmutz E.M., E.L Smith, P.R. Ogden, M.L. Cox,
J.O. Klemmedson, J.J. Norris, and L.C. Fierro.
Dick-Peddie, W.A. 1993. New Mexico Vegetation 1992. Desert Grasslands. In: R.T. Coupland, ed.
– past, present, and future. University of New Ecosystems of the World, 8A; Natural
Mexico Press, Albuquerque, NM. 244 p. Grasslands Introduction and Western
Hemisphere. Elsevier, London, NY. p. 337-362
Epple, A.O. 1995. A Field Guide to the Plants of
Arizona. Falcon Press Publishing Co., Helena, Texas A&M University “Virtual Herbarium”.
Montana. 347 p. Uvalde Research and Extension Center. 2000.
Copyright Texas A&M University System.
Green, G.A. and R.G Anthony. 1989. Nesting http://uvalde.tamu.edu/herbarium/gusa.htm [not
success and habitat relationships of burrowing paged].
owls in the Columbia Basin, Oregon. The
Condor. 91: 347-354. Tirmenstein, D. 1999. USDA Forest Service,
Hatch, S.L. and J. Pluhar. 1993. Texas range Laboratory. Fire Effects Information System,
plants. The W.L. Moody Jr Natural History http://www.fs.fed.us/database/ [not paged].
Series 123. Texas A&M University Press,
College Station, TX. 326 p. Whitson, T.D., ed., L.C. Burrill, S.A. Dewey, D.W.
Cudney, B.E. Nelson, R.D. Lee, and R. Parker.
Humphrey, R.R. 1949. Fire as a means of 1991. Weeds of the West. The Western Society
controlling velvet mesquite, burroweed, and of Weed Science, Western U.S. Land Grant
cholla on southern Arizona ranges. Journal of Universities Cooperative Extension Services and
Range Management. 2: 175-182. the University of Wyoming, Pioneer of Jackson
Hole, Jackson WY. p 132-133.
vascular flora of the United States, Canada, and United States Department of Agriculture, Forest
Service. 1988. Range Plant Handbook. Dover
Publications, Inc. New York. 837 p.
369
Gynerium sagittatum (Aubl.) Beauv. wild cane
POACEAE
Synonyms: Saccharum sagittatum Aubl.

Arundo saccharoides Poir.
Airo gigantea Steud.
John K. Francis
Central America and South America to Paraguay

(Howard 1979). It is not known to have naturalized
elsewhere. Two types coexist in the western
Amazon Basin: a “small” and a “large” type that
differ considerably in physical form and mode of
reproduction (Kalliola and others 1992). The
distinction is not reported elsewhere.
Ecology.—Wild cane grows on sites with moist

soils, usually high in organic matter, often with the
water table near the surface. These sites are
seasonally flooded areas such as lake shores,
swamps, river flood planes, or sand bars. The
species grows at elevations from 10 to 1,600 m
above sea level in Costa Rica (Instituto Nacional
de Biodiversidad 2002). Wild cane resists damage
from moderate flooding and sprouts after being
covered with sediment. “Large type” stands in the
General Description.—Wild cane, also known as western Amazon region vary in density from 0.6 to
bitter cane, uva grass, and caña brava, is a tall 2.6 culms/m2. Forest edges “shade out” portions of
shrub with a grass-like habit. Its culms are usually wild cane stands, and occasional trees grow up
5 or 6 m in height and 2 or 3 cm in diameter but through stands and eventually suppress culms
may reach 10 m in height and 4 cm in diameter in growing under their crowns. The species affects the
Puerto Rico. The species varies from 5 to 14 m in course of forest succession (Kalliola and others
height in the western Amazon Basin (Kalliola and 1991). Apparently, disturbance that creates bare,
others 1992). The culms arise from underground wet soil is necessary for seedling establishment.
rhizomes which also produce weak and flexible
lateral roots, mostly 1 mm or less in diameter. The Reproduction.—In some environments flowering
culms have closely imbricated woody sheaths occurs throughout the year (Croat 1978); in others
around a hard, woody exterior, and a fibrous it occurs near the end of the low water period
interior. They are usually unbranched and taper (Kalliola and others 1992). The species is
little except near the top. The older leaves are shed, apparently wind pollinated (Pohl 1983). There are
leaving a plainer, fan-like group near the apex. The 1.67 million seeds/kg, and they can be expected to
leaf blades are 1 to 2 m long and have sharp germinate between 3 and 7 days following sowing
serrulate margins. The clonal groups of plants are at temperatures between 20 and 30 °C (Association
dioecious. The grayish-white plume-like terminal of Official Seed Analysis 2002). Almost all the
panicles are large, up to 2 m long. The male and seeds of the “short” type from the Amazon Basin
female inflorescences are similar in appearance, germinated within 3 weeks, and 0 to 2 percent of
but pistillate plants have a slightly fuzzy the “large” type germinated (Kalliola and others
appearance because of hairy lemmas. The fruits are 1992). Seeds are dispersed by wind and water
brown and about 1 mm long (Croat 1978, Howard (Kalliola and others 1992). Vegetative propagation
1979, Pohl 1983, Stevens and others 2001). is also important, both for expanding colonies and
establishing new ones. Horizontal runners or
Range.—Wild cane is native to the West Indies rhizomes, surface or underground, are constantly
except the Bahamas, and from Mexico through active and establish new plants or clumps as far as
370
20 m from the parent plants (Pohl 1983). Segments Croat, T.B. 1978. Flora of Barro Colorado Island.
of culm or rhizome, carried by floodwaters and Stanford University Press, Stanford, CA. 943 p.
covered with soil or debris, sprout and start new
colonies. Howard, R.A. 1979. Flora of the Lesser Antilles,
Growth and Management.—Growth of wild cane Arboretum, Harvard University, Jamaica Plain,
is rapid. Nursery seedlings reached 20, 30, and 50 MA. 586 p.
cm after 1, 2, and 4 months (Kalliola and others
1992). How long seedlings take to reach maturity Instituto Nacional de Biodiversidad. 2002.
and how rapidly suckers grow is unknown. Jerarquía taxonómica: Lista de especímenes de
Theoretically, baring catastrophes and invasion and Gynerium sagittatum. http://www.inbio.ac.cr/
shading by trees, clones can endure indefinitely. bims/k03/p13/c046/00159/f01382/g008647/s027
Culms of Amazon Basin plants produced close to 308.htm. 2 p.
200 leaves during their lifetimes, having from 19 to
28 living leaves at a time (Kalliola and others Kalliola, R., M. Puhakka, and J. Salo. 1992.
1992). Unbranched culms die after flowering, but Interspecific variation, and the distribution and
only the branches of branched culms die. If not ecology of Gynerium sagittatum (Poaceae) in the
controlled, wild cane slowly invades wet western Amazon. Flora 186(3-4): 153-167.
bottomland pastures and eliminates forage plants.
Periodic mowing appears to be adequate for control Kalliola, R., J. Salo, M. Puhakka, and M. Rajasilta.
of advancing clumps. 1991. New site formation and colonizing
vegetation in primary succession on the western
Benefits.—Wild cane provides cover for wildlife Amazon floodplains. Journal of Ecology 79(4):
and protects stream banks from erosion (Kalliola 877-901.
and others 1992). Its culms lack the strength and
toughness of hardwoods and bamboo but still are Pohl, R.W. 1983. Gynerium sagittatum (caña
used in rude construction, drying racks, vegetable brava, cane). In: D.H. Janzen, ed. Costa Rican
stakes and fruit props, and for weaving mats, natural history. University of Chicago Press,
baskets, and hats (Kalliola and others 1992). In the Chicago and London. p. 248-249.
Amazon area, arrow shafts are made from the dried
culms (Russo 2002). Plumes are used for dry floral Russo, E.B. 2002. Village life of the Machiguenga.
arrangements. http://montana.com/manu/village.html. 4 p.
References Stevens, W.D., C. Ulloa-U., A. Pool, O.M.

Association of Official Seed Analysis. 2002. Monographs in Systematic Botany Vol. 85, No.
Reference base. http://www.aosaseed.com/ 3. Missouri Botanic Garden Press, St. Louis,
species/speciesG.pdf. [not paged]. MO. p. 1,911-2,666.
371
Hamelia patens Jacq. firebush
RUBIACEAE
Synonyms: Hamelia erecta Jacq.

Hamelia pedicellata Wernh.
Hamelia latifolia Reichb. ex DC.
John K. Francis
modified dichasium with flowers that are tubular,
12 to 22 mm long, and orange to red in color. The
fruit is a berry, spherical to elliptical, 7 to 10 mm
long, turning red and then black at maturity. The
seeds are orange-brown, 0.6 to 0.9 mm long
(Howard 1989, Liogier 1997).
Range.—The native range of firebush extends

from southern Florida and Bermuda, through the
Bahamas, the Greater and Lesser Antilles, Trinidad
and Tobego, and from Mexico through Central
America and South America to Paraguay and
Argentina (Little and others 1974). The species is
also cultivated throughout the moist tropics and
subtropics but is not reported to have naturalized
outside its native range.
Ecology.—Firebush grows in deforested areas, in

thickets with other brushy species, in forest
openings, or in the understory of low basal-area
forest stands. The species is found in moist and wet
areas that receive from about 1600 to about 3000
mm of precipitation. Firebush prefers loamy or
clayey soil. It grows on soils derived from volcanic
General Description.—Firebush, which is also and sedimentary parent materials and is most
known as hummingbird bush, scarletbush, common in areas with limestone rocks.
bálsamo, busunuvo, pata de pájaro, Doña Julia,
fleur-corail, and many other names, is a medium- Reproduction.—Firebush flowers throughout the
sized shrub. This shrub commonly ranges from 1 year. It is a hummingbird-pollinated plant
to 3 m in height, but sometimes reaches 7 m in (Cunningham 1994). The flowers are also visited
height (Howard 1989). It grows as a tree in the by butterflies (Desert-Tropicals 2001). There were
Atlantic tropical lowland of Costa Rica (Paciorek 970,000 seeds/kg in a collection of seeds from
and others 1995). Firebush has a tap and lateral Puerto Rico. Sown on wet peat, germination started
root system with abundant fine roots. The roots are after 20 days and ended at 40 days with 46 percent
red-brown. Stem bark is gray and smooth and the germinated. Although the species is a good
inner bark is light green (Little and others 1974). producer of fruits and seeds in Puerto Rico,
Plants may have single or multiple stems. The seedlings are not common. The fruits are eaten by
twigs are orange to purple. Leaves are opposite or birds (Floridata 2001), which disburse the seeds.
grouped in threes or fours, and finely hairy to Layering occurs whenever stems come in contact
glabrous. The leaves have petioles 1 to 3.5 cm long with the ground. The species can be propagated
and blades that are mostly ovate-elliptic to from seeds, but most commercial ornamentals are
obovate-elliptic with an acute or acuminate tip. The produced from cuttings. In one test, cuttings 15 cm
lateral and especially the midveins are red or pink. long with four leaves, which had been taken from
In temperate areas, as the temperatures turn cool in the shoot tip, treated with 4000 ppm of indol-
the fall, the foliage turns to a brilliant red—hence buteric acid (IBA) and planted in sand, gave the
the common name. The inflorescence is terminal, a best rooting (Bhattacharjee and Balakrishna 1992).
372
Another experiment using an auto-humid chamber Cunningham, S.A. 1994. Measuring the
obtained 100 percent rooting of softwood cuttings relationships between floral duration and fruit
treated with hormones and 40 percent without set for Hamelia patens (Rubiaceae). Biotropica
hormone treatment (Maiti 1974). 26(2): 227-229.
Growth and Management.—In its native and Desert-Tropicals. 2001. Texas firecracker bush,
other frost-free habitat, firebush grows as a typical scarlet bush. http://www.desert-
shrub, but cultivated in warm temperate areas with tropicals.com/Plants/Rubiaceae/Hamelia_patens.
hard frosts, it will freeze to the ground and sprout html. 2 p.
again in the spring. Firebush is also planted as an
annual bedding plant in temperate climates. Floridata. 2001. Hamelia patens.
Ornamental plants need frequent watering during http://www.floridata.com/ref/h/hame_pat.cfm.
the establishment phase, but afterward, they are 3 p.
drought tolerant (Floridata 2001). Pruning of
ornamental plants is recommended to keep them Howard, R. A. 1989. Flora of the Lesser Antilles,
from becoming tall and spindly (Desert-Tropical Leeward and Windward Islands. Vol. 6. Arnold
2001). However, pruning inhibits blooming, which Arboretum, Harvard University, Jamaica Plain,
occurs at the terminals. MA. 658 p.
Benefits.—Firebush is loved and planted as an Liogier, H.A. 1997. Descriptive flora of Puerto
ornamental almost worldwide in warm, moist Rico and adjacent islands. Vol. 5. Editorial de la
areas. The fruit is edible (Little and others 1974). A Universidad de Puerto Rico, San Juan, PR.
phytochemical screening found firebush tissues 436 p.
rich in phytochemicals including alkaloids and
flavonoids (Raintree Nutrition 2001). Firebush is Liogier, H.A. 1990. Plantas medicinales de Puerto
used in herbal medicine to treat athlete’s foot, skin Rico y del Caribe. Iberoamericana de Ediciones,
lesions and rash, insect bites, nervous shock, Inc. San Juan, PR. 566 p.
inflammation, rheumatism, headache, asthma, and
dysentery (Liogier 1990, Mast Arboretum 2001, Little, E.L., Jr., R.O. Woodbury, and F.H.
Raintree Nutrition 2001). Firebush is one of the Wadsworth. 1974. Trees of Puerto Rico and the
woody species foraged by sheep and goats in Virgin Islands. Vol. 2. Agriculture Handbook
brushy pastures in Costa Rica. These animals 449. U.S. Department of Agriculture,
consumed 1.3 percent of their body weight of this Washington, DC. 1,024 p.
species in a free-choice test (Benavides 2001).
Firebush contains 17.5 percent crude protein and Maiti, R.G. 1974. Regeneration of terminal
has an in vitro digestibility of 61.6 percent greenwood cuttings of some ornamental shrubs
(Benavides 2001). under auto-humid chamber. Plant Science 6:
104-105.
References
Mast Arboretum. 2001. Hamelia patens, firebush.
Benavides, J.E. 2001. Arboles y arbustos Steven F. Austin State University.
forrajeros: una alternativa agroforestal para la http://www.sfasu.edu/ag/arboretum/plants/hamel
ganadería. Conferencia electrónica de la FAO iapatens/hamelia%20patens.htm. 3 p.
sobre la producción animal en Latinoamerica.
http://lead.virtualcentre.org/es/ele/conferencia1/ Paciorek, C.J., B.R. Moyer, R.A. Levin, and S.L.
bnvdes23.htm. 22 p. Halpern. 1995. Pollen consumption by the
hummingbird flower mite, Proctolaelaps
Bhattacharjee, S.K. and M. Balakrishna. 1992. kirmsei, and possible fitness effects on Hamelia
Studies on propagation of Hamelia patens Jacq. patens. Biotropica 27(2): 258-262.
and Ixora singaporensis Hort. from stem
cuttings. Progressive Horticulture 24(3/4): 157- Raintree Nutrition. 2001. Scarlet bush. Raintree
164. Nutrition, Inc. http://www.rain-
tree.com/scarletbush.htm. 3 p.
373
Helicteres jamaicensis Jacq. cowbush
STERCULIACEAE
Synonym: Helicteres spiralis Northop
John K. Francis
that are derived from sedimentary, igneous, and

metamorphic rocks, including ultramafics. In
Puerto Rico, it grows between sea level and 400 m
elevation and in areas that receive between about
700 and 1700 mm of annual precipitation. Well-
drained soils are required. The species competes
well with herbs and shrubs in early and middle
secondary forest. It grows on roadsides, in
openings, and in low and open forest canopies, but
not under closed canopies. Some degree of
disturbance appears to be necessary to perpetuate
the species. It is common in many partially forested
rangelands, but disappears under severe
overgrazing or frequent fires.
Reproduction.—Cowbush flowers irregularly

through the year. The fruits in a Puerto Rican
General Description.—Cowbush is a shrub or collection averaged 1.436 g/fruit and contained an
sometimes a small tree 7 m in height and up to 20 average of 152 seeds/fruit. The seeds weighed an
cm in basal diameter. It is also known as cowitch, average of 0.0032 g/seed or 312,000 seeds/kg. The
blind-eye-tree, salzbush, wild salve, screwtree, carpels of the fruits open at the ends to gradually
jeucon, cottonrat, cuernecillo, gato soga, huevo de release the seeds as the fruits shake in the wind. To
gato, gato, majagüilla de costa, and tapaculo (Little collect small quantities, tear the fruits apart and
and others 1974). The shrub has a dense covering pick out the seeds or cut off the ends of the fruits
of soft hairs throughout. The trunk is gray with and tap them to release most of the seeds. Larger
conspicuous warts. A somewhat swollen taproot quantities can be obtained by pulverizing the fruits
with abundant laterals and fines form the root and winnowing the seed. Seeds from the Puerto
system. The tan-colored roots are somewhat stiff Rican collection were sown in commercial potting
and woody. There is usually a single stem unless mix and germinated (99 percent) between 11 and
the plant has been damaged. It develops relatively 22 days after sowing. Germination is epigeal.
few long, stout, spreading or drooping branches. Natural seedlings are usually common wherever a
The alternate leaves have short petioles and ovate seed source is present. However, few progress past
to ovate-lanciolate blades with toothed edges. the small seedling stage.
Flowers are few and form in groups of one to three
at leaf bases. They have five white petals. The fruit Growth and Management.—Growth is relatively
consists of five gray-green hairy carpals that twist rapid after the early seedling stage, but the shrubs
upon themselves to form an ellipse about 2 to 4.5 are not long-lived (5 to 15 years). The use of
cm long. The seeds are blackish and about 1 mm in containerized seedlings for plantations is
diameter (Liogier 1994, Little and others 1974). recommended.
Range.—About 40 tree or shrub species of Benefits.—Cowbush is occasionally planted as an

Helicteres are found in tropical America and Asia ornamental. It is attractive as a large foliage or
(Griffiths 1994). Cowbush is native to the background plant and makes an interesting
Bahamas, the Greater Antilles, the Virgin Islands, curiosity plant. It has been planted in revegetation
and St. Martin (Little and others 1974). Planting or projects (Reforesta 2001). The wood of the species
naturalization in other areas has not been reported. is yellow, hard and heavy, but not available in sizes
sufficient to be used for anything other than fuel.
Ecology.—Cowbush grows on soils of all textures
374
The fibrous bark is strong and was once used for http://jamaicans.com/tourist/bites.htm. 3 p.
making rope (Little and others 1974). Cowbush is
reported to have several uses in herbal medicine. A Liogier, H.A. 1994. Descriptive flora of Puerto
tea made from the wood and leaves is used to treat Rico and adjacent Islands. Vol. 3. Editorial de la
bilious conditions and as an emetic. A decoction of Universidad de Puerto Rico. 461 p.
those same tissues is good for fever and cough, and
the cooked root is able to stop bleeding (Liogier Liogier, H.A. 1990. Plantas medicinales de Puerto
1990). The hairs of the species are reported to Rico y del Caribe. Iberoamericana de Ediciones,
cause serious irritation (Jamaicans 2001), but this Inc., San Juan, PR 566 p.
is not reported by Little and others (1974) and was
found to be untrue by the author, at least for Puerto Little, E.L., Jr., R.O. Woodbury, and F.H.
Rican plants. Wadsworth. 1974. Trees of Puerto Rico and the
References 449. U.S. Department of Agriculture,
Griffiths, M. 1994. Index of garden plants. Timber
Press, Portland, OR. 1,234 p. Reforesta. 2001. Restauración ribereña.
http://www.reforesta.com/projects/Espanol
Jamaicans. 2001. Jamaica: Bites and Stings, etc. /riparian_restoration_esp.htm. 2 p.
375
Hibiscus pernambucensis Arruda seaside mahoe
MALVACEAE
Synonyms: Hibiscus tiliaceus var. pernambucensis (Bertol.) I.M. Johnston

Hibiscus tiliaceus ssp. pernambucensis (Arruda) Castell
Hibiscus arboreus Ham.
Paritium pernambucense (Bertol.) Don
John K. Francis
capsules develop, and upon drying, split open
along five lines to release their seeds.
Range.—Seaside mahoe is native to coastal areas

of Bermuda, southern Florida, the Bahamas, the
Greater and Lesser Antilles, and from Mexico
through Brazil on the Atlantic Coast and in Pacific
coastal areas from Mexico to Peru (Little and
Wadsworth 1964). Seaside mahoe is similar to and
once was classified as a variety of Hibiscus
tiliaceus L. All the New World natives are now
thought to be H. pernambucensis (Howard 1989).
Ecology.—Seaside mahoe usually grows just

inland of mangroves around brackish lakes and
along rivers in the lower reaches of their
floodplains. Thickets have formed from planted
individuals in abandoned subsistence farms in
rainy (usually above 2000 mm of precipitation)
upland sites with poorly drained soils. It also
grows as a beachfront tree in association with
Terminalia catappa L. (Malavasi and Malavasi
1999). Seaside mahoe grows on hummocks and
General Description.—Seaside mahoe is also sand dunes in southern Florida (West and Arnold
known as sea hibiscus and rope mangrove in 1952). The sandy habitat probably has a water
English, emajagua and majagua in Spanish, and table within reach of the roots. Seaside mahoe
mahoe doux, mahaut, and bois flot in French suffers when there are droughts and appears to
(Howard 1989, Liogier 1994). It is a large shrub or require poorly drained sites to form the
occasionally a small tree. The species normally characteristic thickets. Although the species grows
grows in tangled thickets formed by stems that sag near brackish lakes, it is definitely less tolerant of
until they touch the ground and form roots. saline conditions than the mangroves. All textures
Simultaneously, sprouts arise from the horizontal of soil with a wide range of mineral fertility are
stems and grow upward until they in turn become represented in seaside mahoe habitat. The northern
prostrate. Occasionally, especially on dry ground, extent of the range is limited by the frost line.
open-grown individuals grow in a tree-like habit. Seaside mahoe grows in nearly pure
Adult plants have an aggressive lateral root system stands. Even artificially established stands in
with abundant fine roots. The bark of the limbs upland areas maintain themselves for many
and trunks is gray and smooth, but can become decades. This can probably be attributed to its
furrowed and scaly in old stems. The leaves are rapid growth, profuse sprouting, and the dense
nearly round (10 to 15 cm in diameter) except for a crowns that are formed. Seedlings of other species
deeply cordate base and a pointed tip. The petioles rarely survive for long. A number of insects have
are long (6 to 9 cm). Seaside mahoe produces large been collected from seaside mahoe (Martorell
yellow flowers that change to orange or red as they 1975). Although no insect seriously threatens the
age. Soon after, 2-cm long, hairy, cone-shaped species, aphids (Aphidoidea), white flies
376
(Aleyrodidae), and mealy bugs (Pseudococcideae) cleared planting spots with no other treatment. The
sometimes cause chlorosis and slower growth. fast-growing species that were planted overtopped
the sprouting seaside mahoe and eventually
Reproduction.—The principal means of suppressed them.
reproduction is by the layering and sprouting of
prostrate stems. The old prostrate stems eventually Benefits.—The fibrous inner bark was formerly
die and rot, leaving the sprouts as new, isolated used to make ropes, cords, fish nets, floats, cloth,
individuals. Seeds are produced in relative and mats. It is still used for emergency ties and
abundance and occasionally result in seedlings that lashings. The wood is used for fuel in
can eventually establish new clones. Streams and underdeveloped areas (National Academy of
floodwaters are the only known effective vector of Sciences 1983). The flowers make a colorful and
seed dispersal. tasty addition to salads and can be cooked as a
In wetter habitat, seaside mahoe flowers vegetable (Workman 1980). The young leaves,
continuously throughout the year (Little and bark, and roots were reportedly used for food
Wadsworth 1964). The species flowers seasonally during famines in former times (Little and
in habitat with drought stress during part of the Wadsworth 1964). Bees make honey from the
year. The flowers are pollinated by insects, and nectar. The plant is used as an ornamental and to
yield mature seeds about 1 month later. Fruits can stabilize stream banks (Workman 1980).
potentially produce at least 40 seeds per fruit. In
one collection from upland habitat in Puerto Rico, References
fruits averaged only 7.7 filled seeds per fruit. In
two other Puerto Rican collections from upland Howard, R.A. 1989. Flora of the Lesser Antilles,
habitat, the seeds averaged 0.018 + 0.006 and Leeward and Windward Islands. Vol. 5. Arnold
0.024 + 0.003 g/seed. These seed collections Arboretum, Harvard University, Jamaica Plain,
yielded 41 and 53 percent germination, which MA. 604 p.
occurred between 15 and 220 days after sowing.
Germination is epigeal. Seedlings are easily raised Liogier, H.A. 1994. Descriptive flora of Puerto
in the nursery from seed using traditional Rico and adjacent islands. Vol. 3. Editorial de la
techniques. Plants can be established from cuttings Universidad de Puerto Rico. Río Piedras, PR.
and air layers (Workman 1980). In one small 461 p.
planting in Puerto Rico, 40 of 140 untreated
cuttings (45 cm long and 2.5 cm minimum Little, E.L., Jr. and F.L. Wadsworth. 1964.
diameter) placed directly in the planting site rooted Common trees of Puerto Rico and the Virgin
and grew aggressively. Plants can also be started Islands. Agriculture Handbook 249. U.S.
from very large cuttings as living fence posts Department of Agriculture, Washington, DC.
(Little and Wadsworth 1964). 548 p.
Growth and Management.—Seaside mahoe Malavasi, U.C. and M.M. Malavasi. 1999. Tree
grows rapidly from sprouts and can extend its shade preferred by recreational users of a
leader 3 m or more per year. Seedlings grow Brazilian beach sidewalk. Journal of
slowly at first. A group of 6-month-old seedlings Arboriculture 25(2): 85-87.
averaged only 17 cm in height. Probably, 1 year is
required before nursery seedlings are ready for Martorell, L.F. 1975. Annotated food plant catalog
outplanting. Seaside mahoe may occasionally of the insects of Puerto Rico. Agricultural
reach 12 to 15 m in height (Workman 1980), but Experiment Station, University of Puerto Rico,
more typically reaches only 6 m in height and 15 Río Piedras, PR. 303 p.
cm in stem diameter. Because the species is
mainly valued in natural stands for environmental National Academy of Sciences. 1983. Firewood
protection and because it aggressively maintains crops: shrub and tree species for energy
nearly pure stands, little or no management is production. BOSTID Report 40. National
required once it is established. One unwanted Academy of Sciences. Washington, DC. 92 p.
upland stand was converted to timber species by
slashing the seaside mahoe, piling the slash West, E. and L.E. Arnold. 1952. The native trees
between the planting spots, and planting seedlings of Florida. University of Florida Press,
of Eucalyptus grandis Hill ex Maiden, Terminalia Gainsville, FL. 212 p.
ivorensis L., and Hibiscus elatus Sw. in the small
377
Sanibel-Captiva Conservation Foundation, Inc.
Sanibal, FL. 137 p.
378
Holodiscus discolor (Pursh) Maxim. creambush oceanspray
ROSACEAE
Synonyms: Spiraea discolor Pursh

Spiraea ariaefolia Smith in Rees
Schizonotus discolor Raf.
Sericotheca discolor Rydb.
Nancy L. Shaw
on rocky talus slopes with shallow sandy to clay
loam soils. Remnant stands are found on higher
peaks of Great Basin mountain ranges (Hitchcock
and others 1961, Stark 1966, USDA Forest Service
1937).
Ecology.—Creambush oceanspray is a prolific root

sprouter, capable of recovering from fire, grazing,
or mechanical damage. Regeneration from seed
following burns is often limited, and seedlings
develop slowly (Wright and others 1979). However,
regeneration from seed may be important following
extremely hot fires (Morgen and Neuenschwander
1988). The species is a host for vesicular-arbuscular
mycorrhizae (Berch and others 1988) and
Orobanche pinorum Geyer ex Hook. (Harrington
and others 1999). It is shade tolerant and disease
Illustration credits: Hitchcock and others 1961 and E.G. resistant.
Hurd
Reproduction.—Flowers appear from late spring
General Description.—Creambush oceanspray, to mid summer and are insect-pollinated. Fruits
also known as arrowwood, hardhack, rock-spirea, ripen in late summer (Hitchcock and others 1961);
and mountain spray, is a usually erect, deciduous seed is wind dispersed. Fruits are hand harvested
shrub with multiple arching branches. Plants range and conditioned by air drying and screening larger
from 1 to 3 (6) m in height. Large tree-like forms material with a fanning mill. Number of cleaned
grow in coastal areas. Young twigs are finely seed/kg exceeds 11 million (King 1947, Link 1993),
hairy; bark on older stems is gray to deep grayish but few seeds are generally sound. Seed supplies are
red. Leaves are alternate, 2 to 10 cm long, and rare and costly. Moist prechilling at 1 to 2 oC for
ovate to ovate-lanceolate with lobed to doubly 120 days relieves dormancy (Marchant and
toothed edges and prominent veins. Upper surfaces Sherlock 1984). Natural germination may be
are hirsute; lower surfaces are lanate and enhanced by a heat pretreatment or microsite
sometimes glandular. The common name derives conditions provided by exposed mineral soils
from the showy, diffuse, drooping panicles that are (Morgan and Neuenschwander 1988). Seed may be
10 to 17 (30) cm long with numerous, tiny, white broadcast over a rough seedbed and covered by
to pink flowers. Dried flowers and panicle natural soil sloughing. It should be planted
branches turn brown and persist into winter separately or included in seed mixes of other native
(Hitchcock and others 1961). species, particularly if broadcast seeded. It should
not be seeded with competitive introduced species
Range.—Creambush oceanspray grows at (Shaw and others 2001). Bareroot and container
elevations from sea level to 2,150 m. It is most stock are easily propagated (Everett 1957). Achenes
abundant in coastal forests from British Columbia to should be fall sown or artificially prechilled and
southwestern California. It occurs eastward to spring sown in the bareroot nursery. Creambush
Montana in drier conifer types of the interior Pacific oceanspray wildings and layers have also been used
Northwest. It is also common in riparian areas and successfully (Kruckeberg 1982). Plants can be
379
grown from cuttings, but rooting varies widely Ericaceae. University of Washington Press,
among clones, cutting type, and propagation Seattle, WA. 614 p.
technique (Antieau 1987, Link 1993).
King, J.E. 1947. The effect of various treatments
Growth and Management.—Seedlings develop to induce germination of seeds of some plants
slowly, producing spreading, fibrous root systems valuable for soil conservation and wildlife.
(Wright and others 1979). Growth rate is moderate Master’s thesis. University of Idaho, Moscow,
and protection from introduced grass and forb ID. 97 p.
competition is required for one or two growing
seasons. Excessive browsing is rarely a threat to Kruckeberg, A.R. 1982. Gardening with native
developing seedlings, but they may be girdled by plants of the Pacific Northwest. University of
rodents or trampled by big game or livestock. Brush Washington Press, Seattle, WA. 252 p.
fields may be rejuvenated by prescribed burning at
10 to 15 year intervals (Orme and Leege 1980). Link, E. 1993. Native plant propagation techniques
for National Parks interim guide. Rose Lake
Benefits.—Creambush oceanspray provides habitat Plant Materials Center, East Lansing, MI. 240 p.
for a wide variety of organisms. Its palatability and
forage value vary geographically but are generally Marchant, C. and J. Sherlock. 1984. A guide to
low for big game and domestic livestock. Use may selection and propagation of some native woody
be greater in winter when other forage is limited. species for land rehabilitation in British
Palatability may be improved following fire. Columbia. Forest Research Report RR84007-
Creambush oceanspray offers considerable potential HQ. Ministry of Forests, Victoria, BC. 117 p.
for revegetating disturbed areas. Populations
capable of growing on dry, rocky, well-drained sites Morgan, P. and L.F. Neuenschwander. 1988. Seed-
or exhibiting low palatability may be particularly bank contributions to regeneration of shrub
valuable. It has been recommended for use in species after clear-cutting and burning. Canadian
highway plantings, riparian areas, windbreaks, Journal of Botany 66: 169-172.
erosion control, wildlife habitat improvement,
conservation plantings, and low maintenance Orme, M.L. and T.A. Leege. 1980. Phenology of
landscaping (Antieau 1987). Native Americans ate shrubs on a north Idaho elk range. Northwest
the seeds and made digging sticks and arrow shafts Science 54: 187-198.
from the hard, straight branches.
Shaw, N.L., E.G. Hurd, and P.F. Stickney.
References Holodiscus (K. Koch) Maxim. 2001. Ocean-
spray. In: F.T. Bonner and R.G. Nisley, eds.
Antieau, C.J. 1987. Field notes: Holodiscus Woody plant seed manual. U.S. Department of
discolor. American Nurseryman 166: 110. Agriculture, Forest Service, Agriculture
Handbook. Washington, D.C.
Berch, S.M., S. Gamiet, and E. Deom. 1988. http://wpsm.net/index.html. 7 p.
Mycorrhizal status of some plants of
southwestern British Columbia. Canadian Stark. N. 1966. Review of highway planting
Journal of Botany 66: 1,924-1,928. information appropriate to Nevada. College of
Agriculture Bulletin B-7. University of Nevada,
Everett, P.C. 1957. A summary of the culture of Desert Research Institute. Reno, NV. 209 p.
California plants at the Rancho Santa Ana
Botanic Garden. The Rancho Santa Ana Botanic USDA Forest Service. 1937. Range plant
Garden, Claremont, CA. 223 p. handbook. U.S. Government Printing Office,
Washington, D.C. 512 p.
Harrington, C.A., J.M. McGrath, and J.M. Kraft.
1999. Propagating native species: experience at
the Wind River Nursery. Western Journal of
Applied Forestry 14: 61-64.
Hitchcock, C.L., A. Cronquist, M. Ownbey, and

J.W. Thompson. 1961. Vascular plants of the
Pacific Northwest. Part 3: Saxifragaceae to
380
Wright, H.A., L.F. Neuenschwander, and C.M.
Britton. 1979. The role and use of fire in
sagebrush-grass and pinyon-juniper plant
communities: A state of the art review. General
Technical Report INT-58. U.S. Department of
Agriculture, Forest Service, Intermountain
UT. 48 p.
381
Holodiscus dumosus (Nutt. ex Hook.) Heller rockspirea
ROSACEAE
Synonyms: Spiraea dumosa (Nutt.) T. & G.
Schizonotus dumosus Koehne.
Holodiscus microphyllus Rydb.
Sericotheca dumosa Rydb.
Nancy L. Shaw
Range.—Rockspirea grows east of the Cascade and

Sierra Nevada Mountains from north central
Oregon, east to Wyoming, and south to Chihuahua,
Mexico at elevations ranging from 760 to 3,660 m
(Harrington 1964, Hopkins and Kovalchek 1983).
Ecology.—Rockspirea occurs in a wide array of

plant communities including Artemisia spp.,
Pinus spp.--Juniperus spp. L., Cercocarpus
ledifolius Nutt., chaparral, Populus tremuloides
Michx.--Pinus contorta Dougl. ex Loud., Picea
spp.--Abies spp., Pseudotsuga menziesii (Mirbel)
Franco, Abies concolor (Gord. & Glend.) Lindl. ex
Hildebr., and Pinus ponderosa Dougl. (Sutton and
Johnson 1974, USDA 1937). It grows as a pioneer
species in cracks and fissures on rock surfaces, as
a seral species in brush fields of forested areas, and
as a climax species in self-replacing, monospecific
stands. Within its range it commonly occurs on
well-drained, dry to moderately dry sandy or
gravelly soils that may be somewhat alkaline, but
it is also found on finer-textured soils. Rockspirea
Illustration credits: Hitchcock and others 1961 and often grows on dry, rocky ridges, talus slopes, and
E.G. Hurd
basalt outcrops (Hitchcock and others 1961, Welsh
and others 1987). It is a prolific root sprouter,
General Description.—Rockspirea, also capable of recovering from fire, grazing, or
known as gland oceanspray, bush oceanspray, mechanical damage.
mountain-spray, and creambush, is a compact,
rounded, intricately branched deciduous shrub Reproduction.—Panicles and floral buds develop
0.1 to 4 m tall with a crown diameter of 1.5 to 3 in early spring, but flowering is delayed until
m (Hitchcock and others 1961, Welsh and summer or early fall depending upon elevation.
others 1987). Leaf blades are oval and Fruits are wind and gravity dispersed through late
shallowly or coarsely lobed or toothed, but fall (Hitchcock and others 1961). King (1947)
generally without secondary teeth. Petioles are determined that although there are about
decurrent along the stems. Panicles are showy 12,000,000 seed/kg, only about 7 percent were
and terminal, overtopping the shrub. Flowers sound. These required moist prechilling at 5 oC for
are numerous, small, creamy-white, perfect, 18 weeks to release dormancy. Heat generated by
perigynous, and insect pollinated. Fruits are wildfires may also stimulate germination. Seed can
tiny hirsute achenes. Roots are densely fibrous be broadcast seeded on a rough seedbed. Planting
and spreading. The species has n = 18 stock can be grown by fall seeding in the bareroot
chromosomes (McArthur and Sanderson nursery or planting moist prechilled seed in the
1986). greenhouse.
Growth and Management.—Excessive browsing

rarely occurs as the species is present primarily in
382
rocky, inaccessible areas and on summer ranges range shrubs. Journal of Range Management 6:
where other species receive preferential use. Based 309-317.
on results of clipping studies in Oregon, Garrison
(1953) recommended a maximum of 50 to 60 Harrington, H.D. 1954. Manual of the plants of
percent use for sustained production. On some Colorado. Sage Books, Denver, CO. 666 p.
western juniper/big sagebrush habitat types of
eastern Oregon, the species tends to increase when Hitchcock, C.L., A. Cronquist, M. Ownbey, and
other species receive excessive use (Ferguson J.W. Thompson. 1961. Vascular plants of the
1983, Hopkins and Kovalchik 1983). Pacific Northwest. Part 3: Saxifragaceae to
Ericaceae. University of Washington Press,
Benefits.—Rockspirea is a potentially valuable Seattle. 614 p.
species for revegetation as it is drought tolerant
and adapted to sites with dry, rocky, unstable Hopkins, W.E. and B.L. Kovalchik. 1983. Plant
surface conditions in a variety of forested and associations of the Crooked River National
nonforested communities (Stark 1966). Its use has Grassland. R6 Ecology 133-1983. U.S.
been limited by a lack of high quality seed Department of Agriculture, Forest Service,
supplies. Palatability and forage value of Pacific Northwest Region, Portland, OR. 98 p.
rockspirea are low (USDA 1937), but it does
King, J.E. 1947. The effect of various treatments
provide food and cover for small animals. Kufeld
to induce germination of seeds of some plants
and others (1973) reported its use by mule deer
valuable for soil conservation and wildlife. .
(Odocoilus hemionus) was moderate in fall and
Master’s thesis. University of Idaho, Moscow,
light during the remainder of the year. Rockspirea ID. 97 p.
is used in summer by bighorn sheep (Ovis
canadensis) and rabbits (Sutton and Johnson 1974, Kufeld, R.C. 1973. Foods eaten by the Rocky
Todd 1975). It is an attractive, low water use Mountain elk. Journal of Range Management
ornamental. Native Americans made digging sticks 26: 106-113.
and arrow shafts from the hard, straight branches
(Anderson and Holmgren 1969, Daubenmire 1970, McArthur, E.D. and S.C. Sanderson. 1985. A
Hopkins and Kovalchik 1983). Native Americans of cytotaxonomic contribution to the western North
the Great Basin ate the seeds, and pioneers made American rosaceous flora. Madroño 32: 24-28.
nails from its wood.
Stark. N. 1966. Review of highway planting
References information appropriate to Nevada. College of
Agriculture Bulletin B-7. University of Nevada,
Anderson, B.A. and A.H. Holmgren. 1969. Desert Research Institute, Reno. 209 p.
Mountain plants of northeastern Utah. Circular
319. Utah State University, Logan, UT. 148 p. Sutton, R. and C.W. Johnson. 1974. Landscape
plants from Utah’s mountains. EC-368. Utah
Daubenmire, R. 1970. Steppe vegetation of State University, Logan, UT. 137 p.
Washington. Technical Bulletin 62. Washington
State University, Pullman, WA. 131 p. Todd, J.W. 1975. Foods of Rocky Mountain
bighorn sheep in southern Colorado. Journal of
Ferguson, R.B. 1983. Use of rosaceous shrubs for Wildlife Management 39: 108-111.
wildland plantings in the intermountain west. In:
S.B. Monsen, and N. Shaw, comps. Managing USDA Forest Service. 1937. Range plant
Intermountain rangelands--Improvement of handbook. U.S. Government Printing Office,
range and wildlife habitats: Proceedings. Washington, D.C. 512 p.
General Technical Report INT-157. U.S.
Department of Agriculture, Forest Service, Welsh, S.L., N.D. Atwood, S. Goodrich, L.
Intermountain Forest and Range Experiment Higgins, eds. 1987. A Utah flora. Great Basin
Station, Ogden, UT. p. 136-139. Naturalist Memoir 9. Brigham Young
University, Provo, UT. 894 p.
Garrison, G.A. 1953. Effects of clipping on some
383
Hyperbaena laurifolia (Poir.) Urban limestone snakevine
MENISPERMACEAE
Synonyms: Cissampelos laurifolia Poir. in Lam.

Anelasma laurifolia (Poir.) Meirs
Cocculus laurifolia Eggers
John K. Francis
1981). It had been reported in Montserrat (Little

and others 1974), but apparently this was in error
(Howard 1988).
Ecology.—Little and others (1974) and Mathias

and Theobald (1981) reported that this species is
confined to limestone areas. However, the author
knows of two stands near Salinas, Puerto Rico that
are growing on soil residual from igneous rocks.
Limestone snakevine grows from near sea level to
1,200 m in elevation (presumably in Haiti)
(Mathias and Theobald 1981). In Puerto Rico, it
grows to elevations of 600 m (Little and others
1974), and in areas that receive from about 800 to
about 2000 mm of mean annual precipitation.
General Description.—Limestone snakevine is an Limestone snakevine grows in a variety of soils,
uncommon scrambling shrub or vine-like small most of which are very rocky in the currently
tree. It is also known as hyperbaena, following the available habitat. The species is moderately shade
convention among natural resources workers to use tolerant and grows and reproduces mainly under
the Latin genus or species name as a common the crowns of trees in remnant and advanced
name. The vine develops a tap and lateral root secondary forests.
system. Plants of the species have single or
multiple smooth, gray stems with a moderate Reproduction.—Limestone snakevine flowers in
number of branches and stiff, slender, hairless the spring and bears fruits during the spring and
twigs. Limestone snakevine has alternate, oblong summer (Little and others 1974). The fruits are
to lanciolate, hairless, leathery leaves with a short easily detached and presumably dispersed by fruit
petiole and blades from 7 to 16 cm long and 2.5 to bats. Forty fruits in a Puerto Rican collection
6 cm broad. The leaves are shiny, dark green above weighed an average of 9.31 + 0.27 g each. Seeds
and dull, lighter green below, have smooth edges, from that collection averaged 4.18 + 0.12 g/seed or
and may be rounded or pointed at either end. Male 240 seeds/kg. Ninety-eight percent of the seeds
and female flowers grow on different plants sown in potting mix germinated between 45 and 72
(dioecious). The individual flowers are tiny, days after sowing. Two of the stones produced two
greenish-yellow, and grow in clusters at the leaf seedlings each. Germination is hypogeal, a stiff,
bases. The fruits (drupes) that develop are rounded well-branched root developing considerably before
and somewhat flattened, bright red when ripe, 2 to the top emerges. The tops reach 10 to 15 cm in
3 cm broad with a plum-like skin and flesh about 5 height from the reserves in the seeds. The seedlings
mm thick and contain a single seed. The large are ready to transplant within a month of
yellow stone within is grooved and shaped like a emergence. They are hardy and survive well when
crescent curved back upon itself (Liogier 1985, transplanted from nursery beds to pots. Seedlings
Little and others 1974, Mathias and Theobald are not common in the wild.
1981).
Growth and Management.—Limestone
Range.—Limestone snakevine is native to Puerto snakevine may reach 3 m in height, 8 m in lateral
Rico, St. Thomas in the U.S. Virgin Islands extension, and 10 cm of basal stem diameter.
(Liogier 1985), and Haiti (Mathias and Theobold Established plants appear to have a moderate
384
growth rate, and plants known to the author appear Leeward and Windward Islands.
to be several decades old. However, no data on Dicotyledoneae, Part 1. Vol. 4. Arnold
growth rate and longevity or experience on Arboretum, Harvard University, Jamaica Plain,
management have been published. MA. 673 p.
Benefits.—The fruits of limestone snakevine are Liogier H.A. 1985. Descriptive flora of Puerto
slightly acid, bitter, and unpleasant to the taste and Rico and adjacent islands, Spermatophyta. Vol.
smell. The fruit pulp immediately stains skin and 1. Editorial de la Universidad de Puerto Rico,
cloth a lasting reddish-brown and might be useful Río Piedras, PR. 352 p.
as a natural dye in crafts. The wood is heavy and
hard and would certainly be good for small- Little, E.L., Jr., R.O. Woodbury, and F.H.
diameter fuel. The species contributes to the Wadsworth. 1974. Trees of Puerto Rico and the
biodiversity of local forests and in a minor way, Virgin Islands. Vol. 2. Agriculture Handbook
furnishes wildlife food and cover, and protects 449. U.S. Department of Agriculture,
against erosion. Washingtion, DC. 1,024 p.
References Mathias, M.E. and W.L. Theobald. 1981. A

revision of the genus Hyperbaena
Howard R.A. 1988. Flora of the Lesser Antilles, (Menispermaceae). Brittonia 33(1): 81-104.
385
Indigofera suffruticosa P. Mill. wild indigo
FABACEAE
Synonyms: Indigofera anil L.

Indigofera truxillensis HBK.
Indigofera divericata Jacq.
Anila tinctoria vera Kuntze
John K. Francis
Range.—Wild indigo is native to Southern United

States through tropical and subtropical South
America as well as the Caribbean Islands (Howard
1988). The species has naturalized in Hawaii (Neal
1965) and is present in American Samoa, Guam,
and a large number of the Pacific Island groups
(Pacific Island Ecosystems at Risk 2001). Wild
indigo has been introduced into tropical regions of
the Old World (Liogier 1988).
Ecology.—In Hawaii, wild indigo has naturalized

in dry, highly disturbed areas from near sea level to
1,160 m in elevation (Pacific Island Ecosystems at
Risk 2001). In Puerto Rico, the species grows from
near sea level to about 700 m in elevation in areas
that receive from about 750 to 1800 mm of
precipitation annually. Soils derived from volcanic
and sedimentary rocks in a wide range of soil
textures are colonized. Good drainage is required,
but not high fertility. Wild indigo is not shade
tolerant and will not grow under a closed forest
stand.
Reproduction.—In Texas, wild indigo flowers

from July through November (Everitt and Drawe
1993). In Mexico, fruiting occurs between October
and February (Moreno-Casasola and others 1994).
The pods remain open on the branches, exposing
General Description.—Wild indigo, also known but not expelling the seeds. Mechanisms of
as indigo, Guatamala indigo, añil, añil de pasto, transport have not been reported. The seeds in a
and ti cafe, is a short-lived shrub that reaches 1 to 2 collection of fruits in Puerto Rico had been
m in height and 1 to 2 cm in stem diameter. The partially consumed by an unknown insect and
shrub may have multiple stems, especially if it has averaged only 0.9 undamaged seeds per pod. These
been disturbed by grazing or fire. The stems are seeds averaged 0.0039 g/seed or about 257,000
gray-brown, pubescent, and more or less straight. seeds/kg. Of these, 17 percent germinated in 7 to
The light green leaves are pinnately compound 24 days after sowing. Seed from a Mexican
with 9 to 17 narrowly oblong, pubescent leaflets collection gave more than 90 percent germination
that are 1.5 to 2.5 cm long and about 9 mm wide. (in the field) when a mechanical scarification was
There are 6- to 8-mm lanciolate stipules at the base used and nearly 100 percent when wet-dry cycles
of the leaves. Crimson to rust-red flowers grow in were combined with scarification (Moreno-
short, many-flowered racemes. The curved Casasola and others 1994).
legumes are short (1.1 to 2.5 cm) and contain three
to seven seeds that are 1.5 mm wide and 1 mm Growth and Management.—Neal (1965) reports
thick (Howard 1988, Liogier 1988). that in Hawaii, 20,400 kg of wild indigo can be
386
produced per hectare from which 227 kg of indigo HealthLink. 2001. Monograph: indigo naturalis.
paste can be separated. Wild indigo tested for http://www.healthlink.com.au/nat_lib/htm-
revegetation of soil-lignite overburden varied in data/htm-herb/bhp1016.htm. 3 p.
biomass yield from 162 to 2,432 kg/ha and
maintained from 8 to 30 percent cover over a 3- Howard, R.A. 1988. Flora of the Lesser Antilles,
year period (Skousen and Call 1987). Surface Leeward and Windward Islands.
sowing at the start of the rainy season on freshly Dicotyledoneae, Part 1. Vol. 4. Arnold
tilled ground is recommended. Plants usually live 2 Arboretum, Harvard University, Jamaica Plain,
or 3 years in Puerto Rico. MA. 673 p.
Benefits.—Although indigo obtained from other Liogier H.A. 1988. Descriptive flora of Puerto
species of Indigofera was used in the Old World, Rico and adjacent islands, Spermatophyta. Vol.
the use of wild indigo by pre-Columbian natives of 2. Editorial de la Universidad de Puerto Rico,
Mexico to dye cloth and paint in various shades of Río Piedras, PR. 481 p.
blue was passed down to the Spanish colonists
(Haude 1997). Indigofera species became Liogier, H.A. 1990. Plantas medicinales de Puerto
important commercial crops in various tropical and Rico y del Caribe. Iberoamericana de Ediciones,
subtropical areas. The blue dye was produced by Inc., San Juan, PR. 566 p.
fermentation of the leaves, usually with caustic
soda or sodium hydrosulfite, and the exudates Moreira-Casasola, P., J.P. Grime, and M.L.
processed into dry cake. The blue color developed Martinez. 1994. A comparative study of the
as the cake was exposed to the air (Simon and effects of fluctuations in temperature and
others 1984). Indigo was used as a bluing to moisture supply on hard coat dormancy in seeds
counter the yellowing in clothes from washing with of coastal tropical legumes in Mexico. Journal of
soap (Vélez and van Overbeek 1950). In the last Tropical Ecology 10(1): 67-86.
few decades, natural indigo has been almost wholly
replaced by synthetic dyes. Poultices and extracts Neal, M.C. 1965. In gardens of Hawaii. Special
of wild indigo leaves, alone or in combination with Publication 50. Bernice P. Bishop Museum
other ingredients, are used in herbal medicine to Press, Honolulu, HI. 924 p.
treat fever, headaches, hemorrhages, convulsions,
acute cough, skin parasites, and boils (HealthLink Opler, P.A. and V. Malikul. 1992. A field guide to
2001, Liogier 1990). Domestic and wild ruminant eastern butterflies. Peterson field guide 4.
animals browse wild indigo plants. It and another Houghton-Mifflin Co., Boston, MA. 396 p.
Indigofera species are reported to be the sole hosts
of the false dusky wing butterfly [Gesta gesta Pacific Island Ecosystems at Risk. 2001. Invasive
(Herrich-Schäffer)] (Opler and Malikul 1992). plant species: Indigofera suffruticosa Miller,
Wild indigo is useful as a natural cover plant in Fabaceae. http://www.hear.org/pier/insuf.htm.
disturbed areas. The species may become weedy, 2 p.
but is seldom aggressive or common enough to
cause difficulties in croplands. Simon, J.E., A.F. Chadwick, and L.E. Craker.
1984. Herbs: An indexed bibliography. 1971-
References 1980. Scientific literature on selected herbs, and
aromatic and medical plants of the Temperate
Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs Zone. Archon Books, Hamden, CT. 770 p.
and cacti of South Texas. Texas Tech University
Press. 213 p. Skousen, J.G. and C.A. Call. 1987. Grass and forb
species for revegetation of mixed soil-lignite
Haude, M.E. 1997. Identification and classification overburden in East Central Texas. Journal of
of colorants used during Mexico’s early colonial Soil and Water Conservation 42(6): 438-441.
period. Book and Paper Group Annual Vol. 16.
The American Institute of Conservation. Vélez, I. and J. van Overbeek. 1950. Plantas
http://aic.stanford.edu/conspec/bpg/annual/v16// indeseables en los cultivos tropicales. Editorial
bp16-05.html. 26 p. Univeresitaria, Río Piedras, PR. 497 p.
387
Isocoma tenuisecta Greene burro weed
ASTERACEAE
Synonyms: Haplopappus tenuisectus (Greene) Blake

Aplopappus tenuisectus (Greene) Blake
and Hartman 1996).
Range.—Burro weed occurs in southwestern

Texas, southern New Mexico, southern Arizona
and northern Mexico (Kearney and others 1960,
Martin and Hutchins 1980, Ivey 1995).
Ecology.—Burro weed grows on dry slopes and

mesas between 600 and 2,000 m in elevation. It is
a member of various vegetation communities
including desert scrub, desert grassland, and pine-
oak woodland (Bowers and McLaughlin 1987).
Although burro weed can be moderately long
lived, up to 20 years, long-term monitoring studies
indicate as few as 2 percent survive more than 7
years (Goldberg and Turner 1986). Burro weed has
a deep (3 to 6 m long), relatively unbranched, root
system but is susceptible to drought and often dies
during drought periods (Schmutz and others 1992).
However, following droughts burro weed shrubs
often increase in number (Schmutz and others
1992). Similar to other species within the Isocoma
Illustration credit: Robert DeWitt Ivey genus, fire easily kills burro weed, but burned sites
will eventually become reoccupied (White 1969,
General Description.—Burro weed is a compact, Cable 1967). For example, fire reduced a pre-burn
rounded shrub, sometimes referred to as a subshrub density of 3,762 plants per acre to 327 plants
shrub, growing to approximately 1 m high and per acre that was reduced even further to 87 plants
approximately 1 m wide. Its grayish-green, per acre in the following year. Those plants that
glandular leaves are pinnately cleft (deeply lobed) survived were in unburned or light burned patches
into four to eight linear acute lobes. The yellow within the burn area (Cable 1967). However, the
flowers are discoid, having no ray flowers numbers of shrubs gradually increased in
(“petals”), and are arranged in dense terminal subsequent years to number 7,175 shrubs per acre
clusters. The flowers dry and turn a light brown 13 years after the burn. The spatial association
color while remaining on the stems (PCC 2001). between different three-awn grass species and
The bracts around the flowers are also glandular burro weed is an interesting example of the
(Martin and Hutchins 1980). Other members of the tradeoffs between protection and liability of a
same genus are called golden weeds or yellow bush potentially advantageous microclimate and refuge.
(Dayton 1931). Historically, Haplopappus was a Santa Rita three-awn (Aristada glabrata (Vasey)
large and variable genus that was a depository for Hitchc.) grows between burro weed individuals,
loosely related “yellow daisies” that shared a whereas some other three-awn species, such as
limited number of characteristics (Taylor 1998). Aristida hamulosa Henr. and Aristida ternipes
However, with the aid of molecular (DNA) data Cav., grow within the burro weed crowns. The
and more powerful computer analyses, taxonomists latter species are subjected to greater heat in the
have placed members of Haplopappus into smoldering, resinous crowns than in the open areas
different genera, such as Isocoma, Ericameria, and, in combination with the shallow subsurface
Stenotus and Machaeranthera (Nesom 1991, Lane buds that characterize all three-awns, suffer a
388
higher mortality rate than the Santa Rita three-awn and vegetation of the Rincon Mountains, Pima
(Blydenstein 1966, Cable 1967, McPherson 1995). County, Arizona. Desert Plants 8(2): 51-94.
Reproduction.—Burro weed flowers from August Cable, D.R. 1967. Fire effects on semidesert
into October (Martin and Hutchins 1980). The grasses and shrubs. Journal of Range
seeds (achenes) are partially covered by silky hairs Management 20: 170-176.
(Martin and Hutchins 1980). The long hairs that
comprise the pappus at one end of the seed aid Dayton, W.A. 1931. Important western browse
dispersal (Parker 1990). plants. Misc. Publ. 101. U.S. Department of
Agriculture. Washington, DC. 214 p.
Growth and Management.—Burro weed is a
resilient species that can flourish on highway Elmore, F.H. 1987. Shrubs and Trees of the
shoulders, parking lots, dirt roads, vacant lots and Southwest Uplands. Southwest Parks and
other disturbed areas. It tends to be sporadically Monuments Assoc. Tucson, AZ. 214 p.
scattered on range covered by perennial bunch
grasses. However, on disturbed ranges, with even Epple, A.O. 1995. A field guide to the plants of
a good stand of annual grasses, it can become one Arizona. LewAnn Publishing Company, Mesa,
of the principal plant species (USDA FS 1988). AZ. 347 p.
Burro weed foliage is poisonous to livestock
(AVDL 1998, Bowers 1993). Horses are most Goldberg, D.E. and R.M. Turner. 1986. Vegetation
sensitive while sheep are least sensitive. If change and plant demography in permanent
livestock eat large amounts of the vegetation they plots in the Sonoran Desert. Ecology 67(3): 695-
suffer from “trembles,” “alkali sickness,” or “milk- 712.
sickness” and humans can become ill after
drinking the milk from affected cows (Dayton Ivey, R.D. 1995. Flowering plants of New
1931, Epple 1995). The toxic substance Mexico. 3rd Ed. Rio Rancho Printing,
transmitted by the milk is called tremetol (AVDL Albuquerque, NM. 504 p.
1998, Kingsbury 1964). Many other species of
Isocoma contain the same toxic constituent Kearney, T.H., R.H. Peebles, and collaborators.
(AVDL 1998, Kingsbury 1964). As a consequence 1960. Arizona flora. 2nd Ed. University of
of its toxic properties, livestock avoid it unless California Press, Berkeley, CA. 1,085 p.
there is nothing else to eat. In overgrazed land
burro weed behaves as an invasive weed Kingsbury, J.M. 1964. Poisonous Plants of the
(Whitfield and Anderson 1938, Parker 1990). United States and Canada, Prentice-Hall, Inc.,
Englewood Cliffs, NJ. 626 p.
Benefits.—Burro weed provides cover and
stabilizes soils that are denuded by overgrazing or Lane, M.A. and R.L. Hartman. 1996.
other disturbance. It adds to the biodiversity of Reclassification of North American
communities of which it is part and may be Haplopappus (Compositae: Asteraceae)
important for pollinators and other arthropods in Completed: Rayjacksonia gen. nov. American
areas that are depauperate in native species. Journal of Botany 83(3): 356-370.
References Martin, W.C. and C.R. Hutchins. 1980. A Flora of

New Mexico. Strauss & Cramer, Hirschberg,
Arizona Veterinary Diagnostic Laboratory Germany. 2,590 p.
Quarterly. 1998. Equine 3 (4): 2.
McPherson, G.R. 1995. The role of fire in desert
Blydenstein, J. 1966. Root systems of four desert grasslands. In: M.P. McClaran and T.R. Van
grassland species on grazed and protected sites. Devender, eds. The desert grassland. The
Journal of Range Management 19: 93-95. University of Arizona Press, Tucson, AZ. p.
130-151.
Bowers, J.E. 1993. Shrubs and trees of the
southwest deserts. Southwest Parks and Neson, G.L. 1991. Taxonomy of Isocoma
Monuments Assoc., Tucson, AZ. 140 p. (Compositae: Asteraceae). Phytologia 70: 69-
114.
Bowers, J.E. and S.P. McLaughlin. 1987. Flora
389
Parker, K.F. 1990. An illustrated guide to Arizona Taylor, R.J. 1998. Desert wildflowers of North
weeds. The University of Arizona Press, America. Mountain Press Publishing Co.,
Tucson, AZ. 338 p. Missoula, MT. 349 p.
Pima Community College. 2001. Internet site. United States Department of Agriculture, Forest
http://wc.pima.edu/Bfiero/tucsonecology/plants/ Service. 1988. Range Plant Handbook. Dover
shrubs_bw.htm [not paged]. Publications, Inc. New York. 837 p.
Schmutz E.M., E.L Smith, P.R. Ogden, M.L. Cox, White, L.D. 1969. Effects of a wildfire on several
J.O. Klemmedson, J.J. Norris, and L.C. Fierro. desert grassland shrub species. Journal of Range
1992. Desert grasslands. In: R.T. Coupland, ed. Management 22: 284-285.
Ecosystems of the World, 8A; Natural
grasslands: Introduction and Western Whitfield, C.J and H.L. Anderson. 1938.
Hemisphere. Elsevier, London, UK. p. 337-362. Secondary succession in the desert plains
grassland. Ecology 19 (2): 171-180.
390
Itea virginica L. Virginia sweetspire
SAXIFRAGACEAE
Synonyms: None
Kristina Connor
form of a small tree but is generally multistemmed

and reaches a height between 0.9 and 2.4 m. This
species has also been placed in the Grossulariaceae
and Iteaceae (USDA-ARS-GRIN 2001, USDA-
NRCS 2001).
Range.—Virginia sweetspire is found throughout

the Southeastern United States, from the Atlantic
to eastern Texas and Oklahoma, although it is
uncommon in the latter state (Johnson and
Hoagland 1999). It can be found in wetlands as far
north as Maryland, Delaware, and New Jersey and
as far south as Florida. It is also found in southern
Illinois and throughout the Mississippi River
Valley.
Ecology.—Virginia sweetspire grows primarily in

wet areas, in swamps, along stream beds and lake
edges, and in floodplains where it can form
thickets. However, it can thrive in a variety of soil
types and pHs and also shows tolerance to
droughty conditions (Bailey and Bailey 1976,
Odenwald and others 1996, Ohio State University
2002). The species is tolerant of various light
Illustration source: U.S. Department of Agriculture [No
regimes and will grow well in both the sun and the
date] shade; however, it can become leggy and sparse in
the shade. It flowers more prolifically and has
optimal fall coloration when grown in full sun.
General Description.—Virginia sweetspire, also
The species is highly adaptable and tough.
known as sweetspire, tassel-white, Virginia tea, or
Virginia willow, is an upright, multistemmed,
Reproduction.—Virginia sweetspire is named for
deciduous or semievergreen shrub with arching
its drooping to upright 5 to 15 cm-long pubescent,
branches (Bailey and Bailey 1976, Krüssmann
terminal racemes of fragrant, white flowers that
1984, Odenwald and others 1996). The branches
occur from early spring to early summer.
are light green or brown, pubescent when young,
Individual flowers have five petals and five sepals
while old stems are glabrous, gray and striated.
and are small, only 6 to 9 mm across (Johnson and
Twigs are slender and may be red when young
Hoagland 1999). Flowering period is fairly long,
(Krüssmann 1984) or in the fall (Seiler and
lasting from May to early July (Krüssmann 1984).
Peterson 2001). The medium to dark green,
Brown seeds are produced in small elongate
simple, alternate leaves are elliptic to oblong in
capsules with cranesbill-like projections (The Ohio
shape, glabrous above and finely pubescent
State University 2002). Capsules can remain on
beneath, four to 10 cm long, and have finely
the plant long after leaf fall. Seeds are quick to
toothed margins. The plants are noted for their
germinate and reportedly require no pretreatment
racemes of white flowers in early summer and
for germination to occur. Virginia sweetspire can
their brilliant red to reddish purple foliage in
be propagated by rooting cuttings, from seed, or
autumn. Leaves can remain on the plant in a mild
from root suckers (Bailey and Bailey 1976).
winter or in the more southern hardiness zones.
Virginia sweetspire can occasionally attain the
391
Growth and Management.—Virginia sweetspire Odenwald, N.G., C.F. Fryling, Jr., and T.E. Pope.
is semievergreen and fairly long lived. With its 1996. Plants for American Landscapes. Louisiana
long flowering period and attractive fall foliage, State Univ. Press, Baton Rouge, LA. 266 p.
the species can do well in landscape plantings. Its
adaptability to both sun and shade, and to wet and Ohio State University. 2002. Horticulture and Crop
dry areas, gives it a versatility lacking in some Science in Virtual Perspective.
other shrub species. However, it can grow beyond http://www.hcs.ohio-state.edu/hcs/TMI/
desired boundaries because it spreads by root Plantlist/it_inica.html. 4 p.
suckers, and if not pruned it will form thickets. It
grows fairly rapidly and easily from basal shoots Seiler, J.R. and J.A. Peterson. 2001. Sweetspire.
and is relatively free from insect pests and diseases Virginia Tech. Dendrology Webpage
(The Ohio State University 2002). http://www.fw.vt.edu/
dendro/dendrology/syllabus/ivirginica.htm. 1 p.
Benefits.—The dense thickets of Virginia
sweetspire provide cover for wildlife (Silberhorn Silberhorn, G. 1996. Virginia willow (Itea
1996). Although the dry fruit capsules and small virginica). Technical Report. Wetlands Flora. No.
seeds are not a preferred food, they are eaten by 96-5. Virginia Institute of Science, School of
birds. The species can be used for erosion control Marine Science, College of William and Mary,
on wet or sloped sites. The fragrant flowers are Gloucester Point, VA. 2 p.
attractive to butterflies.
U.S. Department of Agriculture. [no date].
References Southern wetland flora: Field office guide to
plant species. U. S. Department of Agriculture,
Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third: A Soil Conservation Service, South National
Concise Dictionary of Plants Cultivated in the Technical Center, Fort Worth, TX. Jamestown,
United States and Canada. McMillan Publishing ND: Northern Prairie Wildlife Research Center
Co., Inc., New York. 1,312 p. Home Page. http://www.npwrc.usgs.gov. [not
paged].
Johnson, F.L. and B.W. Hoagland. 1999. Itea
virginica. Catalog of the Woody Plants of USDA-ARS-GRIN. 2001. Germplasm Resources
Oklahoma, Oklahoma Biological Survey. Information Network, National Germplasm
http://www.biosurvey.ou.edu. 1 p. Resources Laboratory. Plant Sciences Institute,
Beltsville, Agricultural Research Center,
Krüssmann, G. 1976. Manual of Cultivated Broad- Agricultural Research Service, U.S. Department of
Leaved Trees and Shrubs. Volume I, A-D. Timber Agriculture. http://www.ars-grin.gov. [not paged].
Press, Beaverton, OR. 448 p.
USDA, NRCS. 2001. The PLANTS database,
Version 3.1. National Plant Data Center, Baton
Rouge, LA. http://plants.usda.gov. [not paged].
392
Ixora coccinea L. jungle-geranium
RUBIACEAE
Synonyms: Ixora grandiflora Bot.

Ixora bandhuca Roxbg.
John K. Francis
shrubs and herbs but not with dense grass. Little is

known about environmental conditions in its native
range, but it grows naturally in Puerto Rico in areas
that receive around 1800 mm of annual
precipitation. Jungle-geranium grows well in all
textured soils if moisture is adequate and drainage
is good. The species prefers slightly acid soils with
good fertility and becomes chlorotic under alkaline
conditions. It tolerates mild salt spray (Watkins and
Sheeham 1975). Jungle geranium is cold sensitive
and will partially defoliate after chilling (Michaelia
and others 1999). Frosts will kill it to the ground,
but it recovers afterward (Tropilab 2001). Jungle-
General Description.—The common name of this geranium in Florida is damaged by nematodes, root
plant, jungle-geranium, seems to be in slightly rot, leaf spots, and scale insects (Florida Gardener
more common use than others: flame of the woods, 2001).
flame of the forest, jungle flame, burning love, red
ixora, ixora, amor ardiente, cruz de Malta, bola de Reproduction.—Jungle-geranium flowers
coral, equisósea, and rajana (Griffiths 1994, throughout the year in full sunlight and light shade
Liogier 1997). Jungle-geranium is a densely (Whistler 2000). Shrubs in heavy shade survive but
branched shrub to 3 m in height and 3 or 4 cm in rarely flower. A collection of fruits in Puerto Rico
basal diameter. The stems and branches are gray. averaged 0.618 + 0.015 g/fruit. Air-dry seeds from
The plant produces a strong taproot with fine these fruits averaged 0.067 + 0.001 g or 15,000
laterals. Mycorrhizal bifurcation is visible on the seeds/kg. Planted in commercial potting mix, 70
tips of fine roots. The dark green, oblong leaves are percent of these seeds germinated between 30 and
sessile or subsessile and hairless or nearly so 72 days following sowing. Jungle-geranium
(Liogier 1997). The inflorescences are terminal, coppices when cut or burned and prostrate plants
dense corymbs that contain from 15 to 50 flowers send up numerous vertical shoots. Frequent
(Whistler 2000). The flowers are tubular with four suckering occurs from lateral roots out to a
or five calyx lobes. The “wild” flower color is red distance of 30 cm or more from the parent plant. It
or red-orange, but ornamental varieties may have also layers whenever a branch touches the ground.
white, yellow, salmon, or pink flowers. There are Jungle-geranium can be easily propagated from
also dwarf varieties. The fruits are fleshy, globose untreated cuttings with four leaves placed in a
berries that ripen to a dark red or purplish-black sand-peat mixture with bottom heat (Bailey 1941).
(Liogier 1997). When fully developed, there are Jungle-geranium is spreading slowly in Puerto
two seeds per fruit. However, usually one Rico, presumably dispersed by birds.
develops and one aborts in Puerto Rican plants.
Growth and Management.—Suckers and sprouts
Range.—Jungle-geranium is native to India and may grow as much as 1 m the first year, after
Sri Lanka (Griffiths 1994). It is planted worldwide which growth slows. Individual stems live at least
in tropical and subtropical climates and has 15 years; by coppicing and suckering, a plant may
naturalized in at least Puerto Rico (author’s live almost indefinitely. Maintenance of jungle-
observation, Liogier 1997). geranium in hedges requires frequent pruning. The
shrub is difficult to kill by mechanical means.
Ecology.—Jungle-geranium grows well in full sun
and light to moderate shade. It competes well with Benefits.—Jungle-geranium is one of the world’s
393
most popular tropical flowering shrubs. It is used Latha, P.G. and K.R. Panikkar. 2000. Inhibition of
for hedges and borders, for accent plants, in chemical carcinogenesis in mice by Ixora
planters and as an indoor potted plant. The species coccinea flowers. Pharmaceutical Biology 38(2):
may also be shaped into bonsais (Frommer 2001). 152-156.
In laboratory tests, extracts of jungle-geranium
have shown antibacterial (Kumer and others 1997) Liogier, H. A. 1997. Descriptive flora of Puerto
and antitumor (Latha and Panikkar 2000, Serrame Rico and adjacent islands. Vol. 5. Editorial de la
and Lim-Sylianco 1995) activity. From a methanol Universidad de Puerto Rico, San Juan, PR.
extract of the flowers, 13 chemicals were 436 p.
identified, including ursolic acid, which has known
antitumor and antiviral activity (Monteath and Michaelia, R., S. Philosoph-Hadasa, J. Riovb, and
others 2001). In the traditional medicine of India, S. Meira. 1999. Chilling-induced leaf abscission
infusions of the leaves and juice from the roots are of Ixora coccinea plants. I. Induction by
used to treat a wide variety of ailments including oxidative stress via increased sensitivity to
dysentery, ulcers, and gonorrhea (Parrotta 2001). ethylene. Physiologia Plantarum 107(2): 166-
The flowers are visited by butterflies (Collins 173.
2001), and birds eat the fruits.
Monteath, S.A.F.A., V.V. Veiga, A.C. Pinto, A.
References Echevarria, and M.A.M. Maciel. 2001.
Constituentes químicos das flores de Ixora
Bailey, L.H. 1941. The standard cyclopedia of coccinea. Universidade Federal Rural do Rio de
horticulture. The MacMillan Company, New Janeiro. http://www.sbq.org.br/ranteriores/23/
York. 3,639 p. resumos /1147. 3 p.
Collins, M. 2001. Marge Collins flying flowers: Parrotta, J.A. 2001. Healing plants of peninsular
Marge’s story. http://www.ffgc.org/horticulture India. CAB International, Wallingford, UK and
/flgardens/collins/garden_collins.htm. 4 p. New York. 944 p.
Florida Gardener. 2001. Plant of the month: Ixora Serrame, E. and C.Y. Lim-Sylianco. 1995. Anti-
coccinea. http://www.floridagardener.com tumor promoting activity of decoctions and
/ixora.htm. 2 p. expressed juices from Philippine medicinal
plants. Philippine Journal of Science 124(3):
Frommer A. 2001. Growing an indoor bonsai. 275-281.
Arthur Frommer’s Budget Travel Newsletter.
http://www.dummies.com/Lifestyle/ Tropilab. 2001. Ixora coccinea L. Tropilab Inc.
Home_and_Garden/Healthy…/0-7645-5102- http://www.tropilab.com/jungle-ger.html. 1 p.
7_0019.htm. 4 p.
Watkins, J.V. and T.J. Sheeham. 1975. Florida
Griffiths, M. 1994. Index of garden plants. Timber landscape plants, native and exotic. The
Press, Portland, OR. 1,234 p. University Presses of Florida, Gainesville, FL.
420 p.
Kumer, T.K., C.E. Rajini, V.K. Sasidharan, and T.
Krishna-Kumar. 1997. Antibacterial and Whistler, W.A. 2000. Tropical ornamentals, a
antifungal activity of secondary metabolites guide. Timber Press, Portland, OR. 542 p.
from some medicinal and other common plant
species. Journal of Life Science 2: 14-19.
394
Jacquinia arborea Vahl barbasco
THEOPHRASTACEAE
Synonyms: Jacquinia armillaris Jacq.

Jacquinia barbasco Mez in Engl.
Chrysophyllum barbasco Loefl.
John K. Francis
from the J. armillaris group: J. keyensis Mez from
the Northern West Indies (Florida and the
Bahamas), J. arborea in the Central Caribbean, and
J. armillaris Jacq. in the Lesser Antilles, northern
South America, and eastern Brazil. Stearn (1992)
gives the range of barbasco as the Cayman Islands,
Islands, the Lesser Antilles, Trinidad, and Curaçao.
He has J. arborea and J. armillaris coexisting in
the Lesser Antilles. The Missouri Botanical Garden
Herbarium lists specimens for Jamaica, Puerto
Rico, the Virgin Islands, Quintana Roo, Mexico,
and Honduras (Missouri Botanical Garden 2002).
Barbasco is reported to have naturalized on Key
Largo, Florida (Nelson 1996).
Ecology.—Barbasco is relatively common along

undisturbed coasts. Because the species is
intolerant of shade and not very competitive, it
usually grows on rocks and rocky headlands,
exposed ridges and slopes, and dry coastal forests.
In Puerto Rico, barbasco grows in areas that
receive from 750 to 1700 mm of annual
General Description.—Barbasco, also known as 100 m. It is tolerant of heavy salt spray and
torchwood, azúcares, and bizcocho, is an evergreen moderately salty soils. Barbasco grows on well-
shrub or small tree to 4.5 m in height and 15 cm in drained, mildly acid to mildly alkaline soils of all
stem diameter. It has a frequently thickened textures derived from both sedimentary and
rootstalk and one or a few stems. The bark is dark igneous rocks. It is frequently seen growing out of
brown and finely fissured, and the inner bark is cracks in limestone rocks. The species appears to
light brown and bitter. The wood is light brown or have no major insect or disease problems.
yellowish and hard. The crowns are compact and
often wind-hedged in their sea-side habitat. The Reproduction.—Barbasco flowers from winter to
thick and stiff, dull yellow-green leaves are summer and matures fruits from spring to fall
spatulate or obovate, alternate or in threes, and (Little and others 1974). Fruits collected in Puerto
clustered at the ends of the twigs. Terminal Rico averaged 0.254 + 0.005 g/fruit. Air-dried
racemes are several to many-flowered. The small, seeds cleaned from those fruits averaged 0.0363 +
white or yellowish, five-lobbed flowers are 0.0007 g/seed or 27,500 seeds/kg. Sown without
fragrant. The fruit (berries) are globose 8 to 12 mm any pre-treatment in commercial potting mix, 100
in diameter, orange-red, and contain one to four percent of the seeds germinated between 40 and 60
rounded, brown seeds (Howard 1989, Liogier days of sowing. Seeds are easily collected by hand
1995, Little and others 1974). and can be cleaned by maceration and wet sieving.
The seeds are probably dispersed by birds.
Range.—There has been confusion about the Seedlings are not common.
botanic identity of barbasco. Stahl (1992)
concluded that three species should be recognized
395
Growth and Management.—Growth of barbasco Liogier, H.A. 1995. Descriptive flora of Puerto
seedlings in the nursery is very slow. A group of Rico and adjacent islands. Vol. 4. Editorial de la
seedlings averaged only 6.2 cm 1 year after Universidad de Puerto Rico, San Juan, PR.
pricking into containers. However, survival was 617 p.
relatively good (82 percent). Growth of native-
grown plants of all ages is also believed to be very Little, E.L., Jr., R.O. Woodbury, and F.H.
slow. They seem to be long-lived. Probably the Wadsworth. 1974. Trees of Puerto Rico and the
best strategy for management of native stands is Virgin Islands. Vol. 2. Agriculture Handbook
strict protection from fire, harvesting, and 449. U.S. Department of Agriculture,
development. Washington, DC. 1,024 p.
Benefits.—Barbasco helps hold the soil in Missouri Botanical Garden. 2002. Current
topography where there is little soil to lose and specimen list for Jacquinia arborea.
contributes significantly to the aesthetics of island http://mobot.org/cgi-bin/search_vast. 1 p.
coastlines. The fruits were once used to poison or
stupefy fish and because of that are suspected to be Nelson, G. 1996. The shrubs and woody vines of
poisonous to humans (Little and others 1974). The Florida. Pineapple Press, Inc., Sarasota, FL.
sister species J. keyensis is used as an ornamental 391 p.
(Nelson 1996), and probably barbasco has been
used as well. It is a pretty and well-shaped plant, Stahl, B. 1992. On the identity of Jaquinia
suited for borders and low backgrounds. armillaris (Theophrastaceae) and related
species. Brittonia 44 (1): 54-60.
References
Stearn, W.T. 1992. The genus Jacquinia
Howard, R. A. 1989. Flora of the Lesser Antilles, (Theophrastaceae) in Jamaica. Nordic Journal of
Leeward and Windward Islands. Vol. 6. Arnold Botany 12(2): 231-238.
MA. 658 p.
396
Jasminum fluminense Vell. Brazilian jasmine
OLEACEAE
Synonyms: Jasminum azoricum auct., not L.

Jasminum bahiense DC.
John K. Francis
Seychelles, Arabia, Ethiopia, southern Zimbabwe,

Malawi, Mozambique, Angola, Nigeria, South
Africa (Miami-Dade County 2001), the Azores,
and the Canary Islands (Acevedo-Rodríguez 1985).
The species has naturalized and escaped
throughout much of the tropics and subtropics
including southern Florida (Florida Exotic Plant
Council 2001), Puerto Rico (Liogier 1995),
Hawaii, and Guam (Pacific Island Ecosystems at
Risk 2001).
Ecology.—Brazilian jasmine occurs in Puerto Rico

in areas that receive from about 750 to 1800 mm of
annual rainfall. It grows from near sea level to
more than 600 m in elevation. Soils of all textures
and parent material are colonized. However, it does
General Description.—This plant became known not tolerate poorly drained soils. The species is
as Brazilian jasmine because it was first described restricted to areas with minimum temperatures
from Brazil. It is of African origin, having been above 1.7 °C (Florida Exotic Plant Council 2001).
introduced earlier by Portuguese settlers (Florida It will grow on the coast in areas that do not
Exotic Plant Council 2001). The species is also receive salt spray (Florida Exotic Plant Council
known as jasmine, Gold Coast jasmine, jasmín 2001). Brazilian jasmine will grow in partial shade
oloroso, jasmín de canario, jasmín de trapo, jasmin and climb upward to better light. It can survive but
à bouguet, and jasmin blanc (Acevedo-Rodríguez is not aggressive in the denser shade of unbroken
1985, Howard 1989, Pacific Islands Ecosystems at forest canopies. Most plants grow in natural and
Risk 2001). Brazilian jasmine is a woody artificial openings in the forest such as fencerows,
evergreen scrambling and climbing shrub (vine) river banks, roadsides, brushy pastures, and logged
that may extend as much as 12 m along the ground or burnt-over forest.
or into the crowns of trees. The cylindrical stems,
which are commonly about 1 cm in diameter but Reproduction.—Brazilian jasmine blooms
may grow to 5 cm, have gray, fine-furrowed bark throughout the year (Miami-Dade County 2001).
and are flexible and strong. There are often Under favorable conditions, fruits are produced in
multiple stems from a root crown. Foliage is borne abundance by open-grown plants. Shaded plants
only on the current year’s growth. The opposite, produce few or no fruits. A collection of fruits of
compound leaves have three leaflets, the central Brazilian jasmine from Puerto Rico weighed an
one being larger than the lateral two. The average of 0.413 ± 0.066 g each. Air-dried seeds
inflorescences are cymes that arise from leaf axils. from those fruits averaged 0.164 + 0.002 g/seed or
The tubular white flowers are fragrant, especially 6,000 seeds/kg. Sown on commercial potting mix,
at night. The fruits, which form in groups of two, 100 percent of the seeds collected from this sample
are globose, 8 mm in diameter, and dark purple or germinated between 18 and 52 days after sowing.
dark blue to almost black when ripe (Acevedo- Brazilian jasmine roots (layers) whenever stems
Rodríguez 1985, Howard 1989, Liogier 1995). The come in contact with the ground. The seeds are
fruits have a bitter, disagreeable flavor. Each fruit dispersed by birds and mammals (Miami-Dade
contains one spherical gray seed. County 2001) and by the lateral extension of the
stems. Seedlings are common in suitable habitat in
Range.—The African and Middle Eastern native Puerto Rico. Ornamental plants are propagated by
range of Brazilian jasmine includes Mauritius, the
397
means of cuttings (Florida Exotic Plant Council Bailey, L.H. 1941. The standard cyclopedia of
2001). horticulture. The MacMillan Company, New
York, NY. 3,639 p.
Growth and Management.—The stems of
established plants may extend as much as 2 or 3 m Florida Exotic Plant Council. 2001. Jasminum
in 1 year. Seedlings grow much more slowly. fluminense Vell. www.fleppc.
Brazilian jasmine can engulf shrubs, small trees, org/pdf/Jasminum%20fluminense.pdf. 2 p.
and fences and can ascend power poles. Control of
thickets and mats is often needed. Young plants Howard, R.A. 1989. Flora of the Lesser Antilles,
can be pulled up by hand. Older plants should be Leeward and Windward Islands. Dicotyledoneae.
cut at the ground level and the stumps treated with Vol. 6. Arnold Arboretum, Harvard University,
herbicide. Follow-up treatments will probably be Jamaica Plain, MA. 658 p.
required (Pacific Island Ecosystems at Risk 2001).
Benefits.—Brazilian jasmine has been widely Rico and adjacent islands. Vol. 4. Editorial de la
planted as an ornamental, both in greenhouses and Universidad de Puerto Rico, San Juan, PR. 617
outdoors (Bailey 1941, Liogier 1995). Because of p.
its invasive nature, the species should not be
planted where it might escape to the wild (Nelson Miami-Dade County. 2001. Brazilian jasmine-
1996). It is a food source for birds and mammals. Jasminum fluminense. www.co.miami-
The fruits are heavily consumed by raccoons in dade.fl.us/derm/environment/badplants/plant%2
Florida (Miami-Dade County 2001). Goats browse 0…/brazilian_jasmine.ht. 1 p.
the foliage.
Nelson, Gil. 1996. The shrubs and woody vines of
References Florida. Pineapple Press, Inc. Sarasota, FL.
391 p.
Puerto Rico. Vol. 1. General Technical Report Pacific Island Ecosystems at Rick. 2001. Invasive
SO-58. U.S. Department of Agriculture, Forest plant species: Jasminum fluminense Vell.,
Service, Southern Forest Experiment Station, Oleaceae. www.hear.org/pier/jaflu.htm. 2 p.
398
Jatropha gossypiifolia L. red physic nut
EUPHORBIACEAE
Synonyms: Adenoropium gossypifolium Pohl

Jatropha elegans (Pohl) Klotzsch in Seemann
John K. Francis
known, but today it grows wild from Florida

through the West Indies and from Mexico through
tropical South America (Howard 1989). It has
naturalized in most tropical areas of the world
(Burkill 1994).
Ecology.—Red physic nut grows on nearly all

types of soils within its range. In Puerto Rico, it
may be found in areas receiving from 750 to about
2000 mm of annual precipitation. The species is
more common in soils with high base saturation,
such as dry areas, sites near the ocean, and soils
derived from limestone. Red physic nut usually
requires disturbance to establish itself and does not
compete well with tall grass or heavy weed growth.
It is intolerant of shade. Although plants may
survive for a season in moderate shade, they need
full or nearly full sun for longer-term survival and
fruiting. Cattle and horses generally avoid red
physic nut, except in extreme shortages of forage,
General Description.—Red physic nut, also so that the species in often abundant in overgrazed
known as belly-ache bush, figus nut, wild cassava, range and poorly managed pastures. It is also
higuereta cimarrona, tautuba, túatúa, purga del common in waste lands, roadsides, poorly tended
fraile, médicinier noir, médicinier bâtard, agricultural fields, and river overflow areas.
médicinier rouge, is a short-lived perennial shrub
0.5 to 2 m in height and 1 to 3 cm in basal Reproduction.—Flowering in India occurs from
diameter. It may be evergreen or deciduous, February through July (Parrotta 2001). Sometimes
depending on climate. The plant usually has a both flowers and fruits will be present at the same
single stem and is woody at the base. The small time on plants. Upon drying, the capsule valves
number of branches are stout, green, and semi- spring open propelling the seeds a few centimeters.
woody. A cloudy, yellow sap collects when stems It is not known if there are other seed dispersal
and branches are cut. Red physic nut has a short mechanisms. Seeds collected in Puerto Rico
taproot, robust laterals, and many fine tertiary roots averaged 0.017 ± 0.001 g/seed or 59,000 seeds/kg.
and is usually not deeply rooted. The opposite The seeds of this collection were highly variable in
leaves are three- or five-lobed, 3 to 13 by 3 to 19 weight (C.V.= 53.7 percent) and just 4 percent
cm, reddish or purplish tinged when young and germinated. Collecting seed in quantity is difficult
green when fully developed. Small red or purple because capsules are mature for only a short period
flowers are borne in terminal corymbose cymes. before discharging their seeds, and there are only a
The fruits that follow are oblong, three-lobed few capsules per large plant at any time.
capsules that are green, turning tan at maturity.
Within are three mottled, gray-brown or grayish- Growth and Management.—Red physic nut
red seeds (Burkill 1994, Howard 1989, Liogier usually grows without branches during its first year
1988, Parrotta 2001). and adds adventitious branches in the year or two
following. Growth is about 0.5 m per year and
Range.—Red physic nut is a native of the New plants usually live 2 or 3 years. The species is
World (Burkill 1994). The original range is not
399
considered an agricultural weed but can be Horsten, S.F., A.J. van den Berg, J.J. Kettenes-van
suppressed by common weed-control practices. den Bosch, B.R. Leeflang, and R.P. Lababie.
1996. Cyclogossine A: a novel cyclic
Detriments and Benefits.—Red physic nut heptapeptide isolated from the latex of Jatropha
contains toxins in the seeds, sap and other tissues, gossypifolia. Planta Med 62(1): 46-50.
capable of killing humans (Marcano-Fondeur
2002). Apparently, poisoning cases are rare. Howard, R.A. 1989. Flora of the Lesser Antilles,
Extracts of the plant are used as a purgative and Leeward and Windward Islands. Vol. 5. Arnold
emetic, and to treat headache, diarrhea, venereal Arboretum, Harvard University, Jamaica Plain,
disease, skin sores, mouth sores, and cancer MA. 604 p.
(Burkill 1994, Parrotta 2001). The use of the seeds
in herbal medicine is advised against because of Liogier H.A. 1988. Descriptive flora of Puerto
their high toxicity (Liogier 1990). Phytoactive Rico and adjacent islands, Spermatophyta. Vol.
chemicals found in tissues of red physic nut 2. Editorial de la Universidad de Puerto Rico,
include the protease curcain, the alkaloid Río Piedras, PR. 481 p.
jatrophine, the cyclic heptapeptide, cyclogossine A,
saponins, and numerous fatty acids (Burkill 1994, Liogier, H.A. 1990. Plantas medicinales de Puerto
Horsten and others 1996, Marcano-Fondeur 1992, Rico y del Caribe. Iberoamericana de Ediciones,
Ogbobe and Akano 1993). The seeds contain 36 Inc., San Juan, PR. 566 p.
percent oil (Ogbobe and Akano 1993) and are used
as an illuminant in Africa (Burkill 1994). Swaths of Marcano-Fondeur, E. de J. 1992. Las plantas
plants are frequently established around villages in venenosas en la medicina popular. Naturaleza
Africa as fire barriers. The species is also attributed Dominicana. http://marcano.freeservers.com/
magical powers that protect against lightning, nature/conference/venen.html. 6 p.
snakes, and violence (Burkill 1994). The wood is
soft and weak and of little use. A red-leafed Ogbobe, O. and V. Akano. 1993. The physico-
variety is used as an ornamental (Howard 1989). chemical properties of the seed and seed oil of
Jatropha gossipifolia. Plant Foods for Human
References Nutrition 43(3): 197-200.
Burkill, H.M. 1994. The useful plants of West Parrotta, J.A. 2001. Healing plants of Peninsular
Tropical Africa. Vol. 2. Royal Botanic Gardens, India. CABI Publishing, Wallingford, UK and
Kew, UK. 636 p. New York. 917 p.
400
Juniperus communis L. common juniper
CUPRESSACEAE
Synonyms: Juniperus canadensis Lodd. ex Burgsd.

Juniperus alpina (Sm.) S.F. Gray
Juniperus sibirica Burgsd.
listed as extremely rare or critically imperiled in

many Southeastern States, including Georgia
(Georgia Natural Heritage Program 2001) and
Alabama (Alabama Natural Heritage Program
1996), or of likely rarity in South Carolina (South
Carolina Heritage Trust 2000).
Range.—The range of common juniper is

extensive. It is the only circumpolar conifer in the
northern hemisphere (Pojar and Mackinnon 1994).
It is found across the U.S.A. and Canada to
Greenland, through Europe, across Siberia and
Asia. However, within its range there are at least
five sub-species or varieties (FNA 1993). For
example, var. depressa occurs throughout much of
Canada and the United States where it occurs in
the Southwestern States of Arizona, New Mexico,
Colorado, and Utah and extends west to California
and east to North Carolina through Arkansas; var.
megistocarpa occurs in Nova Scotia,
Newfoundland, and Quebec; var. montana occurs
in Greenland, British Columbia, and California,
Oregon and Washington States (FNA 1993); var.
communis and var. nana occur in Britain and
Ireland (Stace 1997). Larger specimens of var.
depressa in the United States have frequently been
misidentified as var. communis (FNA 1993).
Hunt Institute
Ecology.—Common juniper is a hardy shrub that
General Description.—Common juniper is
grows in a wide range of ecological conditions.
known by a variety of common names: dwarf, low,
Typically it grows on dry, open, rocky slopes and
mountain, alpine, prostrate or Siberian juniper,
mountainsides but may be found in stressed and
ground cedar, genevrier commun, and enebro
acid environments where competition with other
(Elmore 1987, FNA 1993). There are several sub-
plants is almost non-existent (Ward 1982). It also
species or varieties, each having several
often grows in partial shade. Depending upon the
synonyms. For a list of the synonyms consult
latitude it can be found from lowland bogs at sea-
Kartesz (1994). Common juniper is a low shrub,
level to sub-alpine ridges and alpine tundra at over
with spreading or ascending branches, that
3,600 m (FNA 1993; Lanzara and Pizzetti 1977;
generally grows no more than 1 m high. Larger
Pojar and Mackinnon 1994). It is also a common
individuals, up to 10 m, have been observed. It
shrub in abandoned lowland fields in Northern
typically forms dense mats 1 to 4 m in diameter.
United States (Steele 1982).
The brown fibrous bark is smooth on small
branches (<1 cm diameter) but exfoliates into thin
Reproduction.—Common juniper has berry-like
strips on larger branches. The leaves are awl-
female cones that mature in 2 years in
shaped, in groups of three and tend to be “prickly”
Mediterranean climates and every 3 years in cooler
standing out from the stems. Common juniper is
401
climates (Lanzara and Pizzetti 1977). Male and Benefits.—Common juniper has low value for
female cones are on separate plants. The dark blue short-term re-vegetation projects but moderate to
seed (female) cones are globose to ovoid in shape, high value for long-term rehabilitation projects
6 to 13 mm in diameter and are resinous to and is useful in preventing soil erosion
obscurely woody with two to three triangular- (Tirmenstein 1999). Common juniper provides
shaped seeds. From pale green the female cones important cover and browse for wildlife such as
ripen to dark blue in late summer and then tend to mule deer (Dusek 1975). The cones are eaten by
whiten with a waxy “bloom.” The staminate cones several species of birds and are important food
are 3 to 6 mm in diameter (Cronquist and others sources for wild turkeys (Decker and others 1991),
1972). Although the cones are generally terminal Cedar and Bohemian waxwings (Catling and
in other species of juniper, in J. communis the Brownell 1998), and others. Domestic livestock
cones are axillary (FNA 1993). The fruits are of rarely utilize common juniper (Tirmenstein 1999).
relatively low nutritional quality and on average The seed cones of common juniper are used to
number 80,500 seeds/kg (Stiles 1980). The seed is flavor gin, and the word for this alcoholic
dispersed by birds, at least in North America, and beverage was derived from the Latin “iuiperus”
commonly require a long maturation and through the Old French and Dutch words
germination period (Crane and Fischer 1986; Stiles “genevre” and “genever” respectively (Elmore
1980). Because seeds do not readily germinate, 1987). The seed cones are also used as a flavoring
the establishment of seedlings, for example for re- for foods. Many Native American tribes used
vegetation projects, can be a problem (Dietz and common juniper for medicinal and ceremonial
others 1980, Stiles 1980). As well as poor purposes (Moerman 1998). It has been used
germination, poor seed dispersal has been blamed medicinally for childbirth and to treat urinary
for the relative rarity of juniper in areas where the infections (Pojar and Mackinnon 1994). However,
majority of potential habitat is not occupied it can also cause miscarriages. The different
(Diotte and Bergeron 1989). Plant age also affects varieties of common juniper make excellent,
reproduction; 40 to 60 percent of old individuals vigorous landscaping shrubs, which are readily
are sterile and up to 94.8 percent of seeds of seed- propagated by cuttings in the horticultural trade
producing old plants are non-viable. In contrast, 80 (Cronquist and others 1972).
percent of seeds produced by young plants are
viable (Diotte and Bergeron 1989). References
Growth and Management.—Common juniper is Alabama Natural Heritage Program. 1996.

often killed by fire (Crane and Fischer 1986). It NatureServe, Member Program--Plants of
has been described as having minimal “fire- Alabama. Website Last Update: 8/1/96:http://
surviving regeneration properties," and resprouting www.abi.org/nhp/us/al/plants.html 1 p.
after fire is rare (Mallik 1995). The foliage is
resinous and flammable (Diotte and Bergeron Catling, P.M. and V.R. Brownell. 1998.
1989). Mortality depends upon fire intensity. For Importance of fire in Alvar Ecosystems-
example, in eastern Canada, older common juniper Evidence from the Burnt Lands, Eastern
shrubs will survive fires of low severity, and some Ontario. Canadian Field Naturalist 112(4): 661-
fire regimes allow common juniper to survive 667.
several fires; individuals of more that 170 years
old have been observed to survive in areas exposed Crane M.F. and W.C. Fischer. 1986. Fire Ecology
to fire (Diotte and Bergeron 1989). However, it of the Forest habitat Types of Central Idaho.
should be noted that the patchy nature of fires General Technical Report INT-218. USDA
leads to refugia for juniper shrubs, and so Forest Service Intermountain Research Station,
particular individuals can escape from harm and Ogden, Utah. 85 p.
provide seed for re-vegetation of the burnt areas
(Diotte and Bergeron 1989). Re-establishment of Cronquist, A., A.H. Holmgren, N. H. Holmgren,
common juniper as a browse species after and J.L. Reveal. 1972. Intermountain Flora.
disturbance and fire has had varying success Volume 1. Vascular Plants of the Intermountain
(Dietz and others 1980). In the Black Hills, South West, U.S.A. New York Botanical Garden.
Dakota, the only successful means for establishing Hafner Publishing Co. New York. p. 238-240.
common juniper was from nursery stock, and seed
germination was particularly poor. Decker, S. R., P. J. Pekins, W. W. Mautz. 1991.
Nutritional evaluation of winter foods of wild
402
turkeys. Canadian Journal of Zoology. 69 (8): Mallik, A. U. 1995. Conversion of temperate
2,128-2,132. forests into heaths: role of ecosystem
disturbance and ericaceous plants.
Dietz, R.D., D.W. Uresk, H.E. Messner, and L.C. Environmental Management. 19(5): 675-684.
McEwen. 1980. Establishment, Survival, and
Growth of Selected Browse Species in a Moerman, D.E. 1998. Native American
Ponderosa Pine Forest. General Technical Ethnobotany. Timber Press, Inc, Portland, OR.
Report RM-219. USDA Forest Service Rocky 927 p.
Mountain Forest and Range Experiment Station.
Fort Collins, CO. 12 p Pojar, J. and A. Mackinnon. 1994. Plants of the
Pacific Northwest Coast. British Columbia
Diotte, M. and Y. Bergeron. 1989. Fire and the Ministry of Forests and Lone Pine Publishing,
distribution of Juniperus communis L. in the Vancouver, Canada. 527 p.
boreal forest of Quebec, Canada. Journal of
Biogeography 16: 91-96. Stace, C. 1997. New Flora of the British Isles.
Second ed. Cambride University Press,
Dusek, L. 1975. Range relations of Mule Deer and Cambridge, England. 1,130 p.
cattle in prairie habitat. Journal of Wildlife
Management. 39(3): 605-616.
Steele, F.L. 1982. At Timberline: A Nature Guide
to the Mountains of the Northeast. Appalachian
Elmore. F.H. 1987. Shrubs and Trees of the
Mountain Club, Boston, MA. 285 p.
Southwest Uplands. Southwest Parks and
Monuments Assoc. Tucson, AZ. 214 p.
South Carolina Heritage Trust. 2000. Website.
http://www.abi.org/nhp/us/sc/speclist.htm. 1 p.
Flora of North America Editorial Committee.
1993. Flora of North America. Vol. 2. Oxford
Stiles, E.W. 1980. Patterns of fruit presentation and
University Press, New York. 475 p. seed dispersal in bird-disseminated woody plants
in the eastern deciduous forest. The American
Georgia Natural Heritage Program. 2001. Naturalist. 116(5): 670-688.
http://www.dnr.state.ga.us/dnr/wild/natural/sppl
_t.htm. 1 p Tirmenstein 1999. Juniperus communis. U.S.
Kartesz, J.T. 1994. A synonymized checklist of the Rocky Mountain Research Station, Fire Sciences
vascular flora of the United States, Canada, and Laboratory. Fire Effects Information System,
Greenland. Vol.1 – Checklist. 2nd Ed. Timber http://www.fs.fed.us/database/feis/ [not paged].
Press, Portland, OR. 622 p.
Ward, L.K. 1982. The Conservation of Juniper.
Lanzara, P. and M. Pizzetti. 1977. Simon and Journal of Applied Ecology. 19: 165-188.
Schuster’s Guide to Trees. Simon and Schuster
Inc., New York. 316 p.
403
Juniperus osteosperma (Torr.) Little Utah juniper
CYPRESSACEAE
Synonyms: Juniperus utahensis (Engelm.) Lemmon

Juniperus californica Carr. ssp. osteosperma (Torr.) E. Murr.
Juniperus californica Carr. var. utahensis Engelm.
Juniperus knightii A. Nels.
Juniperus megalosperma Sudworth
Juniperus occidentalis Hook. var. utahensis (Engelm.) Kent
Sabina osteosperma (Torr.) Antoine
John K. Francis
gray-brown or gray, thin, fibrous, and shredding.

Sapwood is white and the heartwood is yellow-
brown with narrow annual rings. It is relatively
soft, brittle, and has a dry-weight specific gravity
of 0.553 g/cm3 (Paettie 1953). The wood and all
other parts are aromatic. Crowns tend to be
branchy, rounded to flat-topped, and sometimes
dense; old trees often have bleached, dead limbs.
Utah junipers have deep taproots and lateral roots
that may extend as much as 30 m, a few cm below
the soil surface. The roots are colonized by
vasicular-arbuscular mycorrhizae. The
photosynthetic organs are yellow-green branchlets
with opposite (or whorled), reduced scale-like
leaves. The species is monoecious with small,
separate male and female flowers. Fruits are
subglobose, 6- to 12-mm diameter, leathery cones
(“berries”), brownish to blue-purple at maturity.
They have a strong, resiny flavor and contain one
(sometimes two) hard seed (Abrams 1940, Welsh
and others 1987, Zlatnik 1999).
Range.—Utah juniper occurs as large populations

and scattered disjuncts in southern Idaho, southern
Montana, Wyoming, western Colorado, Utah,
Nevada, Arizona, eastern and southern California,
northeastern New Mexico, and Chihuahua and
Durango in Mexico (Natural Resources
General Description.—Utah juniper, also known Conservation Service 2003, Secretaría de Medio
as cedar, desert cedar, sabina, and sabina morena, Ambiente y Recursos Naturales 2003, Tree Guide
has a habit between that of shrub and tree. 2003). The species is not known to have
Although slow-growing and long-lived, rare naturalized outside its native range. It hybridizes
individual plants attain large diameters (record with J. occidentalis Hook. where the species grow
tree: 217 cm diameter breast height and height together (Terry and others 2000).
11.6 m, American Forests 1998). The species is
short statured, (mainly 2 to 4 m tall), (Welsh and Ecology.—Utah juniper grows on a wide variety
others 1987), usually has multiple stems (limbs of well-drained soils. It accommodates all the soil
arising near the ground that become codominant textures, pH’s of 6.5 to 8.2, high free calcium
with the original central stem), and becomes carbonate, but does not tolerate salty soils (Natural
reproductively mature at small sizes. The bark is Resources Conservation Service 2003). It is
404
commonly found on rocky, gravely, and sandy Growth and Management.—Utah juniper grows
sites, usually originating from igneous (Stuart and slowly, usually only 0.13 cm/year in diameter. An
Sawyer 2001) and sedimentary (sandstone and 86-year-old stand in Utah had 17.2 percent canopy
limestone) rocks, at 900 to 2,500 m above sea cover and a basal area of 7.7 m2/ha (Zlatnik 1999).
level (Zlatnik 1999). Mean annual precipitation Poor site conditions may result in 15 cm-tall plants
varies from 300 to 510 mm, but the species can over 50 years old. Individuals of the species may
withstand severe droughts (Natural Resources live 650 years or longer. Closing Utah juniper
Conservation Service 2003) and is a fierce stands eliminate most forage plants; killing trees
competitor for the little available moisture in its can result in a doubling of herbaceous cover
habitat. It has a specific leaf area of 18 cm2/g (Zlatnik 1999). Controlling Utah juniper has been
(Grier and others 1992). Utah juniper also tolerates attempted with fire, but fires on these sites are
cold and heat well. However, the species is difficult to start under humid, still conditions and
intolerant of shade and does not survive under the hard to control under dry, windy conditions.
canopy of taller vegetation. It is alleged to exert an Controlled burns can be done with the greatest
allelopathic effect on some understory grass precision in late winter or early spring when the
species. Utah juniper is a climax species in many wood is low in water content (personal
vegetation types with species such as Artimisia communication with Stanley Kitchen, Shrub
tridentata Nutt., Pinus edulis Engelm., and Pinus Sciences Laboratory, USDA Forest Service,
monophylla Torr. & Frém. The range of Utah Provo, UT). The shrubs and trees can be killed
juniper is known to have expanded since European with chainsaws or herbicides, but the preferred
colonization, and stands have changed from open, method is “chaining,” dragging sections of anchor
savanna types to dense stands probably because of chain between two bulldozers. Many young trees
reduction of fine fuels by livestock grazing, fire are not killed, but the treatment does open stands
suppression, and changes toward warmer and drier and allows greater shrub and herbaceous growth
climates (Tausch 1999). Utah juniper is easily (Plummer and others 1968, Stevens 1999).
killed by fire, both by scorching the top or heat-
girdling through the thin bark. However, the Benefits.—Utah juniper is a major constituent of
habitat seldom has sufficient fine fuels to sustain a semidry ecosystems of the West. It protects the
fire. Stands most at risk are young stands with soil from wind erosion but in some instances may
scattered trees in brush and grass, and continuous aggravate sheet erosion from rains by suppressing
stands of large, decadent trees during high winds understory grasses, forbs, and shrubs. The species
(Zlatnik 1999). Utah juniper is sometimes heavily furnishes cover and reproductive habitat for many
infested by the mistletoes, Phoradendron species and is a major source of food for wildlife,
juniperinum Engelm. ex Gray and P. bolleanum especially during winter. Many birds, rodents, as
(Seem.) Eichl. (Zlatnik 1999). The larvae of well as jackrabbits (Lepus californicus) and
Eurytoma juniperina Marcovitch attack the seeds coyotes (Canis latrans) eat the fruits and mule
(Bonner 2003). deer (Odocoileus hemionus), and elk (Cervus
elaphus) browse the foliage when better food is
Reproduction.—Utah juniper flowers from not available. It is not eaten by cattle and is
January to April (depending on site) and is wind browsed to a limited extent by domestic sheep
pollinated. Fruits ripen in the second year and (Stuart and Sawyer 2001, Zlatnik 1999). The
germinate the following spring or the second forage is of low quality because of low
spring after dispersal (Zlatnik 1999). Fruits may be digestibility (44 percent) and low protein content
collected by hand from the shrubs or low trees or (6 to 8 percent) (Zlatnik 1999). The wood is used
picked up by hand after they have fallen. Seed for firewood, charcoal, and novelties (Secretaría de
should be extracted by maceration with water Medio Ambiente y Recursos Naturales 2003). The
before the fruits dry out. Cleaned seed from bark is used as tinder and formerly was used as a
Arizona ranged from 7,900 to 15,700 seeds/kg. “slow match” to maintain and transport fire (Plants
Seed should be dried to about 10 percent moisture For a Future 2003). Native Americans used the
and can be stored at 5 to -18 °C for several years. bark to make sandals, mats, bedding, baskets,
Moist stratification at 3 to 5 °C for 30 to 180 days thatch, and cordage, and they made meal from the
is necessary for good germination. Drilled or fruits (Peattie 1953). The wood is highly decay
broadcast in a nursery bed, seeds should be resistant, which makes it desirable for fence posts
covered with 6 mm of fine soil or sand. and rude construction despite its poor form
Germination ranges from 50 to 64 percent (Bonner (Zlatnik 1999). Tissues of Utah juniper are not
2003). high enough in cedarwood oil to be commercially
405
valuable. Heartwood contains 1.19 percent oil, Utah. Publication 68-3. Utah Division of Fish
principally thujopsene (40 percent) and cedrol and Game, Salt Lake City, UT. 183 p.
(13.2 percent) (Adams 1987). The species was
used by Native Americans to treat bladder and Secretaría de Medio Ambiente y Recursos
kidney troubles, wounds, and mouth sores (Plants Naturales. 2003. Especies con usos no
for a Future 2003). Utah juniper has been used to a maderables en bosques de Encino, pino y pino-
limited extent in revegetation projects and encino: Juniperus osteosperma (Torr.) Little.
xeriscape gardening. http://www.semarnat.gob.mx/pfnm/Juniperus
Osteosperma.html. 2 p.
References
Stevens, R. 1999. Mechanical chaining and
Abrams, L. 1940. Illustrated flora of the Pacific seeding. In: S.B. Monson and R. Stevens,
States. Vol. 1. Stanford University Press, comps. Proceedings: ecology and management
Stanford, CA. 538 p. of pinyon-juniper communities within the
Interior West; Sep. 15-18, 1997. Provo, UT.
Adams, R.P. 1987. Investigation of Juniperus PMRS-P-9. U.S. Department of Agriculture,
species of the United States for new sources of Forest Service, Rocky Mountain Research
cedarwood oil. Economic Botany 41(1): 48-54. Station, Ogden, UT. p. 281-284.
American Forests. 1998. National register of big Stuart, J.D., and J.O. Sawyer. 2001. Trees and
trees 1998-1999. American Forests 104(1): 24- shrubs of California. University of California
46. Press, Berkeley, CA. 467 p.
Bonner, F.T. 2003. Juniperus L., juniper. In: F.T. Tausch, R.J. 1999. Historic pinyon and juniper
Bonner and R.G. Nisley, eds. Woody Plant Seed woodland development. In: S.B. Monson and R.
Manual. U.S. Department of Agriculture, Forest Stevens, comps. Ecology and management of
Service, Washington, DC. 15 p. pinyon-juniper communities within the Interior
West. Proc. RMRS-P-9. U.S. Department of
Grier, C.C., K.J. Elliot, and D.G. McCullough. Agriculture, Forest Service, Rocky Mountain
1992. Biomass distribution and productivity of Research Station, Ogden, UT. p. 12-19.
Pinus edulis-Juniperus monosperma woodlands
of north-central Arizona. Forest Ecology and Terry, R.G., R.S. Nowark, and R.J. Tausch. 2000.
Management 5093-4): 331-350. Genetic variation in chloroplast and nuclear
ribosomal DNA in Utah juniper (Juniperus
Natural Resources Conservation Service. 2003. osteosperma, Cupressaceae): evidence of
Plants profile: Juniperus osteosperma (Torr.) interspecific gene flow. American Journal of
Little, Utah juniper. http://plants.usda.gov/ Botany 87: 250-258.
cgi_bin/plant_search.cgi?mode=Scientific+Nave
&keywordquery=Juniperus+osteosperma&earl= Tree Guide. 2003. Treeguide species details: Utah
plant_sea. 5 p. juniper. http://www.treeguide.com/Species.
asp?SpeciesID=544. 3 p.
Peattie, D.C. 1953. A natural history of Western
trees. Houghton Mifflin Company, Boston, MS. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
749 p. Higgins, eds. 1987. A Utah flora. Brigham
Young University, Provo, UT. 894 p.
Plants For a Future. 2003. Plants for a future:
database search results: Juniperus osteosperma. Zlatnik, E. 1999. Juniperus osteosperma. In: U.S.
Plants for a Future, Cornwall, UK. http://www. Department of Agriculture, Forest Service,
ibiblio.org/pfaf/cgi-bin/arr_html?Juniperus+ Rocky Mountain Research Station, Fire Sciences
osteosperma&CAN=LATIND. 8 p. Laboratory. Fire Effects Information System.
http://www.fs.fed.us/database/feis/plants/tree/
Plummer, A.P., D.R. Christensen, and S.B. junost/all.html. 33 p.
Monson. 1968. Restoring big-game range in
406
Kalmia latifolia L. mountain laurel
ERICACEAE
Synonyms: Kalmia latifolia var. laevipes Fern.
W. Henry McNab and Barton D. Clinton
in the western United States, throughout Europe,

and in New Zealand and Australia.
Ecology.―Mountain laurel occurs most

commonly on dry sites of acid soils, although it
occasionally occurs on well-drained floodplain
soils, and it will tolerate clay soils that are not
waterlogged. Mountain laurel forms dense, almost
impenetrable patches known as "laurel hells" or
"ivy thickets," particularly on upper slopes and
ridges where the tree canopy is sparse or lacking.
In the southern Appalachians, mountain laurel
occurs at elevations to 1500 m and the probability
of its occurrence increases on sites with a thin soil
A horizon (McNab et al. 1999), and likely low
available water capacity. Lipscomb and Nilsen
(1990) found that mountain laurel has high water-
use efficiency, which could give it a strong
competitive advantage on relatively hot and dry
General Description.―Mountain laurel, known southwest slopes. Mountain laurel adapts to a
also as calico-bush, ivy, ivybush, kalmia, laurel, range of light regimes, from little to moderately
and spoonwood, is a medium to large (2 to 9 m in dense shade, typically beneath an oak (Quercus
height) evergreen shrub, often with contorted stem spp.) or pine (Pinus spp.) canopy. The ratio of
and branching habit. The thick, flat, leathery foliage chlorophylls a to b is lower (1.5) in Kalmia
leaves are shiny, dark-green above and pale-green than for most understory species and may benefit
below; margins are entire. Leaf length ranges from this adaptation (Al-Hamdani and others 2002).
5 to 12 cm and width from 1.5 to 5 cm. The Rosebay rhododendron (Rhododendron maximum
yellowish green petioles range from 0.7 to 3 cm L.) is a common associate of mountain laurel on
long. The thin, reddish-brown bark shreds and subxeric/submesic ecotones in the southern
subdivides into ridges. In the absence of Appalachians. A mycorrhizal association has been
disturbance (usually fire), the species is long-lived: demonstrated for seedlings of mountain laurel
40 to 60 years (Chapman 1950). Botanical (Flemer 1949). Natural stands of mountain laurel
characteristics of the genus Kalmia have been are little affected by insects, although at epidemic
studied intensively (Ebinger 1974). Kalmia is the levels caterpillars of gypsy moths (Lymantria
state flower of Connecticut and Pennsylvania. dispar L.) and elm spanworms (Ennomos
subsignaria Hubner) will feed on the foliage. The
Range.―Mountain laurel is primarily a shrub of most serious disease is phytophthora root rot
the Appalachian Mountain region in the eastern (Phytophthora cinnamomi Rands), which can
U.S. It also occurs in the Atlantic Coastal Plain infect damaged plants (Jaynes 1997). Foliage of
from Maine to Virginia, throughout the mountain laurel near roadsides is particularly
Appalachian Piedmont, southwest into Mississippi, sensitive to sodium chloride (NaCl) used for de-
and west into much of Tennessee, Kentucky, and icing roads in New England (Bryson and Baker
southern Indiana and Ohio. Reports of native 2002). Although not shown in mountain laurel,
mountain laurel in Canada have not been other Kalmia species are allelopathic. Waterman
confirmed and are likely ornamentals or large- and others (1995) found that Kalmia thickets do
leaved specimens of sheep laurel [Kalmia not exclude tree seedlings but inhibit their growth
angustifolia (Small) Fernald] (Jaynes 1997).
Kalmia has been planted widely as an ornamental
407
by reducing light and increasing competition for Benefits and Detriments.―Mountain laurel has
water. long been cultivated as an ornamental--since about
1740 when American naturalist John Bartram sent
Reproduction.―The showy, white to pink 2- to plants to England (Jaynes 1997). Cultivars of
3- cm saucer-shaped flowers form convex clusters mountain laurel are particularly valuable as
at the ends of branches and appear after the ornamentals and are propagated commercially by
foliage, from March to July, depending on latitude many nurseries. Nearly 100 horticultural varieties
and altitude (Olson and Barnes 1974). Purple spots of have been selected for variation in flower color,
mark 10 anther pockets in the petals. The method leaf variegation, and growth habit (Jaynes 1997).
of pollen dispersal in Kalmia is unusual in that the Mountain laurel burls have been used for making
anthers are under tension as the flower matures, pipes for smoking tobacco, and early European
and they spring forward when disturbed by an settlers in America used the fine-grained wood for
insect, forcefully and effectively transferring making eating utensils and weaver's shuttles, and
grains of pollen to the insect (Jaynes 1997). The produced a yellow dye from its foliage. Tinctures
flowers are pollinated by more than a dozen of fresh Kalmia leaves have been used for home
insects, primarily bumblebees, but generally not medical remedies of skin disorders. An
honeybees, likely because the flowers produce unidentified anticancer compound has been
little nectar (Jaynes 1997). Glandular hairs on the extracted from its sap (Jaynes 1997). Mountain
flower stalk, calyx, and corolla may deter laurel provides cover and stability to thin soils,
nonflying insects. Flowers are most abundant on however its presence can reduce water yields
plants in full sunlight and occur rarely on plants in (Johnson and Kovner 1956). Foliage of mountain
dense shade (Kurmes 1961). Fruit ripens in late laurel is toxic to domestic cattle and sheep but
summer and consists of a brown, erect, five-celled, provides subsistence winter food for ruffed grouse
globular capsule, 4 to 7 mm wide. Each capsule (Bonasa umbellus L.) and white-tailed deer
contains about 600 seeds, which are 0.8 to 1 mm (Odocoileus virginianus Boddaert) (Robinette
long and average about 27 million per kilogram 1974). Foliage from new sprouts of burned
(Olson and Barnes 1974). The nonwinged seeds mountain laurel provides greater potential benefit
are dispersed by wind, but only for distances of to wildlife diets compared to unburned foliage
less than 15 m (Robinette 1974). Seeds remain (Thackston and others 1982).
viable for more than a year under field conditions.
Light, cold stratification, and moisture enhance Fire.―Mountain laurel contributes to intensity of
germination (Olson and Barnes 1974). Seeds fires in mountainous terrain and is classified as
germinate on mineral soil and on moss (Robinette "extreme" in rate of spread when it occurs in dense
1974). Kalmia may also regenerate by layering, thickets beneath hardwood stands (Jemison and
sprouts, or suckers (Robinette 1974). Techniques Keetch 1942). Flame length can exceed 30 m in
for commercial production of mountain laurel fires burning in Table Mountain pine (Pinus
plants have been well developed (Jaynes 1997). pungens Lambert) stands with mountain laurel
(Waldrop and Brose 1999). Moisture of Kalmia
Growth and Management.―Growth rate of foliage is lower than other species and is little
mountain laurel is relatively slow; young plants (< affected by seasonal changes of soil moisture
15 years) add about 12 cm in height and 9 cm in (Reifsnyder 1961). The high flammability of
width annually, depending on site moisture Mountain laurel thickets results from low foliate
relations (Monk and others 1985). Older mountain moisture (Richards 1940) and a high proportion of
laurel stems may attain heights up to 9 m and total biomass in small (<5 mm) dead twigs.
diameter at ground level of 15 cm (Jaynes 1997). Caloric content of foliage is estimated at about
Mountain laurel stem density can range from 5,000 cal/g based on similar vegetation (Hough
sparse on mesic sites to over 26,000 per ha in 1969). Allometric equations for estimating dry
thickets on upper, southwest slopes; basal area at 3 biomass of 6- to 8-year old open-grown mountain
cm above ground level can exceed 25 m2/ha. laurel stems were developed by Boring and Swank
Herbicides may be used to control mountain laurel (1986), which can be used to predict fuel loadings
on sites where establishment of tree seedlings is of leaf (log10Y = -0.754 + 1.882 log10X), branch
desired (Neary and others 1984). It is sensitive to (log10Y = -1.222 + 2.359 log10X), and bole (log10Y
juglone, a substance produced by black walnut = -0.636 + 1.948 log10X) components of individual
(Juglans nigra L.) trees. Mountain laurel generally plants, where Y is dry weight (g) and X is stem
does not present a problem to timber management diameter (mm) at 3 cm from the ground. Kalmia is
activities on submesic to mesic sites. killed more readily by headfires than by backfires
408
(Hooper 1969), but almost all topkilled stems Hooper, R.M. 1969. Prescribed burning for laurel
recover by producing basal sprouts (Clinton and and rhododentrol control in the southern
others 1993, Johnson and Kovner 1956). Equations Appalachians. Research Note SE-116. U.S.
for estimating stem, foliage, and total biomass of Department of Agriculture, Forest Service,
young mountain laurel sprouts on recently burned Southeastern Forest Experiment Station,
areas were developed by Elliott and Clinton Asheville, NC. 6 p.
(1993).
Hough, W.A. 1969. Caloric value of some forest
References fuels of the southern United States. Research
Note SE-120. U.S. Department of Agriculture,
Al-Hamdani, S.H., P.B. Nichols, and G.R. Cline. Forest Service, Southeastern Forest Experiment
2002. Seasonal changes in the spectral Station, Asheville, NC. 6 p.
properties of mountain laurel (Kalmia latifolia
L., Ericaceae) in north east Alabama. Castanea. Jaynes, R.A. 1997. Kalmia--Mountain laurel and
67(1): 25-32. related species. Timber Press, Portland, OR.
295 p.
Boring, L.R. and W.T. Swank. 1986. Hardwood
biomass and net primary production following Jemison, G.M. and J.J. Keetch. 1942. Rate of
clearcutting in the Coweeta basin. In: Brooks, spread of fire and its resistance to control in the
R.T., Jr. ed. Proceedings of the 1986 southern fuel types of eastern mountain forests. Technical
forest biomass workshop. Tennessee Valley Note 2. U.S. Department of Agriculture, Forest
Authority, Knoxville, TN: 43-50. Service, Appalachian Forest Experiment Station,
Asheville, NC. 15 p.
Bryson, G.M. and A.V. Barker. 2002. Sodium
accumulation in soils and plants along Johnson, E.A. and J.L. Kovner. 1956. Effect on
Massachusetts roadsides. Communications in streamflow of cutting a forest understory. Forest
Soil Science and Plant Analysis. 33:67-78. Science. 2: 82-91.
Chapman, G.L. 1950. The influence of mountain Kurmes, E.A. 1961. The ecology of mountain
laurel undergrowth on environmental conditions laurel in southern New England. Yale
and oak reproduction. Doctoral dissertation. University, New Haven, CT. 85 p. Doctoral
Yale University, New Haven, CT. Dissertation.
Clinton, B.D., J.M Vose, and W.T. Swank. 1993. Lipscomb, M.V. and Nilsen, E.T. 1990.
Site preparation burning to improve southern Environmental and physiological factors
Appalachian pine-hardwood stands: vegetation influencing the natural distribution of evergreen
composition and diversity of 13-year-old stands. and deciduous ericaceous shrubs on northeast
Canadian Journal Forest Research. 23: 2,271- and southwest slopes of the southern
2,277. Appalachian Mountains. II. Water relations.
American Journal Botany. 77(4): 517-526.
Ebinger, J.E. 1974. A systematic study of the
genus Kalmia (Ericaceae). Rhodora. 76: 315- McNab, W.H., S.A. Browning, S.A. Simon, P.E.
398. Fouts. 1999. An unconventional approach to
ecosystem unit classification in western North
Elliott, K.J. and B.D. Clinton. 1993. Equations for Carolina, USA. Forest Ecology and
estimating biomass of herbaceous and woody Management. 114: 405-420.
vegetation in early-succession Southern
Appalachian pine-hardwood forests. Research Monk, C.D., D.T. McGinty, and F.P. Day, Jr.
Note SE-365. U.S. Department of Agriculture, 1985. The ecological importance of Kalmia
Forest Service, Southeastern Forest Experiment latifolia and Rhododendron maximum in the
Station, Asheville, NC. 7 p. deciduous forest of the Southern Appalachians.
Bulletin Torrey Botanical Club. 112(2): 187-
Flemer,, W., III 1949. The propagation of Kalmia 193.
latifolia from seed. Bulletin of the Torrey
Botanical Club. 76: 12-16. Neary, D.G., J.E. Douglass, J.L. Ruehle, and W.
Fox. 1984. Converting rhododendron-laurel
409
thickets to white pine with picloram and Thackston, R.E., P.E. Hale, A.S. Johnson, and
mycorrhizae-inoculated seedlings. Southern M.J. Harris. 1982. Chemical composition of
Journal of Applied Forestry. 8(3): 163-168. mountain-laurel leaves from burned and
unburned sites. Journal Wildlife Management.
Olson, D.F., Jr. and R.L. Barnes. 1974. Kalmia 46(2): 492-496.
latifolia L., mountain laurel. In: C.S
Schopmeyer, ed. Seeds of woody plants in the Waldrop, T.A. and P.H. Brose. 1999. A
United States. Agriculture Handbook 450. U.S. comparison of fire intensity levels for stand
Department of Agriculture, Forest Service, replacement of table mountain pine (Pinus
Washington, DC. p. 470-471. pungens Lamb.). Forest Ecology and
Management. 113: 155-166.
Robinette, S.L. 1974. Mountain laurel-Kalmia
latifolia. In: J.D. Gill and W.M. Healy, eds. Waterman, J.R., A.R. Gillespie, J.M. Vose, and
Shrubs and vines for northeastern wildlife. W.T. Swank. 1995. The influence of mountain
General Technical Report NE-9. U.S. laurel on regeneration in pitch pine canopy gaps
Department Agriculture, Forest Service, of the Coweeta Basin, North Carolina, U.S.A.
Northeastern Forest Experiment Station, Radnor, Canadian Journal Forest Research. 25: 1,756-
PA. p. 102-105. 1,762.
Reifsnyder, W.E. 1961. Seasonal variation in the

moisture content of the green leaves of mountain
laurel. Forest Science. 7: 16-23.
410
Koanophyllon polyodon (Urban) King & H.E. Robins. polyodon
ASTERACEAE
Synonyms: Eupatorium polyodon Urban
John K. Francis
about 5 to 7. Both sedimentary (including

limestone), igneous, and metamorphic (including
ultramafic) parent materials are colonized. The
species is found in areas that receive from about
1000 to 2500 mm of mean annual precipitation at
elevations ranging from near sea level to 900 m.
Polyodon grows as scattered individuals, diffuse
stands (under forest canopies), or in thickets. The
species has an intermediate tolerance for shade and
is able to grow and reproduce under shade if not
too dense. It depends on taller vegetation for
support in order to reach heights of more than
about 1.5 m. The species is occasional to common
in open forest, brushlands, abandoned pastures, and
old road cuts and banks.
Reproduction.—Polyodon flowers near the end of

the wet season (November-January) and fruits
during the dry season (March-May). It is insect
General Description.—The name polyodon is pollinated. A collection of seeds averaged 0.00019
assigned according to the convention that if no g/seed or 5.2 million seeds/kg. Placed on moist
common name exists, the species (or sometimes filter paper without any pretreatment, 67 percent
the genus) Latin name is used as a common name. germinated between 12 and 68 days after sowing.
Polyodon is a somewhat spindly, upright or arching Germination is epigeal. The fruits are dispersed by
shrub usually 1.5 to 3 m (occasionally to 6 m) in the wind. Seeds are produced in abundance, but
height and 2 to 3 cm (sometimes 5 cm) in basal seedlings are scattered. Plants will resprout when
diameter. The shrub may have a single or multiple damaged.
stems, straight lower sections 0.5 to 1 m long, then
curving and branchy with fine twigs above. The Growth and Management.—Individual stems
plant is supported by an extensive lateral and fine may live for 6 years or more. Plants can survive
root system. The opposite leaves are papery and longer by producing sprouts. Sprouts grow at least
rough with short petioles and three main veins 1 m in their first year. No management experience
arising from or near the base. The leaf blades are has been published. However, maintaining areas in
ovate with a heart-shaped or nearly heart-shaped early and middle secondary forest would probably
base, pointed tip, and serrate edges. The encourage populations of polyodon.
inflorescences are terminal corymbs of white or
rose compound flowers (heads). Each develops Benefits.—Polyodon contributes to biodiversity in
several, five-ribbed, 3-mm achenes tipped with its wildland habitat, helps protect the soil from
numerous, 3 to 4 mm hairs (pappus) (author’s erosion, and furnishes wildlife cover. It is browsed
observation, Liogier 1997). to some extent by cattle and goats.
Range.—Polyodon is endemic to Puerto Rico References
(Liogier 1997) and is not known to have been
planted or naturalized elsewhere. Liogier, H.A. 1997. Descriptive flora of Puerto
Ecology.—Polyodon grows on moderately and Universidad de Puerto Rico, San Juan, PR.
well-drained but rarely excessively drained soils of 436 p.
loamy and clayey textures with pH’s ranging from
411
Krameria ixine L. abrojo colorado
EUPHORBIACEAE
Synonyms: Krameria cuspidata C. Presl.
John K. Francis
others 2001). It is not known to have naturalized

Ecology.—In Puerto Rico, abrojo colorado grows

in areas that receive from 750 to about 1000 mm of
mean annual precipitation and at altitudes from
near sea level to about 400 m above sea level.
Because the shrub is intolerant of shade and low in
stature, it does not withstand serious competition. It
will grow in almost any well-drained, nonsaline
soil. It is most frequently seen on rocky ridges and
south slopes, disturbed sites, and overgrazed range.
It is rarely browsed by cattle or horses.
Reproduction.—Abrojo colorado flowers and

fruits throughout the year in Costa Rica where both
flowers and fruit may be present (Instituto
Nacional de Biodiversidad 2002). Both flowers and
fruits are often present in Puerto Rican plants
(author’s observation). Fruit and seed production is
moderate to abundant. A collection of air-dried
fruits in Puerto Rico weighed an average of 0.0516
General Description.—Abrojo colorado, ± 0.0009 g. The seeds are transported as they cling
sometimes known as retama falsa, is a shrub or to passing animals or humans. After drying
sufruticose shrub to 1 m in height and 1 cm in stem sufficiently, the capsules open, releasing the seeds.
basal diameter. The stems are stiff and woody, the After disturbance, the plants regenerate by
twigs flexible and less woody. Older plants have sprouting.
lateral roots with a small or no taproot and many
fine roots. The roots are brittle, dark brown outside, Growth and Management.—Individual stems of
with red inner bark. There are multiple stems from abrojo colorada grow about 0.5 m/year in the first
the root crown. The branches and leaves are year from sprouts and live 3 or 4 years. By
covered with white hairs, giving the whole plant a continual sprouting, established plants live much
gray-green color. The leaves are narrowly oblong longer. Although no management experience has
to oblong-lanciolate, 1 to 2.5 cm long, with a been published, it probably is safe to say that
spiny-acuminate tip. The inflorescences are overgrazing and disturbance encourages the
compact lateral or terminal racemes bearing several species, and that reestablishment of tree, tall shrub,
flowers. The pink- to purplish-red flowers are or heavy grass cover discriminates against or
about 1.2 cm in diameter, from which develop eliminates it.
globose 5 to 6 mm, woody capsules (burrs) covered
with orange or red, hooked spines. The seeds are Benefits.—Abrojo colorado contributes ground
light tan and 4 mm in diameter (Howard 1988, cover, soil stability, and scenic beauty in rocky and
Liogier 1988, Stevens and others 2001). often disturbed terrain where it grows. On the other
hand, the burrs can be a nuisance to both humans
Range.—Abrojo colorado is native to Hispaniola, and animals.
Puerto Rico, the Lesser Antilles, Mexico through
Costa Rica, Colombia, Venezuela, and Guyana
(Liogier 1988, Missouri Botanical Garden 2002,
New York Botanical Garden 2002, Stevens and
412
References Missouri Botanical Garden. 2002. Manual de la
Flora de Costa Rica. http://www.mobot.org/
Howard, R.A. 1988. Flora of the Lesser Antilles, manual.plantas/019706/S019706.html. 1 p.
Dicotyledoneae, Part 1. Vol. 4. Arnold New York Botanical Garden. 20023. Specimens
Arboretum, Harvard University, Jamaica Plain, search results: Krameria ixine Loefl.
MA. 673 p. http://scis.../wwwspecimen.search_list?taxon=
Krameria+ixine+Loefl.+++++&projcode=VAS.
Instituto Nacional ce Biodiversidad. 2002. Lista de 1 p.
especímenes de Krameria ixine.
http://www.inbio.ac.cr/bims/k03/p13/c045/o013 Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
2/f01646/g008002/s024181.htm. 2 p. Montiel, eds. 2001. Flora de Nicaragua.
Liogier H.A. 1988. Descriptive flora of Puerto 2. Missouri Botanical Garden, St. Louis, MO.
Rico and adjacent islands, Spermatophyta. Vol. p. 945-1,910.
413
Lantana camara L. lantana
VERBENACEAE
Synonyms: Lantana aculeata L.

Lantana antillana Raf.
Lantana camara L.var.aculeata (L.) Mold.
Lantana scabrida Soland.
John K. Francis
Range.—The native range of lantana extends from

Bermuda, the Bahamas, the Greater Antilles, the
Lesser Antilles, through Trinidad and Aruba. On
the mainland, it is native to coastal areas of the
United States from Georgia through Texas and
from northern Mexico to South America including
Brazil and Peru and probably Bolivia, Paraguay
and northern Argentina. Lantana has naturalized in
most suitable habitats in tropical and subtropical
Africa, Asia, and Australia. Most of the world’s
tropical and subtropical islands including the
Hawaiian Islands, Guam, Pitcairn Island,
Madagascar, the Juan Fernandez archipelago,
Reunion, and the Galapagos have naturalized
populations. This colonization has occurred largely
during the last century.
Ecology.—Lantana grows on all types of well-

drained soil in areas that receive from about 250
mm to 2900 mm of rainfall. It resists droughts very
well and tolerates salt spray. Aerial portions of the
plant are killed by temperatures of –2 °C, but
quickly grow back (Anonymous 2000). Large and
General Description.—Lantana has many vigorous plants survive fires and cutting well,
common names including cariaquillo, filigrana, although less vigorous plants are often killed.
mille fleurs, sauge, red sage, yellow sage, prickly Lantana is an intolerant pioneer that colonizes
sage, and lakana. It is a medium-sized aromatic disturbed areas. It will grow under an open forest
shrub with quadrangular stems, sometimes having canopy but quickly disappears when the shade
prickles. The posture may be suberect, scrambling, becomes heavy. Many pests and diseases lightly
or occasionally clambering (ascending into shrubs and incidentally affect the species across its broad
or low trees, clinging to points of contact by means range.
of prickles, branches, and leaves). Frequently,
multiple stems arise from ground level. The leaves Reproduction.—The inflorescence is a capitate,
are generally oval or broadly lance-shaped, 2 to 12 many-flowered head. The corolla may vary widely
cm in length, and 2 to 6 cm broad, having a rough in color depending on the variety but
surface and a yellow-green to green color. The characteristically changes colors between the
flat-topped inflorescence may be yellow, orange, center flowers and older, outer flowers. Lantana
white, pale violet, pink, or red (Howard 1989, blooms almost continuously under favorable
Liogier 1995). The small individual flowers of conditions. Somatic chromosome numbers of 33,
wild Puerto Rican shrubs darken with age so that 44, and 55 were recorded in India, the latter
inflorescences are yellow in the center and orange tetraploid being the most common (Sinha and
toward the perimeter. Fruits form in clusters others 1995). Insects, especially butterflies,
similar in appearance to a blackberry. pollinate the flowers. Clusters of drupes are
produced abundantly. The fruits are blueblack
414
when ripe and contain one seed each. They are In some areas, competition by lantana results in a
eaten by birds and are widely scattered. If not reduction of biodiversity (Kumar and Rohatgi
eaten, they dry and remain on the shrub for weeks. 1999). Despite the establishment of a number of
A sample of seed collected in Puerto Rico natural enemies of lantana into exotic populations,
contained 100,700 seeds/kg of which 75 percent control of lantana populations has been usually
germinated within 7 weeks of sowing. Early limited or a failure (Day and others 1999, Hill
growth is rapid. Also, lantana can be propagated 1999). In thick stands, the shrub increases costs in
with cuttings and air layers. forest management by inhibiting access in stands
for thinning and felling, competes with
Growth and Management.—Ornamental plants reproduction, and increases fire hazards (Graaff
are established as potted seedlings. This is 1986). Lantana leaves contain poisonous
probably also the best method to establish plants triterpines and lantadenes A and B that cause death
for environmental protection projects such as of horses, cattle, sheep, goats, and rabbits by
reclamation of mine spoils. Lantana may reach 3 failure of the liver and other organs (Morton 1994,
m in height within 3 or 4 years. In naturalized Munyua and others 1990). However, most animals
habitat, the species often forms dense thickets. carefully avoid eating this plant when given a
These brush stands may be eliminated at choice. Green fruits also contain the poisons and
considerable expense from pastures and tree have caused illness and death in children (Morton
plantations by cutting and herbicide application. 1994). Lantana leaves and their leachates exert
The preferred methods are to cut the clumps and alelopathic effects in vetro and to a lesser extent in
drench the stumps with herbicide or to later spray soil on seed germination, root elongation, and
the tender regrowth with postemergence herbicide. plant growth of many species (Casado 1995, Sahid
Once established, tree plantations with closed and Sugau 1993).
canopies will keep lantana from reestablishing
itself. References
Benefits.—Lantana, in many horticultural Anonymous. 2000. Floridata, Lantana camara.

varieties, is planted the world over as a flowering www.streetside.com/plants/floridata/ref/l/lant_c.
ornamental. It is grown as an annual bedding plant ht. 1p.
in temperate areas. Lantana oil, an aromatic
mixture that varies by local plant variety, is Casado, C.M. 1995. Allelopathic effects of
exported from at least Brazil (Weyerstahl and Lantana camara (Vergenaceae) on morning
others 1999). In herbal medicine, infusions of the glory (Ipomoea tricolor). Rhodora 97: 264-274.
leaves and other plant parts are used as an
antiinflammatory (Oyedapo and others 1999), a Day, M.D., R.H. Holtkamp, and P. Blackmore.
tonic and expectorant, and added to baths as an 1999. The status of biological control of
antirhumatic. Lantana extracts have also been Lantana camara in Australia. In: Practical weed
shown to be a powerful febrifuge (Liogier 1990). management: protecting agriculture and the
Because the leaves and some other parts of lantana environment. 10th Biennial Noxious Weeds
are poisonous, care must be taken when it is used Conference, 10-22 July 1999, Ballina, Australia.
medicinally. The ripe fruit is benign and heavily p. 257-260. New South Wales Agriculture,
consumed by birds and frequently eaten by Armidale, Australia.
humans in some countries (Herzog and others
1994). Extracts of lantana leaves have shown Graaff, J.L. 1986. Lantana camara, the plant and
strong insecticidal and antimicrobial activity in some methods for its control. South African
numerous experiments. Storing potatoes with Forestry Journal 136: 26-30.
lantana leaves nearly eliminates damage by
Phthorimaea operculella Zeller, the potato tuber
Hill, M.P. 1999. Past and present initiatives on
moth (Lal 1987). Stems and leaves are used as
the biological control of Lantana camara
mulch. Although of inferior quality because of size
(Verbenaceae) in South Africa. In: J. R. Baars,
and form, lantana stems are widely used as fuel in
S. Neser, and T. Olckers, eds. Biological control
less developed countries.
of weed in South Africa 1990-1998. African
Entomology Memoir 1. 21-33.
Detrimental Effects.—Lantana has become a
weedy invader of disturbed forest land and
neglected pasture in much of its naturalized range.
415
Herzog, F., Z. Farah, and R. Amado. 1994. popular garden flower; some cases of poisoning
Composition and consumption of gathered wild in Florida. Economic Botany 48(3): 259-270.
fruits in the V-Baoule, Cote d’Ivoire. Ecology of
Food and Nutrition 32(3-4): 181-196. Munyua, S.J.M., M.J. Nienga, T.P. Karitu, T.P.
Kimoro, J.E. Kiptoon, and I.B.J. Buoro. 1990. A
note on clinical-pathological findings and serum
Howard, R.A. 1989. Flora of the Lesser Antilles.
enzyme activity in sheep, goats and Friesian
Vol. 6. Arnold Arboretum, Harvard University,
calves with acute Lantana camara poisoning.
Bulletin of Animal Health and Production in
Africa 38(3): 275-279.
Kumar, S. and N. Rohatgi. 1999. The role of
invasive weeds in changing floristic diversity.
Oyedapo, O.O., F. C. Sab, and J.A. Olagunju.
Annals of Forestry 7:(1): 147-150.
1999. Bioactivity of fresh leaves of Lantana
camara. Biomedical Letters 59: 179-183.
Lal, L. 1987. Studies on natural repellents against
potato tuber moth (Phthorimaea operculella
Sahid, I.B. and J. B. Sugau. 1993. Allelopathic
Zeller) in country stores. Potato Research 30(2):
effects of lantana (Lantana camara) and siam
329-334.
weed (Chromolaena odorata) on selected crops.
Weed Science 41(2): 303-308.
Sinha, S., B. Sinha, and A. Sharma. 1995.
Chromosome composition of Lantana camara
L.: karyotype, basic number and DNA diversity.
Nucleus Calcutta 38(1-2): 16-22.
Weyerstahl, P., H. Marschall, A. Eckhardt, and C.
617 p.
Christiansen. 1999. Constituents of commercial
Brazilian lantana oil. Flavour and Fragranced
Morton, J.F. 1994. Lantana, or red sage (Lantana
Journal 14(1): 15-28.
camara L. Vergenaceae), notorious weed and
416
Lantana involucrata L. wild sage
VERBENACEAE
Synonyms: Lantana odorata L.
John K. Francis
terminal heads. The corolla may be white, pink to

pinkish purple, or pale blue. The infrutescences are
clusters of purple to blue-black drupes 3 to 4 mm
in diameter containing one seed each. The
chromosome number is 2n = 36 (author’s
observation, Howard 1989, Liogier 1995, Long and
Lakela 1971, Nelson 1996).
Range.—The native range of wild sage includes

southern Florida, the West Indian islands, Mexico
through northern South America bordering the
Caribbean, and the Galápagos Islands (Howard
1989, Liogier 1995). It is naturalized or possibly
native in Bermuda (Britton 1918). Wild sage is
now widely planted as an ornamental and probably
has naturalized in additional areas.
Ecology.—Wild sage grows on most well-drained

soil types derived from both sedimentary
(including limestone), igneous, metamorphic
(including ultramafic) rock types. Mean annual
precipitation in areas where it grows in Puerto Rico
varies from 750 to about 1700 mm. The species
grows from near sea level to 600 m or more in
Puerto Rico. In Mexico, wild sage is reported to
occur from 1,000 to 2,000 m above sea level
(Secritaría del Medio Ambiente y Recursos
Naturales 2002). It requires disturbance to become
established and full or nearly full sun to grow and
reproduce. Wild sage is especially common in
over-grazed range and poorly managed pastures,
General Description.—Wild sage, also known as and may be found in abandoned farmland and early
button sage, common sage, sea sage, white sage, secondary forest as well as road cuts, cliffs, and
orégano, terete, peonía colorada, contite, and rocky sites. It grows in pinelands, hammocks, and
frutilla, is a shrub commonly 1 to 2 m in height shell mounds in Florida (Long and Lakela 1971,
with showy flowers. The plant is supported by Nelson 1996).
many stiff lateral roots and abundant fine roots.
The roots are light brown with a corky bark. Older Reproduction.—Wild sage blooms all year
plants, often with a basal diameter of 5 cm or more, (Nelson 1996) and also is a good fruit and seed
have several main branches arising at or near the producer. The seeds number about 110,000
ground line and a branchy crown. The bark is seeds/kg (air-dried). They may be cleaned by
yellowish and the wood is brittle. Petioles are 3 to maceration and wet sieving. No pretreatment is
5 mm long and the leaf blades are 1 to 4 by 1 to 2.5 necessary, and germination begins in about 15
cm wide and broad, rotund to narrowly elliptic- days. Wild sage sprouts when cut. Seedlings are
oblong, rough and stiff with a crenate or toothed fairly common in favorable habitat near seed
margin. The leaves are aromatic with a spicy, sage- sources.
like smell. The small tubular flowers form compact
417
Growth and Management.—The growth rate is Howard, R.A. 1989. Flora of the Lesser Antilles.
moderate and plants live for 10 years or more. Vol. 6. Arnold Arboretum, Harvard University,
Propagation may be by seed, by rooting of Jamaica Plain, MA. 658 p.
softwood cuttings, or by transplanting wildlings.
Wildlings transplant well if the roots are not badly Liogier, H.A. 1995. Descriptive flora of Puerto
damaged and about one-third of the top is pruned Rico and adjacent islands. Vol. 4. Editorial de
away. Periodic pruning of ornamental plants is la Universidad de Puerto Rico, San Juan, PR.
necessary to prevent them from becoming lank 617 p.
(Workman 1980).
Benefits.—Wild sage contributes to biodiversity, tropical Florida. University of Miami Press,
helps protect the soil, and furnishes wildlife food Coral Gables, FL. 962 p.
and cover. The wood is useful for fuel, but it is
generally too small for anything but campfires. The Malone, K. 2002. Palm Beach County
species is used as a foundation plant and to form (Southeastern Florida) top butterfly nectar
low hedges (Workman 1980). It is listed as one of flowers. North American Butterfly Association.
the best butterfly nectar plants (Malone 2002). The http://www.naba.org/ftp/pbco.pdf. 4 p.
leaves are used as a condiment in cooking, and the
essential oil is used in cosmetics and liquors McNary-Wood, K. 2002. Medicinal plants for
(Secritaría del Medio Ambiente y Recursos what ails you in paradise. http://www.
Naturales 2002). Leaves are added to baths to ease timespub.tc/Natural%History/Archive/Winter%
heat rashes and mild insect bites (McNary-Wood 20200102/medicinal.htm. 2 p.
2002). Extracts of the plant are also said to have a
sedative effect (Beattie and others 2002). Teas and Nelson, G. 1996. The shrubs and woody vines of
decoctions of leaves and twigs are used to control Florida. Pineapple Press, Inc., Sarasota, FL.
colic, vomit, cough, fever, and congestion 391 p.
(Secritaría del Medio Ambiente y Recursos
Naturales 2002). It is reportedly not as poisonous Secritaría del Medio Ambiente y Recursos
as the more widespread L. camara L. (Workman Naturales. 2002. Especies con usos no
1980). maderables en bosques tropicales y
subtropicales: Lantana involucrata L.
References http://www.semarnat.gob.mx.pfnm2.fichas/
lantana_involucrata.htm. 1 p.
Beattie, L., M. Martin, M. Shaposhnik, and K.
Vaga. 2002. Stress treatments. http://members. Workman, R.W. 1980. Growing native. The
tripod.com/~Moiraine/treatment.html. 7 p. Sanibel-Captiva Conservation Foundation, Inc.,
Sanibel, FL. 137 p.
Britton, N.L. 1918. Flora of Bermuda. Charles
Schribner’s Sons, New York. 585 p.
418
Larrea tridentata (Sesse' and Moc. ex DC.) Coville creosote bush
ZYGOPHYLLACEAE
Synonyms: Covillea tridentata (Sesse' & Moc. ex DC.) Vail

Covillea glutinosa (Engelm.) Rydb.
Larrea divaricata Cav.
Larrea glutinosa Engelm.
Larrea mexicana Moric.
James E. Nellessen
bush ancestors arriving in North America 8.4 to

4.2 million years before present (Lia and others
2001) with current distribution occurring since the
last glaciation. Creosote bush is believed to be a
migrant through possible bird dispersal of the L.
divaricata populations (Mabry and others 1977).
There is a ploidy gradient from east to west (from
moister to dryer deserts) across the Southern
United States: Chihuahuan Desert populations are
diploid, Sonoran tetraploid, and Mojave hexaploid.
From a study of plant remains in packrat middens,
the plants appear to have spread slowly from lower
Rio Grande refugia in Texas north and west,
reaching the Southern Mojave about 8,500 years
ago, the Northern Sonoran about 6,400 years ago,
General Description.—Creosote bush is also and the Northern Chihuahuan about 4,000 years
know by the names gobernadora, hediondilla ago (Hunter and others 2001). Larrea divaricata
(Spanish for little stinker), and guamis. It is and diploid L. tridentata have homologous
sometimes erroneously referred to as greasewood. genomes (Mabry and others 1977). Two varieties
It is an evergreen resinous shrub of warm deserts. have been recognized by one author: L. tridentata
It is a well branched shrub with no defined trunk, var. tridentata and L. tridentata var. arenaria L.
typically reaching heights of 1 to 1.5 m, but can Benson (Kartesz 1994).
grow as high as 3 m. It is strongly scented,
especially noticeable after a rainstorm. The stems Range.—Creosote bush occurs from Southern and
are gray with black, somewhat swollen bands at Western Texas through Southern New Mexico
the nodes; the twigs and young branches are (with a band up the Rio Grande valley to
relatively flexible. The leaves are opposite, Albuquerque), into Arizona, and Southern
bifoliately compound, each leaflet joined at the California, as far north as Southern Nevada and
base, divergent, but the tips often point toward Southwestern Utah, and south well into Mexico.
each other. Leaflets generally are olive green in Its elevational range is from below sea level
color, to 1 cm in length, 3 to 4 mm wide, (Death Valley, CA) to about 1,700 m.
asymmetrical, oblong to obovate in shape, and
broader away from the base (Carter 1997, Correll Ecology.—Creosote bush is a characteristic
and Johnston 1970, Hickman 1993, Kearney and dominant evergreen shrub of the Southwestern
others 1951, Martin and Hutchins 1980-81). The warm Chihuahuan, Sonoran, and Mojave deserts.
leaves are often glossy with a thick resinous On the northern fringes of the Mojave its range
coating. Stipules are persistent. The South just touches the Great Basin Desert (Mabry and
American L. divaricata is so similar to L. others 1977). It grows best in gravelly to sandy,
tridentata that some researchers have considered well drained soils of desert plains, benches, mesas,
them the same species (Mabry and others 1977). alluvial fans, and rocky slopes. It does not occur
These two species have been demonstrated to on alkali flats or pans. It roots deeply, to 3 m or
hybridize in experimental gardens. South America more, but most roots occur in the upper soil layers
is the likely origin for the genus, with creosote (but below about 10 cm), and it will tolerate a
419
caliche layer. The shallow roots are able to utilize increases associated with spring temperatures and
small rainfall events. Creosote roots inhibited moisture availability (Mabry and others 1977). It
further elongation of the roots of both adjacent has high water use efficiency and ability to resorb
creosote bushes and the shrub Ambrosia dumosa some nutrients, especially phosphorus, 72 to 86
(A. Gray) Payne (Mahall and Callaway 1992). percent, while nitrogen resorption is within the
Stem angle facilitates water stemflow to the roots, general range for other evergreen shrubs, 47 to 57
which is maximal with stems greater than 65° and percent. Creosote bush stem and leaf orientations
almost none less than 45°. Creosote bushes in the maximize light collection during cooler and
driest part of their range, for example, Death moister early morning periods. Creosote bush has
Valley, generally have steeper angles, while there a greater ability to withstand xylem cavitation
is more variability in stem angle in the Chihuahuan during droughts than riparian species (Pockman
Desert. Shrubs with shallower stem angles take on and Sperry 2000). An Arizona population of
a more hemispherical shape as opposed to an creosote bush experiences freezing-induced xylem
inverted cone shape. Shrub canopy architecture vessel cavitation and consequent disruption of
also affects soil characteristics creating "islands of water transport at temperatures from -11 to -20 °C
fertility." Shrubs with hemispherical shapes collect (Pockman and Sperry 1997). These temperatures
much more litter underneath, hence soil nitrogen is correspond to 20+ year minimum isotherms and
greater under hemispherical shrubs than under also correspond with the northern distribution in
inverted cone shaped shrubs (Whitford 2002). both the Sonoran and Mojave Deserts. A few
Creosote bushes often require a nurse shrub such studies have examined the effects of
as the small shrub A. dumosa for seedling/sapling environmental pollutants on creosote bush. Foliage
establishment, and creosote bush itself will serve demonstrates leaf injury when exposed to acute
as a nurse plant for certain other species (Whitford levels of sulfur dioxide (SO2) (Olyszyk and others
2002). Creosote bush often forms monotypic 1987) and elevated CO2 concentrations may
stands that extend for miles, although its density ameliorate the effects of heat stress (Hamerlynck
varies greatly from site to site, with the greatest and others 2000). Creosote foliage may
densities occurring in the Chihuahuan Desert (up accumulate heavy metals from industrial pollution
to 8,000 plants/ha), and lesser densities in the as demonstrated in a 1980 to 1995 study from El
Sonoran (up to 4,000 per ha) and the Mojave (up Paso, TX, and Ciudad Juarez, Mexico (Mackay
to 1,800 per ha) (Mabry and others 1977). Above- and others 1998).
ground standing crop biomass tends to reflect
these density trends. Chihuahuan Desert Reproduction.—Creosote bush blooms
populations tend to show a greater size or age class periodically during the growing season, blooming
structural diversity than those in other deserts correlated first with seasonality, secondarily with
(Mabry and others 1977). The resinous coating on the timing of moisture availability and other local
the leaves helps retard water loss. During extreme environmental factors. Peak blooming periods
droughts creosote bush will produce smaller, occur in spring (February to May) and again in late
tougher leaves, and will also shed many of its summer into the fall (August to December). A
older leaves. It quickly produces new shoot single mature shrub may produce more than 2,000
growth after substantial summer rains. Creosote flowers (or about 50 g of flowers) during the
bush has the C3 photosynthetic pathway, with growing season. Numerous insects visit the
peak photosynthetic activity (Pmax) in the spring flowers, but bees are the most effective pollinators.
and during summer monsoon seasons especially Many species of bees are known to visit the
when pre-dawn water potentials are high flowers; some are generalists but some are
(Whitford 2002). Pmax is as high as that found in specialists and forage solely on creosote bush
C4 plants (Fitter and Hay 1987), and creosote (Minckley and others 2000). It is an outcrossing
shrubland has a similar community energy balance species but is always self-compatible (Mabry and
as the C4 desert grasslands they have invaded others 1977) with localized populations having
(Dugas and others 1996). Creosote bush has higher levels of inbreeding in the Chihuahuan
demonstrated a threshold for stomatal closure at - Desert. The flowers are borne solitary in the axils,
5.8 MPa, has maintained full turgor at -3.0 MPa, are 2 to 3 cm in diameter, are perfect, with five
with positive net photosynthesis occurring at -8 to yellow clawed petals, obovate to spatulate, 6 to 10
-16 MPa (Fitter and Hay 1987, Mabry and others mm long, and 5 mm wide. There are five
1977). It can tolerate leaf water potentials below - yellowish green, unequal, deciduous sepals, 5 to 8
50 MPa. It is able to maintain a relatively stable mm long. There are 10 stamens with winged
photosynthetic rate all year in Death Valley, with filaments, and the ovary has five locules. The
420
petals twist at a 90 degree angle after pollination. fertilized, irrigated, or unmodified ungrazed black
The fruit is a roundish capsule, about 5 to 7 mm in grama (B. eriopoda) grassland, a creosote
diameter, covered with a dense concentration of shrubland, or a heavily grazed grassland. The only
white to reddish hairs, and separates into five site in which seedlings survived was in the heavily
indehiscent one-seeded carpels at maturity (Carter grazed grassland (Whitford 2002). Prior to heavy
1997, Correll and Johnston 1970, Kearney and livestock grazing creosote bush was generally
others 1951, Martin and Hutchins 1980-81). Both restricted to well-drained gravelly or sandy soils of
fruits and flowers are commonly found on the steep slopes. Creosote bush and mesquite
plant at the same time. Flowering may begin on (Prosopis glandulosa Torrey) invasion likely
plants as young as 4 to 6 years, although heavy occurred in episodes in conjunction with both
flowering and fruiting generally occurs after 8 to reduced grass cover and drought (Grover and
13 years. Greenhouse-raised seedlings may flower Musick 1990). Localized decreases have also been
at 2 years of age. Fruit production ranges from 39 reported. In an area of Big Bend National Park,
to 278 fruits per 100 g of branches or 120 to 1,710 previously overgrazed by livestock, the dry period
per plant (Young and Young 1992). There are from 1960 to 1967 showed increased creosote
about 370 seeds/g. The seeds have mechanical bush cover, while the wetter period from 1967 to
dormancy and scarification aids germination 1981 showed a decrease in creosote bush cover
(Young and Young 1992). Germination success and increased cover of perennial grasses, forbs,
varies by population, latitudinal and local and most other shrubs (Wondzell and Ludwig
environmental conditions with reported success 1995). At a site in southern New Mexico creosote
rates of 2 to 70 percent. Salt (NaCl) at 10,000 ppm shrub size has decreased over a 10-year period,
(-0.78 MPa) resulted in 0 percent germination, and possibly due to longer term climate change (Miller
seed germinated equally well at pH's from 7 to 10 and Huenneke 1996). Certain creosote bush
(Barbour 1968), although some studies have populations in the Sonoran Desert of Mexico
shown some variation from these pH's (reviewed declined by 50 to 90 percent, perhaps due to
in Baskin and Baskin 2001). Leachates from fruit prolonged droughts from 1936 to 1964 (Turner
coats inhibit germination of some plant species 1990). Creosote bush shrublands have been
[Bouteloua eriopoda (Torr.) Torr. but not experimentally treated with a variety of herbicides
Muhlenbergia porteri Scribn. ex Beal], but not of for shrub management. Several liquid-applied
creosote bush seeds (Baskin and Baskin 2001). herbicides (2,4-D, dicamba, picloram, 2,3,6-TBA)
Seeds stored dry in paper bags at room either singly or in combination with other
temperature for as long as 8 years showed some herbicides have demonstrated greater than 50
germinability (Barbour 1968). percent root kill (Herbel and Gould 1995). But
individual shrub treatment with dry herbicides
Growth and Management.—Growth response (bromacil, fenuron) or aerially applied tebuthiruon
and productivity can be rapid after rain, but there pellets have given greater control, 80 to 90
is not a strong pattern of productivity and rainfall, percent. Perennial grass and annual forb
while nitrogen has been shown as an important production increased substantially on tebuthiuron-
factor in creosote bush productivity (Whitford treated plots.
2002). Maximum leaf longevity is 16 months.
Maximal root growth occurs in the fall. Branches Benefits and Disadvantages.—Creoste bush is
will root if covered with soil. Ring counts of stems unpalatable as livestock forage and is usually
have indicated that creosote bush is long-lived, at toxic, sometimes causing death (Gay and Dwyer
least 30 to 50 years, but ring counts may not be 1998, Kearney and others 1951, Stubbendieck and
reliable. Individual shrubs may sometimes others 1997, USDA 1937). Jackrabbits are one of
reproduce by fragmentation or separation of the the few mammals that will consume the plant to a
root crowns along with new basal sprouting, certain degree, largely based on the water content
causing the plant to spread out and away from the of a particular plant (Whitford 2002). Some
center. Such fragmentation may contribute to long- woodrat populations will also consume creosote
term survival estimated at more than 100 years to bush, demonstrating differentially developed
several thousand years. Creosote bush shrubland tolerances to the resin (Mangione and others
has been expanding over the past 100 years or 2000). Woodrats consume creosote leaves and
more due to overgrazing of desert grasslands by stems in the Mojave Desert when necessary to
livestock, although climate trends have also been maintain water balance but suffer ill effects such
implicated. One experiment to test this involved as loss of body mass (Karasov 1989). Both rabbits
transplanting seedling creosote bushes into and woodrats will peel open the stems to get water.
421
Once a particular shrub is browsed by a jackrabbit, principal compounds in creosote bush resin is the
that same shrub is much more likely to be browsed anti-oxidant nordihydroguaiaretic acid (NDGA).
again than another shrub that has never been This is a phenolic aglycone and comprises 5 to 10
browsed (Ernest 1994). Younger leaves have a percent of leaf dry weight, 80 percent of all
higher concentration of resins that affect relative phenolics in the resin, and once was used as an
leaf palatability and digestibility. Resin content anti-oxidant in foods, pharmaceuticals, and
may be as high as 26 percent in young leaves, industrial materials (Mabry and others 1977).
dropping to 11 to 16 percent in older leaves, with NDGA has demonstrated an ability to inhibit
insects showing preference for the older leaves tumors and cancers but does have toxic side effects
(Mabry and others 1977). A phenoloxidase system to both animals and humans. Its use as a
in the leaves appears to make digestibility of the chemoprevention for skin cancer and protection
resinous leaves even more difficult. Chemical against the effects of ultaviolet light has been
treatment of creosote foliage to remove the resins investigated (Gonzales and Bowden 2002). The
results in nutritious edible forage, but due to botanical dietary supplement "chapparal" has been
economics this practice is not often implemented associated with nonviral toxic hepatitis and
(Mabry and others 1977). Numerous species of contains lignans similar to estrogens (Obermeyer
arthropods set up habitation in creosote bush. In and others 1995) and may cause acute liver
particular males of the grasshopper species, damage (Sheikh and others 1997). NDGA has
Ligurotettix coquilletti McNeil, select a particular shown potential to inhibit the human
shrub as their territory and defend it against other immunodeficiency virus (HIV) (Gnabre and others
male grasshoppers (Mabry and others 1977). The 1995). NDGA (masoprocol) lowered glucose and
mounds created by kangaroo rats (Dipodomys triglyceride levels in rats and may have potential in
spp.) appear to have a positive effect on creoste Type II diabetes treatment (Reed and others 1999).
bush survival, flowering, and fruiting, while Numerous flavanoid aglycones, flavanoid
creosote bush itself has a negative effect on glycosides, wax esters, saponins, and volatile
kangaroo rat populations (Chew and Whitford constituents such as terpenes, have been identified
1992). Creosote bush exposure to 0.1 to 0.2 ppm from creosote bush.
ozone consistently reduced the toxic
nordihydroguaiaretic acid content of leaves, which References
could result in potentially greater levels of
herbivory (Gonzalez and others 1988). Powdered Barbour, M.G. 1968. Germination requirements of
creosote leaves added to stored pinto beans made the desert shrub Larrea divaricata. Ecology 49:
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Ecology, Biogeography, and Evolution of
Pharmaceutical Chemistry.—Although a toxic Dormancy, and Germination. Academic Press,
plant for livestock and many wildlife, it has value San Diego, CA. 666 p.
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other potential medicinal uses. Native Americans Bowers, J.E. and B. Wignall. 1993. Shrubs and
of the Southwestern deserts have used this plant in Trees of the Southwest Deserts. Southwest Parks
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diuretic to treat venereal disease, tuberculosis,
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snakebites, menstrual cramps, and chicken pox Chew, R.M. and W.G. Whitford. 1992. A long-
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422
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424
Lasiacis divaricata (L.) A.S. Hitchc. wild bamboo
POACEAE
Synonyms: Panicum bambusoides Desv. ex Ham.

Panicum chauvinii Steud.
Lasiacis harrisii of Britton and Wilson
Lasiacis sloanei of Britton and Wilson
Panicum divericatum L. var. glabrum Kuntze
Panicum divericatum L. var. stenostachyum Griseb.
John K. Francis
lanceolate, 5 to 20 cm long and 5 to 15 cm wide.

Inflorescences are terminal panicles with few
branches. The fruits are ovoid, about 4 mm long,
and covered (at maturity) with a thin, black
pericarp (Hitchcock 1935, Nelson 1996, Stevens
and others 2001). There are 2n = 36 chromosomes
(CromoPar 2002).
Range.—Wild bamboo is native to southern

Florida, the West Indies, and from Central Mexico
to northern Argentina (Stevens and others 2001).
Stevens and others (2001) report two varieties,
divaricata and leptostachya (Hitchc.) Davidse, in
Nicaragua. The species is not reported to have
naturalized outside its native range.
Ecology.—Wild bamboo is intermediate in its

tolerance to shade. It normally grows in the
understory of medium to low-density forest, in
small openings, and in brushy areas. It is common
along shady forest trails and roads, and sometimes
invades brushy pastures. In Puerto Rico, it grows
from near sea level to about 800 m in elevation in
areas that receive from about 1000 to 2400 mm of
mean annual precipitation. Wild bamboo grows up
to an elevation of 1300 m in Nicaragua (Stevens
General Description.—Wild bamboo, also known and others 2001). The species colonizes soils with
as smallcane, pitillo de monte, tibisí, and chico, is the full gamut of textures and pH’s from about 5.5
an evergreen, slender, woody, climbing or to 7.5 in areas of sedimentary (including
scrambling shrub that sometimes reaches 5 m in limestone), igneous, and metamorphic (including
length and is commonly about 6 mm in basal stem ultramaphic) rocks. The soils are usually well
(culm) diameter. There are usually several stems drained.
from the root crown. The plant develops an
extensive and abundant system of fibrous roots. Reproduction.—Wild bamboo flowers throughout
The whole pant is more or less glabrous. The culms the year except during periods of high drought
are hollow between nodes, thin-walled, brittle, and stress. The flowers are assumed to be wind
tough, so that they have low bending strength but pollinated. Collections of air-dried seeds from two
high tensile strength. The lower internodes are areas in Puerto Rico in different years yielded
greenish yellow and smooth. There are few or no averages of 0.0064 ± 0.0001 and 0.0156 + 0.0001
branches on the culms until near the ends. g/seed or roughly 90,000 seeds/kg. Some
Relatively few leaves are located on recently pretreatment is probably needed: one lot yielded 0
grown branches. The alternate leaves are narrowly
425
percent germination and the other 11 percent. The and cattle. The species seems resistant to light and
seeds are somewhat delicate and probably damaged moderate browsing. The tough stems, which often
by scarification. The pericarp can be removed by run across low brush, make it difficult to walk
rubbing with great care with unknown effect on through the forest.
germination. It is assumed that birds and possibly
rodents are seed dispersers. Seedlings are not References
common. Culms layer (root) whenever they come
in contact with the ground and are covered with CromoPar. 2002. Recuentos cromosómicos
litter. Wild bamboo sprouts when cut or damaged foráneos de plantas presentes en Paraguay.
to renew damaged culms. Universidad de Barcelona, Spain. http://
www.ub.es/botanica.cromopar.cro-out/pdf. 24 p.
Growth and Management.—Growth of wild
bamboo from sprouts is rapid. Sprouts from mature Hitchcok, A.S. 1935. Manual of the grasses of the
plants may reach most of their eventual length United States. Miscellaneous Publication 200.
during the first year. Thereafter, branches are U.S. Department of Agriculture, Washington,
added for 1 or 2 years. After old culms die, new DC. 1,040 p.
ones grow from the root crown. It is not known
how fast seedlings grow. Wild cane seldom forms Nelson, G. 1996. The shrubs and woody vines of
thickets and mostly grows in forests and brush Florida. Pineapple Press, Inc., Sarasota, FL.
lands so that control does not seem warranted. If it 390 p.
should be needed, grubbing out the root crowns or
cutting and spot-spraying the resulting sprouts with Stevens, W.D., C. Ulloa-U., A. Pool, O.M.
glyphosate would probably be effective. Montiel, eds. 2001. Flora de Nicaragua.
Benefits.—Wild bamboo furnishes food and cover 3. Missouri Botanic Garden Press, St. Louis,
for wildlife and helps protect the soil. The leafy MO. p. 1,911-2,666.
and more succulent portions are browsed by horses
426
Leandra krugii (Cogn.) Judd & Skean leandra
MELASTOMATACEAE
Synonyms: Calycogonium krugii Cogn.
John K. Francis
about 600 to 1000 m in areas that receive from

about 1800 to 3000 mm of mean annual
precipitation. The soils it grows on are usually
moderately acid, loamy or clayey, deep or rocky,
well drained to somewhat poorly drained Ultisols,
Alphisols, and Inceptisols. These soils develop
from sedimentary, igneous, and metamorphic
(ultramaphic) rocks. Leandra is most common on
side slopes and ridge tops, in forest plantations,
secondary forest, and primary remnants (author’s
observations).
Reproduction.—Leandra blooms during the late

spring and early summer (April through June) and
fruits during late summer and fall (August through
October). The flowers appear to be insect
pollinated. There are several million seeds per
General Description.—Leandra is an evergreen
kilogram. They germinated profusely when
shrub usually 1 to 1.5 m in height and 1 to 3 cm in
smeared on the surface of potting mix and placed
basal diameter, but rarely reaching 3 m in height
in a mist bed. The minute seedlings did not survive,
and 6 cm in basal diameter. Multiple stems from
however. The plant can be propagated by air
basal sprouts are the rule and stems are covered
layering. Thirty-nine of 46 air layers treated with
with finely furrowed brown bark. Young plants
0.3 percent IBA on wild plants rooted in 6 months.
have deep taproots; older plants have deep and
Wild plants layer whenever branches come in
extensive root systems. Branches and twigs are
contact with the ground. Woody cuttings treated
stiff, slender, and tend to grow vertically.
with IBA (indol-butyric acid) and misted failed to
Trichotomous branching is common. The green to
successfully root. Apparently, birds are the
dark-green leaves are opposite, usually oval to
principal dispersers of seeds. Reproduction can be
elliptic, rough, and coriaceous. They are 2 to 5.5
abundant in shaded areas that have been disturbed
cm long and have a 6-mm petiole. Edges are
(author’s observations).
serrulate. Five nerves are raised on the underside,
the three principal ones radiating from a point
Growth and Management.—Leandra grows
above the base of the leaf. The six-parted white
slowly as a seedling and at a moderate rate from
flower is about 3 cm across. Fruits (berries) are
sprouts. Plants appear to be relatively long-lived.
globose, bright red, and about 12 mm in diameter.
Management experience is lacking or has not been
They are edible and contain many minute seeds
published. Disturbance of the understory of forests
(author’s observations, Liogier 1995).
in suitable habitat with low to moderate basal area
in which mineral soil is exposed should result in
Range.—Leandra is endemic to central and
the establishment of seedlings. Stands could be
western Puerto Rico (Liogier 1995). It is not
thickened by ground-layering existing plants.
known to have been planted or naturalized outside
Puerto Rico.
Benefits.—Leandra adds to the aesthetics of the
forests where it grows, helps protect the soil, and
Ecology.—Leandra is intermediate in tolerance. As
furnishes food and cover for wildlife. The
single plants and in thickets, it grows naturally in
convenient size, pretty flowers, and foliage
small openings and in the understory of low to
(W3Tropicos 2002), as well as the attractive fruits
moderate basal-area remnant and secondary
suggest leandra as an ornamental.
forests, and in plantations. It occurs at elevations of
427
References W3Tropicos. 2002. Calycogonium krugii Cogn.,
flowering shrub.http://digitalis.mobot.org/mrsid/
Liogier, H.A. 1995. Descriptive flora of Puerto bin/mosid/mosid.pl?client=203&image=MOA-
Rico and adjacent islands. Vol. 4. Editorial de la 06799_001.sid&title=Calycogonium+krugii+ Cog...
Universidad de Puerto Rico, San Juan, PR. 1 p.
617 p.
428
Ledum groenlandicum Oeder Labrador tea
ERICACEAE
Synonyms: Ledum palustre L. ssp. groenlandicum (Oeder) Hulten

Ledum palustre L. var. latifolium (Jacq.) Michx.
Ledum latifolium Ait.
Rhododendron groenlandicum (Oeder) Kron & Judd
James E. Nellessen
McCance and Burns 1984). Labrador tea has been

classified as an endangered species in Ohio, where
one population occurs in a bog habitat in the
northeastern corner of the state (McCance and
Burns 1984). One isolated collection has occurred
in the Black Hills of South Dakota (Great Plains
Flora Association 1986).
Ecology.—This species grows in acidic soils of

bogs, especially sphagnum (Sphagnum spp.) bogs,
and other swampy and wet shoreline habitats in
association with Picea mariana (Mill.) BSP (black
spruce), Chamaedaphne calyculata (L.) Moench.
(leatherleaf), and Kalmia spp. (laurels). Labrador
tea will also grow on some drier upland soils with
a reasonable moisture regime in association with
Pinus spp. (pines) and Vaccinium spp.
General Description.—Labrador tea is an
(blueberries). Stands of Labrador tea appear to
evergreen, well branched, spreading shrub up to
retard black spruce growth and regeneration
about 1 m in height. The twigs are densely hairy
(Inderjit 1996). Labrador tea has been reported as
and the buds scaly. The leaves are simple,
an undergrowth species on serpentine substrates on
alternate, entire (not toothed), lanceolate,
mountain tops in eastern townships of Quebec,
somewhat narrowly elliptic to oblong, 1.5 to 5 cm
Canada. It has also been observed in shrub and
long, about 0.7 to 2 cm wide, thick, leathery and
sedge dominated serpentine ecotonal communities
evergreen with rolled margins, dark green above,
of Newfoundland with a soil pH of 6.27, and
densely off-white to rusty hairy beneath, and
exchangeable soil mineral concentrations: calcium,
fragrant when crushed (Gleason and Cronquist
13 µg/ml, magnesium 237 µg/ml, and nickel 0.72
1963, Britton and Brown 1913). Several
µg/ml (reviewed in Brooks 1987).
synonyms, as given above, may be found for this
species (Kartesz 1994, Britton and Brown 1913).
Ecology in Human Impacted Environments.—
The genus Ledum appears to be monophyletic, and
Exposure to sulfur dioxide has been shown to
cladistic analysis places it as most closely related
cause visible foliar symptoms and significantly
to genus Rhododendron subsection Edgeworthia,
reduce photosynthetic net assimilation rates, but
hence its synonym placing it in the genus
the rate of reduction was significantly slower than
Rhododendon (Kron and Judd 1990).
that in deciduous species (Addison and others
1984). In a radionuclide study, lead-210 and
Range.—This shrub occurs from Greenland,
polonium-210 tended to accumulate in lichens and
across Canada from Labrador to British Columbia,
mosses, while radium-226 tended to accumulate in
north to Alaska, south to New Jersey,
shrubs such as Labrador tea (Sheard 1986).
Pennsylvania, Massachusetts, Michigan,
Labrador tea may be an indicator of lead
Wisconsin, Minnesota, and Washington, with
contamination in the environment in the vicinity of
isolated sites or occurrences in Ohio and South
lead/zinc mines (Pugh and others 2002). Labrador
Dakota (Gleason and Cronquist 1963, Britton and
tea was among four ericaceous shrub species
Brown 1913, Great Plains Flora Association 1986,
naturally revegetating an abandoned vacuum-
429
mined peatland, despite the absence of new as a folk remedy for lung ailments, dysentery,
sphagnum moss colonization (Berube and Lavoie indigestion, and to kill lice and treat leprosy.
2000). Successful traditional use for gout treatment may
be due to the presence of phenolics and tannins
Reproduction.—Labrador tea flowers are within the plant (Owen and Johns 1999). This
relatively small, about 1 cm wide and 2 cm long, species, as well as others in the genus, has
white, with five to seven stamens, and are grouped generally been regarded as having low palatability,
in terminal clusters. It blooms from May to July, being unpalatable, or even slightly poisonous
sometimes as late as August. The fruit is a slender (USDA 1937). Since this species often grows in
capsule, oval in shape, about 5 to 8 mm long and 2 wet boggy habitats, it is generally not accessible to
to 3 mm wide, with a persistent style (Gleason and most livestock. The plant may serve as reindeer
Cronquist 1963, Britton and Brown 1913). The forage in Alaska. Its unpalatability may also be
seeds are elongated, numerous and small. For due to essential oils of the monoterpene family
germination, they should be placed onto a good (sabinene and limonene) and the sesquiterpene
moisture-supplying substrate and covered with a family (alpha- and beta-selinene and germacrone)
clear plastic film (Young and Young 1992). Seeds (Belleau and Collin 1993). Germacrone has been
germinate best after a 30-day cold stratification shown to be a feeding deterrent for snowshoe
period, with optimal germination temperatures of hares (Lepus americanus Erxleben), although they
20 to 25 °C, and light is required (Calmes and do eat the plant to some extent (Reichardt and
Zasada 1982, Baskin and Baskin 2001). The seeds others 1990, MacCracken and others 1988).
are considered to have non-deep physiological
dormancy because of the relatively shorter cold References
periods needed and that some seeds can germinate
shortly after ripening. The cold stratification Addison, P.A., S.S. Malhotra, and A.A. Khan.
period can decrease the germination temperature 1984. Effect of sulfur dioxide on woody boreal
requirement. Full sunlight and high soil moisture forest species grown on native soils and tailings.
seem to be requirements, including a low soil pH, Journal of Environmental Quality 13(3): 333-
e.g. 5.5 (Karlin and Bliss 1983, Baskin and Baskin 336.
2001). Apparently winter-time cuttings will root
well (Dirr and Heuser 1987, Young and Young Baskin, C.C. and J.M. Baskin. 2001. Seeds:
1992). ecology, biogeography, and evolution of
dormancy, and germination. Academic Press,
Growth and Management.—This shrub is San Diego, CA. 666 p.
generally considered a slow growing, late
successional, pre-climax or climax species. The Belleau, F. and G. Collin. 1993. Composition of
low shrub stratum consisting of Labrador tea and the essential oil of Ledum groenlandicum.
leatherleaf was estimated to have an aboveground Phytochemistry (Oxford) 33(1): 117-121.
productivity of 0.4 and 2.0 t/ha/yr on perched and
raised bogs, respectively, in northern Minnesota Berube, M.E. and C. Lavoie. 2000. The natural
(Grigal and others 1985). Rust fungi of the genus revegetation of a vacuum-mined peatland: eight
Chrysomyxa alternate between spruce and years of monitoring. Canadian Field Naturalist
members of the Ericaceae such as Labrador tea. 114(2): 279-286.
Chrysomyxa reticulata sp. nov. may spread from
Ledum spp. to cultivated rhododendrons (Crane Britton, N.L. and A. Brown. 1913 (1970 Dover
2001). edition). An illustrated flora of the northern U.S.
and Canada, Vol. 2. Dover Publications Inc.,
Benefits.—As the name implies, the dried leaves NY. 735 p.
have been used for tea. Native Americans, such as
the Chippewa, have used this shrub for various Brooks, R.R. 1987. Serpentine and its vegetation:
purposes. The root has been used for medicinal A multidisciplinary approach. Dioscorides
purposes to treat ulcers and the leaves for making Press, Portland, OR. 454 p.
tea (Densmore 1928). As a tea this plant has been
used for treating asthma, colds, stomach aches, Calmes, M.A. and J.C. Zasada. 1982. Some
kidney problems, scurvy, and fevers (Foster and reproductive traits of four shrub species in the
Duke 1990). Externally it has been applied as a black spruce forest type of Alaska. Canadian
wash for burns, ulcers, and stings. It has been used Field Naturalist 96: 35-40.
430
Crane, P.E. 2001. Morphology, taxonomy, and Kron, K.A. and W.S. Judd. 1990. Phylogenetic
nomenclature of the Chrysomyxa ledi complex relationships within the Rhodoreae Ericaceae
and related rust fungi on spruce and Ericaceae in with specific comments on the placement of
North America and Europe. Canadian Journal of Ledum. Systematic Botany 15(1): 57-68.
Botany 79(8): 957-982.
MacCracken, J.G., W.D. Steigers, Jr., and P.V.
Densmore, F. 1928 (1974 Dover edition). How Mayer. 1988. Winter and early spring habitat
Indians use wild plants for food, medicine and use by snowshoe hares, Lepus americanus, in
crafts. Dover Publications, New York. 397 p. south-central Alaska, USA. Canadian Field
Naturalist 102(1): 25-30.
Dirr, M.A. and C.W. Heuser. 1987. The reference
manual of woody plant propagation. Varsity McCance, R.M., Jr. and J.F. Burns, eds. 1984.
Press, Athens, GA. Ohio endangered and threatened vascular plants:
Abstracts of state listed taxa. Department of
Foster, S.A. and J.A. Duke. 1990. A field guide to Natural Resources, Columbus, OH. 635 p.
medicinal plants (eastern/central). Peterson Field
Guide Series, Houghton Mifflin Co., Boston, Owen, P.L. and T. Johns. 1999. Xanthine oxidase
MA. 366 p. inhibitory activity of northeastern North
American plant remedies used for gout. Journal
Gleason, H.A. and A.R. Cronquist. 1963. Manual of Ethnopharmacology 64(2): 149-160.
of the vascular plants of northeastern U.S. and
adjacent Canada. D. Van Nostrand Co., New Pugh, R.E., D.G. Dick, and A.L. Fredeen. 2002.
York. 810 p. Heavy metal (Pb, Zn, Cd, Fe, and Cu) contents
of plant foliage near the Anvil Range lead/zinc
Great Plains Flora Association. 1986. Flora of the mine, Faro, Yukon Territory. Ecotoxicology and
Great Plains. University Press of Kansas, KS. Environmental Safety 52(3): 273-279.
1,392 p.
Reichardt, P.B., J.P. Bryant, B.J. Anderson, D.
Grigal, D.F., C.G. Buttleman, and L.K. Kernik. Phillips, T.P. Clausen, M. Meyer, and K. Frisby.
1985. Biomass and productivity of the woody 1990. Germacrone defends Labrador tea from
strata of forested bogs in northern Minnesota. browsing by snowshoe hares. Journal of
Canadian Journal of Botany 63(12): 2416-2424. Chemical Ecology 16(6): 1961-1970.
Inderjit, A.U.M. 1996. Growth and physiological Sheard, J.W. 1986. Distribution of uranium series
responses of black spruce (Picea mariana) to radionuclides in upland vegetation of northern
sites dominated by Ledum groenlandicum. Saskatchewan, Canada I: Plant and soil
Journal of Chemical Ecology 22(3): 575-585. concentrations. Canadian Journal of Botany
64(11): 2446-2452.
Karlin, E.F. and L.C. Bliss. 1983. Germination
ecology of Ledum groenlandicum and Ledum U.S. Department of Agriculture Forest Service.
palustre ssp. decumbens. Arctic Alpine 1937 (1988 Dover edition). Range plant
Research 15: 397-404. handbook. Dover Publications Inc., New York.
816 p.
Kartesz, J.T. 1994. A synonymized checklist of
the vascular flora of the United States, Canada, Young, J. A. and C. G. Young. 1992. Seeds of
and Greenland, Vol. 1, 2nd Edition. Biota of woody plants in North America. Dioscorides
North America Program of the North Carolina Press, Portland, OR. 407 p.
Botanical Garden. Timber Press, Portland, OR.
622 p.
431
Lepianthes peltata (L.) Raf. monkey’s hand
PIPERACEAE
Synonyms: Lepianthes umbellata (L.) Raf.

Piper peltatum L.
Heckeria peltata (L.) Kunth in L.
Pothomorphe peltata (L.) Miq.
Piper umbellatum L.
Pothomorphe dussii Trel. in Stehlé
John K. Francis
7 cm long. In these “fruits,” after development,

have numerous 0.5 mm-long druplets embedded
(Howard 1988, Liogier 1985). The stems, leaves,
and young fruits have a pine-like aromatic smell.
However, overripe spikes, have a smell reminiscent
of oysters or cow’s breath.
Range.—Monkey’s hand is native to Mexico,

Central America, tropical South America, the
Greater and Lesser Antilles, and Trinidad in the
Americas, and Guinea, Sierra Leone, Ivory Coast,
Ghana, and Nigeria in Africa (Burkill 1997,
Howard 1988, Liogier 1985, Liogier and Martorell
2000). The species is also reported as naturalized in
Florida (Langeland and Stocker 2001).
Ecology.—Monkey’s hand grows in full sunlight

and in light to moderate shade. Continually moist
soil is imperative. Rainfall may vary from about
1500 to over 3000 mm of annual precipitation. The
plant usually grows on loamy or clayey soils with
relatively good fertility. In Puerto Rico, monkey’s
General Description.—Common names for
hand grows from near sea level to over 1,000 m in
monkey’s hand include cowheel bush, cow-foot
elevation. Plants may grow singly or in clumps
leaf, baquiña, chapeau g’leau, bois-anisette, fèy a
along streams and roads, in old fields and other
kè, and many others. It is a weak-stemmed shrub
disturbed areas, in forest tree-fall gaps, and on
up to 2 m in height and 3 cm in basal diameter.
nursery benches. Disturbance and bare soil are
This plant usually has a single stem, but may
necessary for successful establishment.
produce multiple sprouts from the root collar if the
top is damaged. The stems have a vertical habit
Reproduction.—Monkey’s hand shrubs bloom
with no branches until the stems reach 0.5 to 1.0 m
and fruit continuously after reaching about 0.5 m in
in height and then form only second order
height. A collection of fresh fruits from Puerto
branches. A taproot does not develop in clayey
Rico weighed an average of 0.95 ± 0.11 g. Seeds
soil, but the plant develops many robust, semi-
separated from these fruits weighed an average of
fleshy lateral roots. Adventitious roots may form at
0.000061 g/seed or 16,400,000 seeds/kg (air-dried).
the lower nodes. The stems may be glaberous or
For unknown reasons, germination on moist filter
pubescent. The leaves have petioles 8 to 20 cm
paper failed entirely (author’s observations). Seeds
long. The leaf blades may be sub-orbicular to
are produced in large numbers and disbursed by
ovoid, peltate or not, and about 15 to 35 cm long
fruit bats and birds. Apparently, only a small
and broad. They have a cordate base and an acute
percent of the seeds germinate and develop,
tip, with 11 to 15 palmate veins. Spikes of minute
because seedlings are scattered and rarely
flowers in small groups are born on peduncles 1 to
432
abundant. Monkey’s hand plants are easy to in many of the same ways in the West Indies
propagate from cuttings. A small group of stem (Liogier 1990).
cuttings were treated with IBA and placed in a mist
bed. After about 5 weeks, 85 percent had rooted.
Because rooting takes place at the nodes, a node 1
Garlon is named here for identification purposes
should be included in the buried portion of the only. The Forest Service does not endorse any
cutting. commercial product.
Growth and Management.—Monkey’s hand References

plants live about 2 years but may perpetuate
themselves by sprouting from the root collar. Burkill, H.M. 1997. The useful plants of West
Plants also root at the nodes when nodes are buried Tropical Africa. Vol. 4. Royal Botanic Gardens,
with soil or leaf mold, or if laid prostrate. The plant Kew, UK. 969 p.
is a common weed in plantations of cacao and oil
palm in Africa (Burkill 1997). Monkey’s hand may Howard R.A. 1988. Flora of the Lesser Antilles,
be controlled by hand pulling. It is important to Leeward and Windward Islands.
remove the pulled material from the site to prevent Dicotyledoneae, Part 1. Vol. 4. Arnold
it from rerooting. If hand pulling is not possible, Arboretum, Harvard University, Jamaica Plain,
plants may be killed by spraying with a 20 percent MA. 673 p.
solution of Garlon1 applied as a basal bark spray or
by cutting the plants and applying a 50 percent Langeland, K.A. and R.K. Stocker. 2001. Control
solution to the cut stumps (Langeland and Stocker of non-native plants in natural areas of Florida.
2001). University of Florida, Cooperative Extension
Service. http://edis.ifas.ufl.edu/BODY_WG209.
Benefits and Medicinal Use.—Monkey’s hand 56 p.
plants are browsed by cattle and other ruminants.
The species is pretty enough to be used as a border Liogier H.A. 1985. Descriptive flora of Puerto
or background plant in landscaping, but there are Rico and adjacent islands. Spermatophyta. Vol.
no current reports of its use in landscaping. In 1. Editorial de la Universidad de Puerto Rico,
several parts of Africa, the leaves and stems are Río Piedras, PR. 352 p.
cooked and eaten as a vegetable with meat and
fish. It is also used as a condiment and as an Liogier, H.A. 1990. Plantas medicinales de Puerto
ingredient in sauces. The fruits and leaves are used Rico y del Caribe. Iberoamericana de Ediciones,
as bait in trapping fish and small birds (Burkill Inc., San Juan, PR. 566 p.
1997). In Haiti, monkey’s hand leaves are used in
combination with other natural products in a Liogier, H.A. and L.F. Martorell. 2000. Flora of
treatment for hypertension (The Temple of Yehwe Puerto Rico and adjacent islands. 2nd Ed.
2001). In Africa, it is used in various ways in Revised. Editorial de la Universidad de Puerto
magical rites to promote good things and ward off Rico, Río Piedras, PR. 382 p.
the influences of evil fetishes. Leaves, root, and
fruits of the plant are also used to control migraine The Temple of Yehwe. 2001. Treatments.
and general pain, as an antiseptic, to control http://www/vodou.org/treatmen.htm. 3 p.
tapeworms, gonorrhea, urinary problems, and
rheumatism (Burkill 1997). Monkey’s hand is used
433
Lepidospartum burgessii B.L. Turner gypsum scalebroom
ASTERACEAE
Synonyms: none
of the involucre on the flower heads clearly
distinguish it from any species in that genus.
Range.—Gypsum scalebroom is both a geographic

and substrate endemic being restricted to certain
gypsum soils in an approximate 1,000 square km
area in northern Culberson County, Texas, and
southern Otero County, New Mexico.
Ecology.—Gypsum scalebroom is found only on

the gypsum flats at the fringe of the northern
Chihuahuan Desert. Dick-Peddie (1993) designated
the region Chihuahuan desert scrub whereas some
authorities have regarded the whole area as part of
the desert plains grassland (Humphrey 1958). The
latter designation may be more appropriate for the
relatively small areas where gypsum scalebroom
occurs because creosote bush, which is the
dominant shrub of the Chihuahuan desert scrub
vegetation type, does not occur with gypsum
General Description.—The common name for scalebroom. Shrubs and sub-shrubs associated
Lepidospartum burgessii is gypsum scalebroom or with Gypsum scalebroom include Atriplex
Burgess’s broomshrub. The botanical name comes canescens (Pursh) Nutt., Opuntia leptocaulis
from the Greek, “lepidotus” meaning scaly and DeCandolle, and Yucca elata (Engelm.) Engelm. as
“spartium” meaning broom (Gledhill 1992). There well as Tiquilia hispidissima (Torr. & Gray) A.
are two other species in the genus Lepidospartum Richards. Poliomintha incana (Torr.) Gray is a
but neither is sympatric with gypsum scalebroom common associate, and Allenrolfia occidentalis (S.
(Baldwin and others 2002). The range of Wats.) Kuntze is also observed to a lesser extent in
Lepidospartum latisquamum S. Wats. extends from Texas. Associated grasses include Bouteloua
southern California to Utah and that of L. breviseta Vasey, Sporobolus nealleyi Vasey, and
squamatum (Gray) Gray ranges from central Sporobolus airoides (Torr.) Torr. Gallardia
California to Baja California (Baldwin and others multiceps Greene, Senecio warnockii Shinners and
2002). Gypsum scalebroom was relatively recently species of Mentzelia and Isocoma are common
described (Turner 1977). Prior to that time, it is forbs associated with gypsum scalebroom,
likely to have frequently been mistaken for although their contribution to the ground cover is
rabbitbrush, Chrysothamnus nauseosus (Pallas ex low. Plants colonize two habitats, semistabilized
Pursh) Britt. Gypsum scalebroom is a woody shrub gypsum dunes and relatively compacted gypsum
with numerous stems growing up to 1.2 m tall. soils at the edge of dry alkaline lakes (Powell
The stems have multiple branches and are covered 1998). In New Mexico the plants generally grow
with silvery, matted, felt-like hairs out of which on stabilized, microbiotic-covered, gypsum soils
protrude numerous small oil blisters. The leaves with approximately 5 percent basal vegetation/litter
are needlelike, alternate, and 5 to 12 mm long. cover. In Texas and at one site in New Mexico,
There are three or four terminal flower heads on shrubs grow on more mobile gypsum dunes with
stems with three, rarely four, bright yellow flowers an average of 20 percent basal vegetation/litter
per head. The achenes are covered by dense white cover (Ladyman and Gegick 2001). The plants
hairs and topped by a pappus of many slender occur singly or in scattered colonies. The climate in
bristles. Although gypsum scalebroom has been this area is relatively extreme with summer
mistaken for rabbitbrush, the three rounded bracts temperature highs between 34 oC and 41 oC and
434
winter temperature ranging from lows of –2 oC and populations are reproductively compatible. Some
highs of 14 oC. Freezing is common during this protective measures, including erecting exclosures,
time. Precipitation comes primarily during the have been made for some colonies growing in an
summer (ranging from 13.1 to 46.8 cm). However, Area of Critical Concern and managed by the
substantial spring rains have been reported, and Bureau of Land Management (BLM 1997). This
there is a high degree of year-to-year variability species appears to be facing relatively imminent
characteristic of most arid and semiarid climates extinction. From a simple model that assumed the
(Noy-Mier 1973). rate of death was linear, extinction is predicted
within 35 years (Ladyman and others 1999).
Reproduction.—The peak of flowering is during
late July through early September, but within a Benefits.—Gypsum scalebroom is visited by a
population, flowering can extend from late April to wide variety of arthropods and possibly provides
early October. The number of flowers per shrub is cover and food for small wildlife species. It is
highly variable; from one flower to literally unlikely to be palatable to livestock, but they do
hundreds of flower heads on any given shrub. appear to brush against it even if they do not use it
There does not appear to be a close relationship as food (anecdotal information from ranchers in
between the size of the plant and the number of area and author’s personal observation). This is
flowers because the largest shrubs are often in a likely due to the absence of other large shrubs in
decadent condition, that is, with more than 50 the region. It is a beautiful silvery ornamental
percent dead wood. In 1991/1992 and 1997 plants shrub. It has been suggested that one way to slow
were counted in seven colonies in New Mexico its eventual extinction is to introduce it into the
(Huenneke 1991, Ladyman and others 1999). horticultural and landscape trade.
Approximately 15 percent of the individuals
counted within those colonies in 1991/1992 were References
dead in 1997. The number of juvenile plants had
also declined. Recruitment is only by clonal Baldwin, B.G., S. Boyd, B.J. Ertter, R.W.
propagation. No seeds have been observed despite Patterson, T.J. Rosatti, and D.H. Wilken. 2002.
abundant yellow flowers and apparently adequate The Jepson Desert Manual, vascular plants of
pollinators (Ladyman and others 1999). The results southeastern California. University of California
of studies on pollen germination on stigmas Press, Berkeley, CA. 624 p.
indicate an incompatibility system is operating
(Ladyman and others 1999). The prematurely
stunted pollen tube growth observed by florescent Bureau of Land Management. 1997. Areas of
microscopy is indicative of sporophytic Critical Environmental Concern Resource
incompatibility, which is typical of members of the Management Plan Amendment, August 1997.
Asteraceae and other species with trinucleate Bureau of Land Management, Las Cruces
pollen (Ladyman and others 1999, de Nettancourt District, Caballo Resource Area, Otero County,
1977). Las Cruces, NM [not paged].
Growth and Management.—Gypsum scalebroom De Nettancourt, D. 1977. Incompatibility in

responds to irrigation and fertilization and can Angiosperms. Springer Verlag, Berlin,
grow to over 1.5 m with luxurious silvery foliage Heidelberg, and New York. 230 p.
in cultivation (author’s personal observation).
Transplantation is possible, but the success rate is Dick-Peddie, W.A. 1993. New Mexico Vegetation
not known. In 1998, two potential disease problems – past, present, and future. University of New
were identified. Corythuca marmorata, a known Mexico Press, Albuquerque, NM. 244 p.
pathogen of some Asteraceae species, caused leaf
loss and stem necrosis. A potential pathogenic Gledhill, D. 1992. The names of plants. Second ed.
fungus, Alternaria alternata, was observed within Cambridge University Press, Cambridge UK.
some flowers (Rotem 1994, Ladyman and Gegick 202 p.
2001). Genetic diversity appears to exist between
the Texas and New Mexico populations. The Huenneke, L.F. 1991. Biological studies of
involucre length of plants growing in Texas was Lepidospartum burgessii, a rare endemic shrub
significantly longer than from those in New of southern New Mexico – First Progress
Mexico (Ladyman and Gegick 2001). It is Report. Bureau of Land Management, New
unknown if individuals between these two Mexico State Office, Santa Fe, NM [not paged].
435
Humphrey, R.R. 1958. The desert grassland, a Noy-Mier, I. 1973. Desert ecosystems:
history of vegetational change and an analysis of environment and producers. In: Annual Reviews
causes. Botanical Review 24:193-252. of Ecology and Systematics IV: 25-51.
Ladyman, J.A.R., L. DeLay, P. Gegick, M. Bogan. Powell, A. Michael. 1998. Trees & shrubs of
1999. Status and Reproductive Biology of Trans-Pecos and adjacent areas. University of
Lepidospartum burgessii (Burgess broomshrub Texas University Press, Austin, TX (First Ed.
or gypsum broomscale). Unpublished report copyright Big Bend Natural History Assoc.) 498
submitted to the USGS Biological Survey, p.
Albuquerque, NM. 108 p.
Rotem, J. 1994. The Genus Alternaria – Biology,
Ladyman, J.A.R. and Patricia Gegick. 2001. Epidemiology, and Pathogenicity. APS, St. Paul,
Status of Lepidospartum Burgessii (Burgess MN. 325 p.
broomshrub or gypsum broomscale). In:
Southwestern Rare and Endangered Plants. Turner, B.L. 1977. Lepidospartum burgessii
Proceedings of Third Conference. RMRS-P-23. (Asteraceae, Senecioneae), a remarkable new
USDA Forest Service, Rocky Mountain gypsophilic species from Trans-Pecos Texas.
Research Station, Fort Collins, CO. p. 116-127. Wrightia 5: 354-355.
436
Leucaena leucocephala (Lam.) de Wit leadtree
FABACEAE
Synonyms: Leucaena glauca Benth.

Mimosa leucocephala Lam.
Leucaena latisiliqua (L.) Benth.
Mimosa glauca L.
Acacia glauca Willd.
John K. Francis
pinnae each with 9 to 20 pairs of leaflets that are

linear-oblong to lanceolate and 8 to 15 mm long.
Solitary or paired inflorescences are 12- to 20-mm
diameter, axillary heads on 2- to 3-cm-long
peduncles. The heads contain numerous fragrant,
greenish-white flowers. Usually four to 10
flattened, brown legumes, 10 to 15 cm long,
develop per inflorescence. Each legume contains
15 to 20 hard, shiny, brown seeds that are flattened
and tear-drop shaped. Leadtree is a polyploid with
2n = 104 chromosomes (Brewbaker 1997, Howard
1988, Liogier 1988, Little and Wadsworth 1964,
Pennington and Sarukhan 1968, Stevens and others
2001).
Range.—Leadtree is native to southern Texas and

the lowlands of Mexico from about 20° N. south to
Nicaragua at about 12° N (Parrotta 2000). The
General Description.—Leadtree occurs in three Mexican state of Oaxaca derives its name from the
forms or types known as the “Hawaiian” or Native American word “Uaxia,” which means the
common type, the “Salvador” or giant type, and the “place where Leucaena grows” (National Academy
“Peru” type. The common type is widespread and of Science 1977). Most authors treat the range in
shrubby and will be dealt with in this document. vague terms or state that it is uncertain. The species
Leadtree is also known as white leadtree, leucaena, is established so completely in the West Indies and
tantan, wild tamarind, koa haole, ipil-ipil, zarcilla, the tropical lowlands of Central and South America
guaje, monval, macata, and a host of other names. that apparently all adapted sites are occupied and
It commonly reaches 2 to 5 m in height and 5 to 10 the population supports many natural enemies.
cm in stem diameter at breast height. Undisturbed, While it is assumed that leadtree was spread to
closed stands usually are of single-stemmed plants; these areas through early Spanish commercial
disturbed plants, especially dispersed ones, almost activity (Parrotta 2000), earlier introductions by
always develop multiple stems. Leadtrees are humans or animals cannot be ruled out without
supported by tap and lateral root systems. They archeological evidence. From the mid-1500’s
root deeply and extensively and form nodules when onward, leadtree has been spread intentionally and
appropriate strains of Rhizobium bacteria are unintentionally throughout the Tropics and the
available. The stem bark is gray or brownish gray, frost-free and lightly frosted subtropics including
more or less smooth with many lenticels. The Florida, Georgia, Arizona, Hawaii, and the U.S.
sapwood is light yellow, and the heartwood is Territories in the Caribbean and Pacific (Natural
brown, hard, heavy (specific gravity 0.7), and has Resources Conservation Service 2002, Pacific
visible annual rings. The branches and twigs are Islands Ecosystems at Risk 2002).
grey-green to gray-brown and somewhat brittle in
older shrubs. Bipinnate leaves 9 to 25 cm long are Ecology.—Leadtree can grow on most soils
borne alternately. There are three to 10 pairs of derived from most parent materials in a wide
variety of site conditions. It grows fairly well on
437
disturbed, eroded, and partially compacted soils. Individual shrubs live 10 to 20 years or more.
However, the species does not grow well in poorly Plantations are relatively easy to establish with
drained soils or periodically flooded soils, in soils containerized seedlings or by direct seeding
with high salinity or those below pH 5.0 or above scarified seed into prepared seedspots at the
pH 8.0, at temperatures below 10 °C, and in areas beginning of the wet season. Because of its
with mean annual rainfall below 750 mm or above aggressive nature, it is not advisable to introduce
2500 mm (Skerman and others 1988). Leadtree is leadtree into areas where it does not already occur.
intolerant of shade. It does best in the open, but Within the native or naturalized range, disturbed
competes aggressively in low stands of weeds and wildlands are usually quickly colonized by leadtree
grass and will grow in low forests with low basal from the soil seedbank or from nearby seed trees.
areas. It often forms thickets that briefly eliminate However, where seed is not present, planting the
nearly everything underneath but soon self-thin to species may be an excellent way of rehabilitating
fairly open stands with normal understory damaged sites. Grass swards should be cultivated
vegetation. Leadtree is drought tolerant. It folds its or burned or sprayed followed by cultivation before
leaves under water stress preventing water loss planting. About 5,000 to 10,000 plants/ha should
and, under severe conditions, will defoliate until be established for erosion control and sward
the rains return (National Academy of Sciences conversion. Dense stands of leadtree needing
1977). control may be slashed and grazed or sprayed with
broadleaf herbicide.
Reproduction.—Leadtree blooms throughout the
year (Pennington and Sarukhan 1968) and may Benefits.—Leadtree quickly invades disturbed
begin blooming as young as 4 to 6 months of age areas and prevents further erosion while allowing
(National Academy of Sciences 1977). The flowers succession to secondary forest cover. It is one of
are largely self-compatible and self-fertilized. It only a few species that can invade and rehabilitate
takes about 4 months after flowering for pods to dense tall grass swards such as Panicum maximum
mature and liberate seeds (Binggeli 1997). From Jacq. and Imperata cylindrica (L.) Raeusch.
10,000 to 21,000 seeds/kg are reported although it (author’s observation, National Academy of
is sometimes unclear from what variety they were Sciences 1977). Leadtree furnishes food and cover
sampled (Brewbaker 1997, Parrotta 2000). A for wildlife, especially browsing animals. The
sample of seeds from shrubby plants from the north species has been planted widely in agroforestry
coast of Puerto Rico averaged 0.0392 ± 0.0006 applications. It is useful for forming erosion
g/seed or 25,500 seeds/kg (author’s observation). barriers, windbreaks, for shade and support, green
Leadtree seeds have a hard coat that requires manure, and cut fodder. It is known to fix useful
scarification for reliable germination. This can be quantities of nitrogen (Parrotta 2000). Plantations
done by abrasion, or treatments with hot water or are established for pasture and cut for hay. The
concentrated acid. Germination of scarified seed is foliage is highly digestible (60 to 70 percent)
usually between 50 and 98 percent in 6 to 10 days (Brewbaker 1997). Young leaves are reported to
after sowing. Seed can be stored for up to 5 years contain 68 calories/100 g and 80 percent moisture
at temperatures of 2 to 6 °C (Parrotta 2000). Pods (Duke 1983). By dry weight, leaves contain 21
open while still attached to liberate their seeds and percent crude protein, 18 percent crude fiber, 8
are dispersed by granivorous and herbivorous percent ash, 6 percent fat, and 46 percent total
animals (Pacific Island Ecosystems at Risk 2002) digestible nutrients (FAO 2002). Cattle readily eat
and machinery. Detached pods are blown some the forage but suffer symptoms of mimosine
distance by wind and occasionally retain attached toxicity if given a pure diet of the species. Pigs and
seeds. Reproduction is common in disturbed sites chickens must not be given more than 5 to 10
near seed sources. Plants aggressively coppice percent in their diet, but goats have no difficulty
when burned, cut, or broken off. The plant can be with the forage, and sheep can become accustomed
propagated vegetatively from cuttings (Parrotta to diets rich in leadtree foliage (National Academy
2000). of Sciences 1977). The wood is a preferred fuel
that burns slowly with little smoke or ash and
Growth and Management.—Leadtree grows makes an excellent charcoal (Brewbaker 1997).
moderately rapidly (not as fast as the giant variety The stems are used for stakes and tool handles. The
for which most of the data are available). Seedlings seeds are widely employed for making jewelry and
may add about 1 m of height in the first year and placemats (Howard 1988). The young pods and
0.5 to 0.75 m for 2 or 3 years thereafter. Diameter green seeds are cooked as a vegetable, and the
growth may vary from 2 to 10 mm per year. mature seeds are used as a coffee substitute and
438
parched for snacks (Duke 1983). The flowers are a Little, E.L., Jr. and F.H. Wadsworth. 1964.
pollen source for honeybees (Little and Wadsworth Common trees of Puerto Rico and the Virgin
1964). Extracts of the roots and bark are a powerful Islands. Agriculture Handbook 249. U.S.
emetic and have been used as an abortifacient Department of Agriculture, Forest Service,
(Liogier 1990). Washington, DC. 548 p.
References National Academy of Sciences. 1977. Leucaena:

promising forage and tree crop for the Tropics.
Binggeli, P. 1997. Leucaena leucocephala (Lam.) National Academy of Sciences, Washington,
de Wit (Mimosaceae). http://members.lycos.co. DC. 115 p.
uk/WoodyPlantEcology/docs.web-sp7.htm. 5 p.
Brewbaker, J.L. 1997. Leucaena leucocephala--a Plants profile: Leucaena leucocephala (Lam.) de
versatile nitrogen fixing tree. Fact Sheet 97-06. Wit white leadtree. http://plants.usda.gov/
Winrock International, Morrilton, AK. cgi_bin/plant_profile.chi?symbol=LELE10. 5 p.
http://www.winrock.org/forestry/factpub/factsh/l
eucaena.htm. 5 p. Pacific Island Ecosystems at Risk. 2002. Leucaena
leucocephala (Lam.) de Wit, Mimosaceae.
Duke, J.A. 1983. Handbook of energy crops: http://www.hear.org/pier_v3.3/leleu.htm. 3 p.
Leucaena leucocephala (Lam.) de Wit. Center
for New Crops & Plants Products, Purdue Parrotta, J.A. 2000. Leucaena leucocephala (Lam.)
University, West Lafayette, IN. http;/www. de Wit Leucaena, tantan. In: J.K. Francis and
hort.purdue.edu/newcrop/duke_energy/Laucaena C.A. Lowe, eds. Bioecología de árboles nativos
_leucocephala.html. 5 p. y exóticos de Puerto Rico y las Indias
Occidentales. General Technical Report IITF-
FAO. 2002. Sistema de información de los 15. U.S. Department of Agriculture, Forest
recursos del pienso: B81 Leucaena leucocephala Service, International Institute of Tropical
(Lam.) de Wit [L. glauca (L.) Benth.]. Food and Forestry, Río Piedras, PR. p. 308-316.
Agriculture Organization of the United Nations,
Rome. http://www.fao.org/livestock/agap/frg/ Pennington, T.D. and J. Sarukhan. 1968. Arboles
afris/espanol/document/tfeed8/Data/417.htm. tropicales de México. Instituto Nacional de
3 p. Investigaciones Forestales, Secretaría de
Agricultura y Ganadaría. México D.F., México.
Howard, R.A. 1988. Flora of the Lesser Antilles, 413 p.
Dicotyledoneae, Part 1. Vol. 4. Jamaica Plain, Skerman, P.J., D.G. Cameron, and F. Riveros.
MA: Arnold Arboretum, Harvard University. 1988. Tropical forage legumes. 2nd Ed. FAO
673 p. Plant Production and Protection Series 2. Food
and Agriculture Organization of the United
Liogier H.A. 1988. Descriptive flora of Puerto Nations, Rome. 692 p.
2. Editorial de la Universidad de Puerto Rico, Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
Río Piedras, PR. 481 p. Montiel, eds. 2001. Flora de Nicaragua.
Monographs in Systematic Botany, Vol. 85, No.
Liogier, H.A. 1990. Plantas medicinales de Puerto 2. Missouri Botanical Garden, St. Louis, MO.
Rico y del Caribe. Iberoamericana de Ediciones, p. 945-1,910.
Inc. San Juan, PR. 566 p.
439
Lindera melissifolia (Walt.) Blume pondberry
LAURACEAE
Synonyms: Lindera melissaefolium (Walt.) Blume

Benzoin melissaefolium (Walt.) Nees
Laurus melissaefolia Walt.
Margaret S. Devall and Nathan M. Schiff
Carolina and South Carolina; it has apparently

been extirpated from Alabama and Louisiana and
possibly Florida.
Ecology.—Pondberry has probably always been a

rare species (Radford and others 1968, Kral 1983),
and knowledge of its ecology is limited. The
species occurs in lowland habitats with hydric
soils (Morgan 1983), in areas that are usually
flooded in winter. In Mississippi pondberry occurs
in bottomland hardwood forests. In northeastern
Arkansas and southeastern Missouri pondberry is
found on the bottoms and edges of shallow
seasonal ponds in old dune fields, but in
southeastern Arkansas it occurs in low habitat
along a river. In South Carolina the species occurs
in areas with karst topography, around the edges of
sinkholes, and in Georgia it occurs along the
borders of sphagnum bogs. Ambient light at the
different sites ranges from deep shade to almost
full sun. It appears that pondberry can occupy
different habitats as long as its requirements for
water are met (Devall and others 2001). The
General Description.—Pondberry, also known as distribution and abundance of pondberry have
southern spicebush (Morgan 1983), a member of been affected by habitat destruction and alteration,
the Lauraceae, is a stoloniferous, deciduous, such as timber cutting and clearing of land. The
aromatic shrub up to 2 m tall. Pondberry usually species was listed as endangered by the U.S. Fish
occurs in clones of numerous stems with erect or and Wildlife Service in 1986 (U.S. Fish and
ascending shoots and few branches. The alternate Wildlife Service 1986). Many of the existing
drooping leaves are subcordate with prominent colonies are small, and occupy only a portion of
venation and pubescence on the lower surface the apparently suitable habitat. Although the
(Klomps 1980). Lindera benzoin var. pubescens pondberry recovery plan states that there are 36
also is pubescent beneath. The species is extant populations (U.S. Fish and Wildlife Service
dioecious, with small yellow flowers that bloom in 1993), new colonies have been discovered since
spring before leaf-out. The fruit is a red drupe 1993, some in new locations and some near known
about 1 cm long that matures in late summer or populations. Some populations that were thought
fall. Female clones are often smaller than male to be separated by enough distance to preclude
clones and are sometimes absent from stands interbreeding (as on the Delta National Forest,
(Wright 1989, 1990). As in many clonal species, MS) may be linked by recently discovered
seedlings are rarely observed (Wright 1990, Devall colonies.
and others 2001).
Reproduction.—Stems flower in the second to
Range.—Pondberry occurs in the Southeastern fourth year of growth (U.S. Fish and Wildlife
United States. At present there are populations in Service 1993). The flowers are pollinated by
Arkansas, Georgia, Mississippi, Missouri, North several species of small bees and are visited by
440
numerous insect species. Flower and fruit Kral, R. 1983. A report on some rare, threatened,
production are highly variable (Morgan 1983, or endangered forest-related vascular plants of
Tucker 1984). Some years the flowers are the South: Volume 1, Isoetaceae through
damaged by late frosts, while other years flower Euphorbiaceae. Technical Publication R8-TP 2.
and fruit production may be heavy, with up to 100 U.S. Department of Agriculture, Forest Service,
fruits per plant. Southern Region, Atlanta, GA. 718 p.
Growth and Management.—Seedlings are rarely Morgan, S. 1983. Lindera melissifolium: A rare
observed (Devall and others 2001). Stems usually southeastern shrub. Natural Areas Journal 3: 62-
live 6 or 7 years, and when a stem dies it is almost 67.
always replaced by a new stem that grows from
the base of the plant. Clones expand vegetatively, Radford, A.E., H.E. Ahles, and C.R. Bell. 1968.
and a mature colony often includes numerous leafy Manual of the Vascular Flora of the Carolinas.
stems along with some dead stems (U.S. Fish and University of North Carolina, Chapel Hill, NC.
Wildlife Service 1993). Most pondberry colonies 1,183 p.
occur in light shade, but a few grow in almost full
sun. In unshaded conditions, competition may be a Smith, C.G. III, P.B. Hamel, M.S. Devall and
problem. More knowledge is needed about N.M. Schiff. 2003. Hermit thrush is the first
management, and studies are currently being observed dispersal agent for pondberry.
conducted on the Delta National Forest in Castanea, in press.
Mississippi and at other locations. The U.S. Fish
and Wildlife Service (1993) states that pondberry Steyermark, J.A. 1949. Lindera melissifolia.
should be protected from forestry and agricultural Rhodora 51: 153-162.
management actions and protected from grazing
and browsing animals. Tucker, G.E. 1984. Status report on Lindera
melissifolia (Walt) Blume. Provided under
Benefits.—The fruits are eaten by hermit thrushes contract to the U.S. Fish and Wildlife Service,
(Catharus guttatus), cardinals (Cardinalis Southeast Region, Atlanta, GA. 41 p.
cardinalis) and other birds (Smith and others
2003). Steyermark (1949) reported that the fruits U.S. Fish and Wildlife Service. 1986. Endangered
are used as ammunition in pop guns, tubular and threatened wildlife and plants;
contrivances constructed from elderberry determination of endangered status for Lindera
[Sambucus nigra L. ssp. canadensis (L.) R. Bolli] melissifolia. Federal Register 51: 27,495-27,500.
twigs.
U.S. Fish and Wildlife Service. 1993. Recovery
References plan for pondberry (Lindera melissifolia). U.S.
Fish and Wildlife Service. Atlanta, GA. 56 p.
Devall, M., N. Schiff, and D. Boyette. 2001.
Ecology and reproductive biology of the Wright, R.D. 1989. Reproduction of Lindera
endangered pondberry, (Lindera melissifolia melissifolia in Arkansas. Proceedings of the
(Walt) Blume. Natural Areas Journal 21: 250- Arkansas Academy of Science 43: 69-70.
258.
Wright, R.D. 1990. Photosynthetic competence of
Klomps, V.L. 1980. The status of Lindera an endangered shrub, Lindera melissifolia.
melissifolia (Walt.) Blume, pondberry, in Proceedings of the Arkansas Academy of
Missouri. Transactions of the Missouri Academy Science 44: 118-120.
of Science 14: 61-66.
441
Lonicera japonica Thunb. Japanese honeysuckle
CAPRIFOLIACEAE
Synonyms: Nintooa japonica (Thunb.) Sweet
John K. Francis
black seeds (Miller 1999, Munger 2002, Stephens

1973). Japanese honeysuckle is a diploid plant
with 2n = 18 chromosomes (Schierenbeck and
others 1995).
Range.—Japanese honeysuckle is native to Japan,

Korea, China, and Taiwan (Bravo 2003, Pacific
Island Ecosystems at Risk 2003). The species has
naturalized in much of the United States except
Alaska and a few states in the Northwest, northern
Midwest, and Vermont (Natural Resources
Conservation Service 2003). It has also naturalized
in Puerto Rico (Liogier and Martorell 1982),
Ontario, Canada (Munger 2002), Juan Fernandez
archipelago, Chile (Sharma and others 1997),
Australia, New Zealand, Christmas Island (Pacific
Island Ecosystems at Risk 2003), England,
Portugal, Corsica, Brazil, and Argentina (Nuzzo
2003). There are two generally accepted varieties,
chinensis and hilliana. The latter is the variety
commonly encountered in naturalized populations
(Nuzzo 2003).
Ecology.—Japanese honeysuckle grows well in a

wide variety of soils with a few exceptions such as
coarse sands and poor peat soils (Munger 2002). It
General Description.—Japanese honeysuckle,
invades fencerows, roadsides, vacant lots, forest
also known as woodbine and Chinese
openings, understories of open forest stands, and
honeysuckle, is a semi-evergreen woody vine that
abandoned pastures and fields. The species is
forms mounds and mats on open ground and by
present in nearly all the forest associations in the
twining, climbs shrubs, low trees, windfalls, and
Southeastern U.S., the region where it is most
fences. The ropy stems, which have gray-brown,
common. Japanese honeysuckle is adapted to
fissured to shredding bark, can extend 5.5 m or
climbing small-diameter stems and sometimes
more and reach 10 cm in diameter in old plants.
smothers or strangles small trees. It cannot ascend
The wood is white, soft, and fine-grained. Young
trees with trunks larger than 15 cm in diameter
stems are reddish brown to light brown and about
(Nuzzo 2003). It grows best under full sunlight but
3 mm in diameter with hollow pith. Roots are
can still survive with 25 percent of sunlight in a
extensive, reaching 2.5 m laterally and 15 to 100
forest understory. Retaining leaves and
cm in depth. Opposite, light-green leaves are 2.5 to
photosynthesizing through all or part of the winter
12 cm long, ovate to oblong, entire to lobbed, with
after canopy forest trees have defoliated helps
short petioles. The leaves are evergreen in tropical
plants survive through the summer in low-light
and subtropical settings and become late deciduous
conditions (Nuzzo 2003). Northern distribution of
to deciduous in cooler temperate climates. The
naturalized Japanese honeysuckle is limited by
paired axillary flowers are 2 to 3 cm long, white
short growing seasons and late spring frosts, and is
(sometimes tinged with pink or purple) turning
generally south of an isotherm where mean
yellow, tubular with five lobes, fused in two
January temperature is -1 °C. It has not generally
unequal lips. The fruits are black, glossy, globose
naturalized west of the 1000 mm mean annual
berries 4 to 6 mm in diameter containing two to 10
442
precipitation limit (except where exogenous water mentioned herbicides to the cut surfaces (Bravo
is available), roughly corresponding with native 2003). Mowing is ineffective (Nuzzo 2003).
hardwood forests. Japanese honeysuckle survives
fire by sprouting from stem tissue buried under Benefits and Detriments.—In spite of the
litter and mineral soil. It generally disappears from invasive and weedy tendencies of the species,
communities (such as longleaf pine) where fires Japanese honeysuckle is still widely used as an
are frequent (Munger 2002). ornamental. It is especially desirable in arid areas
where it has little tendency to spread. Managed for
Reproduction.—Japanese honeysuckle in the U.S. screens and ground cover, Japanese honeysuckle is
flowers from April through July (sometimes beautiful in and out of flower and especially
through October) varying by local climate (Nuzzo captivating because of the strong and pleasant
2003). The flowers open in late afternoon and aroma of its blooms. Some gardeners favor it
remain open for about 3 days (Nuzzo 2003). The because it attracts hummingbirds. The fruits are
flowers are pollinated by hummingbirds and toxic to humans if large quantities are eaten and
insects (nocturnal hawkmoths and diurnal bees). result in vomiting, diarrhea, rapid heartbeat,
The species requires pollen from genetically respiratory failure, and coma. However, children
distinct individuals for successful pollination and suck the nectar from flowers without ill effects
as a result is pollinator limited. Naturally- (Russell 1997). The species was formerly planted
pollinated flowers produced seeds in 17.4 percent for erosion control and wildlife benefits. Planting
of the resulting fruits while hand-pollinated is currently discouraged or prohibited because the
flowers produced seeds in 78.7 percent of the species has dominated large areas and threatens
fruits (Larson 2001). The seeds are dispersed sensitive native species (Nuzzo 2003). Japanese
primarily by birds. Germination of most seeds honeysuckle is important wildlife cover and an
appears to occur in the spring following dispersal important, sometimes the most important, browse
and occurs at similar rates in mineral soil and species for white tailed deer in Eastern and
under leaf litter (Munger 2002). Japanese Southern United States. It is also eaten by
honeysuckle regenerates vegetatively by sprouting cottontail rabbits, cattle, sheep, and goats. The
from underground stems and by the layering of leaves yield 8 to 12 percent crude protein and have
above-ground stems that come in contact with the an in vivo dry matter digestibility of 35 to 70
soil. Discarded prunings will also take root. percent. Wild turkeys, northern bobwhite, and a
number of songbirds eat the fruits (Munger 2002).
Growth and Management.—Seedlings grow Japanese honeysuckle tissues contain anti-
slowly for the first 2 years but afterwards may complementary polysaccharides and polyphenolic
extend their stems 1.5 m/year (Nuzzo 2003). The compounds. Used in herbal medicine, extracts are
species begins bearing fruit at age 3 in full sun and reported to protect against cellular injury, help
at age 5 when growing in the shade. Fruit maintain vascular health, and prevent infections
production generally peaks between ages 4 and 6 (Nuzzo 2003)
(Munger 2002). By fertilization, researchers were
able to increase browse produced by Japanese References
honeysuckle by over 2,000 kg/ha and increase
crude protein as well (Dyess and others 1994). The Bravo, M.A. 2003. Japanese honeysuckle,
species is easy to propagate with cuttings and most Lonicera japonica Thunb. Alien Plant Working
ornamental plants are multiplied in this fashion. Group, National Park Service, Washington, DC.
The greater challenge is to eliminate infestations http://www.nps.gov/plants/alien/fact/loja1.htm.
of Japanese honeysuckle. Pulling and grubbing 6 p.
individual plants and small infestations can be
effective but will require repeat visits to eliminate Dyess, J.G., M.K. Causey, and H.L. Stribling.
missed sprouts and new seedlings. Tethered goats 1994. Effects of fertilization on production and
can provide effective control for spot infestations quality of Japanese honeysuckle. Southern
(Bravo 2003). Over-the-top spraying with Journal of Applied Forestry 18(2): 68-71.
glyphosate, Dichlorprep + 2,4-D, picloram + 2,4-
D, tebuthiuron, sulfometuron, or triclopyr will kill Larson, K.C. 2001. Lack of pollinators limits fruit
it (Nuzzo 2003) but also tend to eliminate most set in the exotic Lonicera japonica. Botanical
other plant species as well. A more selective Society of America, Botany 2001 Abstracts.
method is to cut stems and apply the above p. 39.
443
Liogier, H.A. and L.F. Martorell. 1982. Flora of Nuzzo, V. 2003. Element stewardship abstract for
Puerto Rico and adjacent islands: a systematic Lonicera japonica, Japanese honeysuckle. The
synopsis. Editorial de la Universidad de Puerto Nature Conservancy, Arlington, VA.
Rico, Río Piedras, PR. 342 p. http://tncweeds.ucdavis.edu/esadocs/documnts/
lonijap.html. 23 p.
Miller, J.H. 1999. Japanese honeysuckle, Lonicera
japonica. Protection Report RX-PR 036. U.S. Pacific Island Ecosystems at Risk. 2003. Lonicera
Department of Agriculture, Forest Service, japonica Thunb., Caprifoliaceae. http://www.
Region 8, Atlanta, GA. 2 p. hear.org/pier/lojap.htm. 3 p.
Munger, G.T. 2002. Lonicera japonica. In: U.S. Russell, A.B. 1997. Poisonous plants of North
Department of Agriculture, Forest Service, Carolina, scientific name: Lonicera japonica.
Rocky Mountain Research Station, Fire Sciences North Carolina State University, Charlotte, NC.
Laboratory. Fire Effects Information System. http://www.ces.ncsu.edu/depts/hort/consumer/
http://www.fs.fed.us/database/feis/plants/vine/ poison/Lonicja.htm. 2 p.
lonjap/all.html. 43 p.
Schierenbeck, K.A., J.L. Hamricks, and R.N.
Natural Resources Conservation Service. 2003. Mark. 1995. Comparison of allozyme variability
Plants profile: Lonicera japonica Thunb. in a native and an introduced species of
http://plants.usda.gov/cgi_bin/plant_search.cgi? Lonicera. Heredity 75(1): 1-9.
mode=Scientific+Name&keywordquery+Lonice
ra+japonica&earl=plant_search.cgi. 6 p. Sharma, A., A. Murayama, T. Osaki, K. Ooi, T.
Yahara, M. Ikegami, Silva-O., M. 1997. Plant
conservation in the Juan Fernandez archipelago,
Chile. Aliso 16(2): 89-101.
Stephens, H.A. 1973. Woody plants of the North

Central Plains. The University Press of Kansas,
Laurence, KS. 530 p.
444
Lotus scoparius (Nutt.) Ottley California broom
FABACEAE
Synonyms: Hosackia scoparia Nutt. in T. & G.

Syrmatium galbrum Vogel
Hosackia glaber Greene
Hosackia crassifolia Nutt., not Benth
Lotus glaber Greene, not Mill.
Arlee M. Montalvo
suffrutescent shrubs grow from 0.4 to 1.5 m tall

but prostrate forms often occur along the coastal
strand. A spreading, prostrate ecotype occurs on
coastal sand dunes at Monterey Bay. The many
flexible green branches arise from a woody base,
and are mostly glabrous except toward tips. The
soft, alternate leaves are pinnately compound with
three (occasionally four to five) oblong to
oblanceolate leaflets, usually 4 to 10 mm long with
short, sparse, appressed hairs, and are subtended
by small, gland-like stipules. The sessile, usually
two- to seven-flowered, umbellate inflorescences
are in the axils of leaves. The yellow pea-like
flowers are 7 to 10 (occasionally to 15) mm long,
have short pedicels, and sometimes bear a red
splotch of varying intensity on the back of the
banner. Depending on the plant, flowers fade to
dark orange or red following pollination. Plants
bear hundreds of inflorescences and are an
attractive flag for native bees. The small
indehiscent pods (figure: top right) bear one to two
(rarely three) seeds that vary from brown to
greenish brown with or without dark brown
General Description.—California broom is a mottling. Seeds of var. scoparius are narrow bean-
diverse and widely distributed subshrub (Munz shaped, 1.2 - 2mm long. Seeds of var. brevialatus
and Keck 1968, Isely 1981, Hickman 1993). The are narrower (ca. 3 x longer than wide). Both
Latin name “scoparius” refers to the broom-like varieties have 2n = 14 chromosomes (Munz and
form which is also evident in the common name Keck 1968, Grant 1995).
“California broom.” The other common name,
“deerweed,” refers to its use as browse by deer. Of Range.—California broom occurs primarily after
five previously recognized subspecific taxa (Ottley fire or in open areas of coastal sage scrub,
1923, Munz and Keck 1968), the three from the chaparral, desert scrub, washes, coastal strand, or
Channel Islands were reassigned to L. dendroideus along roadsides. The var. scoparius is distributed
(Greene) Greene by Isley (1981). His treatment of in cismontane California below 1,500 m from
the genus recognizes two varieties of Lotus Humboldt and Plumas Counties, south into Baja
scoparius (L. s. var. scoparius and L. s. var. California, primarily in shrublands and associated
brevialatus Ottley) and has been adopted in open areas, and primarily in the coastal regions of
Hickman (1993). In the former, the keel of the California and north of Los Angeles. In addition to
corolla is about as long as the wings and the calyx the distinct differences in floral form between the
has broad triangular teeth (figure: top left and two varieties, there is variation in floral form
bottom left). In the latter, the keel is longer than within var. scoparius that is associated with
the shortened wings and the calyx has narrow teeth coastal verses inland habitats from the Transverse
(figure: top center and bottom right). The erect Range of California, northward (author’s
445
observation). Variety brevialatus occurs from Los Jones and Cruzan 1999) primarily by native bees
Angeles Co., south into Baja California primarily in the genera Bombus, Hoplitus, Anthophora,
in the hotter and drier interior regions of Riverside, Habropoda, Osmia, and Anthidium, but flowers
Los Angeles, western San Bernardino, and eastern are also visited by butterflies and non-native
San Diego Counties. Although the two varieties honeybees (author’s observation, Jones and
have somewhat different geographic distributions Cruzan 1999). The indehiscent pods ripen in about
and are easily distinguished by floral characters 4 to 6 weeks and are primarily passively dispersed
(Isely 1981, Steppan 1991), they overlap and short distances. The hard seeds require heat or
hybridize in some areas of Los Angeles, Riverside, mechanical scarification to break dormancy. Of
and San Diego Counties. Putative hybrids have several treatments including control, soil heated to
been observed in contact zones between the two 100 oC for 1 hr, ash /chemical fertilizer, and heat
varieties (Isely 1981, Steppan 1991, Montalvo and plus fertilizer, Christensen and Muller (1975)
Ellstrand 2001). Flowers of synthetic F1 hybrids found that heat treatment yielded the highest
are somewhat intermediate, but have wings as long germination. In addition, Keeley (1987) found that
as the keel. Many areas of overlap may be due to heating seeds in their pods to 120 oC for 5 minutes
natural secondary contact, but some are clearly due increased germination over that of unheated
to seeding projects along highways and utility controls.
corridors. The various floral forms are maintained
when plants are grown from seed in a common Genetics.—There are significant genetic
environment (Montalvo and Ellstrand 2000, 2001). differences among populations of the two varieties
of California broom (Montalvo and Ellstrand
Ecology.—In nature, seeds typically germinate 2000, 2001). An analysis of genetic marker data
after scarification by fire but some germinate in (13 allozyme loci) from three populations of var.
open, disturbed sites (Munz and Keck 1968, brevialatus and nine populations of var. scoparius
Keeley and Keeley 1984). Between 2 to 3 years showed significant population substructure due
after fire in sage scrub vegetation, following a primarily to differences among populations of the
burst of herbaceous species, California broom can two varieties. In an analysis of all populations, 18
become the dominant canopy species, eventually percent of the variation was due to differences
becoming replaced by long-lived shrub species. Its among populations, while analysis of just var.
abundance gradually decreases in 5 to 10 years brevialatus or var. scoparius populations showed
after fire. The branched tap root is relatively only 1 and 8 percent of the variation due to
shallow (< 1 m), and roots form symbiotic differences among populations. Thus, populations
associations with nitrogen fixing bacteria and within a variety are substantially more genetically
arbuscular mycorrhizal fungi (author’s similar to each other than to populations of the
observation). Plants are facultatively drought- other variety. Inbreeding coefficients were low for
deciduous, a trait commonly associated with all populations, a pattern consistent with
shallow-rooted shrubs of coastal sage scrub substantial cross pollination, severe loss of inbred
vegetation in California. Seasonality of leaf progeny, or both (mean f = 0.09, range = 0.00 –
production, nutrient accumulation, and leaf drop in 0.18) (Montalvo, Clegg, and Ellstrand, manuscript
response to summer drought has been studied in preparation).
extensively in L. scoparius var. scoparius (Nilsen
There is also genetically based geographic
and Muller 1980, 1981a,b, 1982, Nilsen and
variation in floral form in this species. In San
Schlesinger 1981, Nilsen 1982). Other researchers
Diego Co., Steppan (1991) detected distinct
have examined response to photoperiod
discontinuities in floral morphology between
(Comstock and Ehleringer 1986).
varieties and moderate correlations among
environmental variables and floral traits of wild
Reproduction.—Seeds of California broom
populations. Montalvo and Weaver (unpublished
germinate in mid to late winter during the rainy
data) measured floral traits on 12 populations of
season. Plants establish quickly with normal
plants raised in a common environment and made
rainfall and typically reach flowering size the
pairwise comparisons of genetic, floral,
second year. Flowering occurs primarily from
geographic, and environmental distances of source
March to June but may start as early as January in
populations. Floral morphological distance
warm, wet winters, and last much longer in more
correlated with environmental distance, genetic
moist, coastal areas and in years with long-lasting
distance correlated with floral distance, but genetic
soil moisture. Flowers are self-compatible and
distance did not correlate with geographic distance
insect-pollinated (Moldenke 1976, Hickman 1993,
446
until var. brevialatus was removed from the July depending on location. Pods or cleaned seeds
analysis. Results were consistent with floral form can be covered with boiling water and left to soak
having a genetic basis and with environmental to break dormancy (Atwater 1980, Young and
factors playing a role in the evolution of floral Young 1986, Emery 1988, Montalvo and Ellstrand
divergence. 2000, 2001), or exposed to dry heat (Keeley 1987).
Whole pods can be dry or wet broadcast in the fall
The substantial genetic differentiation of
for revegetation and restoration. Seedlings plugs
populations has implications for translocation of
can also be used for small projects. In greenhouse
California broom for restoration. Common garden
plantings, seeds germinate best when planted less
experiments testing plants originating from seed
than 2 mm deep (author’s observation). Despite
collected from 12 source populations (both
the strong geographic and morphological
varieties represented) demonstrated a significant
differences, the two varieties have been used
home site advantage (Montalvo and Ellstrand
indiscriminately in many seeding projects. Due to
2000). Furthermore, Montalvo and Ellstrand
the adaptive differences and the observed
(2001) directly tested the potential for
outbreeding depression following hybridization,
“outbreeding depression,” a loss of fitness upon
land managers should use the variety native to the
crossing genetically differentiated populations, by
planting location.
crossing individuals from six populations of the
two varieties in every combination and testing the
Benefits.—California broom is an important, fast
progeny in two common gardens at wild sites.
growing, early successional species used
Seeds per flower and seedling emergence
extensively in erosion control, post-fire mitigation,
decreased significantly with an increase in genetic
and habitat restoration in California, especially in
distance of the crossed parental populations.
the rapidly disappearing coastal sage scrub. The
Among variety crosses were only 70 percent as fit
flowers and seeds are an important food resource
as within variety crosses by the time seedlings
for a variety of insects and seed foraging rodents
emerged, and further fitness differences
and birds (Duncan 1968). The plants provide
accumulated after seedlings were outplanted into
valuable forage for deer, especially in drought
field plots. In these common gardens, success of
years when growth of herbaceous vegetation is
progeny decreased with increasing differences
sparse (Dale 2000).
between parental environments and the transplant
location. Results indicate that careful attention
References
should be given to the similarity of taxonomic
position and environment of source populations Atwater, B.R. 1980. Germination, dormancy and
relative to planting location, especially when morphology of the seeds of herbaceous
choosing source populations for restoration, ornamental plants. Seed Science and Technology
mitigation, and roadside landscaping. 8: 523-573.
Christensen, N.L. and C.H. Muller. 1975. Relative
Interspecific Hybridization.—Putative hybrids
importance of factors controlling germination and
have been reported between California broom and
seedling survival in Adenostoma chaparral. The
L. junceus (Benth.) Greene and L. benthamii Green
American Midland Naturalist 93: 71-78.
in central and northern California, as well as
between other members of the species complex Comstock, J. and J.R. Ehleringer. 1986.
(Isely 1981), but there have not been any genetic Photoperiod and photosynthetic capacity in Lotus
studies confirming introgression. Liston and others scoparius. Plant, Cell and Environment 9: 609-
(1990), however, did genetic studies on San 612.
Clemente Is. and documented hybridization
between the rare L. dendroideus (Greene) Greene Dale, N. 2000. Flowering Plants: The Santa
var. traskiae (Nodden) Isely, formally L. scoparius Monica Mountains, Coastal and Chaparral
Regions of Southern California. California Native
ssp. traskiae (Noddin) Raven, and the more
widespread L. argophyllus (A. Gray) E. Greene Plant Society, Sacramento, CA. 240 p.
var. ornithopus (E. Greene) Ottley. They Duncan, D.A. 1968. Food of California quail on
concluded that genetic assimilation of the rare burned and unburned central California foothill
species by the widespread species is possible. rangeland. California Fish and Game 54: 123-127.
Growth and Management.—The small pods of Emery, D.E. 1988. Seed propagation of native
California broom can be collected from May to California Plants. Santa Barbara Botanical
Garden, Santa Barbara, CA. 23 p.
447
Grant, W.F. 1995. A chromosome atlas and Munz, P.A. and D.D. Keck. 1968. A California
interspecific-intergenic index for Lotus and Flora with Supplement. University of California
Tetragonolobus (Fabaceae). Canadian Journal of Press, Berkeley, CA. 1,681 + 224 p.
Botany 73: 1,787-1,809.
Nilsen, E.T. 1982. Productivity and nutrient cycling
Hickman, J.C., ed. 1993. The Jepson Manual: in the early postburn chaparral species Lotus
Higher Plants of California. University of scoparius. In: C. E. Conrad and W. C. Oechel,
California Press, Ltd., Berkeley, CA. 1,400 p. eds. General Technical Report PSW-58. Pacific
Southwest Forest and Range Experiment Station,
Isely, D. 1981. Leguminosae of the United States III
USDA Forest Service, Berkeley, CA. p. 291-296.
subfamily Papilionoidae: tribes Sophoreae,
Podalyrieae, Loteae. In: Memoirs of the New Nilsen, E.T. and W.H. Muller. 1980. An evaluation
York Botanical Garden. Volume 25. The New of summer deciduosness in Lotus scoparius
York Botanical Garden, Bronx, NY. p. 127-191. (Nutt.) T.& G. The American Midland Naturalist
103: 88-95.
Jones, C.E. and M.B. Cruzan. 1999. Floral
morphological changes and reproductive success Nilsen, E.T. and W.H. Muller. 1981a. The influence
in deer weed (Lotus scoparius; Fabaceae). of low plant water potential on the growth and
American Journal of Botany 86: 273-277. nitrogen metabolism of the native California fire
response shrub Lotus scoparius. American Journal
Keeley, J.E. 1987. Role of fire in seed germination
of Botany 68: 402-407.
of woody taxa in California chaparral. Ecology
68:434-443. Nilsen, E.T. and W.H. Muller. 1981b. Phenology of
the drought-deciduous shrub Lotus scoparius:
Keeley, J.E. and S.C. Keeley. 1984. Postfire
climatic controls and adaptive significance.
recovery of California coastal sage scrub. The
Ecological Monographs 51: 323-341.
American Midland Naturalist 111: 105-117.
Nilsen, E.T. and W.H. Muller. 1982. The influence
Liston, A., L.H. Rieseberg, and O. Mistretta. 1990.
of photoperiod on drought induction of dormancy
Ribosomal DNA evidence for hybridization
in Lotus scoparius. Oecologia 53: 79-83.
between island endemic species of Lotus.
Biochemical Systematics and Ecology 18: 239- Nilsen, E.T. and W.H. Schlesinger. 1981.
244. Phenology, productivity, and nutrient
accumulation in the post-fire chaparral shrub
Moldenke, A.R. 1976. California pollination
Lotus scoparius. Oecologia 50: 217-224.
ecology and vegetation types. Phytologia 34: 305-
361. Ottley, A.M. 1923. A revision of the Californian
Species of Lotus. University of California
Montalvo, A.M. and N.C. Ellstrand. 2000.
Publication in Botany, Berkeley, CA. 10: 189-
Transplantation of the subshrub Lotus scoparius:
305.
testing the home site advantage hypothesis.
Conservation Biology 14: 1034-1045. Steppan, S.J. 1991. Geographic distribution of
flower morphological traits in subspecies of Lotus
Montalvo, A.M. and N.C. Ellstrand. 2001. Non-
scoparius. Journal of Biogeography 18: 321-331.
local transplantation and outbreeding depression
in the subshrub Lotus scoparius (Fabaceae). Young, J.A. and C.G. Young. 1986. Collecting,
American Journal of Botany 88: 258-259. Processing, and Germinating Seeds of Wildland
Plants. Timber Press, Portland, OR. 236 p.
448
Ludwigia octovalvis (Jacq.) Raven primrose willow
ONAGRACEAE
Synonyms: Oenothea octovalvis Jacq.

Jussiaea suffruticosa L.
Jussiaea pubescens L.
Jussiaea octovalvis (Jacq.) Sw.
Jussiaea angustifolia Lam.
(many others Howard 1989, Liogier 1995)
John K. Francis
lateral branches. In the next one or more years,

thelateral branches each produce similar secondary
or tertiary branches. The alternate leaves are linear
to narrowly ovate with blades 1.3 to 16.2 cm long.
The species is highly variable in leaf size, shape,
and pubescence. Primrose willow flowers are
solitary in the upper leaf axils. Four sepals form a
peg-like hypanthium, 5 to 12 mm long, from which
emerge four yellow petals. The twigs, leaf veins,
and hypanthium are often tinged in red. The
capsules, which have a clove-like shape, are 2.5 to
5 cm long and contain many rounded, black seeds.
Chromosome number for the species is 2n = 32, 48
(Howard 1989, Jones 1975, Liogier 1995, Long
and Lakela 1976).
Range.—The original range of primrose willow is

unknown. It grows today from Southern United
States through the West Indies, and from Mexico
through Central and South America to at least
Bolivia (Hodgdon Herberium 2002, Natural
Resources Conservation Service 2002). It has
naturalized in Hawaii (Fish and Wildlife
Information Exchange 2002) and grows in tropical
and subtropical regions almost worldwide (Burkill
1997, Howard 1989, Liogier 1995).
General Description.—Primrose willow, also Ecology.—Primrose willow grows in wet soils

known as Jamaica loostrife, wild clove, many-seed, and shallow water, such as drainage ditches,
kamole, cangá, yerba de clavo, girofle-ma, grand borrow pits, sloughs, marshes, ponds, seasonally
giroflé, and manger mouton, is a suffruticose shrub flooded bottoms, river banks, and rice paddies.
or perennial woody herb to 2 m in height and 1 cm Soil type within those sites is apparently not very
in stem diameter. The stems of older plants are important. Primrose willow grows from near sea
woody through most of their height; branches level to at least 800 m in elevation (Missouri
become increasingly succulent toward the tips. Botanical Garden 2002). It prefers full sun and
Only the lower parts of young plants are woody. tolerates no more than side shade. The species
Primrose willow plants are supported by “sinker” grows singly or in clumps.
roots, one of which may be the taproot, and long
white, lateral roots that grow just under the soil Reproduction.—After primrose willow plants in
surface. These roots have a hard central core and the tropics reach about 0.4 m, blooming and
are surrounded by a cottony mass of root hairs. In fruiting is continuous (author’s observation, Long
the first year, the stem produces a number of fine
449
and Lakela 1976). In Texas, it is reported to bloom Correll, D.S. and M.C. Johnston. 1970. Manual of
from July through October (Correll and Johnston the vascular plants of Texas. Texas Research
1970). Seeds collected in Puerto Rico averaged 54 Foundation, Renner, TX. 1,881 p.
million per kg. Placed on moist filter paper, 28
percent of them germinated within 14 days, most Fish and Wildlife Information Exchange. 2002.
on the seventh day (author’s observation). Local Species moorhen, common, Hawaiian.
dispersal is by wind and water. Long-distance http://fwie.vt.edu/WWW/esis/lists/e101014.htm.
dispersal is probably accidental on birds, 17 p.
machinery, or materials being shipped.
Hodgdon Herbarium. 2002. Dicotyledons
Growth and Management.—Primrose willow associated with Bolivian wetlands.
grows up to 1 m/year. Under favorable conditions, http://unh.edu/herbarium/bolivia/dicots.htm. 5 p.
plants live at least 3 years and possibly more.
There seems to be little reason to plant the species. Howard, R.A. 1989. Flora of the Lesser Antilles,
Control may be necessary in drainage ditches to Leeward and Windward Islands. Vol. 6. Arnold
maintain flow and in rice paddies. Control can be Arboretum, Harvard University, Jamaica Plain,
achieved by cultivation, hand pulling, and by MA. 658 p.
applying commercial weed killers.
Jones, F.B. 1975. Flora of the Texas Costal Bend.
Benefits and Detrements.—Primrose willow Mission Press, Corpus Christi, TX. 262 p.
grows in wet areas where few other plants will
grow and as such it aids in the successional Liogier, H.A. 1995. Descriptive flora of Puerto
process. It furnishes cover for wildlife. Its yellow Rico and adjacent islands. Vol. 4. Editorial de
flowers add a splash of color to areas often devoid la Universidad de Puerto Rico, San Juan, PR.
of colorfully flowering plants. In herbal medicines, 617 p.
preparations of the leaves are used to treat diarrhea,
dysentery, as a laxative and a vermifuge, and to Liogier, H.A. 1990. Plantas mediciales de Puerto
relieve headache, chest pains, and rheumatoid pain Rico y del Caribe. Iberoamericana de Ediciones,
(Burkill 1997, Liogier 1990, Parrotta 2001). A test Inc., San Juan, PR. 566 p.
of a local crude drug from Taiwan made from
primrose willow effectively inhibited the Long, R.W. and O. Lakela. 1976. A flora of
cariogenic bacteria Streptococcua mutans (Chen Tropical Florida. Banyon Books, Miami, FL.
and others 1989). The plant is a major weed in rice 962 p.
in the tropics and sometimes chokes drainage
ditches and irrigation canals (Burkill 1997). Missouri Botanical Garden. 2002. Manual de la
flora de Costa Rica. http://www.mobot.org/
References manual.plantas/028593/S028700.html. 3 p.
Burkill, H.M. 1997. The useful plants of West Natural Resources Conservation Service. 2002.
Tropical Africa. Vol. 4. Royal Botanic Gardens, Plant profile for Ludwigia octovalvis (Jacq.)
Kew, UK. 969 p. Raven ssp. octovalvis. http://plants.usda.gov/
cgi_bin/plant_profile.cgi?symbol=LUOCO. 2 p.
Chen, C.P., C.C. Lin, and T. Namba. 1989.
Screening of Taiwanese crude drugs for Parrotta, J.A. 2001. Healing plants of Peninsular
antibacterial activity against Streptococcus India. CABI Publishing, Wallingford, UK and
mutans. Journal of Ethnopharmacology 27(3): New York. 917 p.
285-295.
450
Lupinus arboreus Sims yellow bush lupine
FABACEAE
Synonyms: None
Christopher Ross
2001). The California Native Plant Society places

it on the “A” List of invasive weeds in Humboldt
County; however, it is apparently only a serious
invader of dune communities (Anderson et al.
1999). Most lupines are poisonous to large
herbivores, but I found no data on this species.
Yellow bush lupine is a strong nitrogen fixer, and
is used in New Zealand as a planted precursor to
tree planting in unstable, nitrogen-deficient dune
environments, where it has been estimated to
contribute 220 kg of nitrogen/ha in 4 ½ years of
growth (Gadgil 1971). In California coastal
prairies it is a facilitator of exotic weed and annual
grass invasions, which establish after the dense
canopies of the short-lived lupines are opened by
fire or insects (Maron and Connors 1996). A
variety of insects prey upon it, sometimes killing
extensive stands. Among the predators are the
subterranean ghost moth Hephialus californicus
and the tussock moth Orgyia vetusta. The nitrogen
General Description.—Yellow bush lupine is a rich litter that results from such mortality makes
small erect shrub, less than 20 dm tall, with establishment of native vegetation uncertain and a
cauline leaves on 2- to 3- cm petioles, and long-term process (Maron and Jefferies 1999).
composed of five to 12 leaflets, on 8 to 12 mm
stipules. Leaf colors range from green and smooth Reproduction.—Inflorescences are 10 to 30 cm,
to silver and hairy. In northern California it grades with flowers sometimes arranged in whorls.
into and coexists with L. rivularis Doug. ex Lindl., Flowers are 14 to 18 mm with upper lip 5 to 9 mm
distinguished by blue flowers (Sholars 1993). and lower lips 5 to 7 mm. Petals are yellow, or
north of central California lilac to purple. Pods are
Range.—Yellow bush lupine is native on coastal hairy, brown or black, 4 to 7 cm long. Eight to 12
bluffs and dunes, or less than 5 km inland seeds per pod are black, tan, or striped, and are 4
(Davidson and Barbour 1977), in central to 5 mm long (Sholars 1993). Flowers are both
California from Ventura to Sonoma Counties, and self-compatible and outcrossing and have
is widely naturalized on the northern California generated considerable interest from genetic
coast (Sholars 1993) and as far north as investigators. Self-pollination requires facilitation,
Vancouver, Canada (Hitchcock and Cronquist usually from insects, but is responsible for only
1973). There has been considerable debate over about 25 percent of seed production (Kittelson and
the native range, although it probably encompasses Maron 2000).
only the southern part of the current range.
Fire Effects.—Although fire may cause
Ecology.—Yellow bush lupine is an aggressive widespread mortality in dense lupine stands, and
colonizer of dunes, where it outcompetes native may be followed by invasion of annual weeds and
dune mat species. It encourages other non native grasses (author’s observation), the long-lived
invaders by fixing nitrogen (Pickart and others lupine seed banks may result in reestablishment of
1998) and creating shade. Outside its former lupine (Maron and Connors 1996). The annual
range, it also hybridizes with native lupines, weeds and grasses that follow the nitrogen
especially L. rivularis (National Park Service
451
enrichment by lupine potentially alter fire regimes, Gadgil, R.L. 1971. The nutritional role of Lupinus
increasing frequency and intensity of burns. arboreus in coastal sand dune forestry. Plant
and Soil 35:113-126.
Growth and Management.—Yellow bush lupine
is fast growing and short lived (maximum 7 years) Hitchcock, C.L. and A. Cronquist. 1973. Flora of
(Davidson and Barbour 1977). Stands may grow the Pacific Northwest: an illustrated manual.
rapidly and die off in as little as 3 years (Strong et University of Washington Press. Seattle, WA.
al. 1995). Restoration of lupine-invaded sites 730 p.
should include litter and duff removal in addition
to direct removal of lupine, because it enriches soil Kittelson, P.M. and J.L. Maron. 2000.
to the detriment of native competitors (Pickart and Outcrossing rate and inbreeding depression in
others 1998). In New Zealand, where it is used as a the perennial yellow bush lupine, Lupinus
soil developer, it is controlled prior to tree planting arboreus (Fabaceae). American Journal of
by crushing and herbicides (Gadgil 1971). Coastal Botany 87: 652-660.
dune restoration in northern California has focused
heavily on removal of yellow bush lupine. Manual Maron, J.L. and P.G. Connors. 1996. A native
removal is accomplished by pulling up small nitrogen-fixing shrub facilitates weed invasion.
plants and chopping larger ones, which rarely Oecologia 105:302-312.
crown sprout. Heavy equipment has also been used
successfully, but all mechanical methods are labor Maron, J.L. and R.L. Jefferies. 1999. Bush lupine
intensive and expensive (Pickart and Sawyer mortality: Altered resource availability and
1998). Although others have noted a long-lived alternative vegetation states. Ecology 80: 443-
seed bank (see Maron and Connors 1996 and 454.
above), Pickart and Sawyer (1998) found no
recruitment by 3 years after removal of established Pickart, A.J., L.M. Miller, and T.E. Duebendorfer.
stands. 1998. Yellow bush lupine invasion in northern
California coastal dunes. Ecological impacts
Benefits.—Yellow bush lupine has been widely and manual restoration techniques. Restoration
cultivated as a sand stabilizer and ornamental in Ecology 6(1): 59-68.
coastal California. It is a strikingly beautiful plant,
both in and out of bloom, although its short life Pickart, A.J. and J.O. Sawyer. 1998. Ecology and
and messy litter probably limit its value as an Restoration of Northern California Coastal
ornamental. Elsewhere it has been used to prepare Dunes. California Native Plant Society.
coastal and dune soils for planting of Sacramento, California. 152 p.
commercially useful trees (Gadgil 1971). It
provides habitat for birds, reptiles, and rodents. Sholars, T. 1993. Lupinus Lupine in Hickman,
J.C., ed. 1993. The Jepson Manual: higher
References plants of California. University of California
Press, Berkeley and Los Angeles. 1,400 p.
Anderson, L.W.J., J. DiTomaso, G.F. Hruss, and
M. Rejmanek. 1999. The CalEPPC List: Exotic Strong, D.R., J.L. Maron, P.G. Connors, A.
Pest Plants of Greatest Ecological Concern in Whipple, S. Harrison, and R.L. Jefferies. 1995.
California. California Exotic Pest Plant High mortality, fluctuation in numbers, and
Council. Petaluma, CA. 12 p. heavy subterranean insect herbivory in bush
lupine, Lupinus arboreus. Oecologia 104: 85-
Davidson, E.D. and M.G. Barbour. 1977. 92.
Germination, establishment, and demography
of coastal bush lupine (Lupinus arboreus) at U.S.D.I. National Park Service, 2001. What is
Bodega Head, California. Ecology 58:592-600. yellow bush lupine? http://www.nps.
gov/redw/y-lupine.
452
Lycium pallidum Miers pale wolfberry
SOLANACEAE
Synonyms: None
oligospermum differs from var. pallidum in having

smaller flowers (corolla tube less than 12mm),
smaller anthers (1 to 1.5mm versus 1.8 to 3 mm)
and fewer, only approximately seven seeds, per
berry (Cronquist and others 1984, Baldwin and
others 2002).
Range.—Pale wolfberry is a common shrub in

Arizona, southern Colorado, southern California,
Nevada, New Mexico, western Oklahoma, western
Texas, and Utah in the U.S.A., and in Sonora,
Chihuahua, Zacatecas, and San Luis Potosí in
Mexico (McGregor and others 1986, Vines 1986,
Carter 1988, Welsh and others 1993). Variety
oligosperma occurs only in Nevada and California
(Integrated Taxonomic Information System 2002).
Ecology.—Pale wolfberry grows at elevations of

910 m to 2,134 m in a variety of habitats and soils,
including those derived from limestone and
igneous formations. It often grows in sandy soils
and is tolerant of saline soils (Kearney and others
1960). It is frequently a member, and may be a co-
Illustration credit: Tonia Masaood dominant, in blackbrush, sagebrush, four-wing
saltbush, mesquite, greasewood, mountain brush,
General Description.—Lycium pallidum is and pinyon-juniper communities (Dick-Peddie
commonly known as pale wolfberry, pale lycium, 1993, MacMahon 1988, Welsh and others 1993).
rabbit thorn (Epple 1995), box thorn (Derig and However, it has never been reported as a dominant
Fuller 2001), chico, cambronera, and tomatillo species. Pale wolfberry grows throughout the
(Elmore 1976). The latter name can be confusing Sonoran, Chihuahuan, and Mohave Deserts. At
because “tomatillo” also commonly refers to higher elevations it grows on dry slopes and
species of Physalis (Tull 1987). The epithet plains, but in the deserts it tends to grow in washes
“pallidum” refers to the pale colored leaves. Pale or other areas where run-off supplements rainfall
wolfberry are thorny, intricately-branched shrubs (Bowers and McLaughlin 1987, Bowers 1993).
that form loose thickets. Unlike other Lycium Wolfberry species have tough and fibrous roots
species it is seldom drought deciduous (Bowers and the root systems are relatively extensive
1993, Kearney and others 1960). The pale green, extending 25 to 30 feet (7.5 to 9.0 m) from the
elongated-oblong leaves are adjacent to short, plant (Matthews 1994) that suggests they may be
sharp thorns or spines. The flowers are greenish important in retarding soil erosion.
cream, sometimes tinged with purple, and
funnelform in shape. The corolla tube is longer Reproduction.—Pale wolfberry generally flowers
than 12 mm. The stamens are usually exerted in April to June but in some regions may flower as
(Vines 1986, Welsh and others 1993). The oval early as February (Powell 1998, Epple 1995). The
fruits are red colored and approximately 1 cm or fruits mature in the late spring and summer
less in diameter (Vines 1986). Typically the plants (Bowers 1993, Vines 1986). The fragrant, creamy-
are 1 m high shrubs but may grow to 2 m (Elmore yellow to yellowish-green flowers are insect-
1976, Carter 1997). Two varieties of Lycium pollinated. The bright, shiny red berries contain 20
pallidum are recognized: var. oligospermum C.L. to 50 seeds.
Hitchc., and var. pallidum (Kartesz 1994). Variety
453
Growth and Management.—Pale wolfberry References
sprouts readily when cut or broken (Van Dersal
1938). Once established, shrub longevity may be Baldwin, B.G., S. Boyd, B.J. Ertter, R.W.
substantial. In one Arizona study 69 percent of Patterson, T.J. Rosatti, D.H. Wilken, M.
individual shrubs of a related species, Lycium Wetherwax. 2002. The Jepson desert manual.
berlandii, survived for at least 50 years (Goldberg University of California Press, Berkeley, CA.
and Turner 1986). Seeds are likely dispersed by 626 p.
birds and other animals that feed on the berries
(Bowers 1993). It can be propagated from Bowers, J.E. 1993. Shrubs and trees of the
hardwood cuttings, suckers, layers, and seeds. Southwest Deserts. Southwest Parks and
Monuments Association Tucson, AZ. 140 p.
Benefits.— Pale wolfberry is an important source
of food for wildlife, especially birds and rodents Bowers, J. E. and S.P. McLaughlin. 1987. Flora
(Elmore 1987, Bowers 1993, Powell 1998). In and vegetation of the Rincon Mountains, Pima
many areas of degraded habitat in the County, Arizona. Desert Plants. 8(2): 50-94.
Southwestern United States it frequently provides
a source of berries and browse, in an otherwise Carter, J.L. 1988 Trees and shrubs of Colorado.
depauperate landscape (authors personal Johnson Books, Boulder, CO. 165 p.
observation). It has been reported to be locally
important and valuable as browse for livestock Carter, J.L. 1997 Trees and shrubs of New
especially on winter range (Van Dersal 1938). The Mexico. Johnson Books, Boulder, CO. 534 p.
berries also are eaten by humans, especially Native
Americans, who have also used other parts of the Cronquist, A., A. H. Holmgren, N.H. Holmgren, J.
plant as medicine and in ceremonials (Moerman L. Reveal and P. K. Holmgren. 1984.
1998, Stevenson 1915). In times of famine the Intermountain Flora, Vascular Plants of the
Native Americans of northern Arizona ate the Intermountain West, U.S.A. Vol. 4, New York
dried berries mixed with saline clay (Kearney and Botanical Garden, Bronx, NY. 279 p.
others 1960). In this form they were called “food
clay” by the Navajo or “potato clay” by the Hopi Derig, B. B. and M. C. Fuller. 2001. Wild berries
(Dunmire and Tierney 1997). As well as being an of the west. Mountain Press Publishing
extender, the clay may have been used to reduce Company, Missoula, Montana. 235 p.
bitterness. With the recent interest in using native
plants, more palatable recipes for sauces and jams Dick-Peddie, W.A. 1993. New Mexico
using pale wolfberry berries are available (Derig Vegetation–past, present, and future. University
and Fuller 2001). Medicinal uses include applying of New Mexico Press, Albuquerque, NM. 244 p.
the soaked leaves to cuts and using the ground-up
roots for toothaches. These hardy shrubs have been Dunmire, W.W. and G.D. Tierney. 1995. Wild
used as ornamentals since 1878 but on a limited plants of the Pueblo Province. Museum of New
scale (Vines 1986, Powell 1998). Stands of pale Mexico Press, Santa Fe, NM. 290 p.
wolfberry are frequently associated with ancient
Anasazi ruins in the Four Corners area of the Dunmire, W.W. and G.D. Tierney. 1997. Wild
Southwestern United States (Dunmire and Tierney plants and native peoples of the Four Corners.
1995). Some stands have been speculated to be Museum of New Mexico Press, Santa Fe, NM.
deliberate plantings and others the result of 312 p.
inadvertent seed dispersal. After a pueblo is
abandoned, moisture tends to collect over Elmore. F.H. 1976. Shrubs and Trees of the
decomposing subsurface floors and plazas and Southwest Uplands. Southwest Parks and
these conditions may make suitable habitat for Monuments Assoc. Tucson, AZ. 214 p.
seeds that were unintentionally dropped by the
inhabitants of the ancient Pueblos (Dunmire and Epple, A.O. 1995. A Field Guide to the Plants of
Tierney 1997). Arizona. Falcon Press Publishing Co., Helena,
MT. 347 p.
Goldberg, D.E. and R.M. Turner. 1986. Vegetation

change and plant demography in permanent
454
plots in the Sonoran Desert. Ecology 67(3): 695- University Press of Kansas, Lawrence KS,
712. U.S.A. 1,402 p.
Integrated Taxonomic Information System. 2002. Moerman, D.E. 1998. Native American Ethno-
Lycium pallidum. Plants, database botany. Timber Press, Portland, OR. 927 p.
http://www.itis.usda.gov/ [not paged].
Powell, A. M. 1998. Trees & shrubs of Trans-
Kartesz, J.T. 1994. A synonymized checklist of the Pecos and adjacent areas. University of Texas
vascular flora of the United States, Canada and Press, Austin, TX. 498 p.
Greenland. Vol.1 - Checklist. 2nd ed. Timber
Press, Portland, OR. 622 p. Stevenson, M. C. 1915. Ethnobotany of the Zuni
Indians. Annual Report 30. Smithsonian
Institution-Bureau of American Ethology. 102 p.
Kearney, T.H., R.H. Peebles, and collaborators.
1960. Arizona flora. 2nd Ed. University of
Tull, D. 1987. Edible and Useful Plants of Texas
California Press, Berkeley, CA. 1,085 p.
and the Southwest. University of Texas Press.
Austin, TX. 518 p.
MacMahon, J.A. 1988. Warm deserts. In: M.G.
Barbour, Billings and W. Dwight, eds. North
Van Dersal, W.R. 1938. Native woody plants of
American terrestrial vegetation. Cambridge
the United States, their erosion-control and
University Press, New York. p 231-264.
wildlife values. Miscellaneous Publication 303.
U.S. Deptment of Agriculture. Washington D.C.
Matthews, R. F. 1994. Lycium pallidum. U.S.D.A.
Forest Service, Rocky Mountain Research
Station, Fire Sciences Laboratory. Fire Effects
of the Southwest. University of Texas Press.
Information System. http://www.fs.fed.
us/databse/feis/plants.
Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
McGregor, R. L., T.M. Barkley, RE Brooks, and Higgins. 1993. A Utah Flora. Second Edition,
EK Schofield. 1986. Flora of the Great Plains. revised. Brigham Young University, Provo,
Utah. 986 p.
455
Macfadyena unguis-cati (L.) A.H. Gentry cat’s claw
BIGNONIACEAE
Synonyms: Bignonia unguis L. emend. DC.

Bignonia unguis-cati L.
Batocydia unguis (L. emend. DC.) Mart. ex DC.
Doxantha unguis (L. emend. DC.) Miers
Doxantha praesignis Miers
Doxantha serrulata Miers
John K. Francis
laterally near the soil surface with finer roots

scattered over its length. Tubers are produced by
both young and old plants (Stockard 2001). The
plant grows and maintains few branches until the
growing tip reaches increased light. The
compound leaves have two leaflets with the
clawed tendril between them. The leaves are
generally ovate to lanceolate in shape but quite
variable in size. Depending on both age and
environment, petioles may vary from 0.25 to 2.5
cm in length, and blades may vary from 1 to 16
cm in length. They are dark green above and
lighter below. Inflorescences are axillary with one
to three flowers. The 4.5 to 10 cm-long tubular
flowers have five lobes and are bright yellow with
red-orange lines in the throat. From them develops
a long (up to 75 cm), narrow (1.0 to 1.5 cm),
flattened capsule that produces brown, flattened,
winged seeds. The species has both diploid (2n =
40) and tetraploid (2n = 80) populations
(Acevedo-Rodríguez 1985, Gentry 1983, Howard
1989, Liogier 1995).
Range.—Cat’s claw is native to the Greater and

Lesser Antilles, Mexico, Central America, and
South America to Argentina (Howard 1989). The
General Description.—Cat’s claw is a woody species is reported to have naturalized and escaped
vine or occasionally a scrambling shrub. The in Florida, Texas, Hawaii (Aquatic Plant Control
name, and its equivalent in Spanish, uña de gato, Operations Support Center 2001), New Caledonia
comes from the tripartite, hooked tendrils (Pacific Island Ecosystems at Risk 2001),
resembling an animal’s claws that enable the vine Australia (Stockard 2001), South Africa (Pest
to adhere to tree bark and other surfaces Cabweb 2001).
(Acevedo-Rodríguez 1985). Old plants are more
or less free hanging. It is also known by the Ecology.—In Puerto Rico, cat’s claw grows from
common names bejuco de gato, paz y justicia, and near sea level to over 600 m in elevation and in
griffe à chatte (Howard 1989, Liogier 1995). Cat’s sites that receive mean annual rainfalls from about
claw has a strong and flexible, cylindrical stem 750 to about 2400 mm. It will tolerate most soils,
that is brown in color with many lenticels. The except very poorly drained and salty soils. The
stems produce adventitious roots to anchor them species can tolerate a few degrees of frost, being
tightly to vertical surfaces. Cat’s claw may exceed killed to the ground but resprouting afterward
8 cm in stem diameter and extend more than 20 m (Watkins and Sheehan 1975). Cat’s claw is
into the crowns of trees. A long main root extends
456
moderately shade tolerant as a young plant and still a handsome ornamental. It is often used to
grows in both full sun and under forest canopies. It cover fences or screen unsightly buildings. Cat’s
may be less shade tolerant as an adult. The species claw is recommended for areas where there is little
is common in savannas, secondary forest, and chance of invasion and especially in desert areas
remnant high forest. It can survive grazing and fire because it requires relatively little water for
but is eliminated by deep grass swards. maintenance (Desert-Tropicals 2001). It was
reported that a preparation made from cat’s claw
Reproduction.—In Hawaii, flowering takes place was used to treat dermatitis from Hippomane
during the spring (Rauch and Weissich 2000) mancinella L. (Michell and Rook 2001). A
whereas there are actually two periods of number of other uses in herbal medicine from
flowering in Puerto Rico, both during dry seasons diverse locations are cited by Liogier (1990).
(Acevedo-Rodríguez 1985). In Costa Rica, nearly
all the plants flower at once and flowering lasts References
just a short time (Gentry 1983). The capsules
mature about 6 months after flowering. The Acevedo-Rodríguez, P. 1985. Los bejucos de
winged seeds are dispersed by the wind. Seedlings Puerto Rico. Vol. 1. General Technical Report
are common and widespread in suitable habitat. SO-58. U.S. Department of Agriculture, Forest
Eleven capsules collected in Puerto Rico had a Service, Southern Forest Experiment Station,
maximum of 212 seeds and a minimum of 106 New Orleans, LA. 331 p.
seeds. These seeds (air dry) averaged 0.0224 ±
0.0005 g/seed or 45,000 seeds/kg. One hundred Aquatic Plant Control Operations Support Center.
percent of the seeds tested germinated between 49 2001. Macfadyena unguis-cati—catclaw vine.
and 95 days after sowing in commercial potting www.saj.usace.army.mil/conops/apc/newtt/cat1
mix. Young plants sprout when damaged and layer maps/macfadyenaunguis-cati.htm. 1 p.
(root) whenever stems touch the ground. Cat’s
claw, for ornamental uses, have been propagated Desert-Tropicals. 2001. Philippe and Sura’s
by seed, cuttings, and layering (Turner and Phoenix Tropical Gardens. http://www.desert-
Wasson 1999). tropicals.com. 3 p.
Growth and Management.—Stem extension is Florida Exotic Plant Council. 2001. Macfadyena
moderately rapid, especially in sprouts. Diameter unguis-cati (L.) A. Gentry. http://www.
growth of cat’s claw stems is slow. The vines are fleppc.org/pdf/Macfadyena%20unguis-cati.pdf.
long-lived, nearly as long as the trees that they 2 p.
claim for support. There are relatively few
problems in establishing cat’s claw. Insect and Gentry, A.H. 1983. Macfadyena unguis-cati (uña
disease problems are rare. The biggest problem is de gato, cat-claw, bignone). In: D.H. Janzen, ed.
controlling ornamental plants. Yearly pruning Costa Rican natural history. University of
after the flowering season is essential to Chicago Press, Chicago and London. p. 272-
maintaining them within the bounds desired. In its 273.
native range cat’s claw vines have only a minor
effect on the trees they parasitize. However, the Howard, R.A. 1989. Flora of the Lesser Antilles,
species is beginning to seriously suppress native Leeward and Windward Islands.
vegetation in parts of Florida (Florida Exotic Plant Dicotyledoneae. Vol. 6. Arnold Arboretum,
Council 2001) and Australia (Stockard 2001). The Harvard University, Jamaica Plain, MA. 658 p.
only means of control recommended at present is
to cut the vines and paint the cut ends with Liogier, H.A. 1990. Plantas medicinales de Puerto
glyphosate herbicide (Stockard 2001). However, Rico y del Caribe. Iberoamericana de Edicones,
Vélez and van Overbeek (1950) report that older Inc. San Juan, PR. 566 p.
plants are killed by simply cutting the stems. A
more practical long-term solution is hoped for in Liogier, H.A. 1995. Descriptive flora of Puerto
the ongoing biological control introductions in Rico and adjacent islands. Vol. 4. Editorial de la
several countries (Pest Cabweb 2001). Universidad de Puerto Rico, San Juan, PR.
617 p.
Benefits.—Despite the propensity to naturalize
and compete with native vegetation, cat’s claw is Michell, J. and A. Rook. 2001. Botanical
457
Dermatology Database. http://bodd.cf.ac.UK/ Stockard, J. 2001. The regeneration of Wingham
BotDermFolder/BotDermB/BIGN.html. 3 p. Brush, NSW. AABR lecture series. http://
www.zip.com.au/~aabr/info/seminars/seminar0.
Pacific Island Ecosystems at Risk. 2001. 4.html. 5 p.
Macfadyena unguis-cati (L.) Gentry,
Bigniniaceae. http://www.hear.org/pier/maung. Turner, R.J. and E. Wasson. 1999. Botanica.
htm. 2 p. Barnes and Noble, Inc., New York. 1020 p.
Pest Cabweb. 2001. Spotlight on biological control Vélez, I. and J. van Overbeek. 1950. Plantas
of weeds. http://pest.cabweb.org/Features/ indeseables en los cultivos tropicales. Editorial
Spotlight/spot00-2.htm. 21 p. Universitaria, Río Piedras, PR. 497 p.
Rauch, F.D. and P.R. Weissich. 2000. Plants for Watkins, J.V. and T.J. Sheehan. 1975. Florida
tropical landscapes, a gardener’s guide. University landscape plants, native and exotic. The
of Hawaii Press, Honolulu, HI. 139 p. University Presses of Florida, Gainesville, FL.
420 p.
458
Machaonia portoricensis Baill. alfilerillo
RUBIACEAE
Synonyms: none
John K. Francis
St. John in the U.S. Virgin Islands (New York

Botanical Garden 2002) where it is endangered
(University of the Virgin Islands 2002). Alfilerillo
is not known to have been planted or naturalized
elsewhere.
Ecology.—Alfilerillo grows on a wide variety of

well-drained soils derived from a variety of rock
types including limestone and ultramafics
(serpentine). These are usually rocky, sandy, or
moderately salty so that competition is somewhat
reduced. It is moderately intolerant of shade and
can inhabit forests with low basal areas. The
species grows in dry habitat that receives from
about 750 to about 950 mm of mean annual
precipitation at elevations ranging from near sea
level to about 400 m. Alfilerillo is relatively
tolerant of fire, being killed to the ground, but
readily sprouting. It seems to be resistant to
grazing by cattle probably owing to the spines.
The species may be found in remnant forests on
ridges and bluffs, in overgrazed brush lands and
General Description.—Alfilerillo, also known as savannas, near mangroves and salt ponds, and in
roseta, is a spiny shrub or small tree that beach strand vegetation (author’s observation,
occasionally reaches 6 m in height and 7.5 cm in New York Botanical Garden 2002, Vázquez and
stem diameter. It is supported by an extensive Kolterman 1998).
lateral root system. Alfilerillo may have single or
multiple stems arising just above the ground level, Reproduction.—Alfilerillo flowers and fruits
but it is always branchy and thorny. The stem bark irregularly through the year (Little and others
is gray and smooth becoming fissured in old 1974). Seeds collected from trees in western
plants. The branches and twigs are brown, slender Puerto Rico weighed an average of 0.00182 g/seed
and stiff and support paired spines 1 to 2.5 cm or 548,000 seeds/kg. Sown on moist filter paper,
long at the nodes and ends of twigs. The plant 24 percent germinated within 54 days.
tends to have a slender, vertical profile. Opposite Germination is epigeal. No specialized means of
or whorled, almost sessile leaves are orbicular to seed dispersal is apparent aside from a weak
elliptic with entire edges and blades 7 to 16 mm tendency to cling to clothing. Seedlings are
long and 5 to 10 cm broad. Tiny, white flowers are uncommon in spite of relatively good seed
grouped in many-flowered terminal panicles. The production. Plants sprout when disturbed.
capsules are turbinate (inverse conical), brown,
quadrangular, 4 to 5 mm long, crowned with four Growth and Management.—Growth of
calyx lobes and contain two seeds (Liogier 1997, alfilerillo is relatively slow. A 3-cm thick stem cut
Little and others 1974). by the author had 11 growth rings. No planting or
management experience has been published.
Range.—Alfilerillo is native to Puerto Rico
(Liogier 1997) from Salinas to Cabo Rojo. A Benefits.—Afilerillo helps protect the soil in the
closely-related, recently-separated species, fragile environments where it grows and furnishes
Machaonia woodburyana Acev.-Rodr., occurs on
459
cover for wildlife. The wood, which is light- New York Botanical Garden. 2002. West Indian
brown, hard, and brittle, is good for fuel. vascular plant types catalog. http://
scisun.nybg.org:8890/searchdb/owa/wwwcatalo
References g.detail_list. [not paged].
Liogier, H.A. 1997. Descriptive flora of Puerto University of the Virgin Islands. 2002. Endangered
Rico and adjacent islands. Vol. 5. Editorial de la and threatened plants and animals in the U.S.
Universidad de Puerto Rico, San Juan, PR. Virgin Islands. http://rps.uvi.edu/CES/
436 p. endangered.html. 4 p.
Little, E.L., Jr., R.O. Woodbury, and F.H. Vázquez, O.J. and D.A. Kolterman. 1998. Floristic
Wadsworth. 1974. Trees of Puerto Rico and the composition and vegetation types of the Punta
Virgin Islands. Vol. 2. Agriculture Handbook Guaniquilla Natural Reserve—Cabo Rojo,
449. U.S. Department of Agriculture, Puerto Rico. Caribbean Journal of Science 34(3-
Washington, DC. 1,024 p. 4): 265-279.
460
Mahonia aquifolium (Pursh) Nutt. Oregon grape
BERBERIDACEAE
Synonyms: Berberis aquifolium Pursh

Berberis piperiana (Abrams) McMinn
Mahonia piperiana Abrams
Odostemon aquifolium (Pursh) Rydb.
John K. Francis
California, and northern Idaho (Abrams 1950,

The species has been planted widely as an
ornamental and medicinal plant and has
naturalized in Michigan, Ohio, Kentucky, New
York, New Jersey, United Kingdom, Germany and
probably other areas (Auge and Brandl 1997,
Natural Resources Conservation Service 2003,
Plants for a Future 2003). Oregon grape hybridizes
naturally with Mahonia repens (Lindl.) G. Don
(many ornamentals are actually this hybrid) and
Mahonia pinnata (Lag.) Fedde (Plants For a
Future 2003). At least three artificial intergeneric
crosses between Oregon grape and Berberis
species have been made (Dirr 1983).
General Description.—Oregon grape, also called Ecology.—Oregon grape is tolerant of shade and
tall mahonia, hollyleaved barberry, mountain normally grows in the forest understory. Natural
grape, Oregon grape-holly, and Oregon holly- stands contain scattered individual plants arising
grape, is an upright evergreen shrub from 0.6 to 3 from seeds and clumps of the species arising both
m in height. The stems are slender, stiff, usually from seed and suckers. The species grows well in
upright, and usually without branches. The wood soils with a wide variety of properties. However,
is whitish and brittle and has a significant pith. ornamentals do not do well in dry, wind-swept
Bark of young stems is green; older bark is rough sites, or soils with high pH’s (above 8.0). The tops
and gray-brown with yellow inner bark. The root are severely injured at –29 °C (Dirr 1983). It
systems produce horizontal rhizomes up to 1.8 m grows in areas with natural precipitation ranging
long from which suckers arise. The foliage from 950 to 5450 mm annually (Las Pilitas
attaches directly to the stems. Alternate, stiff and Nursery 2003). A number of diseases and insects
leathery, pinnately compound leaves have five to attack the leaves and fruits but usually do not
13 leaflets with a form like holly (coarse dentate cause serious damage.
with prickles). Leaves are dark green above,
except in winter, when exposed leaves take on a Reproduction.—Oregon grape blooms from April
purplish color. The species is monoecious with to June (Clark and Trelawny 1976). Flowers are
perfect, bright-yellow flowers in terminal self-fertile and pollinated by insects (Plants for a
(sometimes lateral) racemes. Flowers have a Future 2003). The fruits mature in late July
honey-like fragrance. Fruits (berries), 6 to 10 mm through September and remain on the plants into
long, ripen from green to blue or blue-black in late December (Center for Applied Nursery Research
summer (Abrams 1950, Center for Applied 2003). Fruits collected by the author from
Nursery Research. 2003, Dirr 1983). Oregon grape ornamental plants in Utah averaged 0.312 ± 0.013
is the state flower of Oregon (Paghat.com 2003). g/fruit. Air-dried seeds separated from them
averaged 0.0067 + 0.0002 g/seed or 149,000
Range.—Oregon grape is native to British seeds/kg. The number of seeds/fruit varied from
Colombia, Washington, Oregon, northern zero to nine. Fruits can be harvested in quantity by
hand and can be cleaned by gentile maceration,
461
wet sieving, and screening. Seeds should not be The fruit serves as a safe and gentle laxative
allowed to dry out completely and should be (Plants for a Future 2003).
stratified at 5 °C before sowing. Taken in
November and treated with 0.8 to 1.0 percent of References
IBA (Indolbuteric acid), cuttings root well.
Division of established plants is an alternate Abrams, L. 1950. Illustrated flora of the Pacific
method of vegetative propagation (Dirr 1983). In States. Vol. 2. Stanford University Press,
nature, mostly birds disperse the seeds Stanford, CA. 635 p.
(Paghat.com 2003) and both sexual and asexual
reproduction are effective (Auge and Brandl Auge, H. and R. Brandl. 1997. Seedling
1997). recruitment in the invasive clonal shrub,
Mahonia aquifolium (Pursh) Nutt. Oecologia
Growth and Management.—Oregon grape is Berlin 110(2): 205-211.
moderately slow growing. Individual stems reach a
60- to 90-cm height in 3 to 4 years (Dirr 1983). Center for Applied Nursery Research. 2003.
While individual stems probably have a life of not Mahonia aquifolium—Oregongrapeholly,
more than 10 years, clones can last much longer. Oregon Grapeholly, Oregon Hollygrape, Oregon
Plants are grown for the nursery trade in containers grape. http://www.nobleplants.com/classnotes/
and are usually 1 year old when sold. The species spring/springprofiles/evergreen/mahoniaaquifoli
is planted both in sunny and shady locations. um.htm. 2 p.
Ornamental plants tolerate pruning well (Plants for
a Future 2003). Data on management of the Clark, L.J. and J.G. Trelawny. 1976. Wild flowers
species in natural stands are lacking. of the Pacific Northwest. Gray’s Publishing
Limited, Sidney, B.C., Canada. 604 p.
Benefits.—Oregon grape shrubs contribute to the
diversity and beauty of understory vegetation in Dirr, M.A. 1983. Manual of woody landscape
native habitats, help protect the soil, and furnish plants: their identification, ornamental
food and cover for wildlife. The species is planted characteristics, culture, propagation and uses.
in temperate areas throughout the world as an Stipes Publishing Company, Champaign, IL.
ornamental, for hedges, and foliage, and leafy 826 p.
branches with their fall colors are sometimes used
for Christmas decorations. The fruits, edible but Felter, H.W. 1922. The eclectic material medica,
sour, are made into jellies and juice drinks (Clark pharmacology and therapeutics: Berberis
and Trelawny 1976). Flowers are eaten in salads, (Mahonia) aquifolium. published as a database
cooked in tempuras, and used to make a lemonade- in: http://www.ibiblio.org/herbmed/eclectic/
like drink (Paghat.com 2003). Yellow, green, felter/hahonia-aqui.html. 2 p.
violet, and purplish-blue dyes are made from
various tissues for arts and crafts (Plants For a Gieler, U., A. Von der Weth, and M. Heger. 1995.
Future 2003). Mahonia aquifolium—a new type of topical
treatment for psoriasis. Journal of
Herbal Medicine.—Infusions of Oregon grape Dermatological Treatment 6(1): 31-34.
root and doses of isolated alkaloid have been used
effectively in the past to relieve many of the Las Pilitas Nursery. 2003. Mahonia aquifolium.
symptoms of syphilis (Felter 1922). Ointments Las pilitas Nursery, Escondido, CA.
made from the plant and isolated alkaloids are http://www.laspilitas.com/plants/420.htm. 2 p.
currently enjoying tremendous popularity in
treating psoriasis and similar skin conditions. In Muller, K., K. Ziereis, and I. Gawlik. 1995. The
clinical trials, symptoms improved or disappeared anipsoriatic Mahonia aquifolium and its active
in 81 percent of 443 patients suffering from constituents: II. Antiproliferative activity against
subacute and chronic psoriasis (Gieler and others cell growth of human keratinocytes. Planta
1995). Of the constituent compounds tested, the Medica 61(1): 74-75.
benzylisoquinoline alkaloids berbamine and
oxyacanthine were principally responsible for Natural Resources Conservation Service. 2003.
inhibition of abnormal cell growth, with berberine Plants profile: Mahonia aquifolium (Pursh) Nutt.
showing lesser activity (Muller and others 1995). http://plants.usda.gov/cgi_bin/topics.cgi. 4 p.
462
Plants For a Future. 2003. Plants for a future:
Paghat.com. 2003. Paghat’s garden: Mahonia database search results: Mahonia aquifolium.
aquifolium. http://www.paghat.com/ http://www.ibiblio.org/pfaf/cgi-bin/arr_html
oregongrape.html. 4 p. ?Mahonia+aquifolium. 12 p.
463
Mahonia repens (Lindl.) G. Don creeping barberry
BARBERIDACEAE
Synonyms: Berberis amplectens (Eastw.) L.C. Wheeler

Berberis aquifolium Pursh var. repens (Lindl.) Scoggan
Berberis pumila Greene
Berberis sonnei (Abrams) McMinn
Mahonia amplectens Eastw. (additional synonyms: Natural Resources
Conservation Service 2003)
John K. Francis
four seeds. The seeds are 6 to 10 mm long. There

are 2n = 28 chromosomes (Auger and other 2002,
Harrison 2003, Lesica 2002, Rehder 1951, Welsh
and others 1987). Creeping barberry readily
hybridizes with M. aquifolium (Pursh) Nutt. and
M. pinnata (Lag.) Fedde when growing near them
(Pagahat.com 2003).
Range.—Creeping barberry is native to the

Western United States as far east as Texas and
Minnesota, and to British Colombia and Alberta in
Canada. It is present (probably naturalized) in
Indiana and Pennsylvania (Natural Resources
Conservation Service 2003, Walkup 1991). The
species has been planted widely as an ornamental
and has probably naturalized in many areas outside
its native range.
Ecology.—Low in stature and moderately tolerant

of shade, creeping barberry is not highly
competitive. In Utah, it will not grow in heavy
grass and forb stands. It is most frequently seen in
General Description.—Creeping barberry, also rocky or gravely areas with low vegetative cover
known as Oregon grape, creeping Oregon grape, or under open conifer or hardwood stands with
creeping mahonia, creeping hollygrape, ash sparse understory vegetation. However, the species
barberry, and dwarf Oregon grape, is an upright to is reported to be the dominant understory shrub in
recumbent, evergreen shrub 12 to 40 cm in height closed-canopy Douglas fir in central Idaho
and 3 to 9 mm in basal diameter. The stems are (Walkup 1991). Creeping barberry apparently
solitary, usually unbranched but occur in clonal grows well under conifers that produce a thick
groups arising from rhizomes. The main root is layer of decomposing needles. However, it does
stiff and deep, and the lateral roots are short and not flower when growing in shade (Pahgat.com
fine. Stem bark is slightly rough and gray brown, 2003). Natural populations are affected little by
and the inner bark is bright yellow, especially the treatments such as prescribed fire, clearcutting,
inner root bark. The wood and pith are yellow. The and scarification. It usually survives (by sprouting)
alternate leaves, which have petioles 2 to 3 cm even moderate and high-intensity fires (Walkup
long, are pinnately compound with three to seven 1991). Creeping barberry is an invader of
leathery, sessile, ovate leaflets that are dull on both disturbed sites (Harrison 2003). It will grow in
surfaces, and have spiny-toothed margins. There most well-drained soils and is especially common
are usually two to eight leaves per plant. Several in rocky or gravelly soils. The species is much
flowers with six petals and six sepals, both yellow, more common over limestone or other alkaline
are clustered in terminal racemes 3 to 7 cm long. parent materials than granite or quartzite (Walkup
Fruits are glaucous, blue berries containing one to
464
1991). It tolerates acidic soils but grows poorly State University Research and Extension 1996). It
when concentrations (less than 2.5 mg/L) of boron is included in reclamation plantings (Welsh and
are low (Harrison 2003). Creeping barberry grows others 1987) and has been used especially for
at elevations of 1,125 to 3,300 m in the inland revegetation of mine spoils (Harrison 2003). Mule,
West (Harrison 2003, Walkup 1991). Although it white-tailed deer, and sometimes elk browse
is most common on north-facing slopes, it grows creeping barberry during fall and winter. The
on south-facing slopes and ridge tops at higher protein content ranges from 4.7 to 5.5 percent, and
elevations. the species contains high contents of carotene in
July. Creeping barberry foliage is mildly toxic and
Reproduction.—Creeping barberry flowers in domestic animals normally make little use of it for
May through June and matures fruit in June forage except when other food is scarce (Walkup
through July (Rudolf 1974). The flowers are 1991). In one case, browsing of creeping barberry
pollinated by bees and butterflies (Paghat.com killed cattle in Arizona (Harrison 2003). Many
2003). Flowers not cross pollinated with other mammals (including black bears) and birds eat the
plants will self pollinate, but produce much fewer fruits (Walkup 1991). The fruits become edible to
seeds (Walkup 1991). Ratio weight of fresh fruits humans after they have passed through at least one
to air-dried seeds is 12.5:1 (Auger and others frost. They are used to make jelly, wine, and a
2002). There are 119,000 to 157,000 seeds/kg juice drink (Harrison 2003). Native Americans
(Rudolf 1974). Cleaned seed can be sealed and used the berries and the roots for lavender and
stored for 5 years (Harrison 2003). Rudolf (1974) yellow dyes, respectively, and made teas from the
recommends stratification of 30 days at 1 °C roots to treat a range of afflictions (Lesica 2002,
followed by 60 days at 21 °C, and then 196 days at Paghat.com 2003). Creeping barberry contains the
1 °C. Treated thus, 74 percent of test seeds alkaloids berberine and oxyacanthin (Harrison
germinated. Alternately, a hot water treatment 2003). The species is an alternate host for black
followed by cold stratification for 60 days is stem rust of cereal grains (Walkup 1991).
recommended (Kjelgren 2003). Seeds can be sown
in fall without pretreatment. In nature, seeds are References
dispersed by birds and to a lesser extent by
mammals including black bears and rodents Auger, J., S.E. Meyer, and H.L. Black. 2002. Are
(Auger and others 2002). Once established, plants American black bears (Ursus americanus)
spread through sprouts from rhizomes (Walkup legitimate seed dispersers for fleshy-fruited
1991). shrubs? American Midland Naturalist 147: 352-
367.
Growth and Management.—Growth is slow.
Individual stems may live 10 years or more. It is Harrison, S. 2003. Mahonia repens. Crop
difficult to imagine the need to control this Development Centre, University of
unobtrusive species in its native habitat. On the Saskatchewan, Saskatoon, Saskatchewan,
other hand, it should frequently be planted as part Canada. http://www.usask.ca/agriculture/
of environmental restoration projects. The species plantsci/classes/range/berberis.html. 4 p.
can be artificially propagated by seed, divisions,
cuttings, and layers (Walkup 1991). Direct seeding Kansas State University Research and Extension.
in the fall is the most practical method to introduce 1996. Low water use plants for Kansas
it to disturbed sites. Repellants or poisons may be landscapes. MF2067. Kansas State University,
necessary because the seeds are readily consumed Manhattan, KS. 4 p.
by mice (Kjelgren 2003).
Kjelgren, R. 2003. Mahonia repens. Utah State
Benefits.—Creeping barberry adds beauty to the University, Logan, UT. http://www.hort.usu.
forest, helps protect the soil, provides food for edu/natives/shrubs/repens.htm. 1 p.
wildlife, and makes a useful ornamental. It is
planted as ground cover, especially under light tree Lesica, P. 2002. A flora of Glacier National Park,
canopy cover or on poor, gravelly ground. In the Montana. Oregon State University Press,
summer, the leaves are dark green; in winter they Corvallis, OR. 512 p.
are green, mottled with bright red, or a bonze-
purple. The species is particularly recommended Natural Resources Conservation Service. 2003.
because it requires little or no irrigation (Kansas Plants profile: Mahonia repens (Lindl.) G. Don,
465
creeping barberry. http://plants.usda.gov/ woody plants of the United States. Agric.
cgi_bin/plant_search.chi?mode=Scientific+Nam Handb. 450. U.S. Department of Agriculture,
e&keywordquerry=Mahonia+repens&earl=plant Forest Service, Washington, DC. p. 247-251.
_search.cgi. 5 p.
Walkup, C.J. 1991. Mahonia repens. In: U.S.
Paghat.com. 2003. Creeping mahonia, aka Department of Agriculture, Forest Service,
creeping barberry, aka creeping hollygrape, aka Rocky Mountain Research Station, Fire Sciences
ash barbarry, aka small or dwarf Oregon grape. Laboratory. Fire Effects Information System.
http://www.paghat.com/oregongrape2.html. 4 p. http://www.fs.fed.us/database/feis/plants/shrub/
mahrep/all.html. 10 p.
Rehder, A. 1951. Manual of cultivated trees and
shrubs. The MacMillan Company, New York. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
996 p. Higgins. 1987. A Utah flora. Great Basin
Rudolf, P.O. 1974. Berbers L., barberry, mahonia. University Press, Provo, UT. 894 p.
In: C.S. Schopmeyer. tech. coord. Seeds of
466
Malvastrum americanum (L.) Torr. Indian Valley false mallow
MALVACEAE
Synonyms: Malva americana L.

Malva spicata auct. non L.
Malvastrum spicatum auct. non (L.) Gray
Malva blumeana Steud.
Malva gangetica L.
Malva borbonica Willd. (and others, Institute of Systematic Botany 2003)
John K. Francis
usually ovate blades 2 to 8 cm long with serrate

margins. Inflorescences are dense spikes, terminal
on stems and branches. The flowers are sessile,
subtended by an involucel of three linear bracts.
The petals are pale yellow to yellow-orange and 8
to 9 mm long. The fruits are 5 to 6 mm in diameter
and contain 10 to 15 mericarps without spines with
seeds 1.5 mm long (Howard 1989, Liogier 1994,
Stevens and others 2001). There are 2n = 24
chromosomes (Lavia and others 2000).
Range.—Indian Valley false mallow occurs in

Florida and throughout the Caribbean including
Puerto Rico and the U.S. Virgin Islands and from
Texas through Central and South America to
Argentina (Howard 1989, Stevens and others
2001). The species occurs as a naturalized exotic
in Hawaii (Natural Resources Conservation
Service 2003), Australia, India, and other locations
in the Old World tropics and subtropics (Howard
1989).
Ecology.—Plants observed by the author appear to

be short-lived shrubs that grow yearly without
dieback. The species is reported also growing as
an herb (annual) and subshrub (dieing back to
permanent woody parts each year) (Howard 1989,
Stevens and others 2001). Indian Valley false
mallow is usually not eaten by cattle or other
General Description.—Indian Valley false herbivores. Moderate grazing coupled with fire
mallow, also known as spiked malvastrum, false tends to make the species more abundant. It also
mallow, malva loca, malva silvestre, and mauve grows in areas disturbed by river overflows,
d’Amérique, is a shrub, subshrub, or woody herb farming, road building, and land clearing. The
that often reaches 1.5 m in height and 1 cm in species is intolerant of shade and severe
basal diameter. The semideciduous plant generally competition. It disappears under closed tree
has a single stem that is upright and sparsely canopies or in dense tall grass swards. However,
branched. The wood is light colored, moderately Indian Valley false mallow does grow under a
hard, and moderately strong. Plants are supported broken tree canopy. It grows on most types of
by a tap and lateral root system of ivory colored well-drained soil. Mean annual rainfall in Puerto
roots. The twigs and leaves are stellate-pubescent. Rican habitat ranges from about 750 to 1000 mm
Gray-green leaves have a 1- to 6-cm petiole, (author’s observation). It is known from near sea
467
level to 1000 m in elevation in Nicaragua (Stevens Howard, R.A. 1989. Flora of the Lesser Antilles,
and others 2001). Leeward and Windward Islands. Vol. 5. Arnold
Reproduction.—Indian Valley false mallow is MA. 604 p.
reported to flower and fruit throughout the year in
Nicaragua (Stevens and others 2001). In Puerto Institute of Systematic Botany. 2003. Atlas of
Rico, plants begin flowering at about 15 cm in Florida vascular plants: Malvastrum
height and continue through life except for pauses americanum. http://www.plantatlas.usf.edu/
during dry seasons. Air-dried seeds collected in main.asp?plantID=3662. 2 p.
Puerto Rico averaged 1.19 million seeds/kg.
Placed on moist filter paper without pretreatment, Lavia, G., G. Seijo, A. Fernández, and A.
15 percent germinated over a period of 6 months. Krapovickas. 2000. Los cariotipos de algunas
No specialized means of dispersal is apparent; the especies de Malvastrum A. Gray (Malvaceae).
seeds are small and undoubtedly are moved to Universidad Nacional del Nordeste, Corrientes,
some extent by wind, water, and animals. Argentina. http://www.unne.edu.ar/cyt/2000/
Seedlings are relatively common in favorable sites. 6_biologicas/6_pdf/6_034.pdf. 3 p.
The dry fruits can easily be collected in quantity
by hand but cleaning the tiny seeds is difficult. Liogier, H. A. 1994. Descriptive flora of Puerto
Growth and Management.—Growth rates are Universidad de Puerto Rico, Río Piedras, PR.
moderate (0.3 to 0.5 m/year) and plants live 1 to 5 461 p.
years, depending on competition and growing
conditions. Although control is occasionally Mensah, R., M. Dillon, M. Kahn, C. Tann, and L.
needed in rangelands and disturbed areas, no Wilson. 2003. Cotton insect pests and their weed
specific recommended treatments are known to the hosts. Australian Cotton Cooperative Research
author. Centre. http://www.cotton.pi.csiro.au/Assets/
PDFFiles/IPMGL99/IPMSO4.pdf. 2 p.
Benefits and Detriments.—Indian Valley false
mallow helps protect the soil, furnishes cover for Natural Resources Conservation Service. 2003.
wildlife, and because of its blooms contributes Plants profile: Malvastrum americanum (L.)
moderately to the aesthetics of wildlands where it Torr., Indian Valley false mallow. http://
grows. It is a host for the cotton bollworm, plants.usda.gov/cgi_bin/plant_search.cgi?mode=
Helicoverpa armigera (Hubner) (Mensah and Scientific+Name&keywordquery=Malvastrum+
others 2003). The species also is the larval food- americanum&earl=plant. 3 p.
plant for the Laviana white-skipper butterfly,
Heliopetes laviana (Hewitson) of Texas (Quinn Quinn, M. 2003. Caterpillar food plants for the
2003) and a Cuban butterfly, Strymon columella lower Rio Grande Valley. http://www.naba.
cybira (Hewitson) (Fernández 2001). org/ftp/plants_info.pdf. 8 p.

Montiel, eds. 2001. A Flora de Nicaragua.
Fernández, D.M. 2001. New oviposition and larval Monographs in Systematic Botany Vol. 85, No.
hostplants for twenty-three Cuban butterflies, 2. Missouri Botanical Garden Press, St. Louis,
with observations of the biology and distribution MO. p. 945-1,910.
of some species. Caribbean Journal of Science
37(1-2): 122-125.
468
Mecranium latifolium (Cogn.) Skean camasey almendro
MELASTOMATACEAE
Synonyms: Mecranium amygdalinum (Desr.) C. Wright ex Sauvalle

Melastoma amygdalinum Desr.
John K. Francis
globose, 5.5 mm in diameter, purplish black when

mature, and have a slightly sweet and slightly
bitter flavor. Within are many seeds that are light
brown, tear-drop shaped, and about 0.5 mm long
by 0.25 mm thick (author’s observation, Liogier
Range.—Camasey almendro is native to the

Luquillo Mountains and the Cordillera Central of
Puerto Rico and to St. Thomas in the U.S. Virgin
Islands (Liogier 1995). DNA evidence indicates a
likelihood that this species arose from M.
multiflorum (Desr.) Triana or M. septentrionala
Skean now found in the adjacent island of
Hispaniola (McKenny 2002). The species is not
the native range.
Ecology.—Camasey almendro is widely

distributed throughout the mountain forests in
Puerto Rico from 300 m elevation upward (Little
and others 1974) in areas that receive more than
about 1800 mm of precipitation. Soils are usually
clayey or loamy, developed from igneous and
sedimentary rocks with pH’s from about 5.0 to 6.0.
Camasey almendro flowers and fruits in full sun
and partial shade. It grows well in moderate shade
General Description.—Camasey almendro is a
but eventually succumbs to the heavy shade of a
shrub or small tree sometimes reaching 7.5 m in
fully closed forest canopy. Apparently, at least
height and 13 cm in stem diameter. The plant
mild disturbance is necessary for establishment.
frequently has multiple stems arising near the base.
Such disturbance results naturally from hurricanes
Horizontal stems and branches produce vertical
and other winds that cause tree-fall gaps, rarely
sprouts. Stems, which are covered with gray-
from landslides, and more recently from road cuts
brown, thin, slightly furrowed bark, tend to be
and other construction. However, invasion is slow
straight with relatively few branches. Twigs are
and never with large populations. Thirty years
light green turning brown, slightly four-angled,
after a plane crash removed the elfin forest
and marked with half-round leaf scars. The wood
vegetation at 1,000 m elevation in the Luquillo
is light tan, weak and brittle, and has a 2-mm pith.
Mountains of Puerto Rico, 0.1 percent of the stems
Plants are supported by a shallow system of dark
of the resurgent vegetation was camasey almendro
tan, somewhat brittle roots. Opposite, light green
(Weaver 2000).
leaves are elliptic to ovate, pointed at both ends,
with the margin mostly finely serrate, and 5.4 to
Reproduction.—Camasey almendro flowers and
15.4 cm long. There are three main veins, the two
fruits throughout the year (Little and others 1974).
lateral ones arising 0.3 to 1.3 cm above the base of
The small fruits are produced in relatively large
the blade. Small inflorescences (panicles) are
numbers, and each fruit contains a large number of
axillary and bear several-to-many small, white,
seeds. A collection of fruits from Puerto Rico
four-petaled flowers. The juicy fruits (berries) are
469
averaged 0.138 ± 0.005 g/fruit. The seeds are tiny; ornamental in natural landscaping if it proves
an air-dried sample averaged 25.4 million/kg. The adaptable at lower elevations where the island
seeds can be cleaned by crushing, wet sieving with urban centers are located.
a fine mesh screen, filtering and drying on the
paper filter, and rubbing the seed off the paper. References
Without pretreatment seeds took 4 months to begin
germinating and completed 11 percent germination Liogier, H.A. 1995. Descriptive flora of Puerto
in 11 months. Germination is epigeal, and Rico and adjacent islands. Vol. 4. Editorial de la
seedlings are about 1 mm across the two cotyledon Universidad de Puerto Rico, San Juan, PR.
leaves. It is assumed that birds eat the fruits and 617 p.
disperse the seeds. Seedlings are scattered to
relatively common in suitable, disturbed habitat. Little, E.L., Jr., R.O. Woodbury, and F.H.
Layering (rooting) occurs whenever horizontal Wadsworth. 1974. Trees of Puerto Rico and the
stems come in contact with the ground. Camasey Virgin Islands. Vol. 2. Agriculture Handbook
almendro sprouts when stems are broken or cut. 449. U.S. Department of Agriculture,
Growth and Management.—Camasey almendro
grows moderately slowly and probably lives at McKenny, S. 2002. DNA evidence for the origin
least 2 or 3 decades. No planting or management of Mecranium latifolium (Melastomataceae), a
experience has been published. Removal of the Puerto Rican endemic. Abstract in: Elkin Isaac
forest canopy and soil disturbance near seed Research Symposium, Apr. 18, 2002, Albion
sources would likely result in reproduction. College, Albion, MI. 1 p.
Benefits.—Camasey almendro helps protect the Weaver, P.L. 2000. Elfin woodland recovery 30
soil and furnishes food and cover for wildlife. It is years after a plane wreck in Puerto Rico’s
an attractive plant and adds to the aesthetics of the Luquillo Mountains. Caribbean Journal of
forest. The species might be suitable as an Science 36(1-2): 1-9.
470
Melochia nodiflora Sw. malva colorada
STERCULIACEAE
Synonyms: Mougeotia nodiflora (Sw.) HBK.

Riedlea nodiflora (Sw.) DC.
Visenia nodiflora (Sw.) Spreng. in L.
John K. Francis
Liogier 1994). The species is not known to have

been planted or naturalized outside its natural
range.
Ecology.—Malva colorada grows on all types of

well drained soils with medium or high fertility
(Sánchez, and Uranga 1993) derived from
metamorphic (including ultramafic) rocks. In
Puerto Rico, it grows in areas that receive from
750 to about 2000 mm of mean annual
precipitation. In dry areas, the species is confined
to riparian zones. Malva colorada grows in Puerto
Rico from near sea level to over 600 m in
elevation. The species does not tolerate heavy
shade and seems to require at least minor soil
disturbance to become established. Malva
colorada grows in old fields and neglected
General Description.—Malva colorada, also pastures, on roadsides and fencerows, along river
known as britónica prieta and mauve, is a flood planes and bars, and on neglected
medium-sized shrub to 3 m in height and 5 cm in construction sites and vacant lots. It competes well
stem basal diameter. The species forms a fairly with low grass and broadleaf weeds but does not
robust tap and lateral root system. The species is grow under closed tree canopies.
herbaceous when small, becoming woody before
the end of the first year. Malva colorada may have Reproduction.—Malva colorada flowers over a 2
single or multiple stems that arise from the root or 3 month period. After flowering, capsules take
crown and lateral roots and large lateral branches about 6 months to mature seeds. The species
may form low on the stem. Malva colorada often produces large quantities of seeds that are released
develops only primary and secondary branches. during the dry season. They are small and are
The bark of older stems and branches is reddish carried by wind, water, animals and agricultural
brown. The leaves are alternate with petioles 0.5 equipment (Sánchez and Uranga 1993); no
to 3 cm long and ovate or ovate-lanciolate blades specialized means of dispersal is known. Seeds of
are 2.5 to 10 cm long and 1.2 to 6 cm broad. The malva colorada collected in Puerto Rico weighed
tip is pointed, the margin serrate, and the veins an average of 0.00167 g/seed or 597,000 seeds/kg.
prominent. The inflorescences are axillary in Seeds with no pretreatment were placed to
dense globose clusters in which are situated tiny, germinate on moist filter paper, and 36 percent
dark pink, pink, or white flowers. The germinated over an 11-month period. Germination
inflorescences mature into clusters of reddish- is epigenous.
brown capsules each of which contain, when
completely filled, five 2-mm brown seeds Growth and Management.—Individual stems
(Grisebach 1864, Howard 1989, Liogier 1994). live from 2 to about 4 years. Through suckers and
coppices, plants can live several times longer.
Range.—Malva colorada is native to the Malva colorada is sometimes a weed and is
Bahamas, the West Indies, and from Mexico primarily a problem in perennial plantations such
through Brazil (Grisebach 1864, Howard 1989, as citrus, pasture, coffee, and sugar cane (Sánchez
471
and Uranga 1993). The species is controlled by Howard, R.A. 1989. Flora of the Lesser Antilles,
mowing, cutting with machete, and by spraying Leeward and Windward Islands. Vol. 5. Arnold
with broadleaf weed killers. Arboretum, Harvard University, Jamaica Plain,
MA. 604 p.
Benefits.—Malva colorada is browsed by goats
and to a limited extent by cattle. It protects against Liogier, H.A. 1994. Descriptive flora of Puerto
erosion in recently disturbed areas. Rico and adjacent islands. Vol. 3. Editorial de la
References 461 p.
Grisebach, A.H.R. 1864. Flora of the British West Sánchez, P. and H. Uranga. 1993. Plantas
Indian Islands. J. Cramer-Weinheim, New York. indeseables de importancia economica en los
789 p. cultivos tropicales. Editorial Científico-Técnisa.
La Habana, Cuba. 166 p.
472
Melochia tomentosa L. pyramid bush
STERCULIACEAE
Synonyms: Melochia frutescens Jacq.

Moluchia frutescens (Jacq.) Medikus
Melochia turpiniana Kunth
Moluchia tomentosa (L.) Britton
John K. Francis
pyramidal capsules 6 to 9 mm across. The seeds

are 2 mm long and reddish brown (Howard 1989,
Liogier 1994).
Range.—The native range of pyramid bush

extends from southern Florida and southern Texas
through the West Indies and Central America into
Brazil and Colombia (Howard 1989, Liogier 1994,
Instituto Nacional de Biodiversidad 2002,
Recursos Hidricos 2002). No planting or escapes
have been reported outside its natural range.
Ecology.—Pyramid bush grows in dry areas

(areas receiving from 750 to 1000 mm of annual
precipitation in Puerto Rico), usually within a few
km of the coast. It grows from near sea level to
altitudes of about 400 m in Puerto Rico. The
species tolerates a wide variety of well-drained
soils derived from sedimentary and igneous rocks
and grows on coastal sands. Because there is
relatively little competition, pyramid bush is most
common on sites that are rocky, eroded, or
excessively drained. In Texas, the species is found
on sandy or rocky soil in mesquite thickets, palm
groves, and dry streambeds (San Antonio
General Description.—Pyramid bush, also Botanical Gardens 2002). The species inhabits
known as tea bush, raichie, broom wood, broom pinelands in southern Florida (Long and Lakela
weed, black widow, balsam, bretónica afelpada, 1976). In Brazil, it is part of the pioneer
and bois-champignon, is an erect shrub 0.5 to 4 m community on the sandy soils of the coastal plain
in height and up to 10 cm in basal diameter. (Recursos Hidricos 2002). The species does not
Pyramid bush is supported by a tap and lateral root tolerate heavy shade, and overtopping by trees will
system. Usually, a single stem emerges from the eliminate it. It is not eaten by cattle and tends to be
ground, but it may branch near the base. The common on overgrazed rangeland--probably
branches farther up the stem are slender. The bark because competition from grasses and other plants
is slate-gray with many lenticels. Stemwood is tan is reduced.
colored, moderately hard, and tough. The leaves
are covered with a short wooly hair giving the Reproduction.—Pyramid bush flowers at the end
plant a gray flannel look. Ovate to lanciolate of the wet season (November and December) and
leaves are attached by short petioles and have ripens fruits about 2 months later. It flowers
blades 1.5 to 10.5 cm long and 0.9 to 8.5 cm broad during the summer and early fall in Texas (Everitt
with serrate margins. Its inflorescences are and Drawe 1993). Air-dried capsules of pyramid
axillary umbelliform cymes of perfect flowers bush collected in Puerto Rico weighed an average
with five pink, purple, or blue petals that are 8 to of 0.0135 g. Seeds separated from them weighed
13.5 mm long. The fruits that develop are an average of 0.00104 g/seed or 963,000 seeds/kg.
473
Sown without pretreatment on moist filter paper, Agropecuaria do Tropico Semi-Arido. 28 p.
79 percent germinated within a short period 6
months later. The species can be propagated with Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs,
cuttings (San Antonio Botanical Gardens 2002). and cacti of South Texas. Texas Tech University
There appear to be no specialized means of seed Press. Lubbock, TX. 213 p.
transport; the small seeds are probably disbursed
incidentally by wind, water, machinery, and cattle. Howard, R.A. 1989. Flora of the Lesser Antilles,
Growth and Management.—Nursery-grown Arboretum, Harvard University, Jamaica Plain,
pyramid bush seedlings in Texas reached 60 cm in MA. 604 p.
just 2 months after outplanting (San Antonio
Botanical Gardens 2002). However, adult plants in Instituto Nacional de Biodiversidad. 2002. The
Puerto Rico add only about 30 cm to their height cutting edge. Vol. 7, No. 3. http://www.
each year. Based on ring counts in Puerto Rico, inbio.ac.cr/papers/manual_plantas/jul00/jul00lea
plants appear to live about 10 years. Overgrazing .html. 1 p.
to reduce competition may be the best way of
promoting this species. Pyramid bush is often Liogier, H.A. 1994. Descriptive flora of Puerto
considered a weed in rangeland (Liogier 1990). Rico and adjacent islands. Vol. 3. Editorial de la
Mowing may be sufficient to control it. Universidad de Puerto Rico. Río Piedras, PR.
461 p.
Benefits.—Pyramid bush appears to have great
potential as a flowering ornamental for xeric Liogier, H.A. 1990. Plantas medicinales de Puerto
landscaping (San Antonio Botanical Gardens Rico y del Caribe. Iberoamericana de Ediciones,
2002). In natural habitat, it contributes to Inc., San Juan, PR. 566 p.
biodiversity, soil stability, and wildlife cover.
Although cattle avoid it, sheep and goats in Brazil Long, R.W. and O. Lakela. 1976. A flora of
browse it in mixture with other native species Tropical Florida. Banyon Books, Miami, FL.
(Charles and others 1983). A tea is made from the 962 p.
foliage to treat colds and as an eye wash (Liogier
1990). Recursos Hidricos. 2002. Plano diretor de recursos
hídricos das bacias hidrográficas dos tributaries
References estaduaia dos rios vazas barris e real. Vol. 3, part
4.1. http://hidricosw.mg.gov.br/vazareal/vol3/
Charles, T.N.P., A.M. Maia, C. Guimaraes Filho, 41.htm. 4 p.
L.M.C. Salviano, and E.A.P. de Figueiredo.
1983. Effect of roughage supplementation and San Antonio Botanical Gardens. 2002. Friendly
mineral mixture plus deworming on sheep and natives for Texas. http://www.sabot.org/
goat production. II. Lamb and kid growth rates. natives/native.html. 3 p.
Boletim de Pesquisa No. 20. Centro de Pesquisa
474
Miconia impetiolaris (Sw.) D. Don ex DC. camasey de costilla
MELASOMATACEAE
Synonyms: Melastoma impetiolaris (Sw.) D. Don

Miconia wydleriana DC.
Tamonea impetiolaris (Sw.) Cook & Collins
John K. Francis
tiny brown seeds (Croat 1978, Howard 1989,

Liogier 1995, Little and others 1974, Stevens and
others 2001).
Range.—Camasey de costilla is native to Mexico,

Central America, South America as far south as
Bolivia, and to the Greater and Lesser Antilles
(Liogier 1995, Little and others 1974, Stevens and
others 2001). Three varieties have been described
(Stevens and others 2001). The species is not
its native range.
Ecology.—Camasey de costilla grows from a few

meters above sea level to about 600 m in elevation
(Little and others 1974, Stevens and others 2001).
It requires moist to wet habitat where it rains from
about 1500 to about 3000 mm/year. Although a
wide variety of soils are tolerated, the species is
most frequent on well drained to somewhat poorly
drained, neutral to strongly acid clays and clay
loams, often with the A horizons absent or
partially gone because of erosion. Excessively
General Description.—Camasey de costilla, also drained soils are not colonized. Camasey de
known as camasey colorado, cordobán, oreja de costilla is moderately intolerant of shade. It
mula, dos caras, auguey, hoja de pasmo, punta de succumbs in dense shade, grows but does not fruit
sarvia, maya, and trios côtes, is a shrub or small under a forest canopy, and fruits in broken or full
tree 2 to 8 m in height. The plant is supported by a sun. The species requires disturbance to become
deep taproot and strong, lateral roots. These roots established (Stevens and others 2001). It may be
are light gray and have fissured bark. Camasey de found in early and middle secondary forests,
costilla forms multiple stems if cut or damaged. plantations, fencerows, pastures, and roadsides.
Stem bark is brown and finely fissured. Young The species varies from an occasional plant to
branches are stout, quadrangular, and covered with common as dispersed plants or in clumps.
brown hairs; lower leaf surfaces, petioles, and
flower stalks of camasey de costilla are also Reproduction.—Camasey de costilla flowers
covered with brown pubescence. Leaf petioles are from spring until fall in Puerto Rico (Little and
2 to 7 mm long and mostly obscured by the others 1974). It reportedly flowers during the dry
cordate leaf base. The leaves are opposite and season (February and March) and fruits during the
usually have five main nerves. The blades are wet season (April and May) in Panama (Croat
ovate, minutely serrate at the edges, long pointed, 1978), and flowers and fruits throughout the year
12 to 40 cm long, and roughly half as wide. Large in Nicaragua (Stevens and others 2001). The seeds
terminal panicles bear many tiny, stalkless white are disbursed by birds and primates (Molano and
flowers. The fruits are berries 4 to 7 mm in others 2002). A collection of fruits from Puerto
diameter progressing from red to purple or blue- Rico averaged 0.242 + 0.008 g/fruit. Air-dried
black as they ripen. Each fruit contains numerous seeds from these fruits averaged 0.00044 g/seed or
475
2.3 million seeds/kg. Sown on wet peat, 56 References
percent of the seeds germinated between 13 and
23 days after sowing. The new seedlings are tiny Croat, T.B. 1978. Flora of Barro Colorado Island.
and develop slowly (author’s observation). Stanford University Press, Stanford, CA. 943 p.
Growth and Management.—Camasey de costilla Howard, R.A. 1989. Flora of the Lesser Antilles,
is relatively slow growing. About 0.25 to 0.5 m of Leeward and Windward Islands. Vol. 5. Arnold
height is added annually. Plants live at least 10 Arboretum, Harvard University, Jamaica Plain,
years and probably much longer. No known MA. 604 p.
attempts have been made to plant or manage
camasey de costilla. Probably management Liogier, H.A. 1995. Descriptive flora of Puerto
practices that maintain forests interspersed with Rico and adjacent islands. Vol. 4. Editorial de
pastures, brushlands, and roadsides will ensure at la Universidad de Puerto Rico, San Juan, PR.
least a presence of camasey de costilla. 617 p.
Benefits.—Camasey de costilla is employed as a Liogier, H.A. 1990. Plantas medicinales de Puerto

living fencepost in Colombia when it springs up Rico y del Caribe. Iberoamericana de Edicones,
naturally along fence rows (Molano and others Inc. San Juan, PR. 566 p.
2002). The wood is light colored and hard (Little
and others 1974) and used to a limited extent for Little, E.L., Jr., R.O. Woodbury, and F.H.
stakes and fuel. Camasey de costilla benefits Wadsworth. 1974. Trees of Puerto Rico and the
wildlife by furnishing food (fruits) and cover. It Virgin Islands. Vol. 2. Agriculture Handbook
also contributes to biodiversity and protects the 449. U.S. Department of Agriculture,
soil from erosion. In herbal medicine, leaf extracts Washington, DC. 1,024 p.
are used for aromatic baths, to arrest bleeding,
promote healing, and to treat mouth sores (Liogier Molano, J.G., M.P. Quicero, and C. Roa. 2002. El
1990). The species is pretty enough to be papel de las cercas vivas en un sistema de
employed as an ornamental, but perhaps because produccion agropecuaria en el Pidemonte
of the difficulty of propagation, it has not been Llanero. http://lead.virtualcentre.org/es/ele/
used so far. conferencia2/vbconfe3.htm. 12 p.
Stevens, W.D., C. Ulloa-U., and O.M. Motiel, eds.

2001. Flora of Nicaragua, Angiospermas.
2. Missouri Botanical Garden Press, St. Louis,
MO. p. 945-1,910.
476
Miconia prasina (Sw.) DC. camasey blanco
MELASTOMATACEAE
Synonyms: Melastoma prasina Sw.

Miconia collina DC.
Acinodendrom prasinum (Sw.) Krasser in Engl. & Prantl.
Melastoma parviflora Aublet
Melastoma pendulifolia Rich.
Miconia parviflora (Aublet) Cogn. in A. DC.
John K. Francis
tiny whitish flowers that develop into berries 3 to

8 mm in diameter and blue to black at maturity.
The fruits are juicy, slightly sour but mostly
tasteless. Each fruit contains many minute brown
seeds (Croat 1978, Howard 1989, Liogier 1995,
Little and Wadsworth 1964, Stevens and others
2001).
Range.—Camasey blanco is native to the Greater

and Lesser Antilles, Trinidad, Mexico, Central
America, South America to Bolivia, Brazil, and
Paraguay (Howard 1989, Killeen and others 1993,
Liogier 1995, Little and Wadsworth 1964). The
species is not known to have been planted or
Ecology.—Camasey blanco grows in secondary

and remnant forests, brushy pastures, fencerows,
creek-bottom galleries, coastal thickets, and
disturbed areas. The sites where it grows are moist
or wet and sometimes swampy, receiving from
1600 to 3500 mm of annual precipitation. The
species occupies elevations from near sea level to
650 m in Costa Rica (Missouri Botanical Garden
General Description.—Camasey blanco, also 2002) and up to 1,500 m in Bolivia (Killeen and
known as sardine, granadillo bobo, cenizoso, others 1993). Camasey blanco grows in loamy and
mullaca colorado, waraia, santo selele bélétére, clayey soils derived from both sedimentary and
jacatirão, and mondururu preto, is a shrub or small igneous rocks. Most of these soils have anaerobic
tree usually 2 to 6 m in height. It is an evergreen subsoils. Leaves collected under a Pinus caribaea
with single or multiple stems arising at or near the Morelet plantation had 2.34 percent N, 0.13
ground level. Camasey blanco tends to be percent P, 0.90 percent K, and 10.29 percent ash;
branchy. The sapwood is light brown and the were higher in N, P, and K, but lower in ash than
heartwood is grayish brown. It is a hard and heavy these leaves of plants from nearby natural forest of
wood with a specific gravity of 0.7. The bark is similar age (Lugo 1992). Camasey blanco is a
smooth, gray, and thin. The plant is supported by a shade intolerant pioneer species. It was the second
lateral root system with sinkers and a moderate most abundant species 7 years following
amount of fine roots. The leaves frequently have destruction of forests in Nicaragua by Hurricane
winged petioles and leaf blades that are elliptic to Juana (Granzow-de la Cerdal and others 2002). In
lanciolate, 10 to 30 cm long, entire or with wavy- Puerto Rico, the species appears to invade active
toothed edges, three- to five-nerved, and pointed pastures, taking advantage of disturbance by cattle
at both ends. The terminal panicles contain many
477
and persists as small individuals until the pastures goats. The fruits are eaten by the Puerto Rican
are abandoned, after which they grow rapidly and birds, Mimocichla ardosiaceae portoricensis
dominate for 10 to 20 years (Pascarella 2002). It Bryant, Vircosylva calidris calidris L., Spindalis
usually grows as dispersed individual shrubs or portoricensis Bryant (Whetmore 1916) and
small clumps. Camasey blanco disappears in a few certainly many other species across the range.
seasons after being overtopped by a forest canopy. Camasey blanco serves as a transitional
(successional) species between pasture, farmland,
Reproduction.—Camasey blanco flowers from or disturbed forest and secondary forest.
February to June and fruits from April to
September in Nicaragua (Stevens and others References
2001). The species is reported to flower and fruit
twice each year in Panama (Croat 1978). In Puerto Croat, T.B. 1978. Flora of Barro Colorado Island.
Rico, it is reported to flower and fruit almost Stanford University Press, Stanford, CA. 943 p.
throughout the year (Little and Wadsworth 1964).
Fruits collected in Puerto Rico averaged 0.137 ± Englerth, G.H. and E. Goytía-Olmedo. 1960.
0.003 g/fruit. Seeds separated from them averaged Preservation of Puerto Rican fence posts treated
10.6 million seeds/kg. Sown on wet peat, 57 by cold soaking and hot-and-cold bath method.
percent of the seeds germinated between 20 and Tropical Forest Notes 2. U.S. Department of
63 days after sowing. The seedlings are tiny and Agriculture, Forest Service, Tropical Forest
difficult to manage. In nature, birds are the Recearch Center, Río Piedras, PR. 4 p.
principal dispersers of seeds. Camasey blanco
sprouts readily when disturbed.
Francis, J.K. 2000. Estimating biomass and carbon
content of saplings in Puerto Rican secondary
Growth and Management.—The largest
forest. Caribbean Journal of Science 36(3-4):
camasey blanco recorded in Puerto Rico had a
346-350.
diameter at breast height of 16.3 cm and a height
of 11 m (Puerto Rico Champion Tree List on file
Granzow-de la Cerdal, I., N. Zamora, J.
at the International Institute of Tropical Forestry,
Vandermeer, and D. Boucher. 2002. Diversidad
Río Piedras, PR). The species has a moderate
de especies arbóreas en el bosque tropical
growth rate and is relatively short lived. None of a
húmedo del Caribe nicaragüense siete años
group of 24 camasey blanco stems in a survey in
después del huracán Juana. http://www.rbt.
Puerto Rico were present 24 years later at a
ac.cr/revistas/45-4/granzow.htm. 15 p.
second visit (Weaver 1979). However, by
suckering and ground layering, plants can
Howard, R. A. 1989. Flora of the Lesser Antilles,
perpetuate themselves somewhat longer than
individual stems. Nursery propagation of the
species has not been reported.
MA. 604 p.
Benefits.—Camasey blanco wood is not durable
in the ground and therefore not suitable as fence Killeen, T.J., E. García-E., and D.G. Beck. 1993.
posts (Little and Wadsworth 1964), unless treated. Guía de árboles de Bolivia. Herbario Nacional
Penetration of preservative was about average for de Bolivia and Missouri Botanical Garden, La
52 Puerto Rican species tested (Englerth and Paz and St. Louis, MO. 958 p.
Goytía-Olmedo 1960). The wood is used for fuel
and occasionally as tool handles. Wood specific Liogier, H.A. 1995. Descriptive flora of Puerto
gravity of 30 plants sampled in Puerto Rico was Rico and adjacent islands. Vol. 4. Editorial de la
measured at 0.630 + 0.021. A weighted average of Universidad de Puerto Rico, San Juan, PR. 617
carbon content for above-ground parts was 50.0 p.
percent. Total above-ground biomass in grams
may be predicted by multiplying the stem Little, E.L., Jr. and F.H. Wadsworth. 1964.
diameter in cm squared at 30 cm above the soil Common trees of Puerto Rico and the Virgin
times stem length in meters by 38.344 (R2 = Islands. Agriculture Handbook 249. U.S.
0.981) (Francis 2000). The fruits are edible but Department of Agriculture, Forest Service,
seldom eaten by people. Camasey blanco is Washington, DC. 548 p.
browsed sparingly by cattle and more heavily by
478
Lugo, A.E. 1992. Comparison of tropical tree Monographs in Systematic Botany, Vol. 85, No.
plantations with secondary forests of similar 2. Missouri Botanical Garden, St. Louis, MO.
age. Ecological Monographs 62: 1-41. pp. 945-1,910.
Missouri Botanical Garden. 2002. Flora of Costa Weaver, P.L. 1979. Tree growth in several tropical
Rica: Miconia prasina. http://www.mobot.org/ forests of Puerto Rico. Research Paper SO-152.
manual.plantas/023023/S024494.html. 2 p. U.S. Department of Agriculture, Forest Service,
Pascarella, J.B. 2002. Tropical forest succession in Orleans, LA. 15 p.
Puerto Rico. http://chiron.valdosta.edu/
jbpascar/Research/Prico/puerto_rico.htm. 6 p.
Whetmore, A. 1916. Birds of Porto Rico. Bulletin
326. U.S. Department of Agriculture.
Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
479
Miconia racemosa (Aubl.) DC. camasey de felpa
MELASTOMATACEAE
Synonyms: Melastoma racemosa Aubl.

Melastoma decussata Vahl
Miconia brachypoda DC.
Tamonea racemosa (Aubl.) Cook & Collins
John K. Francis

1974).
Range.—Camasey de felpa is native to

Hispaniola, Puerto Rico, Dominica, St. Lucia,
Granada, Trinidad and Tobago, Venezuela, the
Guianas, and Brazil to the Atlantic forest in the
South (Centro Nordestino de Informaçõas Sobre
Plantas 2002, Fundaçao Andre Tosello 2002,
1974).
Ecology.—Camasey de felpa inhabits areas that

receive 1600 to 3000 mm of annual precipitation.
Common habitat is on moderately well drained,
somewhat poorly, and poorly drained, clayey,
weathered soils, particularly ultisols. Areas of both
sedimentary and igneous rocks are colonized at
elevations from near sea level to 900 m in
elevation. The species is shade intolerant,
requiring partial sunlight to flower and fruit.
Disturbance favors reproduction, but this
requirement does not appear to be absolute.
Seedlings are frequently found under plantations
General Description.—Camasey de felpa in mid-rotation or trees that have colonized old
(meaning “felt” Miconia in Spanish), also known fields. Camasey de felpa may be found in old
as terciopelo and camasey racimoso, is an fields, tree plantations, secondary forests,
evergreen shrub or small tree 2 or 3 m in height roadsides, and landslides (author’s observation,
and 3 to 5 cm in stem diameter. The stems are Little and others 1974).
smooth and gray with an inner bark of light gray.
The plants are supported by an ample lateral root Reproduction.—Camasey de felpa flowers and
system. Multiple stems arising from the root fruits throughout the year (Little and others 1974).
crown and lower trunk are common. The wood is A collection of fruits from Puerto Rico weighed
light brown and hard. Twigs are light green and 0.1315 ± 0.0041 g/fruit. The seeds are tiny
somewhat four-angled and have a ring of hairs at amounting to several million per kg. Placed on
the nodes. The opposite elliptic leaves have moist filter paper at ambient temperature, 33
petioles 8 to 45 mm long, five main veins from the percent germinated within 18 days. The seeds are
base, a saw-toothed, hairy edge, and are pointed at principally dispersed by birds (Devoe 1989). A
both ends. The inflorescences are panicles with study in Puerto Rico showed predispersal losses of
many tiny white, pink, or purple flowers. The seeds to vertebrates and insects accounted for 3
fruits are berries, 3 to 5 mm in diameter, whitish percent of the seeds, fungi destroyed 9 percent,
when immature, dark blue, purple, or black at and postdispersal loss to ants was 15 percent
maturity, and contain numerous tiny brown seeds (Myster 1997). Although seedlings are rarely
480
abundant, the species is common at least in Puerto regeneration in openings and beneath closed
Rico, growing in small clumps or as scattered canopy in sub-tropical wet forest. Ph.D.
individuals. Camasey de felpa coppices readily dissertation, Yale University, New Haven, CN.
when cut. 307 p.
Growth and Management.—Camasey de felpa Fundaçao Andre Tosello. 2002. Base de dados
has a moderate growth rate and probably lives 10 tropicles: Espécies arbóreas da Mata Atlântica.
to 20 years. No management experience has been http://www.bdt.fat.org.br/mata.atlantica/flora/
published. Natural reproduction could be local?SC. 5 p.
encouraged by disturbance of moist and wet forest
and unwanted plants eliminated by grubbing or
spraying with broadleaf herbicides.
Benefits.—Camasey de felpa contributes to the
MA. 604 p.
aesthetics of forests where it grows, helps
revegetate disturbed sites and protect the soil, and
provides food and cover for birds and other
wildlife. The fruits are juicy and edible but small
and almost tasteless (Little and others 1974). The
617 p.
wood is useful for fuel, small fence posts, and
stakes.
References
Centro Nordestino de Informaçõas Sobre Plantas.
2002. Checklist das plantas do Nordeste.
Universidade Federal de Pernambuco.
Myster, R.W. 1997. Seed predation, disease and
http://umbuzeiro.cnip.org.br/db/pnechk/fam/
germination on landslides in neotropical lower
f104.shtml. 6 p.
montane wet forest. Journal of Vegetation
Science 8(1): 55-64.
Devoe, N.N. 1989. Differential seeding and
481
Mimosa aculeaticarpa Ortega wait-a-bit
FABACEAE
Synonyms: Mimosa biuncifera Benth.

Mimosa lindheimeri Gray
Mimosa warnockii B.L. Turner
Mimosopsis biuncifera (Benth.) Britt. & Rose
1997). Barneby (1991) recognized four variants of

Mimosa aculeaticarpa of which var. biuncifera
(Benth.) Barneby is a particularly common one.
However, even within each variety he emphasizes
that substantial variability exists. Varieties
glabrescens and lindheimeri are synonymous with
var. biuncifera (Kartesz 1994) and should not be
confused with the variants described by Barneby
(1991).
Range.—Catclaw mimosa occurs in central and

southern Arizona, southern New Mexico, western
and central Texas, and northern Mexico in Nuevo
León, Tamaulipas, Durango, Coahuila, Puebla,
San Luis Potosí, and Chihuahua (Carter 1997,
Kearney and others 1960, Martin and Hutchins
1981, Vines 1986).
Ecology.—Catclaw mimosa grows at lower

elevations, between 600 to 1,525 m (2,000 to
Illustration source: USDA-Forest Service collection, 5,000 ft), frequently in limestone and igneous rock
Hunt Institute habitats (Powell 1998). It often becomes abundant
on dry, rocky, open slopes (Pase and Brown 1982)
General Description.—Mimosa aculeaticarpa is although it can be equally profuse in washes where
a straggling, thicket-forming shrub typically armed water availability is periodically copious. Catclaw
with curved prickles that are responsible for the mimosa is a member of various communities:
common names, wait-a-bit, Catclaw mimosa, Una Desert shrub, Texas savanna, Southwestern shrub
de Gato and Gatura. The origin of the genus name steppe, Chaparral - mountain shrub, Pinyon –
is likely from the Spanish word Mimoso meaning juniper woodlands, Plains, and Desert grasslands
“sensitive”, which refers to the sensitive, (Dick-Peddie 1993, Cable 1975, Uchytil 1990).
thigmotactic leaf movement (Barneby 1991). An Many authors have mentioned the importance and
equally appropriate derivation may be from the frequency of low shrubs, such as Catclaw mimosa,
Greek mimos that means “to mimic” and refers to in a perennial-grass dominated matrix on rocky
the animal-like response of the leaf movements boulder-ridden terrain in Arizona (Shreve 1915,
(Gledhill 1992, Vines 1986). Although plants may Whittaker and Niering 1965), Mexico (Muller
have straight prickles or be unarmed, they most 1939, 1947), New Mexico (Dick-Peddie and Moir
commonly have two curved spines per node and 1970), and Texas (Gehlbach 1967).
spines on the leaves and pods (Barneby 1991,
Bowers 1993, Carter 1997, Powell 1998). Shrubs Reproduction.—Flowering dates are variable
are generally 1 to 3m tall with deciduous, divided (generally late April to September), and available
leaves that may have as many as 300 tiny leaflets. moisture is the limiting factor for flowering
The species has variable phenotypic and habitat (Carter 1997). The individual flowers are small
characteristics and has been described as (<4 mm) and they are massed in a pale-pink to
genetically and ecologically confusing (Carter whitish globose head. The flowers are fragrant and
482
are pollinated by insects. Only a few flowers in References
each inflorescence produce pods (Bowers 1993).
The fruit is an elongated flattened legume (pod), Ahlstrand, G.M. 1982. Response of Chihuahuan
the margins separate from the valves at Desert Mountain Shrub Vegetation to Burning.
dehiscence. The valves often break transversely Journal of Range Management. 35 (1): 62-65.
into one-seeded sections. The seeds exhibit high
germination rates over a relatively wide range of Barneby, R.C. 1991. Sensitivae Censitae. Memoirs
temperatures (Jordan and Haferkamp 1989). of the New York Botanical Garden Vol. 65.
Livestock and wildlife are likely to disperse the The New York Botanical Garden, Bronx, NY.
seeds (Uchytil 1990).
Bowers, J.E. 1993. Shrubs and Trees of the
Growth and Management.—Catclaw mimosa
Southwest Deserts. Southwest Parks and
forms true brier patches that may become almost
Monumens Association, Tucson, AZ. 140 p.
impenetrable to humans (author’s personal
observation). Plants can profusely re-sprout from
Buechner, H.K. 1950. Life history, ecology, and
protected buds following fire-induced mortality of
range use of the pronghorn antelope in Trans-
aboveground tissue (Cable 1975, Carmichael and
Pecos Texas. American Midland Naturalist.
others 1978, McPherson 1995). However, in
43(2): 257-354.
general, plants must be several years old before
they have the ability to re-sprout. Re-sprouting
Cable, D.R. 1975. Range management in the
decreases as soil-moisture decreases and is less
chaparral type and its ecological basis: the status
common after fires that occur during the growing
of our knowledge. Research Paper RM-155.
season compared with dormant-season fires
(McPherson 1995). Stands can recover from
Rocky Mountain Forest and Range Experiment
burning in 5 years (Ahlstrand 1982, Kittams
Station, Fort Collins, CO. 30 p.
1972). However, at least in Arizona, wildfire does
not appear to stimulate sprouting (Pase and Pond
Carmichael, R.S., Knipe, O.D., Pase, C.P. and
1964). Catclaw mimosa is moderately resistant to
Brady, W.W. 1978. Arizona chaparral: plant
phenoxy herbicides and after only one application
associations and ecology. Research Paper RM-
tend to re-foliate or re-sprout from the crown
202. U.S. Department of Agriculture, Forest
(Hibbert and others 1974). Successive applications
Service, Rocky Mountain Forest and Range
kill the plants.
Experiment Station, Fort Collins, CO. 16 p.
Benefits.—Catclaw mimosa thickets provide
Carr, M.E., Mason, C.T., Jr., and Bagby, M.O.
cover and food for a variety of small wildlife
1986. Renewable resources from Arizona trees
species, especially quail (Graham 1941, Powell
and shrubs. Forest Ecology and Management.16:
1998). The flowers are a good source of nectar for
155-167.
honey bees (Powell 1998, Vines 1986). Deer and
pronghorn browse Catclaw mimosa and its
Carter, J.L. 1997. Trees and Shrubs of New
palatability is rated good (Nichol 1938, Buechner
Mexico. Johnson Books, Boulder, CO. 533 p.
1950). Its livestock palatability is low (Dayton
1931). However, livestock use it when other
Dayton, W.A. 1931. Important western browse
forage is scarce, and by virtue of its abundance in
plants. Miscellaneous Publication 101. U.S.
New Mexico and Arizona it has been described as
important cattle browse (Dayton 1931, Vines
214 p.
1986). Both livestock and wildlife eat the pods
(Dayton 1931, Cable 1975, Uchytil 1990).
Dick-Peddie, W.A. 1993. New Mexico
Because plants have a tendency to form thickets
Vegetation--past, present, and future.
that effectively bind soil, Catclaw mimosa shows
University of New Mexico Press, Albuquerque,
potential for erosion control (Kearney and others
NM. 244 p.
1960, Vines 1986). In a comparison with 100 other
plant species studied for the production of fuels
Dick-Peddie, W.A. and W.H. Moir. 1970.
and chemicals, it yielded substantial amounts of
Vegetation of the Organ Mountains, New
oils, polyphenols, and hydrocarbons (Carr and
Mexico. Range Science Department Science
others 1986).
483
Series 4. Colorado State University, Ft. Collins, Devender, eds. The Desert Grassland. The
CO. University of Arizona Press, Tucson, AZ.
Gehlbach, F.R. 1967. Vegetation of the Guadalupe Muller, C.H. 1939. Relation of the vegetation and
Escarpment, New Mexico-Texas. Ecology 48: climatic types in Nuevo Leon, Mexico.
793-806. American Midland Naturalist 21: 687-729.
Gledhill, D. 1992. The names of plants. Second Muller, C.H. 1947. Vegetation and Climate of
Ed. Cambridge University Press, Cambridge Coahuila, Mexico. Madrono. 9: 33-57.
England. 202 p.
Nichol, A.A. 1938. Experimental feeding of deer.
Graham, E.H. 1941. Legumes for erosion control Technical Bulletin 75. Arizona Agriculture
and wildlife. Miscellaneous Publication 412. Experiment Station. 39 p.
U.S. Department of Agriculture, Washington,
DC. 153 p. Pase C.P. and F.W. Pond. 1964. Vegetation
changes following the Mingus Mountain burn.
Hibbert, A.R., Davis, E.A., and Scholl, D.G. 1974. Note RM-18. U.S. Department of Agriculture,
Chaparral conversion potential in Arizona: Part Forest Service, Rocky Mountain Station, Fort
I: water yield response and effects on other Collins, CO. 8 p.
resources. Research Paper RM-126. U.S.
Department of Agriculture, Forest Service, Pase, C.P. and D.E. Brown. 1982. Interior
Rocky Mountain Forest and Range Experiment chaparral. In: D.E. Brown, ed. Biotic
Station, Fort Collins, CO. 36 p. Communities of the American Southwest—
United States and Mexico. Desert Plants. 4 (1-
Jordan, G.L. and M.R Haferkamp. 1989. 4): 95-99.
Temperature responses and calculated heat units
for germination of several range grasses and Powell, A.M. 1998. Trees & shrubs of Trans-
shrubs. Journal of Range Management. 42(1): Pecos and adjacent areas. (First Ed. copyright
41-45. Big Bend Natural History Assoc.) University of
Texas University Press, Austin, TX 498 p.
vascular flora of the United States, Canada and Shreve, F. 1915. The vegetation of a desert
Greenland. Vol.1-Checklist. Second ed. Timber mountain range as conditioned by climatic
Press Portland, OR. 622 p. factors. Publication 529. Carnegie Institute,
Washington, DC. p. 1-45.
Kearney, T.H., Peebles, R.H. and collaborators.
1960. Arizona flora. 2nd ed. University of Uchytil, R.J. 1990. Mimosa biuncifera. U.S.
California Press, Berkeley, CA. 1,085 p. Department of Agriculture, Forest Service,
Kittams, W.H. 1972. Effect of fire on vegetation of Laboratory. Fire Effects Information System.
the Chihuahuan Desert region. Proceedings: http://www.fs.fed.us/database/feis/.
Tall Timbers Fire Ecology Conf. A Quest for
Ecological Understanding 12: 427-444. Vines, R.A. 1986. Trees, shrubs, and woody vines
of the Southwest. University of Texas Press,
Martin, W. C. and C.R. Hutchins. 1981. A Flora of Austin, TX. 1,104 p.
New Mexico. Volume 2. J. Cramer, Vaduz,
Germany. p. 1,277-2,591. Whittaker, R.H. and W.A. Niering. 1965.
Vegetation of the Santa Catalina Mountains,
McPherson, G.R. 1995. The role of fire in desert AZ: a gradient analysis of the south slope.
grasslands. In: M. P. McClaran and T. R. Van Ecology 46: 429-452.
484
Mimosa arenosa (Willd.) Poir. tepehuiste
FABACEAE
Synonyms: Acacia arenosa Willd.

Acacia malacocentra Mart.
Mimosa caudero L.
Mimosa leiocarpa DC.
Mimosa malacocentra (Mart.) Benth.
Mimosa xantholasia Benth.
John K. Francis
four to 22 pinnae each with 15 to 35 pairs of

leaflets. Inflorescences are 6-cm spikes with
paniculiform branching bearing many tiny, white
flowers. The fruits, which are borne in clusters,
are linear-oblong, flat, brown legumes, 4 to 5 cm
long by 5 to 6 mm broad. The seeds are yellow,
flattened, and about 4.5 by 5 mm wide and long
(author’s observations, Liogier 1988, Stevens and
others 2001).
Range.—Tepehuiste is native to South America

from Colombia and Venezuela through Brazil and
lowland Bolivia. It is possibly native to Mexico
(recorded in Jalisco State) and Nicaragua.
Populations in Puerto Rico and the Dominican
Republic were introduced and have become
naturalized (Liogier 1988, Missouri Botanical
Garden 2002, New York Botanical Garden 2002,
Stevens and others 2001).
Ecology.—Tepehuiste is intolerant of shade. It

competes rigorously with grass, herbs, and low
shrubs but cannot survive overtopping by tall
trees. The Puerto Rican populations occur in areas
that receive from 1200 to 1600 mm of mean
annual precipitation on well-drained to somewhat
General Description.—Tepehuiste is a thorny poorly-drained sites. Tepehuiste does not appear
shrub or tree usually 3 to 5 m (up to 12 m) in to compete or survive on rocky, excessively well-
height and stem diameters to 15 cm or more. drained sites. The species grows at elevations from
Young and undisturbed plants have a single stem near sea level to 500 m in Nicaragua. Tapehuiste
that may branch several times near the ground. As grows in catinga (scrub) forest in Brazil, savannas
trees become older and heavier, they tend to lie in Nicaragua, and vacant lots, roadsides,
down and produce new, vertical sprouts. fencerows, and neglected pastures in Puerto Rico
Sometimes, the mid section of the trunk is thicker (author’s observations, Nunez de Medeiros 2002,
than the base. The hard, somewhat flexible wood Stevens and others 2001).
has light tan sapwood and light brown or brown
heartwood. The plant is supported by a tap and Reproduction.—Tepehuiste blooms June through
lateral root system of brown, strong, and flexible September and fruits September and October in
roots. Stems are covered by dark brown, shallowly Nicaragua (Stevens and others 2001). The author
furrowed bark. The inner bark is green. Twigs are observed stands in Puerto Rico in bloom in March
greenish brown with 3 to 4 mm, curved spines. and other stands with heavy seed crops during that
The alternate, compound leaves commonly have
485
same period. Insects pollinate the flowers. Fruits References
collected in Puerto Rico averaged 7.0 + 0.2
seeds/pod. Seeds separated from them weighed an Liogier H.A. 1988. Descriptive flora of Puerto
average of 0.0054 ± 0.0006 g/seed or 186,000 Rico and adjacent islands, Spermatophyta. Vol.
seeds/kg. One percent of unscarified seed 2. Editorial de la Universidad de Puerto Rico,
germinated in 6 months. but a lot of mechanically Río Piedras, PR. 481 p.
scarified seed germinated at 99 percent between 3
and 5 days after sowing on moist filter paper Missouri Botanical Garden. 2002. VAST
(author’s observation). Germination is epigeal nomenclatural database. http://mobot.mobot.
(Parra 1984). Plants sprout when cut or burned. org/cgi-bin/search_vast?name=Mimosa
+arenosa. [not paged].
Growth and Management.—Tepehuiste grows
rapidly. Because the species is thorny, forms New York Botanical Garden. 2002. Neotropical
almost impenetrable thickets, and aggressively flora mycota catalog. http://scisun.nybg.org:
invades pastures and rangeland, it is a highly 8890/searchdb/owa/wwwspecimin.search_list?
undesirable species in exotic habitats. Every effort taxon=Mimosa+arenosa&projcode=NETR. [not
should be made to prevent it from invading new paged].
areas. Control techniques have not been published.
However, the same measures used against Mimosa Nunez de Medeiros, A. 2002. Caprinocultura de
pigra L. probably would be effective. Corte no Nordeste Brasileiro. Departamento de
Agropecuária, Bananeiras, PB, Brazil.
Benefits.—Tepehuiste helps sites move through http://www.capritio.com.br/art18.htm. [not
the grass stage to secondary forest, protects the paged].
soil, and provides cover for wildlife. It is an
important forage plant for goats in northeastern Parra-G., P. 1984. Estudio de la morfologia
Brazil (Nunez de Medeiros 2002). The wood is externa de plantulas de Calliandra gracilis,
useful for fuel. Mimosa albida, Mimosa arenosa, Mimosa
camporum y Mimosa tenuiflora. Revista de la
Facultad de Agronomía (Maracay) 8(1-4): 311-
350.
Stevens, W.D., C. Ulloa-U., A. Pool, O.M.

Monographs in systematic botany Vol. 85, No.
2. Missouri Botanic Garden Press, St. Louis,
MO. p. 945-1,910.
486
Mimosa ceratonia L. zarsa
FABACEAE
Synonyms: Lomoplis ceratonia (L.) Raf.
John K. Francis
The pods, which are borne in clusters, are straight

or slightly curved, oblong, 4 to 6 cm long and
armed at the edges. The flattened seeds are brown
or black (Acevedo-Rodríguez 1985, Howard 1988,
Liogier 1988).
Range.—The range of zarza includes Hispaniola,

Puerto Rico, the Virgin Islands, Antigua,
Guadeloupe, Martinique, St. Lucia, and Barbados
(Howard 1988, Liogier 1988). Also, populations
of zarza have been identified in Brazil including
the varieties M. ceratonia L. var. pseudo-obovata
(Taubert) Barneby and M. ceratonia L. var.
interior Barneby (Barneby 1985, Paganucci de
Queiroz 2001).
Ecology.—Zarza is moderately intolerant of

shade. It grows in fencerows, roadsides, wooded
drains, brushy pastures, forest edges and openings,
and in low-density secondary forest. The species
often forms compact, nearly impenetrable clumps.
Diffuse stands mixed with tall grass, weeds, and
shrubs form in areas that are mowed or burned on
one- to several-year cycles. Zarza is most common
on soils derived from limestone and serpentine
(Vélez and van Overbeek 1950), but also grows on
General Description.—Zarza, also known as soils overlying other sedimentary and igneous
rocks. It tolerates all textures of well-drained soils
climbing mimosa, black ambret, amourette-grand-
bois, grate-jambe, and croc-chien, is a multi- and a wide range of fertility levels. In Puerto Rico,
stemmed scrambling shrub. It supports itself on the species may be found in areas receiving
rainfalls from 750 mm per year to over 2000 mm
other vegetation by means of curved spines and
may reach 4 to 6 m into the crowns of trees and as per year. Zarza grows from near sea level to at
far laterally. The plant has a weak taproot, a least 600 m in elevation. Frosts do not occur
moderate amount of lateral roots, and abundant within its natural range.
fine roots. Lower stems are gray, scaly, and
cylindrical. The branches tend to be long, almost Reproduction.—Zarsa flowers between June and
straight or evenly curved, with few divisions. The December and fruits December through January in
twigs are green, sometimes reddish with many Puerto Rico (Acevedo-Rodríguez 1985). A
curved 2-mm spines. Lateral twigs usually die collection of zarza pods from Puerto Rico weighed
back after 1 year. The leaves are bipinnately an average of 0.1371 ± 0.0005 g (air dry). They
compound with three to five pairs of pinnae, each averaged 4.18 + 0.12 seeds/pod and ranged from
with three to five pairs of leaflets. The leaflet three to six seeds per pod. The seeds weighed an
blades are obliquely obovate, three-nerved, 1.0 to average of 0.0201 + 0.004 g/seed or 50,000
2.5 cm long and 6 to 10 mm broad, rounded at the seeds/kg. Sown on commercial potting mix, 71
percent germinated between 7 and 165 days after
apex and asymmetrically acute at the base. The
inflorescences are terminal racemes of globose sowing. Germination is epigeal. Stems layer (root)
heads of flowers that are white or cream-colored. if covered by soil or organic debris. Stems sprout
487
when cut and lateral roots sometimes sucker when Barneby, R.C. 1985. The identity and synonymy
damaged. The seeds are disbursed by lateral of Acacia guiladinae, Mimosa obovata, M.
extension of the vines and by the pods clinging to pseudo-obovata and M. laticifera (Mimosaceae).
clothing or to the fur of animals. Seedlings are Brittonia 37(1): 85-87.
common, but only a few survive for more than a
few weeks. Howard R.A. 1988. Flora of the Lesser Antilles,
Growth and Management.—Both seedlings and Dicotyledoneae, Part 1. Vol. 4. Arnold
sprouts grow rapidly, up to 2 or 3 m per year. Arboretum, Harvard University, Jamaica Plain,
Zarza plants may endure for many years. Because MA. 673 p.
of the thorny nature of this species, property
owners often choose to eliminate it from pastures Liogier H.A. 1988. Descriptive flora of Puerto
and plantations. Historically, farmers controlled it Rico and adjacent islands, Spermatophyta. Vol.
by repeated cutting. This method could probably 2. Editorial de la Universidad de Puerto Rico,
be improved by treating the resulting sprouts with Río Piedras, PR. 481 p.
broadleaf weed killers.
Marcano F., E. de J. 1973. La flora apícola de la
Detriments and Benefits.—Thickets and tangles República Dominicana. http://naturalista.
of zarza are almost impenetrable to humans. Paths virtualave.net/estudios/apicola/dicotse.html. 8 p.
may be cut to allow passage, but they soon grow
over. Because of the thorns, cattle will not eat Paganucci de Queiroz, L. 2001. Leguminosas da
zarza (Vélez and van Overbeek 1950), but it is caatinga da Bahia com potencial forrageiro.
browsed by goats. The species is a host for the http://umbuzeiro.cnip.org.br/db/forrag/fam/
diaprepes root weevil (Diaprepes abbreviatus), f19.html. 6 p.
which is of grave concern for growers of citrus
and other agricultural crops (Simpson and others Simpson, S.E., H.N. Nigg, and J.L. Knapp. 2001.
2001). Zarza is a valuable honey plant (Marcano Host plants of diaprepes root weevil and their
1973), protects the soil, and serves as wildlife implications to the regulatory process. Florida
cover. Department of Agriculture and Consumer
Services. http://www.fcprac.ifas.ufl.edu/
References citrustopics/pest%20control/Diaprepes/…/simps
on.hostplant.ht. 16 p.
Puerto Rico. Vol. 1. General Technical Report Vélez, I. and J. van Overbeek. 1950. Plantas
SO-58. U.S. Department of Agriculture, Forest indeseables en los cultivos tropicales. Editorial
Service, Southern Forest Experiment Station, Universitaria. Río Piedras, PR. 497 p.
488
Mimosa pigra L. black mimosa
FABACEAE
Synonyms: Mimosa asperata L
John K. Francis
are flattened, about 6 by 2.5 mm, and brown to

olive green (Howard 1988, Pacific Island
Ecosystems at Risk 2002, Stevens and others
2001).
Range.—The original range of black mimosa is

uncertain. The contention that the native range
extends from Mexico to northern Argentina
(Stevens and others 2001) is supported by the fact
that 440 species of insects and several fungi
species keep it under control there (CSIRO 2002).
On the other hand, a rich native tradition supports
Equatorial Africa as being native range (Burkhill
1995). Perhaps it crossed the South Atlantic in
prehistoric times or early in the colonization of the
New World. More recently, the species has
invaded most humid tropical and subtropical areas
including Florida, Texas, and Puerto Rico (Natural
Resources Conservation Service 2002). There are
two varieties: var. berlandieri (Gray ex Torr.) B.L.
Turner that occurs from the Texas to Costa Rica,
and var. pigra that grows from Mexico to
Argentina, Africa, and in the invaded ranges in the
New and Old Worlds (Stevens and others 2001).
General Description.—Black mimosa is also Ecology.—Presettlement populations of black
known as catclaw mimosa, thorny sensitive plant, mimosa in Costa Rica were found mainly on the
giant sensitive plant, bashful plant, giant mimosa, edges of marshes and on large river floodplains.
dormilona, pigra, agüiste, amouretto, amourette- Today, they are found along roads and in marshy
rivière, and banglin. It is a thorny upright to spots in pastures (Janzen 1983). The species is
scrambling shrub 2 to 4 m in height and up to 12 highly intolerant of shade. It requires full sunlight
cm in basal diameter. The plant usually has a to flower and fruit and nearly full sunlight to
single stem from the ground but may branch near survive. The two principal ingredients for
the ground and form an inverse cone-shaped successful establishment are moist or wet soils and
crown. There are relatively few branches. The disturbance that allows full sunlight. It is a fierce
shrub is supported by extensive lateral roots with competitor with low vegetation and one of the few
numerous fine roots that have occasional nodules. shrubs capable of succeeding in dense, tall grass
Stems, limbs, twigs, and leaves bear curved thorns swards. Black mimosa tolerates a wide variety of
up to 7 mm long. Black mimosa produces a thin soil conditions, short-term flooding, seasonal
crown of touch-sensitive bipinnately compound drought, and grows at elevations from near sea
leaves. Each leaf has four to 14 pairs of pinnae level to 700 m (Janzen 1983). Although it grows
and 20 to 40 pairs of linear-oblong leaflets per mixed with other vegetation in its native habitat,
pinna. The inflorescences are tight, subglobose the species forms pure stands with little understory
mauve or pink heads with about 100 flowers, in many of its exotic habitats (Pacific Island
grouped one to three in the upper axils. The Ecosystems at Risk 2002).
clustered brown legumes are densely bristled, 4 to
12 cm long, and breaking transversally into 14 to Reproduction.—Black mimosa blooms
26 segments, each containing one seed. The seeds
489
continuously in moist habitat, intermittently in environments helps them move on to the brush and
seasonally dry habitat. Bees pollinate the flowers secondary forest stages. It also protects the soil in
and pods mature about 1 month after pollination disturbed sites. Although the cover and diversity it
(Binggeli 1997). Black mimosa grows rapidly and furnishes is probably beneficial for wildlife in
can begin producing seed in as little as 3 months artificially maintained exotic grass swards, the
after emergence (Department of Natural Resources species can reduce wildlife populations in natural
and Mines 2002). One average plant can produce swards such as river floodplains in Australia
more than 9,000 seeds annually (Marko 2002). (Marko 2001). The plant has been used as a green
The pod segments disperse by floating on water manure, a cover crop, for fuel wood, and as
and clinging to clothing and possibly animal fur beanpoles. Although cattle and horses generally do
and feathers. A sample of black mimosa pods in not eat black mimosa because of the thorns
Puerto Rico contained an average of 15.0 (Binggeli 1997), with 22 percent foliar protein, it
seeds/pod. The seeds weighed an average of was found to be perfectly acceptable to rabbits
0.0166 g/seed. Sown on commercial potting mix (Lebas and others 1986). Black mimosa can be
between 6 and 20 days, 81 percent of the seed weedy in row crops and plantations, and a serious
germinated. Germination is epigeal (author’s problem in pastures and hay fields. The thorny
observation). Scarification is required for quick branches make it difficult for people to walk
and uniform germination. In an experiment with through fields and along trails where it grows. In
several treatments of temperature and storage, Africa, it is planted as a barrier around fields and
germination percentages of 75 to 94 percent were cattle enclosures to discourage human intruders
observed (Creager 1992). Seeds can remain viable and animal predators. The species is used in a
in dry soil for more than 2 years. Plants sprout number of herbal remedies and magic rites in
freely following fires (Binggeli 1997). Layering Africa (Burkill 1995).
occurs when stems become prostrate and covered
with litter. References
Growth and Management.—Black mimosa Binggeli, P. 1997. Mimosa pigra L. (Mimosaceae).

grows rapidly. Seedlings grown in a greenhouse http://members.lycos.co.uk/WoodyPlantEcology
reached 25.8 ± 0.5 cm in 30 days, 97.1 + 16.2 cm /docs/web-sp11.htm. 5 p.
in 90 days, and 335.1 ± 15.9 cm in 360 days
(Swarbrick and Mercado 1987). Plants in Puerto Burkill, H.M. 1995. The useful plants of West
Rico in tall, dense grass swards and disturbed Tropical Africa. Vol. 3. Royal Botanic Gardens,
upland sites grow about 1.0 to 1.5 m the first year, Kew, UK. 857 p.
1 m in the second year, and usually live 2 to 3
years. Larger plants tend to lie down due to weight CSIRO. 2002. Mimosa under attack. Global Net
and often do not become taller than 3 m (author’s News.com. http://ausissues.com/ausissues/
observation). Maximum age is about 5 years AINews.nsf/0/DE44CB72D21850A6CA256C10
(Binggeli 1997). In an effort to control the 00169466?OpenDocument. 4 p.
aggressively spreading infestation, nine insect
enemies and two pathogenic fungi have been Creager, R.A. 1992. Seed germination, physical
released in Australia. Most have become and chemical control of catclaw mimosa
established but so far have had little effect on (Mimosa pigra var. pigra) Weed Technology
black mimosa populations (Marko 2002). 6(4): 884-891.
However, a recent release of Malacorhinus
irregularis, a defoliater beetle from Mexico shows Cross, B.T., and H.T. Wiedemann. 1997. Control
early promise (CSIRO 2002). Mowing is of catclaw acacia and mimosa by grubbing.
ineffective (Creager 1992). Grubbing of adult Applied engineering in Agriculture 13(2): 291-
plants achieved only about 82 percent mortality 293.
(Cross and Wiedemann 1997). Until effective
biological control can be achieved, chemical Department of Natural Resources and Mines.
control is probably the best approach for spot 2002. NR & M acts on Mimosa pigra outbreak
infestations (Marko 2002). at Peter Faust Dam, Proserpine. Department of
Natural Resources and Mines, Queensland,
Benefits and Detriments.—Black mimosa Australia. http://www.nrm.qld.gov.au/about/
invades dense grass swards and in some media/apr/mimosa.html. 4 p.
490
Howard R.A. 1988. Flora of the Lesser Antilles, http://www.hort.agri.umn.edu/h5015/99papers/
Leeward and Windward Islands. marko.htm. 10 p.
Arboretum, Harvard University. Jamaica Natural Resources Conservation Service. 2002.
Plain, MA. 673 p. Plants profile: Mimosa pigra L. black mimosa.
Janzen, D.H. 1983. Mimosa pigra (zarza, symbol=MIPI. [not paged].
dormilona). In: D.H. Janzen, ed. Costa Rican
natural history. University of Chicago Press, Pacific Island Ecosystems at Risk. 2002. Mimosa
Chicago and London. p. 277-278. pigra L., Fabaceae. http;//www.hear.
org/pier_v3.3/mipig.htm. 3 p.
Lebas, F., P. Coudert, R. Rouvier, and H. de
Rochambeau. 1986. The rabbit: husbandry, Stevens, W.D., C. Ulloa-U., A. Pool, O.M.
health and production. Food and Agriculture Montiel, eds. 2001. Flora de Nicaragua.
Organization of the United Nations, Rome. Monographs in Systematic Botany Vol. 85, No.
http://fao.org/docrep/x5082e/X5082E06.htm. 2. Missouri Botanical Garden Press, St. Louis,
[not paged]. MO. p. 945-1,910.
Marko, M. 2002. Controlling invasion of the Swarbrick, J.T. and B.L. Mercado. 1987. Weed
exotic shrub (Mimosa pigra) in tropical science and weed control in Southeast Asia.
Australia wetlands. University of Minnesota FAO Plant Production and Protection Paper 81.
Department of Horticultural Science. Food and Agriculture Organization of the United
Nations, Rome. 203 p.
491
Mimosa pudica L. sensitive plant
FABACEAE
Synonyms: Mimosa tetrandra Humb. & Bonpl. ex Willd.

Mimosa pudica L. var. tetrandra (Willd.) DC.
Mimosa unijuga Duch. & Walp.
Mimosa pudica L. var. unijuga (Duch. & Walp.) Griseb.
John K. Francis
defenses against herbivorous insects, leaching loss

of nutrients, or desiccation.
Range.—Sensitive plant was first described from

Brazil (Pacific Island Ecosystems at Risk 2001)
and is perhaps native to much or all of the New
World Tropics (Liogier 1988). Today, it is
pantropical in its distribution (Howard 1988).
Ecology.—Sensitive plant grows on most well-

drained soils, even scalped or eroded subsoils and
soils with low nutrient concentrations. It requires
disturbed soils to establish itself. Repeated
burning may encourage its spread in pastures
(Siregar and others 1990). Sensitive plant is shade
intolerant and does not compete with tall
General Description.—Sensitive plant is a small, vegetation or grow under forest canopies. The
prostrate or ascending, short-lived shrub. Some species’ roots produce carbon disulfide, which
authors consider it a woody herb. It may reach 1 m selectively inhibits colonization of the rhizosphere
in height when supported on other vegetation and by mycorrhizal and pathogenic fungi (Feng and
more than 2 m in horizontal extension. The others 1998). This plant occurs in croplands,
reddish-brown, woody stems are sparsely or orchards, pastures, mowed areas, roadsides, and
densely armed with curved prickles. The root areas disturbed by construction. It may grow as a
system consists of a taproot and extensive fibrous single plant or in tangled thickets. Sensitive plant
roots with nodules. The twigs are fine and flexible grows from near sea level up to 1,300 m in
and support leaves with one or two pairs of pinnae elevation (Holm and others 1977) and in areas
and 15 to 25 pairs of oblong leaflets 3 to 12 mm with annual precipitations from about 1000 to over
long. The flowers are pink and clustered in 2000 mm. The species is frost-sensitive.
globose heads. The legume (pod) is linear-oblong,
1.0 to 1.5 cm long and 3 mm broad, with bristles Reproduction.—In the Philippines, sensitive
on the margins. The pods are born in groups and plant flowers all year and may produce as many as
contain two to four brown seeds (Howard 1988, 675 seeds per plant per year (Holm and others
Liogier 1988, Pacific Island Ecosystems at Risk 1977). The species is both wind (Chieng and
2001). Sensitive plant is also known as dorme Huang 1998) and bee-pollinated (Payawal and
dorme, dormidera, humble plant, marie-honte, others 1991). Air-dry seeds from Puerto Rico
mayhont, morivivi, honteuse, sleeping grass, ti weighed an average of 0.0065 ± 0.0002 g/seed.
mawi, touch-me-not, and many other names With no pretreatment, seeds from this collection
(Holm and others 1977, Howard 1988, Liogier began germinating 7 days after sowing and
1988). The great curiosity of sensitive plant and reached a maximum germination of 17 percent by
the source of most of its names is that when 94 days (author’s observation). In another test, 80
touched, it quickly folds its leaflets and pinnae and percent germination was obtained in 4 weeks with
droops downward at the petiole attachment. The alternating temperatures of 20 and 40 °C (Holm
leaves also droop at night, and when exposed to and others 1977). Bui (2001) recommends a
rain or excessive heat. This response may be pretreatment with hot water followed by overnight
492
soaking. Germination is epigeal. Seeds are Limited, London. 358 p.
transported by means of the bristles on the edges
of their pods that cling to clothing or to the fur of Bui, L-D. 2001. Mimosa pudica. http://bio.
mammals. Most nursery and home propagation is maimi.edu/mimosa/mimosa.html. 7 p.
done using seeds, but summer cuttings may also
be used (Bui 2001). Chieng, H-T. and T.C. Huang. 1998. Aeropollen
of the Pingtung area, South Taiwan. Taiwania
Growth and Management.—In Puerto Rico, 43(2): 73-100.
sensitive plants live 1 to 2 years. Seedlings grow
slowly for 2 or 3 months and then accelerate, Coimbra, A.F. and A. Magnanini. 1953.
reaching 0.5 to 2 m of extension at the end of the Considerations sobre Mimosa pudica no
first year. Growth of plants that survive into the combate a arosao superficial. Anucio Brazilero
second year is much slower. Potted and field- Economico da Floresta 6. Instituto Nacional,
grown individuals are sensitive to overwatering Pinho, Brazil. 131-136.
(Bui 2001). This species has been successfully
tested and recommended for erosion control Feng, Z., P.G. Hartel, R.W. Roncadori, S.J.S.
plantings using potted material at a spacing of 60 Sung, and J.E. Box. 1998. Inhibition of fungal
x 60 cm (Coimbra and Magnanini 1953). colonization on the rhizoplane of the CS2-
production plant, Mimosa pudica L. In: Plant
Benefits and Detriments.—Sensitive plant has and Soil Sciences 82. Kluwer Academic
become a serious weed in fields of corn, soybeans, Publishers, Dordrecht, Netherlands. 115-126.
tomatoes, upland rice, cotton, bananas, sugarcane,
coffee, oil palms, papayas, coconuts, and rubber in Guzmán, D.J. 1975. Especies utiles de la flora
many tropical areas. It is particularly troublesome salvadoreña. Ministerio de Educación, Dirección
where hand pulling of weeds is practiced. The de Publicaciones. San Salvador, El Salvador.
species may be controlled by a number of 703 p.
commercial broad-leaf herbicides (Bui 2001). On
the other hand, it is tolerated or valued as a forage Holm, L.G., D.L. Plucknett, J.V. Paucho, and J.P.
plant in pastures (Holm and others 1977, Turbet Herberger. 1977. The world’s worst weeds.
and Thuraisingham 1948). In fact, sheep grazing East-West Center, University of Hawaii,
is reported to control sensitive plant in pastures Honolulu, HI. 609 p.
and plantations (Simonnet 1990). The root nodules
have been shown to fix nitrogen (Pokhriyal and Howard R.A. 1988. Flora of the Lesser Antilles,
others 1990). Thickets of sensitive plant may be a Leeward and Windward Islands.
fire hazard when dry (Pacific Island Ecosystems at Dicotyledoneae, Part 1. Vol. 4. Jamaica Plain,
Risk 2001). The seeds and other plant parts of MA: Arnold Arboretum, Harvard University.
sensitive plant contain mimonsine, an amino acid 673 p.
that is known to cause hair loss and depressed
growth in mammals (Arora 1983). An unlikely Liogier H.A. 1988. Descriptive flora of Puerto
large dose is necessary to cause problems, Rico and adjacent islands, Spermatophyta. Vol.
however. The pollen is important to honeybees in 2. Editorial de la Universidad de Puerto Rico,
the Philippines (Payawal and others 1991). Río Piedras, PR. 481 p.
Extracts of the plant have been shown in scientific
trials to be a moderate diuretic, depress duodenal Martínez R., G., F.F. Rodríguez L., C.M.
contractions similar to atropine sulphone, promote Contreras, and M. Molina H. 1996. Estudio
regeneration of nerves, and reduce menorrhagia preliminary de las possibles acciones
(Modern-natural 2001). Anitdepressant activity antidepresivas de Mimosa pudica L. In:
has been demonstrated in humans (Martínez and Resumen de ponencias del Primer Congreso
others 1996). Root extracts are reported to be a Nacional de Plantas Medicinales de Mexico.
strong emetic (Guzmán 1975). p. 69. July 24-30, 1996. Tlaxcala, Tlaxcala,
Mexico.
References
Modern-natural. 2001. Mimosa pudica Linn.
Arora, S.K., ed. 1983. Chemistry and biochemistry http://modern-natural.com/mimosa_pudica.htm.
of legumes. Edward Arnold (Publishers) 4 p.
493
Pacific Island Ecosystems at Risk. 2001. Invasive Simonnet, P. 1990. Sheep flock management in a
plant species: Mimosa pudica Mart. ex Colla, tropical environment under coconut.
Fabaceae. http://www.hear.org/pier/mipud.htm. Oleagineux Paris 45(10): 451-456.
2 p.
Siregar, M.E., B. Haryanto, and S. Tjitrosemito.
Payawal, P.C., A.C. Tilde, and A.L. Manimtim. 1990. A review of weed management in
1991. Year round pollen sources of Italian honey Indonesian pastures. BIOTROP Special
bees (Apis mellifera L.) in the Philippines. III. Publication 38. A symposium on weed
Selected areas. Philippine Agriculturist 74(4): mangement. 7-9 June 1989. Bogor, Indonesia. p.
503-509. 229-235.
Pokhriyal, T.C., H.C.S. Bhandari, D.S. Negi, S.P. Turbet, C.R. and K. Thuraisingham. 1948. Feeding
Chaukiyal, and B.B. Gupta. 1990. Identification trials with the sensitive plant Mimosa pudica.
of some fast growing leguminous tree species Tropical Agriculturist, Ceylon 104(2): 81-86.
for nitrogen fixation studies. Indian Forester
116(6): 504-507.
494
Mimulus aurantiacus W. Curtis bush monkeyflower
PHRYMACEAE (formally in SCROPHULARIACEAE)
Synonyms: Diplacus aurantiacus Jeps.

Mimulus glutinosus Wendl.
Diplacus latifolius
Diplacus leptanthus Nutt.
Arlee M. Montalvo and Paul A. McMillan
shortened shoots in the axils of larger leaves. The

lighter underside of leaves and the 2 to 2.5 cm
long green, pleated calyx are covered with sticky
glandular and nonglandular hairs. Flowers have a
long floral tube, spreading limbs, and a large, two-
lobed stigma. Plants are often sticky with
glandular secretions.
Range and Taxonomy.—Recent phylogenetic

work favors placement of woody monkeyflowers
(section Diplacus) in Mimulus and transfers the
genus from family Scropulariaceae to the
Phrymaceae (Beardsley and Olmstead 2002). Bush
monkeyflower (broad sense) ranges from the coast
to coastal ranges to interior foothills from
northwestern Baja California, northward to
General Description.—Bush monkeyflower, or Mendocino County, California below 1,600 m and
sticky monkeyflower, is a beautiful, erect to along the western foothills of the Sierra Nevada
sprawling subshrub, that is named for its woody range. The underlying topography of coastal hills,
habit and typical “monkey-faced” flowers. Bush interior valleys, and foothills is complex, as are the
monkeyflower represents a complex assemblage of west to east and north to south gradients in
hybridizing forms with intergrading floral colors temperature and rainfall. The correspondingly
and morphologies. For this reason, the most recent complex pattern of variation in bush
taxonomic treatment lumps together all the species monkeyflower may reflect once isolated and
of woody monkeyflowers from California, differentiated populations that have come into
excepting M. clevelandii Brandegee, under M. secondary contact and hybridized. The resulting
aurantiacus (Thompson 1993). The various taxa, patterns of genetic diversity do not match the
sensu Munz and Keck (1968), are distinguished by patterns of morphological diversity (Beardsley and
stature, flower size and color, and range, but share others 2003). The taxonomy is necessarily
similar leaf morphology. Given the taxonomic complex and may be resolved with detailed
uncertainty, we describe M. aurantiacus sensu phylogeographic analysis. See Munz and Keck
Munz and Keck. Then under “Range and (1968) and Tulig (2000) for details of the
Taxonomy,” we provide a brief description of geographic distribution of the following color
diagnostic differences and distributions of their forms (narrow sense).
other taxa that were treated as a single species The small, orange-flowered M. aurantiacus
complex by Thompson (1993). occurs in the foothills and coastal ranges north of
Shrubs are erect, 6 to 12 dm tall, and the Santa Barbara County. M. a. ssp. australis
specific epithet “aurantiacus” refers to the often (McMinn) Munz has orange-yellow to buff or
deep orange corollas (sometimes yellowish) that white corollas and occurs in interior San Diego
are 3.5 to 4.5 cm long. The simple, opposite, County to Baja California. M. aridus (Abrams)
leaves are elliptic to narrowly-elliptic, with entire A.L. Grant, the lowbush monkeyflower, has a
to serrulate margins that are often rolled under. yellow corolla, low, spreading habit to 4 dm tall,
Leaves are 2 to 8 cm long and dark green, and and occurs farther east in San Diego County to
resinous above. Smaller leaves are clustered on Baja Calfornia. M. bifidus Pennel has a larger
495
orange corolla with larger, deeply-notched lobes, a hybrid populations that were intermediate between
somewhat spreading habit to 1 m tall, and occurs parental forms. For example, in San Diego County,
from coastal San Luis Obispo to Monterey County putative hybrid populations were intermediate
and along the western base of the Sierra Nevada between M. puniceus (coastal form) and M.
from Placer into Butte and Plumas Counties. M. aurantiacus ssp. australis (inland form), consistent
flemingii Munz has red corollas with limbs about with analysis of Waayers (1996). In the region
half as long as other forms, is low growing from 1 between the San Gabriel and San Bernardino
to 6 dm tall, and is restricted to the Channel Mountains, putative hybrids were intermediate
Islands. M. longiflorus (Nutt.) A. L. Grant, the between M. longiflorus ssp. calycinus (lemon
southern bush monkeyflower (see figure), has an yellow corolla) and M. longiflorus (salmon
orange-yellow to salmon corolla, a long and corolla).
pubescent calyx (to 3.5 cm), is 3 to 12 dm tall, and
occurs from central San Diego County through Ecology.—Bush monkeyflowers occur in areas
Riverside to San Bernardino County. A shorter, with cool moist winters and warm to hot, dry
pale-yellow form [M. l. ssp. calycinus (Eastw.) summers. They grow on rocky hillsides and cliffs,
Munz] occurs primarily farther inland in Riverside usually on the border of chaparral or sage scrub, or
and western San Bernardino Counties. A red- in open foothill-woodland forest. The roots are
corolla form (M. l. var. rutilus Grant) occurs fibrous, less than 2 m deep, with feeder roots
mostly in interior Los Angeles County. Finally, M. concentrated within the first 8 cm of soil (Hellmers
puniceus (Nutt.) Steudel, the red bush and others 1955). There is evidence for
monkeyflower, has a red corolla, short calyx (to differentiation and local adaptation in
2.5 cm), and erect habit from 5 to 15 dm tall. It morphological and physiological characters
occurs on Catalina Island, coastal Orange County, relating to water use. For example, differences in
and inland through the Santa Ana Mountains to wood anatomy among forms and among
southwestern Riverside County south into Baja populations are correlated with habitat (Michener
California. 1983). Plants from mesic habitats have relatively
few, large vessel-elements, while plants from xeric
Hybridization.—All bush monkeyflowers have n habitats have many, small vessel-elements. Under
= 10 chromosomes and crosses among them humid, low-stress conditions, both mesic-adapted
produce viable progeny. Natural, putative hybrids and xeric-adapted populations had high water-use
between forms have been hypothesized based on efficiency and maintained a high rate of
morphology (Munz and Keck 1968, Wells 1980, photosynthesis relative to transpiration (Mooney
Grant 1993, Tulig 2000). Preliminary phylogenetic and Chu 1983). Plants from xeric populations had
work suggests widespread introgression among high water-use efficiency under low relative
forms (personal communication with Paul humidity, while the efficiency of coastal plants
Beardsley, Colorado College, CO). Grant (1993) diminished rapidly with decreasing relative
examined pollinators in relation to several floral humidity.
forms (equals species of Diplacus in his paper) and The primary herbivore of bush monkeyflower
postulated that the populations with different floral is a specialist butterfly Euphydryas chalcedona
forms would be partially reproductively isolated. Doubleday and Hewitson. The timing of growth of
He hypothesized that the red-colored D. puniceus both plants and the butterfly is linked to water
Nutt. and salmon D. longiflorus Nutt. were availability. Emergence of larvae from diapause is
primarily hummingbird-pollinated, and that D. synchronized with the initiation of leaf growth.
calycinus Eastw. (M. l. var. calycinus) with pale The leaves produce a resin that inhibits the growth
yellow flowers and long tube) were hawkmoth- of the larvae. During the growing season the
pollinated. He found that habitats interdigitated youngest leaves have the highest nitrogen (N)
and resulted in D. puniceus and D. calycinus content, the highest carbon gain, and the highest
populations overlapping in distribution, with D. resin content. Growth rate of the larvae increases
longiflorus habitat forming a connecting link with increasing N but decreases with increasing
between the different forms. He hypothesized that resin content (Lincoln and others 1982). Larvae
the intermediate nature of D. longiflorus suggested feed initially on young leaves with high nutritional
past hybridization of D. puniceus and D. calycinus. value, then switch to older leaves with lower resin
Tulig (2000) analyzed floral and vegetative content. During flowering, N is translocated from
variation throughout the geographic range of bush the leaves, and larvae stop feeding (Mooney and
monkeyflowers and found statistical evidence for others 1981).
496
Such studies demonstrate physiological longiflorus was “generally but not exclusively”
trade-offs in resource allocation to growth, pollinated by Black-chinned and Anna's
reproduction, or defense. Han and Lincoln (1994) hummingbirds. Six populations from the Santa
found significant heritability and maternal effects Monica Mountains varied in floral form, sucrose
for resin content within one population, and they content of nectar, and anther-stigma separation.
found negative genetic and phenotypic correlations Populations with high pollinator diversity but low
between resin production and growth rate. This visitation rates had shorter corolla tubes and lower
indicates potential trade-offs between traits during sucrose:hexose ratios in nectar than populations
selection. Leaf resin content varies among with less diverse pollinators. In a different
populations, but little is known about geographic population of M. longiflorus, Eckert (1970) found
differentiation in such trade-offs. that almost all pollination was done by solitary
Hare (2002) found that nearly 30 percent of bees and that Anna's hummingbirds were only
the dry weight of leaves is resin. In a common occasional visitors. Most pollination was effected
garden experiment, he examined the chemistry and by bees in the genus Osmia and carpenter bees in
relative quantities of important resin components Ceratina. Both visited multiple flowers per plant,
for six populations of bush monkeyflower from a and self-pollination appeared common. Eckert also
range of environments and that differed in attack found that selfed flowers produced fewer capsules
by E. chalcedona. The populations differed in per plant and had more aborted ovules than
chemical components of the resins but not in the outcrossed flowers.
pattern of insect attack. This study followed
Thompson’s (1993) classification and did not Horticulture.—The bush monkeyflowers and
mention if there were differences in floral their horticultural hybrids have striking flower
morphology among the populations. The collection colors, produce masses of flowers, and are popular
locations could have represented at least three ornamental plants (Schmidt 1980, Perry 1992,
species or subspecific taxa following Munz and Keator 1994). Plants can be grown from seeds or
Keck (1969) or Tulig (2000). cuttings, but plants grown from seed tend to live
longer (personal communication with Steve
Emergence and Growth.—Seedlings emerge in Morgan, University of California Riverside
the cool rainy season, and most growth is in winter Botanic Garden, CA). The shrubs grow best in full
and spring. Plants can reach reproductive maturity sun to light shade. Flowering and leaf production
in a single growing season, with flowering may be extended by summer watering in more arid
primarily from mid spring to early summer. The regions. However, Atkinson and others (1988)
capsules mature in summer, and the many, tiny found that watering may increase susceptibility to
seeds (about 1 mm long, half as wide and nearly atmospheric pollution in the dry season. In
flat) disperse by gravity and wind. Plants become summer, unwatered plants drop most leaves, are
dormant in the summer dry season and shed many not photosynthetically active, and take up little
leaves by fall. Leaf production begins after the SO2, while watered plants retain their leaves, are
first fall rain and ends near the beginning of July photosynthetically active, and take up SO2.
(Mooney and others 1981). A large and relatively
constant proportion of carbon and nitrogen are Growth and Management.—Bush
allocated to reproduction, and reproductive monkeyflowers have been planted along highways
structures supply some of their own carbon (Alpert corridors and are used in habitat restoration. There
and others 1985). Both growth and reproduction are about 145 million seeds/kg (Mirov and Kraebel
are primarily water limited. 1939). Seeds will germinate without pretreatment
(Mirov and Kraebel 1939, Schmidt 1980), but
Reproduction.—Bush monkeyflowers are seeds need to be exposed to light and leachate
pollinated by diverse flower visitors, and the from charred wood may improve germination
dominant pollinators vary with color form and slightly (Keeley 1987). For pot culture, the tiny
location. This may reflect local differentiation, or seeds should be sown on the surface of the soil and
the relative abundances of pollinators. Fetscher kept moist until seedlings emerge. Outdoors, they
and Kohn (1999) reported Anna’s hummingbird as should be planted with shallow broadcasting
the primary pollinator of a red flower form in San methods. The phylogenetic relationships among
Diego County, California, with black-chinned, populations and the fitness effects of translocations
Costa’s, and rufous hummingbirds as occasional and hybridization are not yet known, and there are
visitors. Bromer and others (1990) reported that M. no published studies on population genetic
497
patterns. However, there is a complex pattern of Fetscher, A.E. and J.R. Kohn. 1999. Stigma
geographic variation, genetic differentiation of behavior in Mimulus aurantiacus
floral form and resin chemistry, local adaptation to (Scrophulariaceae). American Journal of
moisture environments, and extensive ability to Botany 86: 1,130-1,135.
hybridize. Consequently, attention should be made
of flower color and habitat matching when Grant, V. 1993. Origin of floral isolation between
obtaining seeds for large planting projects. Use of ornithophilous and sphingophilous plant species.
regionally local seed sources for planting projects Proceedings of the National Academy of
can mitigate unknown fitness consequences of out Sciences of the United States of America 90:
of range plantings. A careful analysis of hybrid 7729-7733.
zones and the fitness of hybrids is needed to
understand if hybridization following Han, K. and D.E. Lincoln. 1994. The evolution of
translocations will cause decreases in fitness of carbon allocation to plant secondary
adjacent wild populations. metabolites: a genetic analysis of cost in
Diplacus aurantiacus. Evolution 48: 1,550-
Benefits.—Bush monkeyflowers are important 1,563.
plants for water-wise landscaping and revegetation
projects. They provide nectar to hummingbirds Hare, J.D. 2002. Geographic and genetic variation
and large bees and are an important larval host of in the leaf surface resin components of Mimulus
the checkerspot butterfly. aurantiacus from southern California.
Biochemical Systematics and Ecology 30: 281-
References 296.
Alpert, P., E.A. Newell, C. Chu, J. Glyphis, S.L. Hellmers, H., J.S. Horton, G. Juhren, and J.
Gulmon, D.Y. Hollinger, N.D. Johnson, H.A. O'Keefe. 1955. Root systems of some chaparral
Mooney, and G. Puttick. 1985. Allocation to plants in southern California. Ecology 36: 667-
reproduction in the chaparral shrub, Diplacus 678.
aurantiacus. Oecologia 66: 309-316.
Keator, G. 1994. Complete Garden Guide to the
Atkinson, C.J., W.E. Winner, and H.A. Mooney. Native Shrubs of California. Chronicle Books,
1988. Gas exchange and SO2 fumigation studies San Francisco, CA. 314 p.
with irrigated and unirrigated field grown
Diplacus aurantiacus and Heteromeles Keeley, J.E. 1987. Role of fire in seed germination
arbutifolia. Oecologia 75: 386-393. of woody taxa in California chaparral. Ecology
68: 434-443.
Beardsley, P.M. and R.G. Olmstead. 2002.
Redefining Phrymaceae: the placement of Lincoln, D.E., T.S. Newton, P.R. Ehrlich, and K.S.
Mimulus, tribe Mimuleae, and Phryma. Williams. 1982. Coevolution of the checkerspot
American Journal of Botany 89: 1093-1102. butterfly Euphydryas chalcedona and its larval
food plant Diplacus aurantiacus: larval response
Beardsley, P.M., S.E. Schoenig, J.B. Whittall, and to protein and leaf resin. Oecologia 52: 216-223.
R.G. Olmstead. 2003. The radiation of Mimulus
(Phrymaceae) in western North America. Michener, D.C. 1983. Systematic and ecological
American Journal of Botany. In Press. wood anatomy of Californian Scrophulariaceae.
I. Antirrhinum, Castilleja, Galvezia, and
Bromer, W., J. Barnette, J. Lee, D. Green, and V. Mimulus sect. Diplacus. Aliso 10: 471-487.
Ervin. 1990. Genetic variation within and among
populations of Mimulus longiflorus in the Santa Mirov, N.T. and C.J. Kraebel. 1939. Collecting
Monica Mountains, CA: consequences of and handling seeds of wild plants. Civilian
breeding system and pollination. Bulletin of the Conservation Corps, Forestry Publication 5.
Ecological Society of America 71: 103. United States Government Printing Office,
Washington, D.C.
Eckert, J.R. 1970. Pollination studies in Mimulus
longiflorus (Nutt.) Grant. Master’s thesis. Mooney, H.A., and C. Chu. 1983. Stomatal
California State University, Los Angeles. 88 p. responses to humidity of coastal and interior
498
populations of a Californian shrub. Oecologia Thompson, D.M. 1993. Mimulus. In: J.C.
57: 148-150. Hickman, ed. The Jepson Manual: Higher Plants
of California. University of California Press,
Mooney, H.A., K.S. Williams, D.E. Lincoln, and Ltd., Los Angeles, CA. p. 1,037-1,046.
P.R. Ehrlich. 1981. Temporal and spatial
variability in the interaction between the Tulig, M. 2000. Morphological variation in
checkerspot butterfly, Euphydryas chalcedona Mimulus section Diplacus (Scrophulariaceae).
and its principal food source, the Californian Masters Thesis, Department of Biological
shrub, Diplacus aurantiacus. Oecologia 50: 195- Sciences, California State Polytechnic
198. University, Pomona, CA. 82 p.
Munz, P.A. and D.D. Keck. 1968. A California Waayers. 1996. Hybridization, introgression, and
Flora with Supplement. University of California selection in Mimulus aurantiacus ssp. australis
Press, Berkeley, CA. 1,681 + 224 p. and M. puniceus. Master’s thesis. San Diego
State University, San Diego, CA. 77 p.
Perry, B. 1992. Landscape Plants for Western
Regions: an Illustrated Guide to Plants for Water Wells, H. 1980. A distance coefficient as a
Conservation. Land Design Publishing, hybridization index: an example using Mimulus
Claremont, CA. 318 p. longiflorus and M. flemingii (Scrophulariaceae)
from Santa Cruz Island, California. Taxon 29:
Schmidt, M.G. 1980. Growing California Native 53-65.
Plants. University of California Press, Los
Angeles.
499
Mitracarpus portoricensis (Urban) Urban Puerto Rico girdlepod
RUBIACEAE
Synonyms: Mitracarpus frigidus portoricensis Urban
John K. Francis
and Martorell 1982). The species is not known to

have been planted or naturalized anywhere else.
Ecology.—Puero Rico girdlepod is a low-statured,

slow-growing plant that requires full or nearly full
sunlight. It can compete with low herbs and
grasses but cannot grow under a forest or shrub
canopy. Consequently it must grow in areas with
low competition, usually disturbed areas and areas
of low fertility. It finds these conditions in eroded
areas, dirt roads and road cuts, old fields, rocky
hillsides, and coastal sands. Puerto Rico girdlepod
may be found in areas of sedimentary (including
limestone), igneous, and metamorphic (including
ultramaphic) rocks. It tolerates salt spray, low soil
fertility, compacted soils, and moderate physical
abuse. Elevations range from a little above sea
level to about 400 m in areas with rainfall from
about 900 to about 1800 mm of mean annual
precipitation.
Reproduction.—Puerto Rico girdlepod blooms

and produces seed throughout the year. A
collection of seeds from the Susúa State Forest
contained an average of 2,400 seeds/g. Placed on
General Description.—Puerto Rico girdlepod (a moist blotter paper, 29 percent of the seeds
name assigned by the Natural Resources germinated between 4 and 90 days after sowing
Conservation Service) is a low, 20- to 60-cm tall (author’s observation). Seeds are dispersed by
evergreen shrub with basal diameters of 0.5 to 1.0 wind and water. Opening of the forest canopy and
cm. It may have one or more gray stems from the probably bare soil are necessary for establishment.
root crown and a compact, rounded crown with Seedlings are not common.
bifurcate or trifurcate branching. The stems,
branches, and roots are woody but flexible. Plants Growth and Management.—Growth of Puerto
are supported by a tap and lateral root system of Rico girdlepod is slow. Plants observed by the
yellow or tan roots. The glabrous branches and author appeared to have grown about 10 to 25
twigs are four-striate and four-angled. The leaves, cm/year. Life span of the shrubs appears to range
which are crowded on recent twig growth, are from about 2 to 6 years. No planting or
sessile, nearly linear, 30 to 60 mm long and 1 to 3 management experience has been reported. The
mm broad. The 2-cm broad, many-flowered heads species is not common enough or aggressive
are lateral or terminal on long peduncles. Tubular enough to warrant control.
flowers are bright white, about 5 mm long.
Capsules disperse their seeds when the tops come Benefits.—Puerto Rico girdlepod helps protect the
off releasing 1-mm long, ovate, brown seeds soil, revegetates disturbed sites, adds to the
(Liogier 1997). aesthetics of wildland areas (because of its white
flowers), and furnishes cover for wildlife.
Range.—Puerto Rico girdlepod is endemic to
Puerto Rico, particularly to the northern, western,
and southwestern portions of the Island (Liogier
500
References Liogier, H.A. and L.F. Martorell. 1982. Flora of
Puerto Rico and adjascent islands: a systematic
Liogier, H.A. 1997. Descriptive flora of Puerto synopsis. Editorial de la Universidad de Puerto
Rico and adjacent islands. Vol. 5. Editorial de la Rico, Río Piedras, PR. 342 p.
Universidad de Puerto Rico, Río Piedras, PR.
436 p.
501
Morella caroliniensis (P. Mill.) Small bayberry
MYRICACEAE
Synonyms: Myrica caroliniensis P. Mill.

Myrica curtissii A. Chev.
Myrica heterophylla Raf.
Myrica pensylvanica Mirb.
Gerry Moore
plants. The staminate catkins range from 0.4 to 1.8

cm; the pitsillate catkins range from 0.3 to 1.4 cm.
The fruit of the bayberry is technically not a berry
but a globose 3.0- to 6.0-mm drupe (one-seeded
stone fruit) that at maturity is covered by a thick
coat of gray-white wax. The fruits persist
throughout the winter. The globose seed in each
fruit measures slightly less in diameter than the
fruit (author, personal observation).
Range.—Bayberry occurs in eastern Canada (New

Brunswick, Newfoundland, Nova Scotia, Ontario,
Prince Edward Island, and Quebec) and the United
States, from Maine south to Florida and west to
Texas and Arkansas. Inland populations are known
from western New York, Ohio and southern
Ontario (Gleason and Cronquist 1991).
Systematic Botany.—There are taxonomic and

nomenclatural difficulties associated with the
bayberry. Taxonomists disagree regarding
bayberry’s generic placement, with some
Illustration credit: Britton and Brown 1913 (Bornstein 1997, Fernald 1950, Gleason and
Cronquist 1991) placing the bayberry and sweet
General Description.—Bayberry is the name used gale (Myrica gale L.) in the same genus, Myrica,
most often to refer to this common eastern shrub. and others choosing to place the bayberry in its
Other common names include candleberry, own genus, Morella (Wilbur 1994, 2002). Some
tallowshrub, and waxberry. Bayberry is a taxonomists also choose to recognize two species
deciduous (northern part of range) to evergreen of bayberry, a northern bayberry Morella
(southern portion of range) aromatic shrub or pensylvanica and a southern bayberry, Morella
small tree to 4 m. The plant is single or caroliniensis (synonym: Myrica heterophylla
multistemmed. The branchlets and twigs are terete Raf.). The two supposedly can be distinguished
and vary in color from brown-gray-blackish; they based on whether the leaves are evergreen
are dotted with yellow aromatic glands and vary (southern) or deciduous (northern), the fruit wall is
from being glabrous to pubescent. The leaves are hairy (northern) or not (southern), and the twigs
resinous, glandular, aromatic, with short pubescent possess black hairs (southern) or not (northern).
petioles (to 7 mm) and elliptic to oblanceolate, Further complicating matters, bayberry and the
membranaceous-leathery 3.0 to 7.0 cm by 1.5 to wax myrtle [Morella cerifera (L.) Small] can
2.5 cm blades with margins (sometimes slightly hybridize when growing together (Bornstein
revolute) entire or more usually toothed from the 1997). Some authors also treat the name Morella
middle to the apex. The flowers are in unisexual caroliniensis (Miller) Small as a synonym of the
catkins (aments), with the staminate (male) and wax myrtle (M. cerifera). This is due to different
pistillate (female) catkins appearing on different interpretations as to which plant–bayberry or wax
myrtle–Miller (1768) was actually referring to in
502
his original description. The recent work of Wilbur of North America. Vol. 3. Oxford University
(1994, 2002) is followed here; therefore only one Press, New York. p. 430-434.
species is recognized under the name M.
carolininesis. Britton, N.L. and A. Brown. 1913. Illustrated flora
Ecology.—Bayberry can tolerate full sun to dense possessions. Vol. 2, 2nd Ed. Scribner, New York.
shade and can grow in dry to wet conditions. 735 p.
Habitats where it can be found include bogs,
swamps, dunes, fields, heathlands, oak forests, Chittenden, F.J. and P.M. Synge. 1956. Dictionary
pine forests, and the margins of streams and lakes. of Gardening. 2nd Ed. Clarendon Press, Oxford,
The roots of bayberry contain nitrogen fixing UK. 623 p.
nodules, and this allows it to be particularly
tolerant of nitrogen-poor, acidic soils (Morris and Collins, B.S. and J.A. Quinn. 1982. Displacement
others 1974). Results are mixed as to whether of Andropogon scoparius on the New Jersey
bayberry’s nitrogen-fixing properties may assist Piedmont by the successional shrub Myrica
the growth of other plant species growing nearby pensylvanica. American Journal of Botany 69:
(Tiffney and Barrera 1979, Dudley and others 680-689.
1996). Leachate from the leaf litter of bayberry
may have alleopathic properties and affect some Dudley, J.L., B. Michener, and K. Lajtha. 1996.
other plants (Collins and Quinn 1982). The contributions of nitrogen-fixing symbioses
to coastal heathland succession. American
Reproduction.—Bayberry is wind pollinated and Midland Naturalist 135: 334-342.
blooms in the spring. The pollen is allergenic
(Lewis 1977). The seeds of bayberry are dispersed Everett, T.H. 1960. New illustrated encyclopedia
by birds that eat the fruits (McClanahan and Wolfe of gardening. Vol. 3. Greystone Press, New
1993, Place and Stiles, 1992, Ridley 1930). York. 479 p.
Growth and Management.—Bayberry is often Fernald, M.L. 1950. Gray’s manual of botany.
cultivated on dry, sandy, sterile soils where few American Book Co., New York. 1,632 p.
other species can grow (Bailey 1922, Rehder
1940). It thrives best in acid peaty soils; it can not Gleason, H.A. and A. Cronquist. 1991. Manual of
tolerate liming. Propagation can be by seed or vascular plants of the northeastern United States
layering (Chittenden and Synge 1956). and adjacent Canada. 2nd Ed. New York
Propagation can also be done with cuttings taken Botanical Garden, Bronx. 910 p.
in late summer or fall. The cuttings should be
placed in a bed of peat moss and sand (Everett Lewis, W.H. 1977. Medical botany. John Wiley &
1960). Sons, New York. 515 p.
Benefits.—Settlers used the bark of bayberry in McClanahan, T.R. and R.W. Wolfe. 1993.
dentifrice mixtures (Lewis 1977) and the fruits in Accelerating forest succession in a fragmented
candle making (Bornstein 1997). Bayberry candles landscape: the role of birds and perches.
remain popular today; it can take up to 8 kg of Conservation Biology 7: 279-288.
bayberry fruit to yield 1 kg of wax. The fruits are
also often used in decorative arrangements. Native Miller, P. 1768. The gardeners dictionary. 8th Ed.
Americans used the plant for various medicinal Printed for the author, London. 1,329 p.
purposes (Moerman 1986).
Moerman, D.E. 1986. Medicinal plants of Native
References America. 2 Vols. Technical Report 19.
University of Michigan Museum of
Bailey, L.H. 1922. Standard cyclopedia of Anthropology, Ann Arbor, MI. 910 p.
horticulture. Vol. 4. MacMillan Co., London.
660 p. Morris, M., D.E. Eveleigh, S.C. Riggs, and W.N.
Tiffney, Jr. 1974. Nitrogen fixing in the
Bornstein, A.J. 1997. Myricaceae. In: Flora of bayberry (Myrica pensylvanica) and its role in
North America Editorial Committee, eds. Flora
503
coastal succession. American Journal of Botany Tiffney, W.N., Jr. and J.F. Barrera. 1979.
61: 867-870. Comparative growth of pitch and Japanese black
pine in clumps of the N2 –fixing shrub, bayberry.
Place, A.R. and E.W. Stiles. 1992. Living off the Botanical Gazette 140 (Suppl.): S108-S109.
wax of the land: bayberries and yellow-rumped
warblers. Auk 190: 334-345. Wilbur, R.L. 1994. The Myricaceae of the United
States and Canada: genera, subgenera, and
Rehder, R. 1940. Manual of cultivated trees and series. Sida 16: 93-107.
shrubs, Macmillan Publishing Co, New York.
996 p. Wilbur, R.L. 2002. The identity and history of
Myrica caroliniensis (Myricaceae). Rhodora
Ridley, H.N. 1930. The dispersal of plants 104: 31-41.
throughout the world. L. Reeve & Co., Ashford,
Kent, UK. 744 p.
504
Morinda citrifolia L. noni
RUBIACEAE
Synonyms: none
John K. Francis
1997, Little and Wadsworth 1964, Nelson 1996).
Range.—Noni is probably native to maritime

forests of Northern Australia, the Western Pacific,
and Indian Oceans (Stevens and others 2001). It
was widely spread by native peoples beginning
2000 years ago (Smith 2002) and more recently by
Europeans so that it is naturalized in most of the
tropical and tropical coastal forests of the World
(Howard 1989) including Florida, Puerto Rico,
and the U.S. Virgin Islands (Little and Wadsworth
1964).
Ecology.—Noni is most competitive on sands and

loamy sands but will grow on soils of all other
textures if planted and protected from competition.
It also grows well on very rocky soils. Although it
grows naturally only a few meters above sea level,
noni can be cultivated on sites up to 800 m or
more above sea level (Stevens and others 2001).
The species tolerates salty soils and salt spray. It is
intermediate in shade tolerance, growing under the
canopy of forests as well as in the open. Noni
grows naturally on the edges of mangroves, in
coastal forests, and on the landward side of beach
General Description.—Noni, is also known as strand vegetation. The species is grown in
Indian mulberry, great morinda, cheezefruit, plantations, as an ornamental, and as a garden
morinda, mouse’s pineapple, yellow root, jumbie medicinal tree. A symbiotic association in the
breadfruit, hog apple, pain killer, mengkudu, native range with weaver ants (Oecophylla
nono, feyukke friudem rhubarbe caraïbe, bilimbi, smaragdina) provides the ants with food and
pomme-macaque, and pomme de singe. It is a leaves for nesting in exchange for protection from
large evergreen shrub or small tree to 6 m or more insect predators (Tan 2001).
in height and 13 cm or more in stem diameter.
Sapwood is yellow-brown and soft. The bark is Reproduction.—Noni flowers and fruits nearly
gray or brown, smoothish to slightly rough. Twigs throughout the year (Little and Wadsworth 1964).
are light green and four-angled. The opposite Seeds (air dried) in Puerto Rico weighed an
leaves are attached by stout petioles 1 to 2 cm average of 0.0259 g/seed or 38,600 seeds/kg.
long. The blades are dark green and shiny, ovate Without any pretreatment, 24 percent germinated
or elliptic, 14 to 30 cm long by 8 to 18 cm broad, in commercial potting mix beginning 70 days after
and have prominent veins. The white tubular sowing (Francis and Rodríguez 1993). In its native
flowers are grouped in globose heads at the leaf habitat, the seeds of noni are probably dispersed
axils. The five-lobbed flower tubes are about 6 by fruit bats and other mammals. The plant may
mm long. The greenish-white to pale-yellow, be propagated from both seeds and cuttings
fleshy fruits are ovoid or globose syncarps 5 to 7 (Association of Societies for Growing Australian
cm long. They have an unpleasant odor Plants 2000).
resembling cheese. They contain a number of
seeds about 4 mm long (Howard 1989, Liogier Growth and Management.—Published
505
information is lacking on growth and Cambie, R.C. and J. Ash. 1994. Fijian medicinal
management. plants. CSIRO, Canberra, Australia. 365 p.
Benefits.—Although the fruits of noni are Francis, J.K. and A. Rodríguez. 1993. Seeds of
somewhat tasteless and have an unpleasant smell, Puerto Rican trees and shrubs: second
they are eaten as famine food in Indonesia, installment. Research Note SO-374. U.S.
Australia, and the Pacific Islands. The young Department of Agriculture, Forest Service,
leaves are also eaten as a vegetable and contain 4 Southern Forest Experiment Station, New
to 6 percent protein. Noni is planted as a Orleans, LA. 5 p.
windbreak, for support of pepper vines, and shade
for coffee bushes (Tan 2001). The species is Hirazumi, A. and E. Furusawa. 1999. An
useful for coastal erosion control (Association of immunomodulatory polysaccharide-rich
Societies for Growing Australian Plants 2000), substance from the fruit juice of Morinda
and it is planted as an ornamental in some areas. citrifolia (noni) with antitumour activity.
The bark has been the source of a red dye (Little Phytotherapy Research 13(5): 380-387.
and Wadsworth 1964). Roots are used to produce
an orange dye (Nelson 1996). Harvested from both Howard, R.A. 1989. Flora of the Lesser Antilles,
plantations and from the wild, noni is one of the Leeward and Windward Islands.
most important botanical remedies and food Dicotyledoneae. Part 3. Vol. 6. Arnold
supplements traded on the international market. It Arboretum, Harvard University, Jamaica Plain,
is usually the fruit juice that is sold, fresh or dried. MA. 658 p.
Leaves, bark, and roots are also used for a great
many maladies. Pain, arthritis, diabetes, high Legal, L. and M. Plawecki. 1995. Comparative
blood pressure, skin and stomach ulcers, sensitivity of various insects to toxic compounds
depression, senility, diarrhea, arteriosclerosis, from Morinda citrifolia L. Entomological
cancer, AIDS, skin parasites, and bad breath are Problems 26(2): 155-159.
all treated (Association of Societies for Growing
Australian Plants 2000, Cambie and Ash 1994, Liogier, H.A. 1997. Descriptive flora of Puerto
Tan 2001). A number of physiologically active Rico and adjacent islands. Vol. 5. Editorial de la
chemicals, anthraquinones, alkaloids, Universidad de Puerto Rico, San Juan, PR.
asperuloside, caproic, caprylic, and ursolic acids, 436 p.
β-sitosterol, and asperuloside may account for
some of the effects (Cambie and Ash 1994). Little, E.L., Jr. and F.H. Wadsworth. 1964.
Antitumor activity expressed in enhanced survival Common trees of Puerto Rico and the Virgin
of tumor-bearing mice has been demonstrated Islands. Agriculture Handbook 249. U.S.
after treating with juice extracts (Hirazumi and Department of Agriculture, Forest Service,
Furusawa 1999). Aqueous extracts of roots were Washington, DC. 548 p.
shown to have an analgesic effect on mice without
any sign of toxicity, and a sedative effect at high Nelson, G. 1996. The shrubs and woody vines of
doses (Younos and others 1990). Octanoic acid, Florida. Pineapple Press, Sarasota, FL. 391 p.
which is present in the ripe fruits, effectively
poisons a fruit fly (Drosophila melanogaster), Smith, R.S. 2002. History and tradition of
honey bee (Apis mellifera), an ant (Lasius sp.), Morinda citrifolia. http://rsscomp.freeyellow.
and cockroaches (Periplaneta americana and com/morindacitrifoliastory.htm. 2 p.
Blattella germanica) (Legal and Plawecki 1995).
Noni is the principal larval host of the hawk moth, Stampsfiji.com. 2002. The Sphingid (hawk) moths
Macroglossum hirundo vitiensis in Fiji of Fiji stamp issue. http://www.stampsfiji.
(Stampsfiji.com 2002). com/stamps/moths/ 4 p.
References Stevens, W.D., C. Ulloa U., A. Pool, and O. M.

Association of Societies for Growing Australian Monographs in Systematic Botany Vol. 85, No.
Plants. 2000. Morinda citrifolia. 3. Missouri Botanic Garden, St. Louis, MO.
http:/farrer.csu.edu/ASGAP/m-cit.html. 2 p. p. 1,911-2,664.
506
Tan, R. 2001. Mangrove and wetland wildlife at Younos, C., A. Rolland, J. Fleurentin, M.C.
Sungei Buloh Nature Park: Great morinda. Lanhers, R. Misslin, and F. Mortier. 1990.
http://www.naturia.per.sg/buloh/plants/morinda. Analgesic and behavioral effects of Morinda
htm. 3 p. citrifolia. Planta Medica 56(5): 430-434.
507
Murraya exotica L. orange jasmine
RUTACEAE
Synonyms: Murraya paniculata (L.) Jack

Chalcas paniculata L.
Chalcas exotica (L.) Millsp.
John K. Francis
side depending on whether there are one or two

seeds per fruit.
Range.—The native range of orange jasmine

includes China, India, Sri Lanka, the Andaman
Islands, Myanmar, Thailand, Kampuchea, Viet
Nam, Malaysia, northeastern Australia, New
Caledonia, and Taiwan (Parrotta 2001). In
addition, the shrub has been planted throughout
the Tropics and has naturalized in many locations
including Puerto Rico (author’s observation).
Some authors consider M. exotica and M.
paniculata to be separate species (Howard 1988,
Pacific Island Ecosystems at Risk 2002).
Ecology.—Orange jasmine is adapted to a wide

range of conditions. Naturalized plants in Puerto
Rico grow in areas receiving from about 750 mm
to 1900 mm of annual precipitation. The species
grows from nearly sea level to elevations of 1,300
m (Neal 1965). It grows on most well-drained
soils derived from both sedimentary and igneous
General Description.—Orange jasmine is also rocks, although it is said to favor limestone areas
known as mock orange, satin wood, honey bush, (Pacific Island Ecosystems at Risk 2002). Plants
China-box, café de la India, mirto, azahar, naranjo survive temperatures to about –4 °C (Desert-
jazmín, limonaria, and bun (Little and others Tropicals 2002). Ornamental plants are attacked at
1974). It is an evergreen shrub or occasionally a times by white flies, scale insects, nematodes, and
small tree, usually 2 to 3 m in height but reaching sooty mold (Sheehan 1975). It was noted that
7.5 m and 13 cm in stem diameter. Older orange powdery mildew attacked the species in pruned
jasmine normally have multiple stems from the hedges but not in free-growing unpruned plants
ground level. The stems are supported by taproots (Pathak and others 1992). Orange jasmine is
with lateral roots and abundant fine roots. Stem moderately intolerant of shade. Although growing
bark is gray, becoming fissured and rough. Orange well under partial shade, it produces few flowers
jasmine branches and twigs are slender and or fruits. Escaped plants in Puerto Rico are most
abundant at all heights. The alternate leaves are often seen at the edges and in the understory of
pinnately compound with three to nine leaflets dry and moist secondary forests. Seedlings do not
alternating on the rachis. The 1- to 5-cm, leaflets compete well with grass or herbaceous vegetation.
are dark-green, stiff, ovate, and smell of citrus
when crushed. The shrub produces fragrant, five- Reproduction.—Orange jasmine flowers
petaled, white flowers borne in small clusters near irregularly throughout the year (Little and others
the branch ends. Later, shiny, red elliptic fruits 1974). In Hawaii, it flowers from June to
about 1 cm long develop. One or two light green September (Neal 1965). Fruits collected from
seeds are embedded in the bitter, watery pulp plants in Puerto Rico averaged 0.370 + 0.012
(Liogier 1988, Little and others 1974). The seeds g/fruit. Air-dried seeds from these fruits averaged
are tear-drop shaped, rounded or flattened on one 0.0568 + 0.0012 g/seed or 17,600 seeds/kg. Sown
508
on commercial potting mix, 65 percent germinated http://www.desert-tropicals.com/Plants/
between 25 and 60 days of sowing. Germination is Rutaceae/Murraya_paniculata.html. 2 p.
hypogenous. Seedlings quickly develop deep root
systems and grow at a moderate rate. Plants Gonzalez, G. 2002. Bonsai gallery photos of
coppice vigorously after disturbance. Nursery Murraya paniculata bonsi. Tropical
production is by seed, cuttings (Little and others Bonsai.com. http://www.Tropicalbonzai.com/
1974), and air-layers (Woman’s Club of Havana MurrayaPaniculata.htm. 1 p.
1952).
Harris, E.J. and C.Y.L. Lee. 1986. Seasonal and
Growth and Management.—Orange jasmine annual occurrence of Mediterranean fruit flies
plants can live at least 15 years (author’s (Diptera: Tephritidae) in Makaha and Waianae
observation). Once established as an ornamental, valleys, Oahu, Hawaii. Environmental
they need little care. The species has not yet been Entomology 15(3): 507-512.
reported to be a weed in any area. Orange jasmine
can be killed (with moderate success) with Howard R.A. 1988. Flora of the Lesser Antilles,
herbicides recommended for broad-leaf weeds, by Leeward and Windward Islands.
girdling (Negreros-Castillo and Hall 1994), or by Dicotyledoneae, Part 1. Vol. 4. Arnold
grubbing out the plants. The species is the Arboretum, Harvard University. Jamaica Plain,
preferred host of the citrus psyllid, Diaphorina MA. 673 p.
citri, the vector for “citrus greening” disease
(Pacific Island Ecosystems at Risk 2002), and is a Kinoshita, T. and K. Firman. 1996. Highly
host of Mediterranean fruit flies (Diptera: oxygenated flavonoids from Murraya
Tephritidae) (Harris and Lee 1986). paniculata. Phytochemistry 42(4): 1207-1210.
Benefits.—Orange jasmine is a popular hedge Liogier H.A. 1988. Descriptive flora of Puerto
plant in the tropics. It can be used as a background Rico and adjacent islands, Spermatophyta. Vol.
plant or an accent plant. It can be pruned into 2. Editorial de la Universidad de Puerto Rico,
animal and other shapes (topiary), pruned into tree Río Piedras, PR. 481 p.
form, grown as a potted plant (Whistler 2000), or
cultivated as a bonsai (Gonzalez 2002). The Little, E.L., Jr., R.O. Woodbury, and F.H.
sapwood is light yellow, and the heartwood is Wadsworth. 1974. Trees of Puerto Rico and the
light brown and heavy, hard, fine-textured and Virgin Islands. Vol. 2. Agriculture Handbook
good for small turned articles (Little and others 449. U.S. Department of Agriculture,
1974). Branches or stems 2 to 5 cm in diameter in Washington, DC. 1,024 p.
the Indian Himalayan region were evaluated for
fuelwood suitability. Results were as follows: Neal, M.C. 1965. In gardens of Hawaii. Special
oven-dry density 0.72, dry-weight caloric value Publication 50. Bernice P. Bishop Museum
20.2 kj/g, ash 0.6 percent, moisture 54.7 percent, Press, Honolulu, HI. 924 p.
and nitrogen 0.42 percent (Negi and Todaria
1993). In the wild, orange jasmine contributes to Negi, A.K. and N.P. Todaria. 1993. Fuelwood
the biodiversity, protects the soil, and furnishes evaluation of some Himalayan trees and shrubs.
food and cover for wildlife. The leaves and other Energy 18(8): 799-801.
tissues have both stimulant and astringent
properties and are used to treat diarrhea, Negreros-Castillo, P. and R.B. Hall. 1994. Four
dysentery, cuts, joint pain, body aches (Parrotta methods for partial overstory removal in tropical
2001), venereal disease (Kinoshita and Firman forests of Mexico. Journal of Environmental
1996), and as an abortive (Xiao and Wang 1991). Management 41(3): 237-243.
In addition to essential oils, tissues of orange
jasmine contain the indole alkaloid yuehchukene Pacific Island Ecosystems at Risk. 2002. Invasive
(Xiao and Wang 1991) and at least eight highly plant species: Murraya paniculata (L.) Jack,
oxygenated flavones (Kinoshita and Firman 1996). Rutaceae. http://www.hear.org/pier3/mupan.
htm. 2 p.
References
Parrotta, J.A. 2001. Healing plants of Peninsular
Desert-Tropicals. 2002. Orange jasamine. India. CABI Publishing, Wallingford, UK and
509
New York. 917 p. guide. Timber Press, Portland, OR. 542 p.
Pathak, R.K., Madhulika-Mahajan, and S.N. Woman’s Club of Havana. 1952. Flowering plants
Sachan. 1992. Powdery mildew and cultural from Cuban gardens. Criterion Books, New
practices. Indian Journal of Forestry 15(1): 73. York. 365 p.
Sheehan, M.R. 1975. Florida landscape plants, Xiao, P.G. and N.G. Wang. 1991. Can
native and exotic. The University Presses of ethnopharmacology contribute to the
Florida, Gainesville, FL. 420 p. development of anti-fertility drugs? Journal of
Ethnopharmacology 32(1-3): 167-177.
Whistler, W.A. 2000. Tropical ornamentals, a
510
Myrica gale L. sweet gale
MYRICACEAE
Synonyms: Gale palustris A. Chev.

Myrica gale var. subglabra (A. Chev.) Fernald
Myrica gale var. subarctica J. Rousseau.
Myrica gale ssp. tomentosa (C. DC) E. Murray
Myrica gale var. tomentosa C. DC
Gerry Moore
appearing on different plants. Individual plants
have been known to change sex from year to year
(Burges 1993). The catkins are stalkless and borne
on the upper portions of the preceding year’s
branchlets. These branchlets die after flowering.
The staminate catkins range from 6 to 10 mm long,
are crowded on the branchlet, with the individual
flowers overtopped by the broad, shining,
subtending, brown bracts. The pistillate catkins
range from 7 to 10 mm long, each flower
subtended by two wing-like bracts that remain
fused to the fruit. The fruit of the sweet gale is a
three-pointed, compressed, ovoid nutlet that at
maturity is dotted with shining red to yellow resin
glands.
Range.—Sweet gale is found in the New and Old

World temperate regions. It occurs in the
Northeastern (Connecticut, Maine, Massachusetts,
New Hampshire, New Jersey, Michigan,
Minnesota, New York, North Carolina,
Pennsylvania, Rhode Island, Vermont, and
Wisconsin) and the Northwestern (Oregon and
Washington) United States (USDA 2003). The
Illustration source: Britton and Brown 1913
species is also known throughout much of Canada
(Alberta, British Columbia, Manitoba, New
General Description.—Sweet gale is the name Brunswick, Newfoundland, Northwest Territory,
used most often to refer to this shrub species. It is Nova Scotia, Ontario, Prince Edward Island,
also known as bog myrtle, Dutch myrtle English Quebec, Saskatchewan, and Yukon) and Alaska
myrtle, and meadow fern, although this species is (Bornstein 1997). In Eurasia it is known from
neither a true fern nor myrtle. Sweet gale is an northwest Europe to Portugal, Poland and
aromatic, deciduous, shrub up to 2.5 m tall. The northwestern Russia (Belgium, Britain, Denmark,
plant is single or multistemmed, with strongly Finland, France, Germany, Ireland, Netherlands,
ascending branches. The twigs are terete, brown Portugal, Norway, Poland, Russia, Spain, and
with scattered yellow glands. The leaves are 2 to 6 Sweden).
cm long (including the short petiole), oblanceolate
(i.e., widest toward apex) with cuneate bases, Systematic Botany.—Taxonomists disagree as to
toothed only toward the apex, glabrous and shining whether the sweet gale and the waxy-fruited
above but usually pubescent on the blade’s bayberry [Morella caroliniensis (Mill.) Small] and
underside. Like the twigs, the blades on both sides wax myrtle [Morella cerifera (L.) Small] should
are often beset with yellow glands. The flowers are be placed in the same genus (Bornstein 1997,
in unisexual, unbranched catkins (aments), with Fernald 1950, Gleason and Cronquist 1991) or in
the staminate (male) and pistillate (female) catkins separate genera (Wilbur 1994, 2002). Sweet gale
511
can be readily distinguished from the waxy fruited Baker, A. and R. Parsons. 1997. Rapid
species (bayberry, wax myrtle) on the bases of assimilation of recently fixed N2 in root nodules
flowers and fruits. If sweet gale is to be recognized of Myrica gale. Physiologia Plantarum 99: 640-
as generically distinct from bayberry and wax 647.
myrtle, as is done here, the generic name Myrica is
to be retained for sweet gale, with the bayberry Britton, N.L. and A. Brown. 1913. Illustrated flora
and wax myrtle being placed in the genus Morella of the northern states, Canada, and the British
(Wilbur 1994, 2002). Various varieties and possessions. Vol. 2, 2nd Ed. Scribner, New York.
subspecies [e.g., M. gale var. subglabra Fernald, 735 p.
M. gale var. tomentosa C. DC., M. gale ssp.
tomentosa (C. DC) E. Murray] have been Bornstein, A.J. 1997. Myricaceae. In: Flora of
recognized within sweet gale based on the degree North America Editorial Committee, eds. Flora
of pubescence present on the leaves. of North America Vol. 3. Oxford University
Press, New York. p. 430-434.
Ecology.—Sweet gale can tolerate full sun to
dense shade and is usually found in moist to wet Burges, N.A. 1993. Myrica. In: T.G. Tutin, N.A.
areas. Habitats where it can be found include bogs, Burges, A.O. Chater, J.R. Edmondson, V.H.
fens, swamps, and the margins of streams and Heywood, D.M. Moore, D.H. Valentine, S.M.
lakes. The roots of sweet gale contain nitrogen- Walters, and D.A. Webb, eds. Flora Europaea.
fixing nodules, which allow it to be particularly 2nd Ed.. Vol. 1. Cambridge University Press,
tolerant of nitrogen-poor, acidic soils (Baker and Cambridge, UK. p. 66-67.
Parsons 1997). Nitrogen fixation is accomplished
by the actinomycetous fungal genus, Frankia. Chittenden, F.J. and P.M. Synge. 1956. Dictionary
of Gardening 2nd Ed. Vol. 1. Clarendon Press,
Reproduction.—Sweet gale is wind pollinated Oxford, UK. 623 p.
and blooms in the spring. The pollen is allergenic
(Lewis 1977). The fruits of sweet gale mature in Everett, T.H. 1960. New illustrated encyclopedia
the fall and are dispersed by water (Ridley 1930, of gardening. Vol. 3. Greystone Press, New
Wilbur 1994). York. 479 p.
Growth and Management.—Sweet gale is often Fernald, M.L. 1950. Gray’s manual of botany.
cultivated on moist acidic soils where few other American Book Co., New York. 1,632 p.
species can be grown (Bailey 1922, Rehder 1940).
It thrives best in peaty soils and cannot tolerate Gleason, H.A. and A. Cronquist. 1991. Manual of
liming. Propagation can be by seed, layering, or vascular plants of the northeastern United States
root suckers (Chittenden and Synge 1956). and adjacent Canada 2nd Ed. New York
Propagation can also be effected through cuttings Botanical Garden, Bronx, NY. 910 p.
taken in late summer or fall. The cuttings should
be placed in a bed of peat moss and sand (Everett Lewis, W.H. 1977. Medical botany. John Wiley &
1960). Sons, New York. 515 p.
Benefits.—The branches and bark have been used Moerman, D.E. 1986. Medicinal plants of Native
in making a gale beer and in tanning; a decoction America. 2 Vols. Technical Report 19.
has been used for an insecticide and to kill vermin University of Michigan Museum of
(Rook 1998). The branches of sweet gale were Anthropology, Ann Arbor, MI. 910 p.
used by the Native Americans, Bella Coola, to
prepare decoctions to be used as a diuretic or as a Rehder, A. 1940. Manual of cultivated trees and
treatment for gonorrhea (Moerman 1986). shrubs, Macmillan Publishing Co, New York.
996 p.
References
Ridley, H.N. 1930. The dispersal of plants
Bailey, L.H. 1922. Standard cyclopedia of throughout the world. L. Reeve & Co., Ashford,
horticulture Vol. 4. MacMillan Co., London. Kent, UK. 744 p.
660 p.
512
Rook, J.S. 1998. A boundary waters compendium. Wilbur, R.L. 1994. The Myricaceae of the United
http://www.rook.org/earl/bwca/nature/shrubs/ States and Canada: genera, subgenera, and
myricagale.html [notpaged]. series. Sida 16: 93-107.
USDA, NRCS. 2003. The PLANTS Database, Wilbur, R.L. 2002. The identity and history of
Version 3.5 Baton Rouge, LA. Myrica caroliniensis (Myricaceae). Rhodora
http:plants.usda.gov. [not paged]. 104: 31-41.
513
Neea buxifolia (Hook. f.) Heimerl nia
NYCTAGINACEAE
Synonyms: Eggersia buxifolia Hook. f.
John K. Francis
Ecology.—Nia has an intermediate tolerance for

shade. It usually grows in the understory of low to
medium-density remnant and late secondary
forests. The species sometimes grows in semi-
exposed positions with other shrubs and low trees
on the tops of ridges. These are moist and dry
forests over sedimentary (especially limestone),
igneous, and metamorphic rocks. It is rare to
infrequent in most habitats except the moist
limestone hills in northern Puerto Rico where it is
locally common. Nia is sometimes found in coastal
thickets. A wide variety of well-drained soils are
colonized. Mean annual precipitation ranges from
750 to about 2200 mm at elevations from a little
above sea level to 500 m. The minimum
temperature for potted plants is reported to be 4.5
°C (Zane 2002).
General Description.—Nia, also known as Reproduction.—Nia flowers in the spring and

saltwood, is a fine-leafed, upright, evergreen shrub early summer and matures fruits in mid- to late
from 2 to 7 m in height and 2 to 8 cm in basal stem summer. The fruits, which are produced in sparing
diameter. There is usually a single stem emerging quantities, ripen a few at a time over a period of a
from the ground with multiple branches low on the few weeks. A group of fruits collected in Puerto
stem. Nia plants have a weak taproot with lateral Rico averaged 0.0291 ± 0.0011 g/fruit. However,
and fine roots, all brownish orange in color. The the seeds from this collection that averaged 0.0021
few slender branches form a diffuse crown. Bark is ± 0.0001 g/seed (air dried) failed to germinate.
smooth and gray. The inner bark is bitter. The Seeds of this species are probably dispersed by
wood is whitish, soft, and does not have birds. Natural seedlings are rare to scattered.
discernable annual rings. Oblong to oblanceolate, According to Zane (2002), nia can be propagated
entire leaves are opposite or in whorl-like groups by seed, branch cuttings taken in March through
along the twigs. The 9- to 24-mm long and 3- to 9- June, and by air layering.
mm broad blades are supported by 1- or 2-mm
petioles. Small light yellow flowers are borne on Growth and Management.—Nia is relatively
separate plants (dioecious). The elliptic, red, slow growing. Open-grown ornamentals reached 2
slightly fleshy fruits are 4 to 5 mm long and m in height in about 8 years. Understory saplings
contain one seed each (Liogier 1985, Little and add just a few centimeters of height per year. Life
others 1974). span in forests may reach several decades.
Although methods of propagation and culture in
Range.—Nia is native to Puerto Rico, its offshore pots as bonsais is well understood, no wildland
island of Culebra, St. Thomas and St. John of the plantings have been documented. Probably the
U.S. Virgin Islands, and Virgin Gorda of the best management is protection of the stands where
British Virgin Islands (Clubbe 2000, Liogier 1985, the species occurs.
Little and others 1974). It has been widely used as
a bonsai plant and planted somewhat as an Benefits.—Nia contributes to the diversity and
ornamental but is not known to have naturalized aesthetics of the forests where it occurs, helps
anywhere. protect the soil, and furnishes food and cover for
wildlife. It is one of the best broadleaf tropical
514
(room temperature) bonsai species available and 1. Editorial de la Universidad de Puerto Rico,
offers promise as an ornamental, especially for Río Piedras, PR. 352 p.
hedges and other shaped ornamentals.
References Wadsworth. 1974. Trees of Puerto Rico and the
Clubbe, C. 2000. British Virgin Islands: 449. U.S. Department of Agriculture,
conservation and training. Kew Scientist 17:5. Washington, DC. 1,024 p.
Liogier H.A. 1985. Descriptive flora of Puerto Zane, T.L. 2002. Neea buxifolia—Neea buxifolia.
Rico and adjacent islands, Spermatophyta. Vol. http://www.bonsai-bci.com/species/neea.html.
2 p.
515
Odontonema cuspidatum (Nees) Kuntze cardinal’s guard
ACANTHACEAE
Synonyms: Odontonema tubiforme (Bertol.) Kuntze

Odontonema strictum (Nees) Kuntze
Thrysacenthus cuspidatus Nees in DC.
Thyrsacanthus strictus Nees in DC.
John K. Francis
moist. It is moderately tolerant of shade and will
bloom in full sun, broken sun, and moderate shade
(author’s observation, Blomber 2000). In Central
America, it grows from near sea level to 1,400 m
in elevation (Stevens and others 2001). Cardinal’s
guard is not salt tolerant (Watkins 1975) and dies
to the ground if subjected to frost. In areas subject
to frost, it is grown as an annual or herbaceous
perennial (Blomber 2000, Watkins 1975). The
plants are mostly free of pests and disease but may
occasionally be attacked by mealy bugs
(Pseudococcidae) (Woman’s Club of Havana
1952).
Reproduction.—In tropical countries, cardinal’s

guard blooms throughout the year (Whistler
General Description.—Cardinal’s guard, also 2002), but in warm temperate areas, it blooms in
known as firespike, is an evergreen shrub 1 to 2 m the fall (Watkins 1975). Although the species is
in height and 1 to 2.5 cm in basal stem diameter. It common in Puerto Rico, the plants produce few
grows in patches and clumps that generate from viable seeds. Capsules fall off before drying and
root suckers. The smooth, green stems, which are liberating seeds. Fresh capsules collected in Puerto
woody below and semiwoody above, are Rico weighed an average of 0.0095 + 0.0002
supported by stiff lateral roots. The stems develop g/capsule. One hundred of these capsules were
few branches. The opposite leaves are dark green, sown on moist potting mix and only a single
glabrous, elliptic, 10 to 30 cm long, and acuminate seedling emerged. Cardinal’s guard is most
at the tip. The inflorescences are terminal racemes, common in disturbed areas (Stevens and others
sometimes branched, composed of tubular flowers 2001), along streams and drains, and in extra-
of scarlet or other colors including pink, white, moist places in secondary forests. Most of the
and lavender. Fruits are clavate (club-shaped) stands of cardinal’s guard in Puerto Rico have
capsules 1 to 2 cm long that contain flattened originated from abandoned gardens or errant
semiorbicular seeds (Liogier 1997). pieces of stem or root that have been transported
by streams or dumped in the woods with garden
Range.—Cardinal’s guard is native from Mexico prunings. Once established, plants spread by root
to Panama (Stevens and others 2001). The species suckers. The stems also layer (root) readily when
has naturalized and escaped in at least Puerto they become prostrate.
Rico, Hispaniola, and Cuba (Liogier 1997), and is
widely grown as an ornamental in Florida Growth and Management.—Cardinal’s guard
(Watkins 1975). Some authors propose separating grows rapidly from suckers and must be pruned
O. cuspidatum and O. tubiforme into individual frequently when used as hedges (Woman’s Club
species (Whistler 2000) or using only the name O. of Havana 1952). Normally, little care is needed
tubiforme (Stevens and others 2001). once the plant is established (Whistler 2000).
Ecology.—Cardinal’s guard prospers on fertile Benefits.—In natural stands, cardinal’s guard adds
and moderately fertile soils that are continually to biodiversity and protects against soil erosion.
516
The species is widely cultivated as an ornamental Universidad de Puerto Rico, San Juan, PR.
in tropical and subtropical areas for its striking red 436 p.
flowers. It is used as an accent and background
plant and sometimes employed to form hedges Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
(Watkins 1975, Whistler 2000, Woman’s Club of Montiel, eds. 2001. Flora de Nicaragua.
Havana 1952). It is also popular in gardens Monographs in systematic botany Vol. 85, No. 1.
because it attracts butterflies and hummingbirds Missouri Botanical Garden Press. 943 p.
that feed on the nectar (Blomber 2000, Watkins
1975). Watkins, J.V. 1975. Florida landscape plants,
native and exotic. The University Presses of
References Florida, Gainesville, FL. 420 p.
Blomber, M. 2000. In the garden: tropical plants Whistler, W.A. 2000. Tropical ornamentals, a
can add diversity to gardens. guide. Timber Press, Inc., Portland, OR. 542 p.
http://gainesvillesun.com/news/marina/02-26-
00marina.shtml. 4 p. Woman’s Club of Havana. 1952. Flowering plants
from Cuban gardens. Criterion Books, New
Liogier, H.A. 1997. Descriptive flora of Puerto York. 365 p.
517
Olyra latifolia L. carricillo
POACEAE
Synonym: Olyra paniculata Sw.

Olyra cordifolia Kunth.
John K. Francis
base rounded and the other straight, forming a 50 °

angle with respect to the leaf axis for about 2 cm.
Inflorescences are panicles 10 to 15 cm long,
terminal or upper axillary, and contain few to
many flowers. The 5-mm fruits are smooth, shiny,
ivory-colored, and hard (author’s observation,
Croat 1978, Howard 1979, Stevens and others
2001).
Range.—Carricillo is native to southern Mexico,

Central America, South America to northern
Argentina, the Antilles, moist Tropical Africa, and
Sri Lanka (Burkill 1994, Howard 1979, Judziewicz
and others 1999). There is speculation that the
species may be exotic in the Old World
(Judziewicz and others 1999).
Ecology.—Carricillo is moderately intolerant to

intermediate in tolerance to shade. It is widespread
and relatively common in the understory of thin
canopy forests, brushy forests, small openings, and
edges of high forest. These are most often medium
to late-secondary forests but can be primary forests
and remnants. Carricillo grows on a wide variety
of well-drained to somewhat poorly drained soils
with pH’s from near neutral to about 5.5 over
General Description.—Carricillo (or carrucillo), sedimentary (including limestone), igneous, and
also known as lintentwa, cortadora, lambedora, metamorphic (including ultramafic) rocks.
and sonadora, is a slender, arching bamboo-like Elevation may vary from near sea level to 1,100 m
grass. Unsupported culms reach about 1.5 m in areas that receive from about 1000 to more than
before sagging to horizontal. Climbing into the 3000 mm of mean annual precipitation (author’s
crowns of shrubs and low trees, plants may reach 3 observation, Croat 1978, Shaka and others 1997,
to 7 m in height and sometimes as much laterally. Stevens and others 2001).
The culms (stems) are thin-walled, brittle but
tough, and can reach 1.5 cm in diameter. Each Reproduction.—Carricillo flowers and fruits
plant usually has several culms arising from a tight primarily during the rainy season in Panama
root crown composed of brief rhizomes that (Croat 1978) and throughout the year in Nicaragua
support abundant fine roots. The culms are green, (Stevens and others 2001). Seed production is not
sometimes mottled with purple, throughout their abundant in Puerto Rico, but may be in Costa Rica
length and are enveloped with up to 8-cm sheaths (Missouri Botanical Garden 2002). Seeds in two
arising from the nodes. The culms are lineal except collections made in Puerto Rico on the same site
for vertical sprouts on reclining stems and on different years averaged 0.0210 + 0.0003 and
“brooms” or diffuse multiple branches that form 0.0225 + 0.0003 g/seed or about 46,000 seeds/kg.
near the ends of older culms. The leaves are Placed in moist potting mix, only 1 percent had
ribbon-like, 15 to 25 cm long and 3 to 6 cm broad germinated after 18 months. Scarification,
with long pointed tips and one edge of the leaf alternate wetting and drying, and heat treatment all
518
failed to stimulate germination (author’s Croat, T.B. 1978. Flora of Barro Colorado Island.
observation). Birds disperse the seeds (Burkill Stanford University Press, Stanford, CA. 943 p.
1994, Judziewicz and others 1999). New plants are
uncommon. Attempts to cultivate the species in Howard, R.A. 1979. Flora of the Lesser Antilles,
temperate greenhouses have failed (Judziewicz and Leeward and Windward Islands. Vol. 3. Arnold
others 1999). Arboretum, Harvard University, Jamaica Plain,
MA. 586 p.
Growth and Management.—Carricillo is a
relatively fast growing plant. Individual culms Judziewicz, E.J., J.G. Clark, X. Londoño, and M.J.
grow 2 m or more during their first year and 1 m Stern. 1999. American bamboos. Smithsonian
or so thereafter and live 2 to 4 years. By continual Institution Press, Washington, DC.
sprouting, plants may live for many years. http://email.uwsp.edu/publicanon/Course%20Inf
Artificial vegetative propagation has not been ormation/200120%-%20Spring%202002/
explored. Until vegetative methods are developed Biology/PF-BIOL345_xF8FF_545.1_200120/
or the extended dormancy problem of seeds is American%20Bamboos%20book.doc?Cmd=
solved, planting is not advised. Eradication can be open. 175 p.
done by grubbing out individual plants or probably
by cutting and spraying the sprouts with Missouri Botanical Garden. 2002. Manual de la
glyphosate or other grass herbicides. flora de Costa Rica. Missouri Botanical Garden,
St. Louis, MO. http://www.mobot.org/manual.
Benefits.—Carricillo contributes to the diversity plantas/050531/S050880.html. 7 p.
of the forest, helps protect the soil, and furnishes
food and cover for wildlife. Cattle eat the leaves North Carolina Botanical Garden. 2002. The biota
and fine twigs. Birds eat the fertile florets and of North America program: Olyra. North
fruits, although there is speculation that they may Carolina Botanical Garden, Chapel Hill, NC.
not be able to digest the hardened seeds (Burkill http://www.funet.fi/pub/sci/bio/life/plants/magno
1994). The species is the food plant for larva of the liophyta/magnoliophytina/liliopsida/poaceae/
moth Eryphanis reevesii (Doubleday) (North olyra/ 2p.
Carolina Botanical Garden 2002). The hollow
culms are used for flutes (Judziewicz and others Shaka, J.M., W. Kabushemera, and A. Msangi.
1999) and drinking straws. Leaves, culms, roots, 1997. Soils and vegetation of Semdoe proposed
and seeds are used in a number of applications in forest reserve Bombwera Division, Muheza
herbal medicine (Burkill 1994). Blades made from District, Tanga. Tech. Paper 36. Ministry of
the culms are used by the Cuiba tribe in Colombia Agriculture, National Soil Service, Agricultural
to cut the umbilical cords of newborns (Judziewicz Research Institute, Mlingano, Tanganyika. 14 p.
and others 1999).
References Montiel, eds. 2001. Flora de Nicaragua.
Burkill, H.M. 1994. The useful plants of West 3. Missouri Botanic Garden Press, St. Louis,
Tropical Africa. Vol. 2. Royal Botanic Gardens, MO. p. 1,911-2,666.
Kew, UK. 636 p.
519
Opuntia leptocaulis DC. desert Christmas cactus
CACTACEAE
Synonyms: Cylindropuntia leptocaulis (DC.) D.M. Knuth

Opuntia ramulifera Salm Dyck
Cylindropuntia leptocaulis var. stipata Coulter
Cylindropuntia leptocaulis var. longispina Berger
that is usually less than 1.5 m tall but can grow up
to 2.8 m in shady, protected environments such as
under trees (Earle 1990). Although relatively
compact, it is extensively branched from a main
trunk that is covered by scaly bark in old age. It
has pencil-like woody joints that are 12 to 40 cm
long and 5 mm in diameter and can be easily
detached from the plant. The tubercules, usually
less than 1 cm long, may be indistinct and the
joints smooth. The areoles, which are the small,
clearly defined raised areas on the surface, are oval
or almost diamond-shaped with short white wool.
The areoles may be spineless or have from one to
three spines that are from 19 to 51 mm long. There
are few, usually short, yellowish-brown glochids,
or minute bristles, in one to three small clusters in
the upper part of the areole. The leaves are green,
less than 1.3 cm long, and are early deciduous. The
greenish-yellow flowers are approximately 1.3 to
1.9 cm in diameter and up to 2.5 cm tall. The fruits
are up to 2.5 cm long, and smooth with brown
glochids in the areoles. There are usually less than
12 seeds per fruit (Earle 1990, Weniger 1991,
Vines 1986). Desert Christmas cactus is a diploid
species where n = 11 (McGregor and others 1986).
General Description.—The common name for Range.—Desert Christmas cactus occurs in

Opuntia leptocaulis is desert Christmas cactus. Arizona, New Mexico, Oklahoma, and Texas in
This name is likely to have originated because the the U.S.A. and in Sonora and Chihuahua in
green -colored fruits turn a very festive red color in northern Mexico (Bowers 1993, McGregor and
December just in time for Christmas. Other others 1986, Powell 1998).
common names include slender-stem cactus,
turkey pear, tasajillo, aguijilla and garrambulla Ecology.—Desert Christmas cactus is a water
(Taylor and others 1997, Weniger 1991). In the storer and employs a type of photosynthesis called
indigenous language of Nahuatl it is called crassulacean acid metabolism that allows them to
tzazahuistli (Crook and Mottram 1999). A widely make efficient use of available water (Burgess
applied name that includes all the long jointed 1995, Nobel 1988). The species is found at
cylindropuntia cacti, of which desert Christmas elevations below 1,524 m. It grows in most soil
cactus is one, is cholla. The genus name, Opuntia, types but appears to be most successful in
is taken from the name given to succulent plants gravelly, sandy, and heavier bottomland soils
that were originally from the old Greek town of (Taylor and others 1997, Epple 1995, Rondeau and
Opus. The species name, leptocaulis means others 1996, Bowers and McLaughlin 1987,
slender stemmed in Greek (Crook and Mottram Kearney and others 1960). It is commonly in the
1999). Desert Christmas cactus is an erect shrub same habitat with creosotebush, Larrea tridentata
520
(DC) Colville, with which it has a cyclical the fourth year (Bunting and others 1980). Desert
relationship in the Chihuahuan desert (Yeaton Christmas cactus recovery appeared particularly
1978). Creosotebush colonizes sites and modifies sensitive to the amount of precipitation.
microhabitat and edaphic conditions for desert Precipitation was required before plants would re-
Christmas cactus establishment and is eventually sprout. Plants tolerate minimum temperatures of
replaced by it. However, a combination of 7.3 oC in cultivation (Innes and Glass 1991). In
burrowing rodents and soil erosion increases the cultivation, desert Christmas cactus is susceptible
mortality rate of desert Christmas cactus while to the cochineal bug and periodic oil spray is
having little effect on creosotebush. The openings recommended (Earle 1990).
left by the dead cactus are then re-colonized by
creosotebush. Benefits.—Weniger (1991) describes it as a major
pest and “probably one of the most hated cacti in
Reproduction.—Desert Christmas cactus our area [Texas].” This is an unfortunate sentiment
reproduces both sexually by seed and vegetatively. because, although the spines can impale people
Flowers generally appear in May to July and livestock that venture too close, it provides
depending upon regional and environmental significant food, protected nesting sites, and cover
conditions. In response to wet weather, they can for a variety of wildlife (Earle 1990, Dayton
flower as early as April and again in August. The 1931). Most birds, including the bobwhite quail
flowers open wide in late evening (Taylor and and wild turkey, as well as small mammals, favor
others 1997). The fruit ripens in December and the fruit (Powell 1998, Taylor and others 1997).
persists on the plant until approximately February. Some animals, such as white-tailed dear, consume
Seeds may sprout while in the fruit and have a the joints as well as the fruit (Everitt and Drawe
shoot of up to 7 cm long while still on the plant 1993). There is apparently no seasonal variation
(Weniger 1991). Detached joints root and grow in the crude protein content of the stems, which is
into plants. The terminal stem segments detach similar to the fruits; both are 8 percent (Taylor and
particularly easily and readily take root near the others 1997). The larger shrubs provide cover and
parent plant so that natural populations that appear nesting sites, especially for the cactus wren. In
to be dense monocultures of clonal individuals addition, there is now a commercial value to
develop (Rebman and Pinkava 2001). Cuttings maintaining these shrubs. Land management for
taken from woody stems do not root easily (Earle wildlife has become increasingly important to
1990). Hybrids between cholla species are livestock producers in the Western U.S.A. because
common in nature (Rondeau and other 1996). An of fluctuations in livestock prices. In some parts of
example herbarium specimen of such a hybrid, O. Texas, income generated from consumptive (e.g.,
leptocaulis x O. spinosior (cane cholla), can be hunting) and non-consumptive (e.g., bird
seen on the Internet (Pinkava 2001). watching) wildlife activities already exceed
revenue obtained through livestock operations
Growth and Management.—Desert Christmas (Hernandez and others 1999). Under these
cactus is frequently abundant on overgrazed range circumstances upland gamebirds, such as northern
(Powell 1998). It tends to form thickets that can bobwhites and Rio Grande wild turkeys, have
become almost impenetrable to humans and become particularly important. Recommendations
livestock. Individual plants can be long-lived and have been made that low, thorny brush composed
may be more than 53 years old (Goldberg and of species such as lotebush [Ziziphus obtusifolia
Turner 1986). In a study over a 72-year period, 40 (Hook. ex Torr. & Gray) Gray] and desert
percent of plants were determined to survive at Christmas cactus should be protected in order to
least 7 years (Goldberg and Turner 1986). enhance nesting cover (Hernandez and others
Survival after fire is dependent upon the intensity 1999) and provide a food source. Desert Christmas
and frequency of the fire (Thomas 1991, Cave and cactus also provides refugia to native plant species.
Patten 1994). Plants can re-sprout after the fire A clear example of this on a landscape scale is in
kills the above ground parts (Bunting and others Arizona where clumps of tobosa grass are almost
1980). However, desert Christmas cactus appears completely restricted to refugia that are most
more susceptible to direct damage from fire than commonly under the desert Christmas cactus
some other Opuntia species (Bunting and others canopy where it is inaccessible to livestock
1980). After a fire in a mixed-grass prairie in (McAuliffe 1995). Native Americans gathered the
Texas, mortality was 65 percent in the first year fruits and ate them raw or cooked them into a jam
and reached a cumulative value of 80 percent by (Earle 1990). The Apache tribes mixed the
521
crushed fruits with a beverage to produce narcotic semiarid environments. Proceedings from
effects (Moerman 1998). Northern Texas and South Texas Wildlife
Conservation and Management Workshop,
References Saltillo, Mexico. August 19-20. Internet site:
http://cnrit.tamu.edu.
Bowers, J.E. 1993. Shrubs and trees of the
Southwest Deserts. Southwest Parks and Innes, C. and C. Glass. 1991. Cacti. Portland
Monuments Assoc. Tucson, AZ. 140 p. House, New York. 320 p.
Bowers, J.E. and S.P. McLaughlin. 1987. Flora Kearney, T.H., R.H. Peebles, and collaborators.
and vegetation of the Rincon Mountains, Pima 1960. Arizona flora. 2nd Ed. University of
County, Arizona. Desert Plants 8(2): 51-94 California Press, Berkeley, CA. 1,085 p.
Bunting, S.C., H.A. Wright, and L.F. Neuen- McAuliffe, J.R. 1995. Landscape evolution, soil
schwander. 1980. Long-term effects of fire on formation, and Arizon’s desert grassland. In:
cactus in southern mixed prairie of Texas. M.P. McClaran and T.R. Van Devender, eds.
Journal of Range Management 33(2): 85-88 The desert grassland. The University of Arizona
Press, Tucson, AZ. p. 100-129.
Burgess, T.L. 1995. The dilemma of coexisting
growth forms. In: M.P. McClaran and T.R. Van McGregor, R.L., T.M. Barkley, RE Brooks, EK
Devender, eds. The desert grassland. University Schofield. 1986. Flora of the Great Plains.
of Arizona Press, Tucson, AZ. p. 31-67 University Press of Kansas, Lawrence, KN.
1,402p.
Cave, G.H. and D.T. Patten. 1994. Short-term
vegetation responses to fire in the upper Sonoran Moerman, D.E. 1998. Native American ethno-
desert. Journal of Range Management 37(6): botany. Timber Press, Portland, OR. 927 p.
491-496
Nobel, P.S. 1988. Environmental biology of
Crook, R. and R. Mottram, 1999. Opuntia Index – agaves and cacti. Cambridge University Press,
Part 5: Nomenclatural note and I-L. Bradleya New York. 288 p.
17: 109-131
Pinkava, D. 2001. Biologist profile, specimens at
Dayton, W.A. 1931. Important western browse Arizona State University herbarium.
plants. Misc. Publication 101. U.S. Department http://lsvl.la.asu.edu/askabiologist/profiles/
of Agriculture, Washington, DC. 214 p. pinkava/pink.html.
Earle, W.H. 1990. Cacti of the Southwest. Desert Powell, A.M. 1998. Trees & shrubs of Trans-
Botanical Garden, Phoenix, AZ. 210 p. Pecos and adjacent areas. University of Texas
University Press, Austin, TX . 498 p.
Epple, A.O. 1995. A field guide to the plants of
Arizona. Falcon Press Publishing Co., Helena, Rebman, J.P., and D. J. Pinkava. 2001. Opuntia
MT. 347 p. cacti of North America–An overview. Florida
Entomologist 84 (4): 474-483
Everitt, J.H. and D.L. Drawe. 1993. Trees, Shrubs
and Cacti of South Texas. Texas Tech Rondeau, R., T.R. Van Devender, C.D. Bertelsen,
University Press, Lubbock, TX. 213 p. P. Jenkins, R.K. Wilson, and M.A. Dimmitt.
1996. Annotated flora and vegetation of the
Goldberg, D.E. and R.M. Turner. 1986. Tucson Mountains, Pima County, Arizona.
Vegetation change and plant demography in Desert Plants 12(2): 3-46
permanent plots in the Sonoran Desert. Ecology
67(3): 695-712 Taylor, R.B., J. Zrutledge, and J.G. Herrera. 1997.
A field guide to common south Texas shrubs.
Hernandez, F., C. Kleberg, and A. Radomski. Texas Parks and Wildlife Press, Austin, TX.
1999. Northern bobwhite and Rio Grande turkey p. 106.
management in relation to livestock grazing in
522
Thomas, P.A. 1991. Response of succulents to Weniger, D. 1991. Cacti of Texas and neighboring
fire: A review. International Journal of Wildland states. University of Texas Press, Austin, TX.
Fire 1(1): 11-22. 356 p.
Vines, R.A. 1986. Trees, shrubs, and woody vines Yeaton, R.I. 1978. A cyclical relationship between
of the Southwest. Sixth printing. University of Larrea tridentata and Opuntia leptocaulis in the
Texas Press. Austin, TX. 1,104 p. northern Chihuahuan desert. Journal of Ecology.
66: 651-656.
523
Palicourea crocea (Sw.) J.A. Schultes cachimbo
RUBIACEAE
Synonyms: Palicourea coccinea Poiteau ex DC.

Psychotria crocea Sw.
Palicourea reparia Benth.
Palicourea croceoides Ham.
Palicourea crocea DC. var. riparia (Benth.) Griseb.
Palicourea brevithyrsa Britt. & Standl.
John K. Francis
growing tips. The plant is glabrous or nearly so.

The leaves are elliptic, 7.0 to 14.5 cm long and 3.5
to 5.5 cm broad, and pointed at both ends with a
petiole 0.5 to 1.0 cm long. The tubular flowers are
grouped in cymes. Flower color ranges from
yellow to red. The fruits are ovoid to globose, 4 to
6 mm in diameter and dark red, purple, or black.
They are faintly sweet with a grassy flavor. There
are two seeds per fruit when perfectly developed.
The seeds are black or grayish with a hard, bony
shell, and hemispherical, with a groove on the flat
side and three ridges on the rounded side.
Range.—The natural range of cachimbo extends

from southern Mexico through Central America
and South America to southern Brazil and
Paraguay on the mainland and includes the
Greater and Lesser Antilles in the Caribbean
(Howard 1989, Liogier 1997)
Ecology.—Cachimbo grows in areas receiving

precipitations ranging from 1500 to over 4000
mm/yr. Soils where it grows are usually loams to
General Description.—Cachimbo is a medium to clays derived from volcanic or sedimentary parent
large shrub also known as red palicourea, yellow material. Elevations may be a few meters above
palicourea, yellow-cedar, tapa camino, ponasí, sea level up to 1,000 m (Little and others 1974).
bois de l’encore, bois cabrit, and bois fou-fou Cachimbo requires disturbance to establish itself.
(Little and others 1974). There is some ambiguity It grows well in tree-fall gaps, landslides, artificial
in the classification of this species. Liogier (1997) openings, and under thin forest canopies. It
separates P. crocea and P. croceoides on the basis competes aggressively with forbs, other shrubs,
of flower color; Howard (1989) treats them as a and young trees. When the canopies close and
single species under P. crocea. We will follow shade becomes dense, existing plants decline and
Howard for the purposes of this description. The eventually disappear. Labrón (1977) calls the
stems of cachimbo are green when young, species a “gap opportunist.” Cachimbo was a
changing to gray. The inner bark remains green. principal species in abandoned pastures in the
The wood is moderately hard and brittle. There are Luquillo Mountains, Puerto Rico, 60 years ago,
usually several sprouts from the base and long but not in abandoned shade coffee plantations
straight branches arising from major stems. (Zimmerman and others 1995). The species grows
Cachimbo develops a robust, shallow lateral root slowly on nutrient-poor substrate (as in landslide
system with abundant fine roots. Foliage is areas), but responds to N and P fertilization under
relatively sparse and concentrated near the those conditions (Fetcher and others 1996).
524
Reproduction.—Cachimbo flowers and fruits References
irregularly throughout the year (Little and others
1974). The fruits in one Puerto Rican sample Devoe, N.N. 1989. Differential seeding and
weighed an average of 0.1488 + 0.0055 g/fruit. regeneration in openings and beneath closed
Seeds cleaned from the sample averaged 0.0314 + canopy in sub-tropical wet forest. Ph.D.
0.0048 g/seed or 32,000 seeds/kg. Thirty-five dissertation. Yale University, New Haven, CN.
percent of these seeds germinated between 39 and 307 p.
84 days after sowing in commercial potting mix.
Palicourea flowers are pollinated by humming Fetcher, N., B.L. Haines, R.A. Cordero, D.J.
birds and the fruits are dispersed by frugiverous Lodge, L.R. Walker, D.S. Fernandez, and W.T.
birds (Taylor 1996). Bees were also observed to Lawrence. 1996. Responses of tropical plants to
visit the flowers (Labrón 1977). Cachimbo suffers nutrients and light on a landslide in Puerto Rico.
relatively little seed loss to pathogens and Journal of Ecology 84(3): 331-341.
predators and germinates at a higher percent after
natural dispersal than without it (Myster 1997). Howard, R. A. 1989. Flora of the Lesser Antilles,
Seedlings are common and widespread in suitable Leeward and Windward Islands. Vol. 6. Arnold
habitat. There is no significant difference between Arboretum, Harvard University, Jamaica Plain,
seasons in seedling appearance, but regardless of MA. 658 p.
season, seed germination is always higher in the
open than in closed canopy areas (Labrón 1977). Labrón, M. L. 1977. An autoecological study of
Transplanted wildlings survive and grow poorly Palicourea riparia Bentham (RUBIACEAE): an
(personal communication with F.N. Scatena, IITF, ecologically important species in the recovery of
Río Piedras, PR) a disturbed tropical rain forest in Puerto Rico.
Ph.D. Dissertation. University of North
Growth and Management.—Cachimbo grows Carolina, Chapel Hill, NC. 238 p.
slowly (a few cm) the first year and rapidly (1 m
or more per year) for about 2 years in openings Liogier, H. A. 1997. Descriptive flora of Puerto
(Labrón 1977). After about 3 years, seedlings Rico and adjacent islands. Vol. 5. Editorial de la
sown under an open canopy had accumulated Universidad de Puerto Rico, San Juan, PR.
about 20 times more biomass than seedlings sown 436 p.
under closed canopy conditions. The above-
ground to below-ground ratio of the open-canopy Little, E.L., Jr., R.O. Woodbury, and F.H.
plants was 4.8 (Labrón 1977). Cachimbo may Wadsworth. 1974. Trees of Puerto Rico and the
reach 6 m in height and 7.5 cm in stem diameter Virgin Islands. Vol. 2. Agriculture Handbook
(Howard 1989, Little and others 1974). No annual 449. U.S. Department of Agriculture,
rings are visible. However, single stems last about Washington, DC. 1,024 p.
10 years, and plants can continue to regenerate
with new sprouts. Plants cut during plantation Myster, R.W. 1997. Seed predation, disease and
cleaning and trail clearing quickly regain their germination on landslides in neotropical lower
former height (author’s observation). Scatena and montane wet forest. Journal of Vegetation
others (1993) present an equation to predict oven- Science 8(1): 55-64.
dry above-ground biomass (To) from diameter at
breast height (D) and height (H): To = exp(0.752 Scatena, F.N., W. Silver, T. Siccama, A. Johnson,
ln (D2H) – 2.362) where R2 = 0.861. and M.J. Sánchez. 1993. Biomass and nutrient
content of the Bisley Experimental Watershed,
Benefits.—Cachimbo is a food source for several Luquillo Experimental Forest, Puerto Rico,
species of song birds (Devoe 1989). It is higher in before and after Hurricane Hugo, 1989.
the major nutrients in its tissues than most of the Biotropica 25(1): 15-27.
trees and shrubs with which it is associated
(Scatena and others 1993) and is probably a good Taylor, C.M. 1996. Palicourea (Rubiaceae).
recycler of nutrients and a good forage plant. www.mobot.org/MOBOT/Staff/Research/taylor
People in some areas of Puerto Rico use crushed //palihome.html. 9 p.
cachimbo leaves to stop bleeding (Labrón 1977).
525
Zimmerman, J.K., T.M. Aide, M. Rosario, M.
Serrano, and L. Herrera. 1995. Effects of land
management and a recent hurricane on forest
structure and composition in the Luquillo
Experimental Forest, Puerto Rico. Forest
Ecology and Management 77(1-3): 65-76.
526
Parathesis crenulata (Vent.) Hook. f. rascagarganta
MYRSINACEAE
Synonyms: Ardisia crenulata Vent.
John K. Francis
annual precipitation. In the dry end of the
precipitation range, it only occurs near streams
and extra moist hollows. The species is found
mainly on slightly- to moderately-acid soils with
loamy to clayey texture. Although areas with both
igneous and sedimentary rocks are colonized, it is
most abundant in valleys in limestone formations.
Rascagarganta grows from a little above sea level
to 760 m in elevation (Little and others 1974). The
species is shade tolerant and prefers to grow under
closed forests with moderate to light understory
vegetation. It sometimes forms its own understory
canopy under which little vegetation grows. There
appear to be few insect or disease problems in
natural stands.
Reproduction.—Rascagarganta is reported to
flower from May to July and fruits from July to
December (Little and others 1974). Fruits are
often present in quantity and are easy to pick.
General Description.—Rascagarganta, also They may be cleaned by maceration and wet
known as secagarganta, jalapón, and raisin sieving. A collection of fruits from Puerto Rico
marron, is a shrub or small tree 1 to 3 m weighed an average of 0.407 g/fruit. Air-dried
(occasionally reaching 6 m) in height and 1 to 4 seeds cleaned from that collection weighed an
cm (occasionally to 7.5 cm) in trunk diameter. The average of 0.054 g/seed or 18,000 seeds/kg. Sown
common names in Spanish imply that it will make in peat, 45 percent germinated between 48 and
one’s throat raspy or dry. The species, unless 104 days after sowing. Many seedlings died from
disturbed, has a single stem and is supported by a damping-off; therefore, a looser germination
taproot and lateral roots that are orange-brown in medium than peat is recommended. Seedlings
color. Most of the branches arise from the main grew slowly in the nursery, reaching less than 10
stem, often in whorls of three’s or four’s. The cm in 6 months (author’s observation). Birds
alternate, dark-green, 7- to 23-cm leaves are disperse the seeds. Seedlings are relatively
narrowly elliptic, pointed at both ends, and often common and widespread near fruit-bearing stands.
have wavy-toothed edges. The small pink flowers
are borne in large numbers in terminal panicles. Growth and Management.—Rascagarganta
The 6- to 8-mm fleshy fruits progress from green grows slowly and appears to live several decades.
to pink to red to black as they ripen. Each fruit No management experience has been published.
contains one seed (Howard 1989, Liogier 1995, However, the best strategy for establishment of
Little and others 1974). natural stands may be to maintain plantations or
natural forest stands in moist areas for long
Range.—Rascagarganta is native to Hispaniola, periods with closed canopies and relatively clear
Puerto Rico, and Martinique (Howard 1989, understories to allow rascagarganta to build up.
Liogier 1995, Little and others 1974). Some
taxonomists suggest that P. serrulata (Sw.) Mez, Benefits.—Rascagarganta contributes to
of Cuba and Hispaniola is synonymous with P. biodiversity, cover for wildlife, and protects the
crenulata (Howard 1989). soil from erosion in the forests where it grows.
Although not usually heavily used by wildlife, the
Ecology.—Rascagarganta grows in moist forests fruits are produced in large quantity and are
that receive from about 1500 mm to 2800 mm of available after hurricanes when overstory fruits are
527
scarce. The wood is hard, but due to its small size Liogier, H.A. 1995. Descriptive flora of Puerto
is useful mainly for stakes and fuel. Rico and adjacent islands. Vol. 4. Editorial de la
References 617 p.
Howard, R. A. 1989. Flora of the Lesser Antilles, Little, E.L., Jr., R.O. Woodbury, and F.H.
Leeward and Windward Islands. Vol. 6. Arnold Wadsworth. 1974. Trees of Puerto Rico and the
Arboretum, Harvard University, Jamaica Plain, Virgin Islands. Vol. 2. Agriculture Handbook
MA. 658 p. 449. U.S. Department of Agriculture,
528
Parkinsonia aculeata L. Jerusalem-thorn
FABACEAE
Synonyms: none
John K. Francis
one to several in number, ellipsoid, gray, and

about 4 by 8 mm. The species has a haploid
chromosome number of 14 (Pantulu 1942).
Jerusalem-thorn is also known as horse bean, palo
verde, retama, palo de rayo, espinillo, sulfato,
cina-cina, mataburro, madam naiz, and arrête-
boeuf (Correll and Johnston 1970, Howard 1988,
Range.—Jerusalem-thorn is native to Texas,

Arizona, Mexico, Central America, and South
America as far south as the north of Uruguay
(Holdridge and Poveda 1975, Little and
Wadsworth 1964, Piaggio 2001). It has been
planted and has naturalized in many of the islands
of the Caribbean, including Puerto Rico and the
U.S. Virgin Islands (Liogier 1988). The species
has been introduced into the Old World and has
become widespread in Australia. Jerusalem-thorn
is planted as an ornamental in Florida, and the
Southern and Western United States.
Ecology.—Jerusalem-thorn will grow on most

General Description.—Jerusalem-thorn occupies types of soils, including sand dunes, clay soils,
the transition zone between shrub and tree life- strongly alkaline, chalky, and mildly salty soils. It
forms. In difficult sites and portions of its range, it withstands a great deal of heat and can survive in
is very shrubby, but in moist and fertile areas, it areas that receive less than 30 cm of annual
becomes a small tree up to 10 m in height. rainfall. Jerusalem-thorn is damaged by
Jerusalem-thorn often has multiple stems, temperatures below –8 °C (Floridata 2001).
especially if burnt or cut, and usually has a Individual plants are attacked and damaged by
branchy, low, and diffuse crown. The branches termites, scales, and a number of other insect
and foliage are often somewhat pendulous. It has species, but populations are generally relatively
a taproot, carrot-like in young plants, and slender free of pests. It requires full or nearly full sunlight
laterals radiating out from it at all depths. Old to survive and reproduce. In most environments,
trunks are brown and fissured or scaly. The bark Jerusalem-thorn needs some kind of disturbance to
of smaller stems and twigs is thin, smooth, and become established.
yellow-green or blue-green. There are paired
spines at the nodes and a larger spine (1 to 2 cm Reproduction.—Jerusalem-thorn flowers and
long) at the end of the leaf axis. The leaves are fruits throughout the year in Puerto Rico (Little
alternate, bipinnately compound (20 to 40 cm and Wadsworth 1964). It often flowers and fruits
long) with a flattened rachis and many tiny (2 to 4 at 2 m in height or less. The flowers are pollinated
mm long) leaflets that are shed during the winter by bees that are attracted to and orient themselves
or the dry season. The yellow or yellow and to the banner petal, which absorbs UV radiation
orange fragrant flowers are grouped in racemes (Jones and Buchmann 1974). Seeds collected in
and have five sepals and five petals. The fruits are Puerto Rico by the author numbered 13,300
legumes 3 to 10 cm long, slightly flattened with seeds/kg. Little and Wadsworth (1964) report
constrictions between each seed. The seeds are 12,300 seeds/kg. Seeds from the former collection
529
were scarified and germinated at 59 percent, References
beginning 2 days after sowing (Francis and
Rodríguez 1993). Jerusalem-thorn produces two Correll, D.S., and M.C. Johnston. 1970. Manuel of
kinds of seeds. About 25 percent of them have thin the vascular plants of Texas. Texas Research
testae and will germinate readily without Foundation, Renner, TX. 1,881 p.
pretreatment; the rest have hard seed coats and
must be scarified before they will germinate Everitt, J.H. 1983. Seed germination
(Floridata 2001). Mechanical scarification was characteristics of two woody legumes (retama
used by the author; soaking for 45 minutes in and twisted Acacia) from South Texas. Journal
concentrated sulfuric acid worked equally well in of Range Management 36(4): 411-414.
another test (Everitt 1983). In nature, seeds are
transported by water, birds, and animals (Pacific Everitt, J.H. and D.L. Drawe. 1993. Trees, shrubs
Island Ecosystems at Risk 2001). Artificial and cacti of South Texas. Texas Tech University
propagation is routinely done with seeds followed Press. 213 p.
by ordinary nursery culture in containers. In vitro
propagation using nodal explants has been Floridata. 2001. Plant profile: Parkinsonia
demonstrated (Jaideep and others 1992). aculeata. http://www.floridata.com/ref/p/
Jerusalem-thorn will continue to resprout after park_acu.cfm. 3 p.
many cycles of annual disturbance.
Francis, J.K. and A. Rodríguez. 1993. Seeds of
Growth and Management.—The growth rate is Puerto Rican trees and shrubs: second
moderate, from 0.5 to 1 m per year in early years. installment. Research Note SO-374. U.S.
Jerusalem-thorn plants live about 30 years (Pima Department of Agriculture, Forest Service,
Community College 2001). As an exotic, Southern Forest Experiment Station. New
Jerusalem-thorn can be problematic in rangeland, Orleans, LA. 5 p.
as it has become in Australia. Biological control
attempts are ongoing, but have not yet been Harris, S.C. 1982. Nitrogen fixation by tropical
successful. Jerusalem-thorn can be controlled with woody legumes: potential source of soil
tractors by pulling or dozing the large plants out, if enrichment. In: J.P. Roskoski, J. Monano, C. van
the seedlings are also controlled. Good control has Kessel, G. Castilleja, and P.H. Graham, eds.
been achieved with several herbicides as well Biological nitrogen fixation technology for
(Pacific Island Ecosystems at Risk 2001). tropical agriculture. Centro International de
Agricultura Tropical. Cali, Colomabia. p. 447-
Benefits.—The principal use of Jerusalem-thorn 454.
today is for landscaping. Its abundant yellow floral
display alone justifies its use. Perhaps its best use Hoekstra, D.A., P.J. Wood, and F. Anap Sang.
is in xeric gardens, where it contrasts well with 2001. Proposed second phase dryland
succulents and other desert plants (Floridata agroforestry research project.
2001). It is also useful in hedges and for living http://www.idrc.ca/library/document/031091/
fenceposts (Hoekstra and others 2001). Jerusalem- 31 p.
thorn is also planted for environmental restoration
in desert areas. Besides being hardy, it has the Holdridge, L.R. and L.J. Poveda A. 1975. Arboles
added benefit of fixing nitrogen (Harris 1982). de Costa Rica. Vol. 1. Centro Científico
The wood is moderately hard, heavy (specific Tropical, San José, Costa Rica. 546 p.
gravity 0.6), and brittle. It is used principally for
fuel. Native Americans harvested the seeds, which Howard, R.A. 1988. Flora of the Lesser Antilles,
they parched before eating (Floridata 2001). Leeward and Windward Islands.
Various extracts of leaves, flowers, fruits, and Dicotyledoneae, Part 1. Vol. 4. Jamaica Plain,
bark are used in herbal medicine to treat arthritis MA: Arnold Arboretum, Harvard University.
and fever, and as a nerve stimulant and abortive 673 p.
(Liogier 1990). Livestock eat the foliage and fruits
(Little and Wadsworth 1964). Jerusalem-thorn is Jaideep. M., S. Mukunthakumar, and J. Mathur.
occasionally browsed by white-tailed deer and the 1992. Micropropagation of Bauhinia variegata
seeds are eaten by bobwhite quail (Everitt and and Parkinsonia aculeata from nodal explants of
Drawe 1993). mature trees. Plant Cell, Tissue and Organ
530
Culture 28(1): 119-121. Department of Agriculture, Forest Service,
Jones, C.E. and S.L. Buchmann. 1974. Ultraviolet
floral patterns as functional orientation cues in Pacific Island Ecosystems at Risk. 2001. Invasive
hymonopterous pollination systems. Animal plant species: Parkinsonia aculeata L. Fabaceae.
Behavior 22(2): 481-485. http://www.hear.org/pier/paacu.htm. 2 p.
Liogier H.A. 1988. Descriptive flora of Puerto Pantulu, J.V. 1942. Chromosome numbers of some
Rico and adjacent islands, Spermatophyta. Vol. Caesalpiniaceae. Current Science 11: 152-153.
Río Piedras, PR. 481 p. Piaggio, M. 2001. Plantas natives del Uruguay.
Red Academica de Uruguay.
Liogier, H.A. 1990. Plantas medicinales de Puerto http://www.rau.edu.uy/uruguay/flora/Uy.flora3.
Rico y del Caribe. Iberoamricana de Ediciones, htm. 8 p.
Pima Community Collage. 2001. Desert ecology
Little, E.L., Jr., and F.H. Wadsworth. 1964. of Tucson: mexican paloverde (Parkinsonia
Common trees of Puerto Rico and the Virgin aculeata). http://wc.pima.edu/Bfiero/
Islands. Agriculture Handbook 249. U.S. tucsonecology/plants/trees_mpv.htm. 1 p.
531
Parthenocissus quinquefolia (L.) Planch. Virginia creeper
VITACEAE
Synonyms: Amphelopsis latifolia Tausch

Amphelopsis quinquefolia (L.) Michx.
Hedera quinquefolia L.
Vitis quinquefolia (L.) Lam.
Parthenocissus hirsute (Pursh) Graebn.
Parthenocissus inserta (Kerner) Fritsch (and other synonyms: Natural
Resources Conservation Service 2003)
John K. Francis
yellowish-green flowers are borne in terminal

cymes or panicles positioned under the foliage.
Fruits are berries, somewhat flattened at the apex,
4 to 6 mm in diameter, bluish-black with a definite
bloom when ripe. Fruits usually contain two to
three seeds (Dirr 2003, Nelson 1996, Rehder 1951,
Welsh and others 1987).
Range.—Virginia creeper is native to all the

conterminous States except Washington, Oregon,
California, Nevada, Arizona, New Mexico, Utah,
Idaho, Montana, Wyoming, and North Dakota
(Natural Resources Conservation Service 2003). It
is also native to northern Mexico and southeastern
Canada from Nova Scotia to Ontario (Coladoanto
1991, Nelson 1996). It has naturalized in at least
Utah, England, and France (Compton 2003,
Services Culture Editions Ressources 2003, Welsh
and others 1987). The species is cultivated as an
ornamental in many moist temperate areas of the
world.
Ecology.—Virginia creeper is found in most of the

vegetation associations in the Eastern and
Midwestern United States (Coladoanto 1991). It
General Description.—Virginia creeper, grows in most moist soils types. Virginia creeper
occasionally referred to as woodbine, thicket grows well in full sun but also tolerates shade.
creeper, American ivy, and five-leaved ivy, is a When growing under a forest canopy, it attempts
deciduous woody vine that may reach 15 m or to ascend the tree trunks or obstacles it encounters.
more into the crowns of trees or up masonry walls. It is a mid- to late-seral species. It is sensitive to
Single or multiple stems are slender, round, light fire and is top-killed by most fires. Root crowns
brown, and have prominent lenticels. Young and occasional surviving stems sprout and partially
growth is reddish. The vines adhere to surfaces by renew burned stands (Coladoanto 1991).
means of five- to eight-branched tendrils ending in
cup-like, adhesive tips. The alternate leaves are Reproduction.—Virginia creeper flowers from
palmately compound with five (sometimes three or June to August, matures fruits from August to
four) leaflets. The leaflets are lanceolate, October and drops fruits (or they are eaten by
oblanceolate or obovate, 3 to 15 cm long, and 1.6 birds) from September to February. Good fruit and
to 8 cm broad with singly or doubly serrate seed crops are frequent. There are 21,600 to
margins. Petioles are 15 to 20 cm long, and leaflets 58,000 air-dried seeds/kg of which 44 to 99
have distinct petiolules. Small, greenish-white to
532
percent are sound (Gill and Pogge 1974). The diarrhea and difficult urination, infusions for
seeds are dispersed by birds (Dirr 2003). Seeds jaundice, and applied decoctions and poultices
germinate the first or second spring after dispersal; topically for swelling and lockjaw (Moerman
germination is epigeal (Gill and Pogge 1974). It 1986). The fruits are reported to be poisonous by
layers readily at the nodes when stems come in reason of the oxalic acid they contain (Russell and
contact with the soil (Crandall and Crandall 1995). others 1997). However, consuming more than a
small amount is unlikely due to their bad flavor
Growth and Management.—Once well (author, personal observation).
established, Virginia creeper grows rapidly. It
often must be pruned to prevent it from getting out References
of control in ornamental settings. It can seed into
nearby flower beds and wildlands. Virginia Coladoanto, M. 1991. Parthenocissus
creeper should not be planted to climb up quinquefolia. In: U.S. Department of
buildings with wooden siding because it is difficult Agriculture, Forest Service, Rocky Mountain
to remove and because it increases the humidity Research Station, Fire Sciences Laboratory, Fire
and hastens rot (Gilman 1999). The species Effects Information System. http://www.fs.fed.
occasionally interferes with reproduction of pines us/database/feis/plants/vine/parqui/all.html.
and hardwoods and may be controlled with 10 p.
herbicides. Fire can also be an effective control
agent (Coladoanto 1991). Fruits can usually be Compton, G. 2003. Cambridgeshire flora records
picked in quantity by hand. Seeds are separated since 1538. http://mnlg.com/gc/species2pz/
from fruit tissue by maceration followed by p/par_qui.html. [not paged].
floating off the pulp and empty seeds with water.
However, maceration must be gentle because seed Crandall, C. and B. Crandall. 1995. Flowering,
coats are soft and easily damaged. There are 16 to fruiting, and foliage vines. Sterling Publishing
27 g of seeds/100 g of fruits. The seeds may be Co., Inc., New York. 192 p.
fall-sown 1 cm deep in soil and mulched.
Alternatively, seeds may be stratified in moist peat Dirr, M.A. 2003. Parthenocissus quinquefolia—
or sand at 5 or 6 °C for 60 days and sown in the Virginia creeper, also called woodbine.
spring. Forty-one to 94 percent of the seeds will http://www.nobleplants.com/classnotes/spring/
germinate after 30 days. Seedlings are ready for springprofiles/vines/parthenoquinque.htm. 1 p.
outplanting when they reach 15 cm in height after
1 or 2 years in the nursery. The species can also be Gill, J.D. and F.L. Pogge. 1974. Parthenocissus
propagated with cuttings and layers (Gill and Planch., creeper. In: C.S. Schopmeyer, tech.
Pogge 1974). coord. Seeds of woody plants in the United
States. U.S. Department of Agriculture, Forest
Benefits.—Virginia creeper is important to the Service, Washington, DC. p. 568-571.
aesthetics of forests, helps protect the soil, and
furnishes food and cover for wildlife. Cattle and Gilman, E.F. 1999. Parthenocissus quinquefolia.
white-tailed deer sometimes browse the foliage. Fact Sheet FPS-454. Cooperative Extension
Songbirds are the principal consumers of the fruits. Service, University of Florida, Gainesville, FL.
The foliage, both as a ground cover and climber, 3 p.
provides cover for small mammals and birds
(Coladoanto 1991). It has been planted for Moerman, D.E. 1986. Medicinal plants of Native
watershed protection and erosion control (Gill and America. Technical Reports. 19. University of
Pogge 1974). The principal attraction of Virginia Michigan Museum of Anthropology, Ann
creeper is its fall color. The leaves turn fiery Arbor, MI. 534 p.
shades of purple, red, and scarlet after the first
frost (Dirr 2003). The blue-black fruits and their Natural Resources Conservation Service. 2003.
stems (which turn red also) add color and interest Plants profile: Parthenocissus quinquefolia (L.)
after the leaves have fallen. The species is planted Planch. http://plants.udsa.gov/cgi_bin/
to cover building walls, run up tree trunks, and plant_search.cgi?mode=Common+Name&keyw
form trellises and arbors. It also makes a fine ordquery=Vi.... 4 p.
ground cover (Crandall and Crandall 1995). Native
Americans took decoctions of the roots for
533
Nelson, G. 1996. The shrubs and woody vines of hort/consumer/poison/Parthqu.htm. 2 p.
391 p. Services Culture Editions Ressources. 2003.
Parthenocissus quinquefolia (Linné) Planchon.
Rehder, A. 1951. Manual of cultivated trees and http://crdp.ac-besancon.fr/ressourc/flore/flore/
shrubs hardy in North America. The MacMillan especes/parth_quinq.htm. 1 p.
Company, New York. 996 p.
Russell, A.B., J.W. Hardin, L. Grand, and A. Higgins, eds. 1987. A Utah flora. Great Basin
Fraser. 1997. Poisonous plants: Parthenocissus Naturalist Memoirs 9. Brigham Young
quinquefolia. North Carolina State University, University, Provo, UT. 894 p.
Raleigh, NC. http://www.ces.ncsu.edu/depts/
534
Passiflora edulis Sims passionfruit
PASSIFLORACEAE
Synonyms: none
John K. Francis
filled with aromatic yellow or orange juice. The

fruits of the purple passion fruit are smaller but
more aromatic than those of the yellow form
(Acevedo-Rodríguez 1985, Liogier 1994, Morton
1987).
Range.—The purple passionfruit is a native of

southern Brazil through Paraguay to northern
Argentina. The yellow form of passionfruit is
probably native of the Amazon region of Brazil
(Morton 1987). Passionfruit is cultivated today
throughout the tropics and subtropics and has
naturalized and escaped in many areas including
Florida, Puerto Rico, U.S. Virgin Islands, Hawaii,
Guam, and Western Samoa (Acevedo-Rodríguez
1985, Morton 1987, Pacific Island Ecosystems at
Risk 2002).
General Description.—Passionfruit is also
known as parcha, granadilla, maracuyá, ceibey, Ecology.—Altitude and latitude do not appear to be
lilikoi, and linmangkon (Morton 1987). The a constraint other than through the temperatures
English prefix “passion” derives from the passion associated with them. Purple passionfruit grows
of Christ suggested by the prominent four- best in a subtropical climate, and the yellow
branched style that appears in a few of the flowers. passionfruit prefers a tropical climate with full-
Passionfruit is a woody, perennial vine that bears a season warm days and nights. However, even
delicious fruit and occurs in purple- and yellow- yellow passionfruit can survive temperatures down
fruited forms (Passiflora edulis Sims f. edulis and to – 5 °C (Sentelhas and other 1996). Generally,
P. edulis f. flavicarpa) known as purple and annual rainfall should be at least 900 mm. Rainfall
yellow passionfruits. The plants have a weak in Indian areas that grow passionfruit ranges from
taproot and extensive ivory-colored lateral roots. 1000 to 2500 mm/year (Morton 1987). The species
The stem is usually solitary, up to 7 cm in basal is shallow-rooted but withstands drought by
diameter, extends 5 to 10 m or more into the defoliating. Passionfruit tolerates a wide variety of
crowns of trees, and is covered by a thin, flaky, soils and grows best on well-drained sandy loams
light brown bark. The stemwood is light and with pH’s of 6.5 to 7.5 (Morton 1987). Passionfruit
brittle. The twigs are yellow-green, turning brown, is moderately intolerant of shade, requires trees,
and support themselves on vegetation by means of brush, or fences for support, and benefits from but
tendrils that arise at the leaf axils. The leaves are does not require soil disturbance for reproduction.
alternate, green to yellow-green, three-lobed (on Wild plants are found in broken forests, stream-
mature plants) with serrate edges. The petioles are bottom galleries, fencerows, abandoned farms, and
3 to 6 cm long and the blades are 5 to 11 cm long neglected city lots. Young plants are eaten by
by 4 to 10 cm broad. Solitary flowers arise at the livestock, so passionfruit is almost never found in
leaf axils. The flowers measure 5 to 7 cm across moderate to heavily grazed areas. A large number
with five greenish-white sepals and five white of insects, nematodes, fungi, and viruses attack the
petals topped with a fringe-like corona of straight species (Morton 1987).
purple and white rays. There are five stamens with
large anthers and a triple-branched style. The fruit Reproduction.—The purple passionfruit blooms
is globose or ovoid, purple or yellow and 4 to 7 in spring and early summer and again for a shorter
cm in diameter. Inside a thick rind are many dark- period in fall and early winter (Morton 1987).
brown to black seeds enveloped in small sacs
535
Yellow passionfruit in Puerto Rico flower from inoculation with superior strains of fungi
April to September and yield fruits from June to (Cavalcante and others 2001). Wild plants are
October (Acevedo-Rodríguez 1985). In some usually scattered and attempts at management
areas, plants fruit twice each year (Popenoe 1920). have not been reported.
Plants usually begin blooming and fruiting in their
second year. Yellow passionfruit flowers have Benefits.—By far the greatest benefit of
both male and female parts but are self-sterile. passionfruit to humankind is its fruit and the
They rely mainly on carpenter bees (Xylocopa delicious juice made from it. In addition to being
spp.) for pollination. Other insects and collected by local people in the forests, the fruit is
hummingbirds also visit the flowers. The flowers now grown in vineyards in dozens of countries. It
of purple passionfruit can self-pollinate (Morton is condensed, frozen, and shipped worldwide. The
1987). Fruits of the naturalized yellow-fruited fruit pulp contains 2.2 percent protein, 0.7 percent
form range from about 45 to 120 g in Puerto Rico. fat, and 21.2 percent carbohydrates. In addition,
There is a large variation between plants in size the seeds contain 23 percent oil similar to
and shape of fruits. Small fruits are sometimes sunflower or soybean oil, and the rind residue is
completely devoid of seeds, and large fruits may used for cattle feed (Morton 1987). The fruits of
have over 200 seeds. A collection of seeds from native and naturalized stands furnish food for
naturalized plants in Puerto Rico averaged 0.0251 numerous species of wild mammals and birds. The
+ 0.0004 g/seed or 40,000 seeds/kg. Passionfruit is whole plant, especially the leaves, contains
usually propagated from seeds but can be started alkaloids and a number of other phytoactive
from cuttings, layers, and grafts (Morton 1987). chemicals. Among these is passiflorine, a known
Seeds germinate best if allowed to ferment for a sedative and tranquilizer (Morton 1987). Extracts
few days in the fruit pulp before cleaning and are of the leaves have been used for centuries as
lightly scarified by clipping or sandpapering sedatives by native Brazilians. They prepare a
(Morton 1987). A group of seeds in Puerto Rico drink from the flower to treat asthma, bronchitis,
were sown without pretreatment in commercial and whooping cough. The plant is also used as a
potting mix and began germinating in 14 days and diuretic to treat urinary infections (Rain-tree
completed germination in 24 days with 61 percent 2002).
germinated. Plants are grown in beds or pots and
transplanted when they reach about 25 cm in References
height. The seedlings are heavily watered after
outplanting (Morton 1987). Seeds are disbursed in Acevedo-Rodríguez, P. 1985. Los bejucos de
the wild by humans, animals especially pigs, and Puerto Rico. Vol. 1. General Technical Report
birds, and by vine extension. SO-58. U.S. Department of Agriculture, Forest
Growth and Management.—Plants in fertile soil New Orleans, LA. 331 p.
extend their stems about 3 m per year. Each year
during the annual dry season, the leaves fall off Anonymous. 2001. Exotic ornamental. The Hindu.
and the twigs die, leaving the main stem and a few http://www.hinduonnet.com/thehindu/2001/04/
important branches alive to rebuild the crown after 22/stories/1322045j.htm. 3 p.
the rains begin. Because fruiting takes place on
new wood, light pruning does not reduce yield. Bailey, L.H. 1941. The standard cyclopedia of
Plants live from 3 to 8 years and do not resprout. horticulture. Vol. 3. The MacMillan Co. New
Commercial stands are managed in vineyards, York. p. 2,423-3,639.
somewhat like grapes. They are planted in trellised
rows 4.5 m apart and spaced 4.5 m apart within Cavalcante, U.M.T., L.C. Maia, C.M.C. Costa, and
rows. The orchards are replanted every 4 to 6 V.F. Santos. 2001. Mycorrhizal dependency of
years (Bailey 1941). Alternately, vineyards may passion fruit (Passiflora edulis f. flavicarpa).
be established using small trees or bamboo as Fruits 56: 317-324.
standards (Heenkenda and Punchikumarihami
1991). Fruiting plants can even be grown in pots Heenkenda, H.M.S. and S.M.A.
under glass (Anonymous. 2001). Fruits fall after Punchikumarihami. 1991. Alternate trellising
ripening on the vines and are picked up from the materials for passion fruit (Passiflora edulis
ground at least twice per week. The roots normally Sims) vines. Tropical Agriculturist 147: 11-18.
form mycorrhizal associations and benefit from
536
Liogier, H.A. 1994. Descriptive flora of Puerto Popenoe, W. 1920. Manual of tropical and
Rico and adjacent Islands. Vol. 3. Editorial de la subtropical fruits. The MacMillan Company.
Universidad de Puerto Rico, Río Piedras, PR. New York. 474 p.
461 p.
Rain-tree. 2002. Maracuja, “passion fruit”.
Morton, J.F. 1987. Fruits of warm climates. http://rain-tree.com/passionf.htm. 3 p.
Creative Resources Systems, Inc. Winterville,
NC. 505 p. Sentelhas, P.C., C.T.P. Junior, J.M.M Sigristi, R.
Kavati, and M.T. Parodi. 1996. Freezing points
Pacific Island Ecosystems at Risk. 2002. Invasive of various tropical fruits. Bragantina 55(2): 231-
plant species: Passiflora edulis Sims, 235.
Passifloraceae. http://hear.org/pier/paedu/htm.
2 p.
537
Paullinia pinnata L. bejuco de costilla
SAPINDACEAE
Synonyms: none
John K. Francis
dry (less than 1000 mm precipitation) forest in

Puerto Rico (Acevedo-Rodriguez 1985). Soils
derived from limestone and other sedimentary
rocks, and various igneous rocks at low and
moderate elevations (below 400 m) are colonized.
The species appears to be rare or absent from
serpentine. It is present in both primary remnant
and secondary forest stands.
Reproduction.—In Puerto Rico, the species

blooms from June to November and fruits from
July to December (Acevedo-Rodriguez 1985). The
fruits are capsules that split in three valves when
ripe to expose one to three black seeds partially
covered by a white aril. A collection of fruits in
Puerto Rico weighed 1.674 + 0.0564 g. The fresh
seeds recovered from those fruits weighed 0.150 +
0.008 g and gave 5 percent germination. The seeds
are recalcitrant (drying kills them). A later
collection carefully protected from drying gave
General Description.—Bejuco de costilla is also 100 percent germination. Germination is
known in Spanish-speaking areas as bejuco de hypogenous. The seedlings grew rapidly and
paloma and bejuco de guajanilla and in English- reached a maximum of 18 cm in 18 days after
speaking areas as supple jack. It is a stout and emergence. The seedlings quickly develop a deep
strong woody vine that forms mats in forest and extensive tap and lateral root system.
openings and ascends rocks and trees by means of Moderate amounts of seed can be obtained by
forked tendrils to access full sunlight. It may reach clipping the fruit clusters with a pruning pole and
10 m of extension (Vélez and Overbeek 1950) and shelling out the seeds by hand. Seeds should not be
12 cm or more of stem diameter. The name bejuco picked up from the ground. Seed should be planted
de costilla (rib vine) arises from the appearance of immediately or stored refrigerated in a closed
young stems. The young, green stems are plastic bag containing a moistened paper towel.
angularly striated; the large, lower stems have a Natural reproduction in forest stands is common to
rounded triangular cross section. The plant is rare. Good survival and long life enable the
easily recognized from the leaves that have five species to maintain itself. More than one stem may
serrated leaflets with prominent veins and a arise from a horizontal root segment. Stems layer
winged rachis and petiole. The fruits are red or when they come in contact with the ground.
dark pink when ripe (Acevedo-Rodriguez 1985,
Howard 1989, Liogier 1994). Growth and Management.—If needed for
establishment during environmental restoration
Range.—Bejuco de costilla is native to Cuba, projects, containerized plants can be easily and
Hispaniola, Puerto Rico, the Lesser Antilles, quickly grown from seed. However, no planting
Trinadad, Central America, South America, and projects are known. In forest management activity,
tropical Africa (Howard 1989, Liogier 1994). it is more often desirable to eliminate vines than
establish them. An effective method in closed
Ecology.—Bejuco de costilla is found primarily in forest stands is to cut the stems of vines near the
the moist (1000 to 2000 mm of annual rainfall) ground a year before harvest. Although the vines
forests. It is also reported to occur in subtropical resprout, most of the species, being shade
538
intolerant, die or are seriously weakened and do Howard, R.A. 1989. Flora of the Lesser Antilles,
not pose a serious threat to the reproduction stand. Leeward and Windward Islands. Vol. 5. Arnold
Herbicides can also be applied to the sprouts to Arboretum, Harvard University, Jamaica Plain,
make the method more effective, especially in MA. 604 p.
more open stands. Bejuco de costilla is one of the
alternate hosts of the tsetse flies (Glossina palpalis Liogier, H.A. 1990. Plantas medicinales de Puerto
and G. tachnioides) and has become the target of Rico y del Caribe. Iberoamericana de Ediciones,
eradication measures in some parts of tropical Inc., San Juan, PR. 566 p.
Africa (Morris 1944).
Benefits.—The stems of bejuco de costilla are Rico and adjacent Islands. Vol. 3. Editorial de la
used to make baskets. Ground leaves and seeds are Universidad de Puerto Rico, Río Piedras, PR.
used to stupefy fish and in preparing herbal 461 p.
remedies (Acevedo-Rodriguez 1985). The seeds
and leaves of this species are poisonous to humans Morris, K.R.S. 1944. A large-scale experiment in
and are reported to have been used in arrow the eradication of tsetse (Glossina palpalis and
poisons (Liogier 1990). Aqueous extracts of G. tachninoides). Farm and Forest 5: 149-156.
bejuco de costilla from Africa demonstrated
inhibitory effects against several important Souza, C. de, K.K. Amegavi, K. Koumaglo, M.
infectious organisms (Souza and others 1993). The Gbeassor, and C. De Souza. 1993. Study of the
white aril associated with the seed is reported to be antimicrobial activity of the total aqueous
edible (Vélez and Overbeek 1950). extracts of ten medicinal plants. Revue de
Medecines et Pharmacopees Africaines 7: 109-
References 115.
Acevedo-Rodriguez, P. 1985. Los bejucos de Vélez, I. and J. van Overbeek. 1950. Plantas
Puerto Rico. Vol. 1. General Technical Report indeseables en los cultivos tropicales. Editorial
SO-58. U.S. Department of Agriculture, Forest Universitaria, Río Piedras, PR. 497 p.
539
Pavonia spinifex (L.) Cav. cadillo espinoso
MALVACEAE
Synonyms: Hibiscus spinifex L.

Malache spinifex (L.) Kuntze
John K. Francis
Argentina (Griffiths 1994, New York Botanical

Garden 2002). Howard (1989) states that
Southeastern United States and the West Indies
are the only range; other claims are based on
unsubstantiated reports or misidentifications of
other species. Finally, Nelson (1996) refers to the
populations in Florida as escapes from cultivation.
Ecology.—Cadillo espinoso grows in dry areas. It

is found in Puerto Rico in habitat that receives
from 750 to about 1100 mm of precipitation and at
elevations from near sea level to about 600 m. The
species grows in well-drained soils with a wide
variety of textures and fertility levels. Cadillo
espinoso is more common in areas of igneous
rocks but does occur over limestone and other
sedimentary rocks. Cadillo espinoso is moderately
intolerant of shade. It grows in open areas but
more frequently under low, open stands of small
trees. The species competes well with tall grass,
herbs, and brush, except on the most fertile sites
General Description.—Cadillo espinoso, also where forest stands are dense. Probably because of
known as spiny abutilon, abutilon espinoso, mahot competition, the plant is rarely found in bottoms
jaune, and coquelicot, is a suffruticose shrub and north slopes. Most of the areas where the
usually 1 to 1.5 m in height, but occasionally species is common are recovering from past
reaching 5 m. The plant is supported by a shallow disturbance such as fire, severe overgrazing, or
root system consisting of a weak taproot, lateral clearing. Cadillo espinoso tolerates all but the
roots, and abundant fine roots. Older plants have most severe grazing. It sprouts after fires. The
multiple stems arising from the root crown and the species occurs as occasional plants or in mixture
lower 15 cm of stem. The plant dies back to the with other species, not as pure or dense stands.
main stem or sometimes to the root crown during
the dry season. The branches are slender and Reproduction.—Cadillo espinoso flowers during
relatively few in number. The alternate leaves are the summer (July and August). Fruits mature at
ovate with rounded or cordate bases and pointed the start of the dry season (January-February) and
tips. The leaf blades are 4 to 12 cm long and have are dispersed by sticking to the clothing or fur of
serrate edges. Bright, 2- to 3-cm yellow flowers animals. In addition to attaching directly from the
are usually axillary and solitary and develop into shrub, fruits rest on the ground so that the central
fruits that are schizocarps, dividing into five spine points up, so they may be picked up by the
mericarps. Each mericarp has three barbed spines, feet of passing animals. Seed production is reliable
the central spine, the largest, being about 1 cm and moderately abundant. Air-dried fruits
long (Howard 1989, Liogier 1994). collected in Puerto Rico averaged 0.0258 + 0.0005
g. Incubated in a plastic bag with moist peat, only
Range.—There is considerable confusion about 4 percent germinated in 10 months. Germination is
the extent of the native range of cadillo espinoso. epigeal.
It is reported to be native from Bermuda through
the West Indies and from Mexico to Peru and Growth and Management.—Annual growth is
540
rapid (about 1 m) from sprouts arising from References
established suffruticose plants. Growth begins
during the early summer rains and continues Griffiths, M. 1994. Index of garden plants. Timber
through the fall rainy season. The leaves and Press, Portland, OR. 1,234 p.
branches dry out and die during the late winter-
early spring dry season (in Puerto Rico). Howard, R.A. 1989. Flora of the Lesser Antilles,
Individual plants probably live at least a decade. Leeward and Windward Islands. Vol. 5. Arnold
Except for the spiny fruits that cling to clothing, Arboretum, Harvard University, Jamaica Plain,
there seems to be little need to control cadillo MA. 604 p.
espinosa in range or forest land. When necessary,
the species probably could be controlled by spot Liogier, H.A. 1994. Descriptive flora of Puerto
spraying with broadleaf weed killer to eliminate Rico and adjacent Islands. Vol. 3. Editorial de la
individual plants. The development of a closed Universidad de Puerto Rico, Río Piedras, PR.
canopy forest eventually eliminates the shrub. 461 p.
Benefits.—Cadillo espinoso is planted to a limited Liogier, H.A. 1990. Plantas medicinales de Puerto
extent as an ornamental for its handsome yellow Rico y del Caribe. Iberoamericana de Ediciones,
flowers. It contributes to the recovery of disturbed Inc., San Juan, PR. 566 p.
areas, furnishes cover for wildlife, and is eaten by
grazing animals. In herbal medicine, infusions of Nelson, G. 1996. The shrub and woody vines of
the flowers are used to treat sore throats and skin Florida. Pineapple Press, Inc., Sarasota, FL.
problems, and infusions of leaves and twigs are 391 p.
used to treat stomach problems, gall stones, and
liver pain (Liogier 1990). New York Botanical Garden. 2002. NYBG species
search results. http://s…/wwwspecimen.
search_list?taxon=Pavonia+spinifex+(L.)+Cav.+
+++&projcode=SEB. 1 p.
541
Pennisetum purpureum Schumacher elephant grass
POACEAE
Synonyms: none
John K. Francis
2n = 27, 28, or 56 (Burkill 1994, Holm and others

1977, Long and Lakela 1971, Skerman and
Riveros 1990).
Range.—Elephant grass is native throughout

humid, tropical mainland Africa and the island of
Bioko (Burkill 1994). It has been planted for
forage and has naturalized in many tropical areas
in Asia, the America’s, and Oceania. It grows wild
in the U.S. territories of Florida, Texas, California,
Hawaii, Guam, American Samoa, Puerto Rico,
and U.S. Virgin Islands (Natural Resource
Conservation Service 2002, Pacific Island
Ecology.—Elephant grass will grow on poorly

drained clay soils through the gamut of soil types
to excessively drained sandy soils. Growth is best
on rich, moist, well-drained medium-textured
soils. Soil reaction should range from pH 4.5 to
8.2 (Center for New Crops and Plant Products
2002). Rainfall should be in excess of 1500 mm
per year and temperatures for optimum growth
should be from 25 to 40 °C (Skerman and Riveros
1990). In its native range, the species is a fire sub-
climax to broadleaf tropical forest (Center for
New Crops and Plant Products 2002). Elephant
General Description.—Elephant grass is also grass will grow in light shade but is does not
known as napier grass, marker grass, Uganda survive under a closed tree canopy. In turn, it will
grass, yerba elefante, capim elefante, herbe à suppress most grasses, herbs, and tree seedlings. A
elephants, fausse-cane à sucre, and a host of native frost will kill the above-ground parts, but the soil
African names (Burkill 1994, Holm and others must be frozen to kill the rhizomes (Center for
1977). The species is a robust grass with New Crops and Plant Products 2002).
perennial stems. The plants produce short,
creeping rhizomes 15 to 25 cm long with fine Reproduction.—Flowering takes place mainly in
roots at the nodes and culms that are from 2 to 8 m the fall and winter (Long and Lakela 1971, Holm
in height, up to 2.5 cm in diameter at the base, and and others 1977). Because of asynchrony of male
have a solid center. Older culms may branch and female flower parts, the plant relies on cross-
several times. Leaf blades are 50 to 90 cm long pollination by wind. Elephant grass is an
and 1 to 3 cm wide, flat, and have a white midrib. inconsistent seed producer and rarely develops
Leaves of new, vigorous growth have wide, robust seeds in some habitats. When seeds are produced,
leaves; older culms have finer, narrow leaves. they are often of low viability (Holm and others
Leaf margins are rough (fine-toothed). The 1977). About 3.8 million seeds per kg were
inflorescence is a compact, erect, bristly tawny or reported for a United States source (Skerman and
purplish spike 8 to 30 cm long and 1.5 to 3 cm Riveros 1990). A collection of seeds from Puerto
wide. Spikelets are arranged around a hairy axis, Rico numbered 1.78 million per kg (author’s
and fall at maturity. The chromosome number is observation). The seeds are wind-dispersed.
542
Colonization of new habitats is slow (Skerman young shoots are eaten in soups and stews (Burkill
and Riveros 1990). 1994). Elephant grass is used for mulch in East
Africa where a 25-cm depth of mulch is needed
Growth and Management.—The growth of for good weed control (Nishimoto 1994). Extracts
culms is vertical continually, but their weight of the plant are strongly diuretic and are used for
bends them in the middle and lower part, causing a that purpose in Africa. It is also used in a number
j-shaped habit. As the lower stem makes contact of other herbal remedies (Burkill 1994). The seeds
with moist soil, it roots at every node. Individual are eaten by many bird species. However, because
culms live more than 1 year, and by layering, they of the aggressive spread of the species, it is a
can continue until disturbed or a barrier is reached. menace to native vegetation in the Galapagos
New plantations are established by planting stem Islands (Mauchamp 1997) and at the margins of
pieces with at least three nodes or with root swamps and streams in Florida (Miami-Dade
cuttings (Skerman and Riveros 1990). In an County 2002).
agricultural research plot in Puerto Rico, elephant
grass reached the height of 4 m in 3 months References
(Barrett 1925). The record yield for heavily
fertilized elephant grass is 84,800 kg/ha/yr Agus, F., D.K. Cassel, and D.P. Garrity. 1996.
(Skerman and Riveros 1990). Not-withstanding its Soil-water and soil physical properties under
value as forage, elephant grass has become one of contour hedgerow systems on sloping Oxisols.
the worst weeds in the tropics because of the Soil and Tillage Research 40(3-4): 185-199.
difficulty of controlling it in croplands and fallow
areas. Cultivation alone is usually insufficient to Barrett, O.W. 1925. The food plants of Puerto
control it in croplands (Skerman and Riveros Rico. Journal of the Department of Agriculture
1990). Frequent mowing will cause it to be of Puerto Rico 9(2): 193.
replaced by other grasses. The herbicide
glyphosate provides acceptable control in at least Burkill, H.M. 1994. The useful plants of West
aquatic sites (McCann and others 1996). Tropical Africa. Royal Botanic Gardens. Kew,
UK. 636 p.
Benefits.—Elephant grass makes up the bulk of
the diet of forest elephants in West Cameroon Center for New Crops and Plant Products. 2002.
(Tchamba and Seme 1993). The species is an Pennisetum purpureum K. Schumach. Perdue
important forage and pasture grass in its native University. http://hort.purdue.edu/newcrop/
Africa and throughout the Tropics, especially for duke_energy/Pennisetum_purpurium.html. 4 p.
cattle. It is also cut for hay and fermented for
silage. A number of forage samples of different Holm, L.G., D.L. Plucknett, J.V. Pancho, and J.P.
ages of grass from several countries varied from 4 Herberger. 1977. The World’s worst weeds.
to 15 percent in crude protein, from 28 to 40 East-West Center. Honolulu, HI. 609 p.
percent in crude fiber, from 10 to 16 percent in
ash, from 0.9 to 3.8 percent in fat, and from 39 to Karschon, R. and D. Heth. 1958. Wind speed,
49 percent in nitrogen-free extract (Skerman and wind-borne salt and agricultural crops as
Riveros 1990). The species has been employed affected by windbreaks. La Yaaran 8(3/4): 8-13,
successfully using sown cuttings to replace 32-38.
Imperata swards in the Phillippines (Agus and Long, R.W. and O. Lakela. 1971. A flora of
others 1996, Skerman and Riveros 1990). It is Tropical Florida. University of Miami Press,
planted as hedgerows for erosion protection and Coral Gables, FL. 962 p.
forage production in the alley cropping system of
agroforestry (Magcale-Macandog and others 1998, Magcale-Macandog, D.B., C.D. Predo, K.M.
Menz and others 1999). The plant is also effective Menz, and A.D. Calub. 1998. Napier grass strips
as a windbreak for agricultural crops (Karschon and livestock: a bioeconomic analysis.
and Heth 1958). Lines of plants are used to mark Agroforestry Systems 40(1): 41-58.
boundaries between plots and properties. Elephant
grass is planted on riverbanks to prevent erosion. Mauchamp, A. 1997. Threats from alien plant
In Africa, the plant is used for thatch, and the thick species in the Galapagos Islands. Conservation
culms are made into fences, screens, and Biology 11(1): 260-263.
reinforcement for mud huts. The young leaves and
543
McCann, J.A., L.N. Arkin, and J.D. Williams. Natural Resources Conservation Service. 2002.
1996. Noniddigenous aquatic and semi-aquatic Plants profile: Pennisetum purpureum
plants in freshwater systems. University of Schumacher. Washington, DC. http://plants.
Florida, Center for Aquatic Plants. usda.gov/cgi_bin/plant_profile.cgi?symbol=PEP
http://aquat1.ifas.ufl.edu/mctitle.html. [Not U2. 3 p.
paged].
Nishimoto, R.K. 1994. Weed control in coffee
Menz, K.M., D. Magcale-Macandog, and I.W. plantations. In: R. Labrada, J.C. Caseley, and C.
Rusastra, eds. 1999. Improving smallholder Parker, eds. Weed management for developing
farming systems in Imperata areas of Southeast countries. FAO Plant Production and Protection
Asia: alternatives to shifting cultivation. Paper 120. Food and Agriculture Organization
Australian Centre for International Agricultural of the United Nations. Rome. p. 354-359.
Research. Canberra, Australia. 280 p.
Pacific Island Ecosystems at Risk. 2002. Invasive
Miami-Dade County. 2002. Napier grass- plant species: Pennisetum purpureum
Pennisetum purpureum. http://co.miami- Schumacher, Poaceae. http://www.hear.org/
dade.fl.us/derm/environment/badplants/plant%2 pier3/pepur.htm. 2 p.
0desc…/napier_grass.ht. 1 p.
Skerman, P.J. and F. Riveros. 1990. Tropical
grasses. FAO Plant Production and Protection
Series 23. Food and Agriculture Organization of
the United Nations. Rome. 832 p.
Tchamba, M.N. and P.M. Seme. 1993. Diet and

feeding behavior of the forest elephant in the
Santchou Reserve, Cameroon. African Journal
of Ecology 31(2): 165-171.
544
Penstemon ambiguus Torr. sand penstemon
SCROPHULARIACEAE
Synonym: Leiostemon ambiguus (Torr.) Greene
James E. Nellessen
Ecology.—Sand penstemon occurs on alluvial

plains, valleys, and in sandy hill country. It is
considered one of the major shrub species
comprising Plains-Mesa Sand Scrub vegetation
that occurs along drainages of the Rio Grande,
Pecos River, San Juan River, and areas within the
Llano Estacado ("staked" plains) of western Texas
and eastern New Mexico (Dick-Peddie 1993). It is
common in the Painted Desert of Arizona
(Kearney and others 1951). A couple of hybrid
swarms between P. ambiguus and P. thurberi Torr.
(P. ambiguus x thurberi Nisbet and Jackson) are
known from: 1) an area near Magdalena, Socorro
General Description.—Sand penstemon is also County, and 2) another near Carrizozo, Lincoln
known by the names bush penstemon, prairie or County, both in New Mexico (Nisbet and Jackson
pink plains penstemon, moth penstemon, gilia 1960). P. thurberi has a similar shrubby habit and
penstemon, phlox penstemon, and cow-tobacco. occurs in sandy soils. Hybrids have shown a wide
This species occurs in sandy soils of plains, and range of characteristics between the two species.
mesas, and somewhat hilly terrain. The plant Although generally considered a separate species,
generally ranges from 20 to 60 cm in height, P. thurberi has also been classified as a variety of
sometimes to 1 m, is highly branched, the stems P. ambiguus (P. ambiguus Torr. var. thurberi
are slightly hairy, and become distinctly woody Gray).
well above the bases. Leaves are simple, linear,
opposite, 6 to 30 mm long, enrolled, smooth to Biochemical Ecology.—Several iridoid glucosides
slightly hairy, with slightly rough edges (Carter have been isolated and identified from sand
1997, Epple 1995, Great Plains Flora Association penstemon: catalpol, speciocide, nemoroside,
1986, Kearney and others 1951, Martin and ambiguuside; as well as two phenylpropanoid
Hutchins 1981). There are two recognized glycosides: verbascoside and martynoside
varieties: P. ambiguus var. ambiguus and P. (Arslanian and others 1990). Iridoids are a group
ambiguus var. laevissimus (Keck) N. Holmgren of bitter tasting terpenoid toxins also referred to as
(Kartesz 1994, Allred 2002). Both varieties have monoterpene lactones (Harborne 1988). These
also been classified as subspecies. Variety compounds may be toxic to birds and insects,
laevissimus differs from the typical in that it has hence functioning as a chemical defense against
smooth stems, with the leaf edges smooth or very herbivory.
slightly rough. The chromosome number is
reported as n = 8, which is common to many Reproduction.—Each plant has numerous many-
members of this genus (Nisbet and Jackson 1960). flowered inflorescences, and each inflorescence is
narrow, with one to two flowers per peduncle, the
Range.—This species occurs from eastern flowers spreading. The calyx is 2 to 3 mm long,
Colorado, western Kansas, Oklahoma, and Texas, the corolla 15 to 25 mm long, pale to deep pink,
through eastern and central New Mexico, then the five corolla lobes whitish, the upper ones
west into Arizona, Utah, and Nevada, at elevations reflexed at an oblique angle to the tube, the lower
from about 850 m to 2,000 m. Variety ambiguus one projecting forward, making it distinctive from
occurs in the northern and eastern portions of this most other penstemons. The opening of the tube is
range while variety laevissimus occurs in the bearded with short hairs, in two prominent
southern and western portions of this range. guidelines at the base of the throat (Nisbet and
Jackson 1960, Martin and Hutchins 1981, Carter
545
1997). The sterile stamen (staminode) is beardless Baskin, C.C. and J.M. Baskin. 2001. Seeds:
(hairless), a feature different from most ecology, biogeography, and evolution of
penstemons (beardtongues), hence the alternate dormancy, and germination. Academic Press,
genus name Leiostemon, leio meaning smooth and San Diego, CA. 666 p.
stemon for stamen (Kirkpatrick 1992, Weber and
Wittman 2001). It blooms from May to as late as Carter, J.L. 1997. Trees and shrubs of New
October, depending on local environmental Mexico. Johnson Books, Boulder, CO. 534 p.
conditions, although mass blooming generally
occurs from June through July. The hybrid, P. Dick-Peddie, W.A. 1993. New Mexico
ambiguus x thurberi has smaller corollas, 12 to 15 Vegetation: Past, Present and Future. University
mm long, corolla lobes white to blue-purple, the of New Mexico Press, Albuquerque, NM. 244 p.
throat and tube pale pink to deep reddish purple.
The fruit is a capsule from 5 to 10 mm long that Epple, A.O. 1995. A field guide to the plants of
contains many very small (1.2 to 2 mm) seeds. Arizona. Falcon Publishing Inc., Helena, MT.
Seed germination characteristics (for example, the 347 p.
amount of cold stratification) vary as a function of
the elevation of the source population (Meyer and Great Plains Flora Association. 1986. Flora of the
others 1995). Sand penstemon seeds have a Great Plains. University Press of Kansas, KS.
physiological dormancy requiring 8 to 15 weeks of 1,392 p.
cold stratification before germination will occur
(Kitchen and Meyer 1991, Baskin and Baskin Harborne, J. B. 1988. Introduction to ecological
2001). biochemistry, 3rd Ed. Academic Press, London.
356 p.
Growth and Management.—Seedlings take a
few years becoming established before this long- Haflin, J. 1997. Penstemons: the beautiful
lived plant becomes larger and more conspicuous breadtongues of New Mexico. Jackrabbit Press,
in the landscape. Only a few flowers may be Albuquerque, NM. 50 p.
produced the first year of blooming, before the
plant gains enough mass and root reserves for the Kartesz, J.T. 1994. A synonymized checklist of
large floral displays that make this plant so the vascular flora of the United States, Canada,
characteristic. and Greenland. Vol. 1, 2nd Edition. Biota of
North America Program of the North Carolina
Benefits.—Since this species’ typical habitat is in Botanical Garden. Timber Press, Portland, OR.
sandy, unstable type soils, it acts as a soil 622 p.
stabilizer. Because of its conspicuous and colorful
floral displays, it is a welcome addition to habitats Kearney, T.H., R. Peebles, and Collaborators.
that might otherwise appear drab. This is an 1951 with 1960 supplement. Arizona flora.
attractive plant for native plant landscaping University of California Press, Berkeley.
(Morrow 1995, Warnock 1974), and in full bloom 1,085 p.
with the numerous stems and flowers per stem, it
makes for an attractive display of pink bouquets. Kirkpatrick, Z.M. 1992. Wildflowers of the
Western Plains. University of Texas Press.
References Austin, TX. 240 p.
Allred, K.W. 2002. A working index of New Kitchen, S.G. and S.E. Meyer. 1991. Seed
Mexico vascular plant names. Available on a germination of intermountain Penstemons as
New Mexico State University web page. influenced by stratification and GA3 treatments.
http://web.nmsu.edu/~kallred/herbweb. [not Journal of Environmental Horticulture 9: 51-56.
paged].
Martin, W.C. and C.E. Hutchins. 1981. (reprinted
Arslanian, R.L., T. Anderson, and F.R. Stermitz. 2001). A flora of New Mexico. Vol. 2. Bishen
1990. Iridoid glucosides of Penstemon Singh Mahendra Pal Singh, India and Koeltz
ambiguus. Journal of Natural Products--Lloydia Scientific Books, Germany) p. 1,277-2,591.
53(6): 1,485-1,489.
546
Meyer, S.E., S.G. Kitchen, and S.L. Carlson. 1995. Warnock, B.H. 1974. Wildflowers of the
Seed germination timing patterns in Guadalupe Mountains and the Sand Dune
intermountain Penstemon (Scrophulariaceae). Country, Texas. Sul Ross State University,
American Journal of Botany 82: 377-389. Alpine, TX. 176 p.
Morrow, B.H. 1995. Best plants for New Mexico Weber, W.A. and R.C. Wittman. 2001. Colorado
gardens and landscapes. University of New flora: Eastern Slope. University Press of
Mexico Press, Albuquerque, NM. 267 p. Colorado, Boulder CO. 521 p.
Nisbet, G.T. and R.C. Jackson. 1960. The genus

Penstemon in New Mexico. The University of
Kansas Science Bulletin 41(5): 691-759.
547
Philadelphus lewisii Pursh Lewis mock orange
HYDRANGEACEAE
Synonyms: Philadelphus gordonianus Lindl.

Philadelphus cordatus Petz. & Kirchn.
Philadelphus grahami Petz. & Kirchn.
Philadelphus columbianus Koehne
Nancy L. Shaw
It is most common at mid-elevations but grows
from near sea level to 2,100 m (USDA Forest
Service 1937). Lewis mock orange exhibits wide
ecological amplitude, growing in communities
ranging from Pseudotsuga menziesii (Mirbel)
Franco and Sequoia sempervirens (Lamb. ex D.
Don) Endl. to Artemisia L., chaparral, and Pinus
contorta Dougl. ex Loud and Pinus ponderosa P.
& C. Lawson.
Ecology.—Lewis mock orange grows on soils

ranging from dry, rocky, gravelly loams on open
hillsides, to well-drained deep, rich alluvial loams
near riparian zones (USDA Forest Service 1937).
It is found in early to late seral as well as in climax
communities. It occurs as small thickets, isolated
plants, or in association with other shrubs on talus
Illustration credits: E.G. Hurd, and Hitchcock and others slopes, cliffs, canyons, or rocky hillsides and in
1961 transition zones of riparian areas (Hitchcock and
others 1961, Hopkins and Kovalchik 1983). It is
General Description.—Lewis mock orange, also most common on northern and eastern exposures
known as Indian arrowwood, Lewis syringa, mock (USDA Forest Service 1937). The species is
orange, syringa, or western syringa, was named for classified as fire resistant because it resprouts from
Captain Meriwether Lewis, the famous Western the root crown following burning. It also
explorer, who made the first collection of the regenerates from seeds that accumulate in the soil
species in 1806. The state flower of Idaho, Lewis seed bank (Kramer and Johnson 1987). Lewis
mock orange is a long-lived and highly variable mock orange exhibits moderate drought and shade
deciduous species, ranging from densely branched tolerance.
and rounded to erect, open shrubs 1.5 to 2.5 m in
height. Clusters of arching stems develop on older Reproduction.—Lewis mock orange flowers in
specimens. Bark on young branches is red to late spring to early summer. Capsules mature in
chestnut brown, turning gray and exfoliating with late summer and dehisce in September or October.
age. Leaves are 2 to 8 cm long and 1 to 4 cm wide, Seeds are dispersed by wind and gravity. They are
opposite, and ovate to oblong with acute to harvested by hand-stripping the capsules before
acuminate tips. Terminal cymes of showy, perfect, the valves begin to open. Dried capsules are
four-merous, fragrant white flowers develop on crushed to release the seeds, and trash is removed
lateral branches. The fruit is a woody, four-celled with an aspirator or fanning mill. There are
capsule containing numerous tiny, fusiform seeds 7,716,000 to 17,637,000 seeds/kg of clean seed
(Hitchcock and others 1961, Welsh and others (Stickney and others 2001), but seed fill is often
1987). low. Seed may be stored in airtight containers for
up to 1 year. Wet prechilling for 8 weeks at 5 °C is
Range.—Lewis mock orange is distributed from required to release embryo dormancy (Stickney
British Columbia south to northern California and and others 2001). Intermittent exposure to light
east to southwestern Alberta and western Montana. may be required for germination. Seeds may be
548
broadcast seeded on a rough seedbed and covered shafts from the long woody shoots (USDA Forest
using a Brillion seeder or similar device. They Service 1937).
may also be spot seeded in selected, prepared areas
that are well-drained and free of herbaceous References
competition. Seeds may be mixed with other shrub
seeds that require shallow or surface planting. Everett, P.C. 1957. A summary of the culture of
Bareroot stock may be produced by fall seeding or California plants at the Rancho Santa Ana
by seeding moist, prechilled seeds in spring. Botanic Garden. The Rancho Santa Ana
Seedlings develop rapidly and can be transplanted Botanic Garden, Claremont, CA. 223 p.
as 1-year-old stock (Stickney and others 2001).
Container stock is grown from seed or propagated Hitchcock, C.L., A. Cronquist, M. Ownbey, and
from hardwood or softwood cuttings (Marchant J.W. Thompson. 1961. Vascular plants of the
and Sherlock 1984). Rooted suckers and crown Pacific Northwest. Part 3: Saxifragaceae to
divisions are also used. Ericaceae. University of Washington Press,
Seattle, WA. 614 p.
Growth and Management.—Lewis mock orange
seedlings establish well on a wide variety of soils Hopkins, W.E. and B.L. Kovalchik. 1983. Plant
but should be protected from competition and associations of the Crooked River National
browsing. Plants grow at a moderate rate, flower Grassland. R6 Ecol. 133-1983. U.S.
reliably, and are generally free of insect and Department of Agriculture, Forest Service,
disease problems. Seedlings, however, are Pacific Northwest Region, Portland, OR. 98 p.
sometimes susceptible to damping off. Plants
grown from seed may begin flowering in the Kramer, N.B. and F.D. Johnson. 1987. Mature
second or third year (Everett 1957). Flowers are forest seed banks of three habitat types in
produced on twigs of the previous year; hence, central Idaho. Canadian Journal of Botany 65:
landscape plants should be pruned after flowering. 1961-1966.
Benefits.—Seedlings or larger planting stock of Kufeld, R.C. 1973. Foods eaten by the Rocky
Lewis mock orange are transplanted into steep, Mountain elk. Journal of Range Management
rocky, unstable slopes where they provide soil 26: 106-113.
stabilization and vegetative cover. The species is
also useful in transitional areas of degraded Leege, T.A. 1968. Prescribed burning for elk in
riparian zones. Lewis mock orange is usually not northern Idaho. Proceedings: Annual tall
grazed heavily by livestock, but in some areas it timbers fire ecology conference. 8: 235-253.
does receive fair amounts of use by cattle (Bos
spp. L.) and sheep (Ovis spp. L.) (Leege 1968, Marchant, C. and J. Sherlock. 1984. A guide to
USDA Forest Service 1937). It frequently occurs selection and propagation of some native
with other species that are more palatable to big woody species for land rehabilitation in British
game, and consequently, it may receive little use, Columbia. Forest Research Report RR84007-
except under severe conditions. However, it can HQ. Ministry of Forests, Victoria, BC. 117 p.
provide good browse for deer (Odocoileus spp.
Rafinesque) and elk (Cervus canadensis Stickney, P.F., N.L. Shaw, and E.G. Hurd. 2001.
Erxleben), especially on winter ranges (Kufeld Philadelphus L. Mockorange. In: F.T. Bonner
1973, Leege 1968, Marchant and Sherlock 1984, and R.G. Nisley, eds. Woody plant seed
USDA Forest Service 1937). New growth is manual. Agriculture Handbook. U.S.
generally highly palatable to big game (Leege Department of Agriculture, Forest Service,
1968). The species provides food and cover for Washington, D.C. http://wpsm.net/index.html.
birds and other small animals. Lewis mock orange 5 p.
is a valued landscape species because of its showy,
white flowers. It is used in borders, screens, Taylor, S. 1972. Philadelphus lewisii Pursh ‘mock
hedges, and as specimen plants (Marchant and orange’ or ‘syringa’. Davidsonia 3: 4-7.
Sherlock 1984). Several commercial cultivars of
Lewis mock orange have been developed. The USDA Forest Service. 1937. Range plant
flowers are used in preparing perfumes and teas handbook. U.S. Government Printing Office,
(Taylor 1972). Native Americans made arrow Washington, D.C. 512 p.
549
Welsh, S.L., N.D. Atwood, L.C. Higgins, and S.
Goodrich. 1987. A Utah flora. Great Basin
Naturalist 9. Brigham Young University, Provo,
UT. 894 p.
550
Philadelphus microphyllus Gray littleleaf mock orange
HYDRANGEACEAE
Synonyms: Philadelphus occidentalis A. Nels.

Philadelphus argenteus Rydb.
Philadelphus minutus Rydb.
Philadelphus nitidis A. Nels.
Philadelphus stramineus Rydb.
Nancy L. Shaw
numerous black, short-caudate seeds. The

chromosome number is 2n = 26 (Cronquist and
others 1997, Hickman 1993, Munz and Keck 1973,
Range.—Littleleaf mock orange is distributed

from southern California east into southern and
central Nevada; most of Utah; southwestern
Wyoming; western, central, and southern
California; Arizona; New Mexico; central Texas;
and central Mexico (Cronquist and others 1997).
The species exhibits considerable morphological
variability and has been subdivided into a number
of species or subspecific taxa by various authors.
Ecology.—Littleleaf mock orange grows in Pinus

spp. L.-Juniperus spp. L., Quercus spp. L.,
mountain brush, Populus tremuloides Michx.,
Pinus contorta Dougl. ex Loud., Pseudotsuga
menziesii (Mirb.) Franco, and Abies concolor
Lindl. communities at elevations from 1,200 to
3,100 m (Cronquist and others 1997, Hitchcock
1943, Welsh and others 1987). A drought-tolerant
species, it is often found growing in cracks or
fissures in rocky surfaces and in other dry, sandy,
gravelly, or rocky areas.
Illustration source: Cronquist and others 1997
Reproduction.—Plants flower in May to
General Description.—Littleleaf mock orange, September depending upon location. The capsules
also known as desert mock orange, is a small, mature in late summer. Mature capsules are
rounded, multistemmed shrub 0.9 to 2.1 m tall harvested by hand just after they have begun to
with a crown diameter of about 1.2 m. New woody dehisce. Dried capsules are crushed with a barley
growth is strigose, while bark of mature stems is debearder to permit seed extraction. Seed is
gray to yellowish and exfoliating. Some stems may separated from debris using an aspirator or air-
be subspinose. Leaves are opposite, entire, short screen machine. Littleleaf mock orange is readily
petiolate, 8 to 25 mm long, 2 to 13 mm wide, propagated from seed (Sutton and Johnson 1974,
elliptic to lanceolate or linear, and sometimes Swingle 1939). Germination of a New Mexico
revolute. Flowers are solitary or in clusters of three seed collection was 12 times greater when
at the ends of branches produced the previous incubated at 15 compared to 20/10 oC (8 hrs/16
year. They are showy, white or creamy, fragrant, hrs) for 28 days (Stickney and others 2001).
and four-merous with numerous stamens. Fruits Germination was improved by a 28-day moist
are four-chambered loculicidal capsules containing
551
prechill at 3 to 5 oC when seeds were subsequently Hickman, J.C. 1993. The Jepson manual: higher
incubated at 20/10 oC, but not at 15 oC. plants of California. University of California
Germination is epigeal. Press, Berkeley, CA. 1,400 p.
Growth and Management.—Little data are Hitchcock, C.L. 1943. The xerophyllous species of
available on the propagation of littleleaf mock Philadelphus in southwestern North America.
orange, but species of the genus Philadelphus are Madroño 17: 35-56.
generally easily propagated from softwood or
hardwood cuttings, rooted suckers, divisions, or Macdonald, B. 1986. Practical woody plant
layers (Hartmann and others 1990, Macdonald propagation for nursery growers. Vol. I. Timber
1986). Littleleaf mock orange is easily Press, Portland, OR. 669 p.
transplanted (Sutton and Johnson 1974). Seeds of
Philadelphus spp. are tiny and may be broadcast Munz, P.A. and D.D. Keck. 1973. A California
seeded on a rough seedbed and covered lightly or flora and supplement. University of California
spot seeded on prepared seedbeds (Stevens and Press, Berkeley, CA. 1,681 p.
others, in press). Seeds may also be surface-
planted using a Brillion or similar seeder. Best
results are obtained if seeds are planted in well- Patton, D.R. and M.G. Ertl. 1982. Run wild
drained sites free of herbaceous competition. wildlife/habitat relationships. Wildlife Unit
Seeds may be mixed with other shrub seeds that Technical Series. U.S. Government Printing
require surface or shallow planting. Growth of Office, Washington, D.C. 49 p.
littleleaf mock orange is moderate to rapid (Sutton
and Johnson 1974). Rehder, A. 1940. Manual of cultivated trees and
shrubs. 2nd Ed. The MacMillan Co., New York.
Benefits.—Littleleaf mock orange provides good 996 p.
cover for small animals. It is used to some extent
by mule deer (Odocoileus hemionus Rafinesque) Stevens, R., S.B. Monsen, and N.L. Shaw. [in
(Patton and Ertl 1982). Although the species press]. Shrubs of other families. In: S.B. Monsen
receives little use in revegetation, it has potential and R. Stevens, comps. Restoring western
for planting on disturbances on steep, rocky, ranges and wildlands. INT-GTR. U.S.
unstable slopes within its native range (Stevens Department of Agriculture, Forest Service,
and others, in press). Seedlings or larger stock are Rocky Mountain Research Station, Ogden, UT.
recommended for such use. The species might
also be used to advantage in drier areas of Stickney, P.F., N.L. Shaw, and E.G. Hurd. 2001.
degraded riparian zones. Littleleaf mock orange is Philadelphus L. Mockorange. In: F.T. Bonner
an attractive ornamental because of its showy and R.G. Nisley, eds. Woody plant seed manual.
flowers and fall coloration. It was first cultivated U.S. Department of Agriculture, Forest Service,
in 1883 (Rehder 1940). It can be used in borders, Agriculture Handbook. Washington, D.C.
screens, hedges, or as isolated specimens in sunny http://wpsm.net/ index.html. 5 p.
areas. It can also be used for low maintenance
landscaping and in recreational area plantings. The Sutton, R. and C.W. Johnson. 1974. Landscape
small seeds were eaten by Native Americans plants from Utah’s mountains. EC-368. Utah
(Cronquist and others 1997). State University, Logan, UT. 137 p.
References Swingle, C.F. 1939. Seed propagation of trees,

shrubs and forbs for conservation planting. SCS-
Cronquist, A., N.H. Holmgren, and P.K. TP-27. Washington, DC: U.S. Department of
Holmgren. 1997. Intermountain flora. Volume 3, Agriculture, Soil Conservation Service. 198 p.
Part A. New York Botanical Garden, Bronx,
NY. 446 p. Welsh, S.L., N.D. Atwood, L.C. Higgins, and S.
Goodrich. 1987. A Utah flora. Great Basin
Hartmann, H.T., D.E. Kester, and F.T. Davies, Jr. Naturalist 9. Brigham Young University, Provo,
1990. Plant propagation principles and practices. UT. 894 p.
5th Ed. Prentice-Hall, Inc., Englewood Cliffs,
NJ. 657 p.
552
Phoradendron quadrangulare (Kunth) Griseb. guacimilla de canario
VISCACEAE (sometimes placed in LORANTHACEAE)
Synonyms: Phoradendron randiae (Bello) Britt.

Loranthus quadrangularis Kunth in Humb.
Viscum quadrangulare (Kunth) DC.
Phoradendron rubrum sensu Griseb.
Viscum trigonum D. Dietr.
John K. Francis
Ecology.—Guacimilla de canario colonizes a

number of tree species, especially Guazuma
ulmifolia Lam. in Puerto Rico (author’s
observation). The former species, as well as
Amphitecna latifolia (Mill.) A.H. Gentry, Cordia
stellifera I.M. Johst., Erythrina sp., Malvaviscus
arboreus Cav., and Persea Americana Miller, are
mentioned as hosts in a Costa Rican herbarium
collection (Missouri Botanical Garden 2002).
Most of the guacimilla de canario plants occur in
the mid-crown of the affected trees. After the
parasite becomes general throughout a tree crown
(a process that takes many years), trees are
weakened and sometimes die as a result. More
often, parasitized trees die of other causes or old
General Description.—Guacimilla de canario is age before they succumb to the parasite. Trees in
an evergreen, epiparasitic shrub that obtains water, forest, rangeland, farmland, roadside, and cities
minerals, and support from its tree hosts and are attacked. Guacimilla de canario grows at lower
produces stems up to 1 m long and 1 cm thick. and middle elevations in Puerto Rico (Liogier
The woody stems are much branched, brown, and 1985), that is, from sea level to about 600 m.
lightly furrowed in their lower parts with a These areas range from 750 to about 2400 mm of
swelling at the point of attachment to the host. The annual precipitation. In Ecuador, the plant may be
bark of branches and twigs is green. The young found from near sea level to 1,000 m in elevation
branches especially are square or rhombic in cross (Jorgensen and León-Yánez 1999).
section. The stem wood is light-colored and
brittle. The dull green leaves have a leathery- Reproduction.—Fruits collected in Puerto Rico
fleshy texture. They are linear, lanciolate, or weighed an average of 0.0294 + 0.0004 g/fruit.
oblanciolate with a tapered base and acute to Seeds cleaned from those fruits weighed (air-
obtuse apex. Inflorescences are small spikes, one dried) an average of 0.0017 + 0.0000 g/seed or
to three in number at the leaf axils, and support 600,000 seeds/kg. Sown without any pretreatment
small flowers. The yellow, globose fruits are 3 to on moist filter paper, 59 percent germinated within
4 mm in diameter (Howard 1988, Liogier 1985). 35 days of sowing. After 4 days, mesoscopic roots
However, fruits in Costa Rica range from pale emerged followed at 17 days by a shoot that
yellow to red-orange (Missouri Botanical Garden extended hypogenally (author’s observation).
2002). Within the pigmented skin is a layer of Birds transport the seeds (the fruits are sticky and
sticky jelly and a flat, green seed with longitudinal adhere to their feet or beaks) to new trees or
striations (author’s observations). stands. After fruits stuck to the bark of a suitable
host tree, they require a rainy period long enough
Range.—Guacimilla de canario is native to to allow the roots of the germinating seed to
Central America, South America, and the West penetrate the phloem of the host, or the seed or
Indies (Howard 1988, Liogier 1985). plantlet will desiccate. Once established in a new
tree, the guacimilla de canario spreads by seeds
553
within the crown of the host tree. It does not References
spread vegetatively.
Growth and Management.—The growth rate of Leeward and Windward Islands.
guacimilla de canario is not known. Although it is Dicotyledoneae, Part 1. Vol. 4. Arnold
a parasite and does harm its hosts, the plant takes Arboretum, Harvard University, Jamaica Plain,
many years to become sufficiently abundant to MA. 673 p.
seriously weaken a host tree. Guazuma ulmifolia
and most of the other hosts are not used as Jorgensen, P.M., and S. León-Yánez. 1999.
ornamentals and are not commercially valuable. If Catalogue of the vascular plants of Ecuador.
control is desired, probably the best method is to Missouri Botanical Garden, St. Louis, MO.
clip the guacimilla de canario plants from the 1,181 p.
limbs each year or until they die. This will prevent
it from bearing fruit and spreading within the tree. Liogier, H.A. 1985. Descriptive flora of Puerto
Alternately, one may prune out the infected limbs Rico and adjacent islands, Spermatophyta. Vol.
of the parasitized tree. 1. Editorial de la Universidad de Puerto Rico,
Benefits and Detriments.—Guacimilla de
canario contributes in a minor way to biodiversity, Missouri Botanical Garden. 2002. Manual de la
to tree canopy density, and to biomass production. flora de Costa Rica. http://www.mobot.
It furnishes fruits for wildlife food. On the other org/manual.plantas/040567/S040679.html. 4 p.
hand, a few trees/km2 are attacked, some are
weakened, and an occasional tree dies as a result.
Parasitized trees are often unsightly.
554
Phragmites australis (Cav.) Trin. ex Steud. common reed
POACEAE
Synonyms: Phragmites communis Trin.

Phragmites phragmites (L.) Karst.
John K. Francis
flowered. The panicles are initially grayish purple,

becoming whitish, and finally weathering to light
brown. There are 2n = 36, 48, and 54
chromosomes (Duke 1998, Gould 1975,
Harrington 1964, Uchytil 1992, Welsh and others
1987).
Range.—Common reed occurs in every state in

the Continental United States except Arkansas
(probably present but undocumented), in Puerto
Rico (Natural Resources Conservation Service
2003), southern Canada, and Mexico. In fact it is
found on all continents except Antarctica
(Harrington 1964). It is native to the United States
and is known from the archeological record.
However, genetic analysis has shown that more
aggressive European genotypes have replaced the
native genotype in much of the former American
range and have invaded new areas (Blossey 2002).
Ecology.—Common reed is a warm-season grass

that starts its growth after the last frost has
occurred but remains green until frost in temperate
areas (Uchytil 1992). Wide climatic tolerance is
indicated by its range that stretches from the
tropics to cold temperate areas and habitat from
sea level to elevations of at least 1,980 m
(Harrington 1964, Welsh and others 1987).
Common reed typically forms pure stands that
Drawing source: Britton and Brown 1913 contain up to 205 stems/m2 (Uchytil 1992). In Gulf
Coast marshes, it is often codominant with
General Description.—Common reed is also Spartina cynosuroides (L.) Roth. (Duke 1998).
known as giant reed, giant reedgrass, roseau, The species grows on the shores of streams and
roseau cane, yellow cane, and cane (Uchytil 1992). lakes, and in marshes, both fresh and brackish.
It is a shrub or perennial woody graminoid herb Although common reed grows best in clay, it
that grows annually from woody rhizomes to occurs in a wide variety of soils. The species
heights of 2 to 4 m (occasionally to 6 m). The tolerates flooding, frost, high pH, salt, and weeds
normally unbranched, hollow, jointed stems (Duke 1998). Stands in the Nile River Delta
(culms) reach nearly 2.5 cm in diameter but tolerate soil pH’s of 7.0 to 9.3 (Serag 1996).
usually measure 1.0 to 1.5 cm in diameter. In Standing dead material often totals twice as much
addition to rhizomes, plants occasionally produce biomass as current growth, allowing stands to burn
stolons. Alternate leaves are glabrous except for even during the growing season. New stands arise
being strongly scabrous at the margins. They are from rhizomes that are under the soil or water. The
flat, 1 to 5 cm wide, and up to 50 cm long. species also tolerates late season drought (author,
Terminal plume-like panicles are 15 to 50 cm long personal observation). Moderately deep rooting (to
with branches ascending to nodding and densely 1 m, Uchytil 1992) undoubtedly imparts drought
555
resistance. A large number of fungal species and at sooth lung pain, especially during pneumonia, and
least one nematode have been isolated from the to treat stomach problems and diarrhea (Moerman
species, none of which cause extensive damage 1986). There are numerous other applications in
(Duke 1998). herbal medicine throughout the world. The stems
were used as arrow shafts by Native Americans
Reproduction.—Common reed blooms mostly (Duke 1998). Native people of Tasmania used the
from July through November (Diggs and others hollow stems to make rafts, jewelry, baskets, and
1999). It apparently blooms and fruits throughout light spears (Australian National Botanic Gardens
the year in frost-free areas (Gould 1975). Under 2003). Common reeds are harvested today in
favorable artificial conditions, germination begins Britain for thatching (Tyler-Walters 2002). They
1 day after sowing and may reach nearly 100 were once harvested for fuel where better sources
percent (Ekstam and Forseby 1999). Although were unavailable and may hold promise as an
common reed is a relatively good seed producer energy crop in the future. Stands yield 415
and seeds are added annually to the soil seed bank, calories/100g (Duke 1998). Early in the growing
successful seedling establishment is rare (Uchytil season, common reed is high-quality forage for
1992). New stands can also be started by flood- cattle and horses and may be cut for hay. It
borne pieces of rhizome. Once a new plant is contains 11.4 percent protein, 2.3 percent fat, 42.1
established, it spreads by vegetative means. percent carbohydrates, 31.1 percent crude fiber,
and 10.8 percent ash (Duke 1998). It becomes
Growth and Management.—Because stems are woody and unpalatable later in the season. The
renewed each growing season, annual height seeds are eaten by waterfowl and the roots are
growth is from 2 to 4 m per year. Growth can eaten by muskrats. The edges are occasionally
reach 4 cm/day. In southern Manitoba, stems reach used for nesting by ducks and the interior of the
their full height by the end of July (Uchytil 1992). stands are nesting habitat for snowy egrets, black-
During periods of falling water, colonies can crowned night herons, and yellow-headed
increase in width by as much as 15 m in a single blackbirds. A few species such as deer use the
season (Fewless 2003). Annual productivity ranges stands for escape cover (Uchytl 1992). It is
from 7.5 to 63 metric tons/ha (Duke 1998). If important habitat for wildlife in Britain and
harvested during the dormant season, productivity supports such species as the bittern, the reed
is not affected, except through compaction. New bunting, and the marsh harrier (Tyler-Walters
stands are established by planting rhizome 2002). Throughout the world the stems have been
segments (Duke 1998). This is normally not used for building materials, lattice work, baskets,
recommended because of the invasive nature of the and mats. The young shoots and rhizomes are
species. Eliminating stands is difficult. The best sometimes eaten, and the rhizomes processed for
approaches are heavy grazing by cattle, and starch. The stems are processed into pulp, paper,
spraying with amatrol, dalapon, or glyphosate with and fiberboards. Stalks contain over 50 percent
follow-up treatments. Stands can be degraded by cellulose and have fibers 0.8 to 3.0 mm long by
cultivation or summer mowing (Uchytil 1992). 5.0 to 30.5 µm in diameter. The dried plumes are
used for decorations and a variegated variety is
Benefits and Detriments.—Common reed employed as an ornamental (Duke 1998).
aggressively colonizes large areas of shoreline and
shallow marsh, and with its thick sod and heavy References
stand of stems effectively prevents wave- and
current-caused erosion (Uchytil 1992). It also traps Australian National Botanic Gardens. 2003.
sediment and pollutants. In fact the species has Aboriginal plant use in South-eastern Australia:
been used to aid in settling and drying sewage Phragmites australis. http://www.anbg.gov.
sludge (Graduate College of Marine Studies 2003). au/aborig.s.e.aust/phragmites-australis.html. 2 p.
The dense single-species stands completely
displace other native marsh communities (Fewless Blossey, B. 2002. Native to North America or
2003) and many of the fauna they support introduced (or both)? Ag Web, Cornell
(Weinstein and Balletto 1999). On the other hand, University, Ithaca, NY. http://www.
strong declines in populations in many European invasiveplants.net/phragmites/phrag/natint.htm.
countries have occurred during the last two 3 p.
decades (Clevering 1998). Native Americans used
the sap from green stems to loosen phlegm and
556
Britton, N.L. and A. Brown. 1913. Illustrated flora Harrington, H.D. 1964. Manual of the plants of
of the northern states, Canada, and the British Colorado. Sage Books, Denver, CO. 666 p.
possessions. Vol. 2, 2nd Ed. Scribner, New York.
735 p. Moerman, D.E. 1986. Medicinal plants of Native
America. Technical Reports. 19. University of
Clevering, O.A. 1998. An investigation into the Michigan Museum of Anthropology, Ann
effects of nitrogen on growth and morphology of Arbor, MI. 534 p.
stable and die-back populations of Phragmites
australes. Aquatic Botany 60(1): 11-25. Natural Resources Conservation Service. 2003.
Plants profile: Phragmetes australis (Cav.) Trin.
Diggs, G.M., Jr., B.L. Lipscomb, and R.J. ex Steud. http://plants.usda.gov/cgo_bom/
O’Kennon. 1999. Thinner’s and Mahler’s plant_search.cgi?mode=Scientific+Nave&keyw
Illustrated flora of North Central Texas. ordquery=Phragmites+australis&earl=plant_s…
Botanical Research Institute of Texas, Fort 6 p.
Worth, TX. 1,622 p.
Serag, M.S. 1996. Ecology and biomass of
Duke, J.A. 1998. Handbook of energy crops: Phragmites australes (Cav.) Trin. ex Steud. In
Phragmites australis (Cav.) Trin. ex Steud. the north-eastern region of the Nile Delta,
Center for New Crops and Plants Products, Egypt. Ecoscience 3(4): 473-482.
Purdue University, West Lafayette, IN. http://
www.hort.purdue.edu/newcrop/duke_energy/ Tyler-Walters, H. 2002. Phragmites australis,
phragmites_australis.html. 5 p. common reed. Marine Life Information
Network, Plymouth, UK. http://www.marlin.
Ekstam, B. and A. Forseby. 1999. Germination ac.uk/species/Phraus.htm. 4 p.
response of Phragmites australis and Typha
latifolia to diurnal fluctions in temperature. Seed Uchytil, R.J. 1992. Phragmites australis. In: U.S.
Science Research 9(2): 157-163. Department of Agriculture, Forest Service,
Fewless, G. 2003. Invasive plants of Wisconsin: Laboratory, Fire Effects Information System.
Phragmites australis: common reed. http:// http://www.fs.fed.us/database/feis/plants/gramin
www.uwgb.edu/biodiversity/herbarium/invasive oid/phraus/all.html. 19 p.
_species/phraus01.htm. 2 p.
Weinstein, M.P. and J.H. Balletto. 1999. Does the
Gould, F.W. 1975. The grasses of Texas. Texas common reed, Phragmites australis, affect
A&M University Press, College Station, Texas. essential fish habitat? Estuaries 22(3B): 793-
653 p. 802.
Graduate College of Marine Studies. 2003. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
Optimized reed (Phragmites australis) function Higgins, eds. 1987. A Utah flora. Great Basin
in sludge drying beds: producing a more Naturalist Memoirs 9. Brigham Young
effective genotype. University of Delaware, University, Provo, UT. 894 p.
Newark, DL. http://www.ocean.udel.edu/
level1/facultystaff/faculty/dseliskar/sludgebeds.
html. 3 p.
557
Physocarpus malvaceus (Greene) Kuntze mallow ninebark
ROSACEAE
Synonyms: Spiraea oputifolia var. pauciflora Torr. & Gray

Neillia malvaceae Greene
Opulaster malvaceus Kuntze
Physocarpus pauciflorus Piper
John K. Francis
Range.—Mallow ninebark is native to British

Columbia, Alberta, Washington, Oregon, Idaho,
Montana, Wyoming, Colorado, Utah, and Nevada
(Abrams 1950, Natural Resources Conservation
Service 2003). It is not known to have naturalized
Ecology.—Mallow ninebark grows on a wide

variety of soils derived from igneous,
metamorphic, and calcareous and noncalcareous
sedimentary rocks. Textures range from sandy
loams to silty clay loams and surface pH ranges
from 5.5 to 7.1. In Idaho and Montana, the average
duff layer thickness was 6.5 cm with an effective
rooting depth of 38 cm. In northern areas, mallow
ninebark is found mainly in areas with no exposed
rock on mountain slopes, ridges, and streambanks
(Habeck 2003); in Utah, it often grows in skeletal
soils and rubble in steep canyons and
mountainsides. It occurs at elevations of 1,600 to
3,300 m at the southern end of its Utah range
(Welsh and others 1987) and at somewhat lower
elevations farther north. The species is associated
General Description.—Mallow ninebark, also with Abies lasiocarpa (Hook.) Nutt., Abies grandis
known as ninebark, mallow-leaved ninebark, (Dougl. ex D. Don) Lindl., and Picea engelmannii
mountain ninebark, few-flowered ninebark, and Perry ex Engelm. in cool/moist habitat and with
shallow ninebark, is a multistemmed, deciduous Pseudotsuga menziesii (Marbel) Franco and Pinus
shrub 1 to 2 m (rarely 3 m) in height. The stems ponderosa P. & C. Lawson in warmer, moist sites
and branches are slender and covered with a brown (Habeck 2003). While associated with these trees,
bark that exfoliates in papery strips on older mallow ninebark usually grows in disturbance-
branches. The 3- to 5-cm, alternate leaves are caused openings or diffuse stands. It also grows on
round to ovate, three to five lobed, and doubly nonforested shrubland sites. Some common shrub
serrate or crenate. The petioles are 1 to 2 cm long. associates are Holodiscus discolor (Pursh)
Inflorescences are terminal, corymbose with five Maxim., Symphoricarpos albus (L.) Blake,
to 30 flowers. The hypanthium of the flowers is Symphoricarpos oreophilis Gray, Spirea butilifolia
stellate-pubescent; twigs, petioles, and leaf blades Pallas, Amelanchier alnifolia (Nutt.) Nutt. ex M.
vary from pubescent to glabrous. The flowers have Roemer, and Mahonia repens (Lindl.) G. Don
five sepals and five white, rounded petals 3.5 to (Habeck 2003). Mallow ninebark is fire resistant,
6.5 mm long. Each produces two inflated, brown sprouting after topkill from surviving root crowns
follicles united for two-thirds their length. Each or horizontal rhizomes (Palouse Prairie Foundation
fruit may contain one to several shiny, pair- 2003). Having 36 to 99 percent of its roots buried
shaped, straw-colored seeds (Abrams 1950, Welsh in mineral soil enables it to escape the heat of
and others 1987). surface fires (Habeck 2003). Mallow ninebark
558
stands often owe their existence to logging or fires. sometimes appear later and will have to be sprayed
The species is an obligate pioneer that increases again (Habeck 2003).
and grows rapidly following disturbance, but as
competition, particularly overhead shade, Benefits.—Mallow ninebark helps protect the soil
increased, it decreases in mean height and during secondary succession until the shrub is
gradually disappears (Habeck 2003). The species replaced by taller vegetation. It adds beauty to
is generally free of insect and disease problems, forest openings and shrublands when its leaves
although it is occasionally attacked by foliage- turn russet-red in the fall (Clark 1976). The species
eating and seed-eating insects (Youngblood and provides cover for both large and small animals. It
others 2003). is one of the principal nesting sites of the dusky
flycatcher (Empidonax oberholseri) (Sedgwick
Reproduction.—Flowering takes place between 1993). The shrub is not browsed by cattle and is
late May and late July, depending on altitude, and utilized only to a limited extent by deer, bighorn
fruits ripen between late August and late sheep, and domestic sheep. The upper third of the
September (Habeck 2003). Seed weights are annual growth of plants from a burned area
reported (geographic source unspecified) at contained 1.3 percent N, 0.3 percent P, and 1.2
165,000 seeds/kg (Youngblood and others 2003). percent Ca. Plants in an unburned site contained
Seeds collected by the author in Utah averaged even lower levels (Habeck 2003). Early summer
158,000 seeds/kg. Viability is usually less than 50 sheep grazing increases the nutritional quality of
percent (Youngblood and others 2003). The fruits mallow ninebark for fall and winter browse (Alpe
split open at maturity but most of the seeds do not and others 1999).
immediately fall. Dispersal is by gravity, wind,
and probably by browsing animals. Seeds in the References
soil seedbank are reported to be 11 percent viable
and will germinate when the surface is scarified Abrams, L. 1950. Illustrated flora of the Pacific
(Habeck 2003). Ripe fruits can be picked by hand States. Vol. 2. Stanford University Press,
or beaten off shrubs onto a tarp. Fruits may be Stanford, CA. 635 p.
threshed by rubbing for small quantities or passed
through a hammer mill for larger quantities and Alpe M.J., J.L. Kingery, and J.C. Mosley. 1999.
screened and blown to clean them. They may be Effects of summer sheep grazing on browse
stored cool and dry for at least 5 years. Seeds may nutritive quality in autumn and winter. Journal
be sown in the fall, or in the spring after 30 days of of Wildlife Management 63(1): 346-354.
prechilling. Seeds should be sown to a depth of 3
mm and mulched with another 6 mm of sawdust Clark, L.J. 1976. Wild flowers of the Pacific
(Youngblood and others 2003). Mallow ninebark Northwest. Gray’s Publishing Limited, Sidney,
can easily be vegetatively reproduced with misted BC, Canada. 604 p.
softwood cuttings or by planting hardwood
cuttings directly in the field (Youngblood and Habeck, R.J. 2003. Physocarpus malvaceus. In:
others 2003). U.S. Department of Agriculture, Forest Service,
Growth and Management.—Mallow ninebark Laboratory. Fire Effects Information System.
shrubs in northern Idaho reached their maximum http://www.fs.fed.us/database/feis/plants/shrub/
height by the fourth growing season after a fire phymal/all.html. 17 p.
(Habeck 2003). Although stems are added and lost
continuously, shrubs do not grow taller. Individual Natural Resources Conservation Service. 2003.
plants may live for many years if growing Plant profile: Physocarpus malvaceus (Greene)
conditions permit. Mallow ninebark is sometimes Kuntze. Plants database. http://plants.usda.gov/
planted for soil stabilization and other cgi_bin/plant_profile.chi?symbol=PHMA5. 3 p.
conservation objectives. Killing the shrubs to
make room for more desirable species is Palouse Prairie Foundation. 2003. Palouse Prairie
sometimes attempted. Several herbicides can be Foundation native species search results:
effective if sprayed early in the season from late Physocarpus malvaceus. http://www.
foliar development to flowering or in early fall palouseprairie.org/readppf.pl?entry=PHMA5.
before leaves change color. Sprouts will 3 p.
559
Sedgwick, J.A. 1993. Reproductive ecology of Youngblood, A.Y., J.D. Gill, and F.L. Pogge.
dusky flycatchers in western Montana. Wilson 2003. Physocarpus (Camb.) Raf. U.S.
Bulletin 105(1): 84-92. Department of Agriculture, Forest Service,
Washington, DC. http://wpsm.net/Physocarpus.
Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C. pdf. 5 p.
Higgins, eds. 1987. A Utah flora. Great Basin
560
Picramnia pentandra Sw. Florida bitterbush
SIMAROUBACEAE
Synonyms: Tariri pentandra (Sw.) Baill.

Picramnia antidesmoides Griseb.
Picramnia cubensis Turcz.
Picramnia micrantha Tulasne
Picramnia oblongifolia Turcz.
John K. Francis
on different trees (dioecious) in branched terminal

clusters. The tiny flowers are five-parted, green
with a reddish tinge. Fruits (berries) are scarlet to
almost black when ripe, oblong to obovoid, 9 to 15
mm long, and contain one to three shiny brown
seeds 6 to 10 mm long (Howard 1988, Liogier
1988, Little and Wadsworth 1964, Long and
Lakela 1976).
Range.—Florida bitterbush is native to southern

Florida, the Bahamas, the West Indian Islands, and
Colombia and Venezuela in northern South
America (Howard 1988, Little and Wadsworth
1964). It is not known to have naturalized outside
its native range.
Ecology.—Florida bitterbush occurs in coastal

hammocks in southern Florida and the Keys (Long
and Lakela 1976). It grows in both remnant forests
and secondary forests (logged, pastured, and
abandoned fields) in Puerto Rico. During old-field
or pasture succession, Florida bitterbush invades
General Description.—Florida bitterbush is also during the brushy stage. The species adapts to a
known as bitterbush, doctor-bar, wild-coffee, wide variety of well-drained soils derived from
guarema, hueso, aguedita, palo de peje, palo de most parent materials. It usually does not grow on
paz, roble agalla, quina del país, bois poison, the most fertile sites (where competition is severe)
vaillant garcon, and bois montagne. It is an or on the worst, such as eroded, rocky ridge tops.
evergreen shrub or small tree usually less than 6 m Elevations vary from near sea level to 600 m or
in height and 10 cm in stem diameter. The plant more, and mean annual precipitation ranges from
usually has a single slender stem with smooth, 750 mm to about 2400 mm. Florida bitterbush is
gray bark and relatively few, slender branches. The moderately intolerant to shade, being able to grow
inner bark is brown. The wood is light colored, under the canopy of low basal-area forests as well
hard, and heavy. The inner bark, roots, leaves, as in openings. Seedlings are apparently more
fruits, and seeds have a strongly bitter taste. The tolerant than adults and are able to progress from
alternate, compound leaves are 10 to 30 cm long the understory into the canopy of low forests
and contain five to nine, usually seven, 5- to 10- (Pascarella 1996).
cm leaflets. The elliptic to ovate, papery leaflets
have short stalks, smooth edges, and long points. Reproduction.—Florida bitterbush flowers and
The side leaflets are subopposite. The leaflets are fruits nearly throughout the year in Puerto Rico
green tinged with red, have a net-like structure of (Little and Wadsworth 1964). It blooms March
fine veins when viewed against the light, and are through July in Florida. Flowers of both sexes
shiny when dried. Male and female flowers occur attract small, generalist insect pollinators with
scent, pollen, and small amounts of nectar. Peak
561
ripening occurs in November and December and fungi were treated with ethanol extracts of
(Pascarella 1996). Fruit and seed production is various tissues. Growth reductions of Bacillus
moderately abundant and consistent from year to cereus, Bacillus subtilis, and Pseudomonas
year. Fresh fruits collected in Puerto Rico weighed aeruginosa were observed which was enhanced by
an average of 0.231 + 0.043 g/fruit. Air-dried exposure to long-wave ultraviolet radiation
seeds separated from them averaged 11,900 (O’Neal and others 2002).
seeds/kg. Placed on moist filter paper without
pretreatment, 100 percent germinated, beginning References
14 days after sowing (Francis and Rodríguez
1993). Pascarella (1996) noted fruits collected in Francis, J.K. and A. Rodríguez. 1993. Seeds of
Florida weighed an average of 0.483 + 0.034 g and Puerto Rican trees and shrubs: second
that 85 to 90 percent of the fruits contained just installment. Research Note SO-374. U.S.
one seed. Most seeds fall under the parent trees but Department of Agriculture, Forest Service,
some are dispersed by birds. Seedlings are Southern Forest Experiment Station, New
relatively common in forests where the species Orleans, LA. 5 p.
grows.
Howard R.A. 1988. Flora of the Lesser Antilles,
Growth and Management.—Growth rate of Leeward and Windward Islands.
Florida bitterbush is slow to moderate. Weaver Dicotyledoneae, Part 1. Vol. 4. Arnold
(1979) followed the growth of two plants in a Arboretum, Harvard University, Jamaica Plain,
forest in Puerto Rico with characteristic slow MA. 673 p.
growth and found diameter growth rates over the
24-year period of about 0.2 cm/year. Florida Institute of Food and Agricultural Sciences. 2003.
bitterbush in Puerto Rico live about 30 to 60 years Native landscape plants for South Florida:
(author’s observation). Plants begin flowering and shrubs and small trees (P-Z). http://miami-
fruiting when they are about 1.5 to 2 m in height if dade.ifas.ufl.edu/programs/fyn/publications/nati
sufficient light is available. Fruits are easily veplantlist/npl-shrubs-small-trees-p-z.htm. 5 p.
collected in quantity by hand and with short
pruning poles. They may be cleaned by macerating Liogier H.A. 1988. Descriptive flora of Puerto
or by working them against a screen under running Rico and adjacent islands, Spermatophyta. Vol.
water. Production of seedlings, at least to the 2. Editorial de la Universidad de Puerto Rico,
pricking-out stage, is straightforward. Where a Río Piedras, PR. 481 p.
seed source is available in favorable habitat,
natural reproduction is usually adequate. Florida Liogier, H.A. 1990. Plantas medicinales de Puerto
bitterbush has not been reported to cause any Rico y del Caribe. Iberoamericana de Ediciones,
problems in forests and therefore control measures Inc., San Juan, PR. 566 p.
are probably not needed.
Little, E.L., Jr. and F.L. Wadsworth. 1964.
Benefits.—Florida bitterbush helps protect the soil Common trees of Puerto Rico and the Virgin
and contributes to the biodiversity and aesthetics Islands. Agriculture Handbook 249. U.S.
of forest stands. A number of bird and animal Department of Agriculture, Washington, DC.
species eat the fruits. It furnishes larval food for 548 p.
the bush sulphur butterfly, Urema dina helios
(Institute of Food and Agricultural Sciences 2003). Long, R.W. and O. Lakela. 1976. A flora of
Florida bitterbush is a honey plant (Little and Tropical Florida. University of Miami Press,
Wadsworth 1964). The wood is little used, except Coral Gables, FL. 962 p.
occasionally for fuel. Florida bitterbush is planted
in Florida and Cuba as an ornamental (Little and O’Neal, C.B., J. Salazar, and E. Rodríguez. 2002.
Wadsworth 1964). A bitter tonic is made from A preliminary analysis of the anti-microbial and
tissues of the species in the Caribbean to treat phytochemical properties of Picramnia
fevers, anorexia, and stomach problems and as an pentandra (Simaroubaceae) and its potential
enema to treat for worms (Liogier 1990). relationship to gastrointestinal health.
Qualitative tests indicated the presence of Emanations 4: 57.
antioxidants, phenols, terpenoids, and alkaloids in
ethanol extracts of tissues. A number of bacteria
562
Pascarella, J.B. 1996. Reproductive ecology of Weaver, P.L. 1979. Tree growth in several tropical
Picramnia pentandra (Picramniaceae) in South forests of Puerto Rico. Research Paper SO-152.
Florida. Caribbean Journal of Science 32(1): 99- U.S. Department of Agriculture, Forest Service,
104 Southern Forest Experiment Station, New
Orleans, LA. 15 p.
563
Piper aduncum L. bamboo piper
PIPERACEAE
Synonyms: Artanthe adunca Miq.

Piper hebecarpum C. DC. in Urban
Piper martinicense C. DC. in Briq.
Piper stehleorum Trel. in Stehlé
Piper subrectinerve C. DC. in Urban
John K. Francis
or black seeds (Howard 1988, Liogier 1985, Little

and Wadsworth 1964, Stevens and others 2001).
Range.—Bamboo piper is native to the Greater

and Lesser Antilles and on the mainland from
Mexico to northern Argentina. The species has
been widely planted as an ornamental and has
spread by the movement of equipment between
land masses. It has escaped and established itself
in Florida, Southeast Asia, and a number of Pacific
Islands (Instituto de Botánica Darwinion 2002,
Howard 1988, Pacific Island Ecosystems at Risk
2002, Plant Protection Service 2001, Stevens and
others 2001).
Ecology.—Bamboo piper grows in areas that

receive from about 1500 to over 4000 mm of
mean annual rainfall. It grows in Puerto Rico from
near sea level to over 400 m in elevation. Most
soils are colonized including fill dirt, but not salty
soils and excessively well-drained soils at the dry
end of the range. Disturbance, which is required
General Description.—Bamboo piper is also for establishment, includes removal of the forest
known as jointwood, spiked pepper, false kava, canopy to allow light to enter and the creation of a
cow’s foot, higuillo, higuillo de hoja menuda, bare soil surface, although plants occasionally
cordoncillo, anisillo, guayayo, Santa María negra, establish themselves on rotting logs and stumps.
aperta ruão, and matico. It is a multistemmed Bamboo piper is moderately intolerant of shade. It
evergreen shrub or small tree 1 to 6 m in height can survive and grow slowly under a moderate
and up to 10 cm or more in stem diameter. The overstory but requires at least partial exposure to
wood is whitish, hard, and brittle with a thick, grow large and flower. The species commonly
solid pith, and no annual rings. The stems have grows in clearcuts, tree-fall gaps, shade-grown
swollen nodes and bark that is smooth and gray or coffee and other tree plantations, brushy pastures,
green. Plants are supported by abundant lateral roadsides, and landslides. Bamboo piper grows as
roots and, at times, proproots arising from nodes individual plants or in thickets. As many as 2,632
near the base. The green twigs are not abundant. plants/ha were reported in areas invaded in the
The alternate leaves have short petioles and Philippines (Philippine Council of Agriculture
elliptic to lanciolate blades 12 to 20 cm long. All 2002).
plant parts have a peppery taste and odor. The
cordlike spikes are borne opposite leaves and Reproduction.—Bamboo piper flowers and fruits
contain many, minute flowers that soon develop throughout the year (Little and Wadsworth 1964).
into numerous tiny, imbedded drupes with brown Air-dried seeds from a Puerto Rican source
averaged 0.000237 g/seed or 4.2 million/kg. Only
7 percent germinated when placed on moist filter
564
paper (author’s observation). The seeds are References
dispersed by bats, birds, and possibly arboreal
rodents (Plant Protection Service 2001). Seedlings Bourke, R.M. 1997. Management of fallow species
are fairly common on disturbed ground. composition with tree planting in Papua New
Established plants thicken into clumps by suckers Guinea. http://coombs.anu.edu.au/Depts/
arising from the root crown (Pacific Island RSPAS/RMAP/bourke.htm. 8 p.
Ecosystems at Risk 2002). Plants are established
for agroforestry purposes in Papua New Guinea by Gómez, P., D. Cubillo, G.A. Mora, and L. Hilje.
shoving cuttings into moist soil (Bourke 1997). 1997. Evaluation of possible repellents for
Bemisi tabaci. II. Botanical substances. Manejo
Growth and Management.—The growth of Integrado de Plagas 46: 17-25.
bamboo piper is moderately rapid after the early
seedling stage. Sprouts and suckers grow more Howard R.A. 1988. Flora of the Lesser Antilles,
than a meter in their first year. Producing Leeward and Windward Islands.
seedlings by means of seed is difficult. Dicotyledoneae, Part 1. Vol. 4. Arnold
Propagating plants with cuttings, which need no Arboretum, Harvard University, Jamaica Plain,
hormonal treatment, in the nursery or directly in MA. 673 p.
the field, is recommended. Individual stems live 2
to several years; by sprouting, plants live much Ibrahim, J. A. Abu-Said, A. Abdul-Rashih, M.A.
longer. Infestations of bamboo piper can be Nor-Azah, M.Z. Zaridah, A.K. Azizol, Norhara-
controlled by uprooting young plants and spraying Hussein, P.S. Bacon, and K.C. Khoo. 1996.
older plants with broadleaf weed killers such as Essential oils of selected Malaysian plants and
2,4-D. Treated areas should be checked in a few their potential uses. Forestry and Forest Products
weeks and treated again if regrowth has occurred Research, Proceedings of the Third Conference,
(Plant Protection Service 2001). Oct. 3-4, 1995; Kuala Lumpur, Malaysia. p. 97-
103.
Benefits.—Bamboo piper helps revegetate
disturbed areas and contributes to the biodiversity Instituto de Botánica Darwinion. 2002. Catálogo
and biomass of forests. It also is a source of food de las plantas vasculares de la Argentina.
and cover for wildlife. Bamboo piper established http://www.darwin.edu.ar/Catalogo/indicevascul
in contour rows for soil erosion brakes helps ares.htm. [n.p.].
facilitate agroforestry on steep land in Papua New
Guinea (Bourke 1997). The wood is useful for Liogier H.A. 1985. Descriptive flora of Puerto
fuel, stakes, fences, and rude construction Rico and adjacent islands, Spermatophyta. Vol.
(Philippine Council of Agriculture 2002). 1. Editorial de la Universidad de Puerto Rico,
However, exposed wood rots quickly (Vélez and Río Piedras, PR. 352 p.
van Overbeek 1950). The species is planted as an
ornamental. The peppery fruits have been used to Liogier, H.A. 1990. Plantas medicinales de Puerto
season food (Little and Wadsworth 1964). Rico y del Caribe. Iberoamericana de Ediciones,
Essential oil content of bamboo piper tissue Inc., San Juan, PR. 566 p.
(leaves and twigs), of which dillapiole is the major
component, ranged from 1.2 to 3.4 percent (Maia Little, E.L., Jr. and F.H. Wadsworth. 1964.
and others 1998). Teas and other extracts of the Common trees of Puerto Rico and the Virgin
leaves and roots of bamboo piper are used in Islands. Agriculture Handbook 249. U.S.
herbal medicine as a tonic to ease diarrhea, Department of Agriculture, Forest Service,
dysentery, vomiting, ulcers, and to control Washington, DC. 548 p.
bleeding (Liogier 1990). The chemical 2’, 6’-
dihydroxy-4’-methoxychalcone isolated from Maia, J.G.S., M.G.B. Zohhbi, E.H.A. Andrade,
bamboo piper inhibited 98 percent of the growth A.S. Santos, M.H.L. da Silva, A.I.R. Luz, C.N.
of Leishmania amazonensis parasites in vitro with Bastos, and M.H.L. da Silva. 1998. Constituents
low host cell toxicity (Torres-Santos and others of the essential oil of Piper aduncum L. growing
1996). In addition, the essential oils have shown wild in the Amazon region. Flavour and
strong insecticidal, molluscicidal, and antibacterial Fragrance Journal 13(4): 269-272.
effects (Gómez and others 1997, Ibrahim and
others 1996, Orjala and others 1992). Orjala, J., A.D. Wright, T. Rali, and O. Sticher.
565
1992. Three new prenylated benzoic acid Stevens, W.D., C. Uloa-U., A. Pool, O.M.
derivatives and molluscicidal sesquiterpenoids Montiel, eds. 2001. Flora de Nicaragua.
from Piper aduncum leaves. Medica 58(7): 714. Monographs in Systematic Botany Vol. 85, No.
3. Missouri Botanical Garden Press. p. 1911-
Pacific Island Ecosystems at Risk. 2002. Invasive 2666.
plant specie, Piper aduncum L. Rutaceae.
http://www.hear.org/pier3/ piadu.htm. 2 p. Torres-Santos, E.C., V.C. Moura, G.M. Sperandio,
D.L. Moreira, M.A. Kaplan, and B. Rossi-
Philippine Council of Agriculture. 2002. Research Bergmann. 1996. Anti-leishmanial effect of a
highlights 1997. http://www.pcarrd.dost.gov.ph/ pure chalcone isolated from Piper aduncum
division/FERD/new/highlights97.html. 18 p. (Piperaceae). Memórias Vol. 91 (Supplement).
Instituto Oswaldo Cruz, Rio de Janeiro, RJ,
Plant Protection Service. 2001. Pest alert: false Brazil. 18 p.
kava. Pest Alert No. 19. http://www.spc.org.nc/
pps/PestAlerts/PestAlertNo19-False_kave.pdf. Vélez, I. and J. van Overbeek. 1950. Plantas
2 p. indeseables en los cultivos tropicales. Editorial
Universitaria, Río Piedras, PR. 497 p.
566
Piper amalago L. higuillo de limón
PIPERACEAE
Synonyms: Enckea amalago (L.) Griseb.

Enckea seiberi Miq.
Piper medium Jacq.
Piper seiberi (Meq.) DC.
John K. Francis
thickened in older plants. The alternate, dark
green, hairless leaves are 4 to 14 cm long and 2 to
7 cm broad. They are pointed at the tip, ovate to
elliptic, and have palmate venation with five main
veins. Tiny flowers are borne in compact, cord-
like, gray-green spikes 6 to 12 cm long. At
maturity, the small (1.5 mm) fruits (drupes) are
closely packed along the spike (Liogier 1985,
Range.—Higuillo de limón is native to tropical

Mexico, Central America, South America, and the
Greater and Lesser Antilles (Howard 1988).
Ecology.—Higuillo de limón is primarily an

understory species of old growth and secondary
forests. It is also found in openings and along
roads. The species will endure all but the densest
shade under stands, but needs openings or at least
intermediate crown positions in open forest or
brush thickets to reproduce. However, higuillo de
limón is rarely among the pioneers that colonize
disturbed sites, tending instead to invade after a
low stand of trees or brush has become
established. It endures competition with herbs,
General Description.—Higuillo de limón is a
trees, and shrubs, but does not survive in deep
Spanish name meaning lemon pepper. Although
grass swards. The species will grow in soils with
the species is a pepper with the characteristic
textures from sand to clay and in all drainage
peppery taste of foliage and seeds, the pungent,
classes except excessively drained and very poorly
disagreeable odor of crushed leaves, stems, and
drained. It does not appear to be sensitive to
fruits hardly suggests lemon. Other common
differences in soil parent material. In Puerto Rico,
names include joint wood, soot-soot, guayuyo,
higuillo de limón grows in areas receiving from
cordoncillo, anisillo, alcotán, anisetto, and
about 1200 to 2500 mm of annual precipitation
malimbé (Little and others 1974).
and from near sea level to more than 600 m in
Higuillo de limón is a shrub that
elevation.
sometimes becomes a small tree. It is recognized
by its enlarged, ringed nodes on smooth, green
Reproduction.—Higuillo de limón flowers and
stems with many brown lenticels. Stems on older
fruits throughout the year (Little and others 1974).
plants become gray. Higuillo de limón often has
In Costa Rica, frugivorous bats are the most and
multiple stems from the base, but usually does not
perhaps the only important dispersers of higuillo
branch further until the crown is reached. The
de limón seeds (Elizondo 2000, Fleming and
crowns have relatively few branches and may be
others 1977). One sample of fruits from Puerto
flattened in older individuals. The wood is of
Rico yielded 890,000 seeds/kg. On moist filter
moderate density and brittle. A taproot forms in
paper, these seeds gave 77 percent germination
seedlings, but secondary roots that fan out and
between 8 and 37 days after sowing.
down later match it in size, and even become
567
Growth and Management.—In Puerto Rico, Interactions between crown structure and light
higuillo de limón reaches 5.5 m in height and 8 cm environment in five rain forest Piper species.
in diameter (Little and others 1974). The species American Journal of Botany 75(10): 1,459-
may become a tree to 15 m in high forests in 1,471.
Mexico, although it remains a shrub or small tree
in successional forests and gaps (Chazdon and Elizondo C., L.H. 2000. Artibeus toltecus
others 1988). Its stems have no visible annual Saussure. http://www.Inbio.ac.cr/bims/ubi/
rings and are therefore difficult to age. Judging mamiferos/ubiespejo/ubiid=1557&-find.html.
age by stem internodal segments, individual stems 5 p.
in Puerto Rico appear to reach 5 to 10 years old.
Because resprouting occurs from the roots, Fleming, T.H., E.R. Heithaus, and W.B. Sawyer.
individual plants may live much longer. Natural 1977. An experimental analysis of the food
regeneration of higuillo de limón occurs readily in location behavior of frugivorous bats. Ecology
early secondary forest and in open stands of older 58(3): 619-627.
forest when seed sources are present. Stands can
be easily manipulated to encourage it. Small Grieve, Mrs. M. 2001. A modern herbal: pepper.
quantities of seed have been separated by hand http;//www.botanical.com/botanical/mgmh/p/
with tweezers after mashing the fruit. Larger pepper24.html. 5 p.
quantities probably can be obtained by gentle
maceration and wet sieving. Germination on the Howard R.A. 1988. Flora of the Lesser Antilles,
surface of wet soil or peat is recommended. Leeward and Windward Islands.
Plantations could be established using Dicotyledoneae, Part 1. Vol. 4. Arnold
containerized seedlings, although no plantings Arboretum, Harvard University, Jamaica Plain,
have been reported. MA. 673 p.
Benefits.—Higuillo de limón is eaten by cattle Liogier H.A. 1985. Descriptive flora of Puerto
after the more desirable grasses and forbs are Rico and adjacent islands, Spermatophyta. Vol.
gone. Infusions of the leaves are said to alleviate 1. Editorial de la Universidad de Puerto Rico,
colic and intestinal gas. The roots are used as a Río Piedras, PR. 352 p.
diuretic and to treat water retention (Liogier
1990). The fully ripe fruits and their seeds have Liogier, H.A. 1990. Plantas medicinales de Puerto
the same taste as black pepper, Piper nigrum L., Rico y del Caribe. Iberoamericana de Ediciones,
and are sometimes used as a substitute for it, Inc., Río Piedras, PR. 566 p.
especially in Jamaica. The fruits are usually dried
whole and ground as needed (Grieve 2001). Little, E.L., Jr., R.O. Woodbury, and F.H.
Chazdon, R.L., K. Williams, and C.B. Field. 1988. Washingtion, DC. 1,024 p.
568
Piper swartzianum (Miq.) C. DC. higuillo
PIPERACEAE
Synonyms: Ariantha swartziana Miq.
John K. Francis
and fruits are grouped in 5- to 11-cm cord-like

spikes, 4 mm thick borne singly opposite leaf
attachments near the branch tips. The spikes are
white during flowering, turning light green or
yellow-green as the fruits mature. The individual
fruits are imbedded within the spike in spirals
(Liogier 1985, Little and others 1974).
Range.—Higuillo is native to Puerto Rico and

Hispaniola (Liogier 1985). It is relatively common
to rare in the moist and wet portions of Puerto
Rico, specifically the public forests of Guajataca,
Maricao, and Río Abajo (Little and others 1974),
and private lands nearby. Higuillo was
misidentified as P. tuberculatum Jacq. in Little
and others (1974) (Liogier 1985).
Ecology.—Higuillo inhabits middle and late

secondary and remnant forests. It is shade tolerant
and is usually found in the understory, often in
fairly deep shade. Higuillo does not compete well
in tall, dense thickets of vigorous shrubs and
young trees. At the lower end of the rainfall range
General Description.—This shrub, known by its (about 1200 mm/yr), the species is confined to
Spanish name higuillo in Puerto Rico (Little and extra-humid situations such as deep ravines. It is
others 1974), has been assigned the name Spanish most common in forests that receive from 1600 to
elder in English (Natural Resources Conservation 3000 mm of precipitation. Higuillo grows on sites
Service 2002). The species is an evergreen shrub from a few meters above sea level to more than
that may take on vine-like or tree-like form. 1,000 m in elevation on a wide variety of soils
Higuillo usually has heights of 1.5 to 3 m but may derived from all types of parent material including
reach 4.5 m in height and 7.5 cm in stem diameter ultramafics and limestone.
(Little and others 1974). Plants are supported by
relatively abundant lateral and fine roots. White Reproduction.—In order to flower and fruit,
mycorrhizal fungi associated with the fine roots mature higuillo plants need breaks in the forest
are clearly visible. The shrubs usually have canopy that admit over-head sun for part of the
multiple stems that have arisen as sprouts from the day. Vigorous plants flower and fruit continuously
root crown. The stems are gray with many raised (Little and others 1974). Air-dried seeds weighed
lenticels and have enlarged nodes and a thick pith. an average of 0.00156 g/seed or 641,000 seeds/kg
The wood is light brown and brittle. The twigs are (author’s observation). The seeds are probably
light green turning gray and often have a zig-zag disbursed by fruit bats that eat the fruits. Seedlings
form. The inner bark, twigs, and leaves have a are widely scattered and uncommon.
pleasant spicy odor when crushed, and a peppery
taste. The leaves are dull green to dark green, Growth and Management.—New seedlings
alternate in two rows, with minute gland dots. The grow slowly. Well-established plants and sprouts
leaf blades are oblong to lanceoate, 11 to 24 cm grow about 0.5 m/year in height. Individual stems
long by 3 to 7 cm broad, long-pointed at the apex live 3 or 4 years and are replaced by other sprouts
and unequally rounded at the base. The flowers so that established plants can live indefinitely. No
569
management experience has been published. References
Natural reproduction could probably be
encouraged by disturbance that exposes mineral Liogier H.A. 1985. Descriptive flora of Puerto
soil under a moderately shady tree canopy. Rico and adjacent islands, Spermatophyta. Vol.
Benefits.—Higuillo contributes to the Río Piedras, PR. 352 p.
biodiversity, scenic beauty, and stability of the soil
of the forests where it grows. Like other pipers, its Little, E.L., Jr., R.O. Woodbury, and F.H.
fruits are food for fruit bats. Wadsworth. 1974. Trees of Puerto Rico and the

Plants profile: Piper blattarum Spreng.
http://plants.usda.gov/cgi_bin/checklist.cgi. 2 p.
570
Piptocoma antillana Urban Antilles velvetshrub
ASTERACEAE
Synonyms: Piptocoma rufescens auct. non Cass.
John K. Francis
seeds are straw-colored achenes about 2 mm long,

tipped with a pappus about 4.5 mm long (author’s
observation, Liogier 1997).
Range.—Antilles velvetshrub is native to Puerto

Rico and the Virgin Islands (Liogier 1997).
Previous reports (Liogier and Martorell 1982) that
it occurred in Hispaniola were based on
misidentification of P. rufescens Cass. Antilles
velvetshrub is not known to have been planted or
Ecology.—Antilles velvetshrub grows in coastal

and upland sites from near sea level to 500 m or
more in elevation. Mean annual rainfall varies
from about 750 mm to 2000 mm. It is most
common on ridges and hillsides of areas underlain
with ultramafic rocks and limestone. Antilles
velvetshrub is sometimes seen on road cuts and the
center between the tracks of abandoned roads.
Soils are excessively well or well drained, have
pH’s from about 5.5 to 7.7, and vary widely in
texture and quantity of stones in the profile.
Antilles velvetshrub is drought tolerant--leaves
wilt but the plants usually do not defoliate. The
species tolerates competition from shrubs, herbs,
and a moderate amount of grass. It grows in
openings or under partial shade of low basal-area
forests and may be found in remnant stands and
General Description.—Antilles velvetshrub is an mid- to late-secondary forests. The species
evergreen, narrow-crowned, upright shrub usually probably requires bare soil or small-scale
1 to 2 m in height but sometimes reaching 4 m in disturbance to establish itself. Antilles velvetshrub
height and 5 cm in basal diameter. Mature plants occurs as scattered to occasional plants.
often have several stems arising from the root
crown or low on the stem. Lower stems are Reproduction.—Antilles velvetshrub blooms
furrowed and gray; branches are relatively few, annually, apparently near the end of the wet
slender, straight, and covered with gray bark. The season. Seeds were collected by the author in
pale yellow wood is of moderate density, Puerto Rico in March (dry season). The air-dried
moderately strong, and has a dark brown pith at seeds averaged 0.00046 g/seed or 2,150,000
the center of the stem. Plants are supported by seeds/kg. Placed on moist filter paper without
flexible, weak roots covered by corky, brown bark. pretreatment, 8 percent germinated between 11 and
The species is easily recognized by the narrow 14 days after sowing. Germination is epigeal.
leaves that are dark green above and bright white Seeds are dispersed by wind. Seed crops can be
beneath. The narrowly elliptic to lanceolate leaves moderately abundant, but seedlings are
are 3 to 8 cm long and 0.5 to 2.5 cm broad and uncommon.
have petioles up to 5 mm long. Inflorescences are
corymbs of numerous campanulate heads. The
571
Growth and Management.—Antilles velvetshrub for background and accent in naturalistic
grows 0.5 to 1.0 m per year. The stems do not live landscaping.
long--perhaps 2 to 5 years--but plants can renew
themselves and live longer by sprouting from the References
root crown. Planting and management experience
have not been published. Because the species is Liogier, H.A. 1997. Descriptive flora of Puerto
not abundant or aggressive, control should not be Rico and adjacent islands. Vol. 5. Editorial de la
necessary. Universidad de Puerto Rico, Río Piedras, PR.
436 p.
Benefits.—Antilles velvetshrub helps protect the
soil, furnishes cover for wildlife, and adds to the Liogier, H.A. and L.F. Martorell. 1982. Flora of
aesthetics of the forest. If means of propagation Puerto Rico and adjacent islands, a systematic
can be developed, the species would be valuable synopsis. Editorial de la Universidad de Puerto
Rico, Río Piedras, PR. 342 p.
572
Pisonia aculeata L. pull-back-and-hold
NYCTAGINACEAE
Synonyms: Pisonia helleri Standley

Pisonia yaguapinda D. Parodi
John K. Francis
species may be evergreen or deciduous depending
on the climate. The leaves are variable, ovate to
elliptical and 2.5 to 15 cm long by 1.5 to 6 cm
broad. They are light to dark green, leathery to
somewhat fleshy. The flowers are small axillary
cymes that are succeeded a few months later by 11
to 15 mm long fruits (anthocarps) with five single
or double length-wise rows of sticky glands
(Acevedo-Rodríguez 1985, Howard 1988, Liogier
1985).
Range.—Pull-back-and-hold is native to most of

the humid tropical and subtropical areas of the
world including five continents, most of the
islands of the Caribbean and many of the Pacific
islands. Its range includes southern Florida and the
Rio Grande Valley of Texas (Florida 4-H
Foundation 2001, Friend 2001, Howard 1988).
Ecology.—Pull-back-and-hold grows from near

sea level in coastal areas to as much as 1,000 m in
elevation in continental areas (Múlgura 2000). In
Puerto Rico, the species grows in sites receiving
from 750 mm to over 2000 mm of precipitation.
These shrubs grow well in soils with textures from
sand to clay, with drainage from somewhat poorly
General Description.—Pull-back-and-hold (a drained to excessively well drained in limestone,
name used in Florida) is a species known by a igneous, and alluvial areas. In Florida, the species
large number of common names worldwide. Other grows on hummocks and pinelands (Florida 4-H
names used in U.S. territories are devil’s claw Foundation 2001). Seedlings and young plants
(Florida), wild bougainvillea (Texas), escambrón survive in broken sunlight, but to reproduce, a
(Puerto Rico), and prickly mampoo (U.S. Virgin plant must be relatively large and must have
Islands). Pull-back-and-hold is a large, scrambling reached nearly full sunlight. This usually means
and climbing woody shrub (vine) that rarely growing in the crowns of trees or sometimes on
becomes a small tree. It is reported as a medium to rocks or fences
large tree in Costa Rica (Enquist and Sullivan
2001). Its stems and branches are smooth, dark Reproduction.—In Puerto Rico, pull-back-and-
gray, and usually support opposite, stout, hooked hold flowers from December to May (the dry
and branched spines. The spines aid in season) and fruits in August (just before the wet
clinging to tree crowns. Pull-back-and-hold may season) (Acevedo-Rodríguez 1985) although
have one to several branches arising near the base, fruiting of occasional individuals may occur at
and then the stems are relatively free of branches almost any time of the year. Plants in Costa Rica
until they reach increased light, usually in the fruit late in the wet season (Enquist and Sullivan
canopy of trees where they branch profusely. 2001). A large plant can produce thousands of
Seedlings have a taproot and finer laterals. In older seeds during a fruiting episode. Fruits from a
plants the lateral roots become extensive and Puerto Rican collection averaged 0.077 + 0.028
diffuse. The fine roots are tough and fibrous. The
573
g/fruit. Seeds extracted by hand, which have the References
appearance of grains of wild rice, averaged 0.041
+ 0.011 g/seed. It is not necessary to extract seed Acevedo-Rodríguez, P. 1985. Los bejucos de
because they germinate satisfactorily within the Puerto Rico. Vol. 1. General Technical Report
fruit. Seventy-five percent of seeds from the above SO-58. U.S. Department of Agriculture, Forest
Puerto Rican collection, sown as fruits in Service, Southern Forest Experiment Station,
commercial potting mix, germinated between 21 New Orleans, LA. 331 p.
and 49 days after sowing. Pull-back-and-hold is
susceptible to damping-off as seedlings in Enquist, B.J. and J.J. Sullivan. 2001. Vegetative
germination beds. Seedlings are ready to prick out key and descriptions of tree species of the
into containers about 1 month after germination. tropical dry forests of upland Sector Santa Rosa,
The sticky seeds can cling to birds while on the Area de Conservación, Guanacaste, Costa Rica.
trees; then, after they fall, still attached to the http://www.acguanacaste.ac.cr/paginas_ especie/
infrutescences, stick to the clothing or fur of plantae_online/EnquistSullivanTreeKey.pdf.
passing humans and animals. 68 p.
Growth and Management.—Although pull-back- Florida 4-H Foundation. 2001. Florida forest
and-hold has only a moderate growth rate, it can plants: Devil’s claws (Pisonia aculeata).
live for several decades and become quite large. http://www.sfrc.ufl.edu/4h/Devils_claw/
Plants may reach 20 m of extension (Acevedo- deviclaw.htm. 2 p.
Rodríguez 1985) and 20 cm of stem diameter in
Puerto Rico. It is sometimes desirable to remove Friend, W.H. 2001. Plants of ornamental value for
pull-back-and-hold from forest stands. This the Rio Grande Valley of Texas. http://
increases safety during timber harvest and helps www.aggiehorticulture.tamu.edu/plantanswers/
ensure that the regenerating stand will not be publications/riograndeornamentals.htm. 16 p.
suffocated by resurgent vines. A time-honored
method is to cut the vines hanging from trees 1 Howard R.A. 1988. Flora of the Lesser Antilles,
year before harvest and then to cut or spray the Leeward and Windward Islands.
few surviving sprouts just before harvest. Dicotyledoneae, Part 1. Vol. 4. Arnold
Benefits.—Pull-back-and-hold may be used as a MA. 673 p.
hedge or a foundation plant in landscaping, but it
requires regular pruning to keep it under control Liogier, H.A. 1985. Discriptive flora of Puerto
(Friend 2001) An occasional plant occurs entirely Rico and adjacent islands. Vol. 1. Editorial de la
free of spines. Perhaps these could be propagated Universidad de Puerto Rico, San Juan, PR.
vegetatively for ornamental use. The wood of wild 352 p.
pull-back-and-hold is sometimes used for fuel. It
is hard, heavy, and often available in good Múlgura, M.E. 2000. Catálogo de las plantas
diameters and considerable lengths. Root extracts vasculares de la República Argentina.
have been used in herbal medicine as a purgative, http://www.Darwin.edu.ar/Catologo/
and extracts from leaves and bark are used to treat nyctaginaceae.pdf. 4 p.
arthritis (Acevedo-Rodríguez 1985).
574
Pithecellobium unguis-cati (L.) Benth. bread-and-cheese
FABACEAE
Synonyms: Mimosa unguis-cati L.

Zygia unguis-cati (L.) Sudw.
John K. Francis
terminal or axillary racemes or panicles in heads.

The legume, 5 to 10 cm long, is coiled or curved
and splits open to reveal 4- to 6-mm shiny, black
seeds surrounded by white to red fleshy arils
1974). Bread-and-cheese is easily confused with P.
dulce (Roxb.) Benth., which grows as an exotic in
many Caribbean islands. The latter is a tree with
somewhat smaller leaves, coarser twigs, larger
fruits, and less pronounced rings at the nodes.
Range.—Bread-and-cheese ranges from Florida

through the Greater and Lesser Antilles, Trinidad
and Tobago, Bonaire, Curacao, Aruba, Guyana,
and Venezuela (Liogier 1988, Little and other
1974). Planting has not been reported outside its
native range.
Ecology.—Bread-and-cheese grows in Puerto

Rico from near sea level to about 450 m in
elevation on soils derived from both igneous and
sedimentary rocks (author’s observation). The
General Description.—Bread-and-cheese is the species prefers well-drained soils, but all soil
common name used in the Virgin Islands and the textures appear to be tolerated. With few
English-speaking Lesser Antilles. The species is exceptions, it grows in areas receiving less than
also know as cat’s claw and black bead in Florida 1000 mm of annual precipitation. Because of
and doctor long, groven eye, uña de gato, rolón, reduced competition from trees, the species tends
bois crabbe, griffe-chatte, collier-diable, tendra à to grow on sand dunes, coastal strands and keys,
caulilou-rivière, and bébèl elsewhere (Howard and on shallow rocky soils, sometimes forming
1988, Liogier 1988, Little and others 1974). thickets (West and Arnold 1952). It does survive
Bread-and-cheese is a shrub or small tree usually 3 and grow in the understory of open, dry forests,
to 9 m in height and 4 to 13 cm in stem diameter but maintains thin crowns and seldom flowers in
(Little and others 1974). However, it occasionally these situations. Bread-and-cheese appears to
grows to impressive proportions. The national tolerate salt spray and salty groundwater.
record tree, found in Camino Real, Florida,
measures 22 m in height and 121 cm in diameter Reproduction.—Bread-and-cheese flowers and
(Champion Tree Project 2001). The plant usually fruits irregularly through the year (Little and
has multiple stems arising at or below ground others 1974). Seed yield can be heavy but
level. The stems and branches are gray and nearly inconsistent from year to year. A sample of seed
smooth with lenticels and rings at the nodes. The collected in Puerto Rico contained 14,300
paired spines are persistent. Bread-and-cheese seeds/kg and germinated at 31 percent, beginning
produces a taproot and abundant lateral and fine 3 days after sowing (Francis and Rodríguez 1993).
roots. Alternate hairless leaves have petioles 2 to 5 Artificial propagation is by seed; it is one of the
cm long and leaf blades 2 to 6 cm long. There are easiest shrubs to grow (Workman 1980). Natural
four obliquely obovate or oblong leaflets per leaf. seedlings are seldom abundant. The shrubs
The greenish-yellow, yellow, or pink flowers form resprout readily after cutting or fire.
575
Growth and Management.—Plants started from installment. Research Note SO-374. Southern
seed grow relatively slowly at first and somewhat Forest Experiment Station, U.S. Department of
faster after deep rooting. Supplemental watering Agriculture, Forest Service, New Orleans, LA.
and fertilization are unnecessary. As a landscaping 5 p.
plant, it can be easily pruned and shaped
(Workman 1980). In the poor habitat where the Howard R.A. 1988. Flora of the Lesser Antilles,
species usually occurs in the wild, growth is very Leeward and Windward Islands.
slow and plants live for several decades. Dicotyledoneae, Part 1. Vol. 4. Arnold
Benefits.—Although it is difficult to harvest MA. 673 p.
because of the spines, the wood of bread-and-
cheese is used for fuel. It is heavy and hard and Island Resource Foundation. 2001. Threatened and
has an oven-dry heat of combustion of 19.05 endangered birds of the insular Caribbean:
megajoules/kg (Timyan 1996). The fleshy aril is Yellow-shouldered amazon, Amazona
edible, although not highly desirable because of barbadensis. http://www.irf.org/bbarden.htm.
the lingering flavor. The seeds of bread-and- 13 p.
cheese are used to make necklaces. The species is
planted to form impenetrable hedges (UVI Liogier H.A. 1988. Descriptive flora of Puerto
Wetlands Reserve 2001). Bread and cheese is Rico and adjacent islands, Spermatophyta. Vol.
listed as a nitrogen fixing species (Winrock 2. Editorial de la Universidad de Puerto Rico,
International 2001). The fruits are one of the food Río Piedras, PR. 481 p.
plants of the endangered yellow-shouldered
amazon parrot (Amazona barbadensis) in Little, E.L., Jr., R.O. Woodbury, and F.H.
Venezuela, and the islands of Margarita, La Wadsworth. 1974. Trees of Puerto Rico and the
Blanquilla, and Bonaire (Island Resource Virgin Islands. Vol. 2. Agriculture Handbook
Foundation 2001). The large orange sulfur 449. U.S. Department of Agriculture,
(Phoebis agarithe Boisduval) and the Miami blue Washington, DC. 1024 p.
(Hemiargus thomasi Clench) butterflies both use
bread-and-cheese as rearing plants for their larva Timyan, J. 1996. Bwa yo: important trees of Haiti.
(Association of Florida Native Nurseries 2001). A South-East Consortium for International
small treehopper insect (Homoptera) has Development, Washington, DC. 418 p.
developed an extended thorax that mimics a
bread-and-cheese thorn and serves as protective UVI Wetlands Reserve. 2001. A list of plants
camouflage (Workman 1980). tagged along the trail at the Reserve, and some
of their uses. http://rps.uvi.edu/
References VIMAS/plantlist.htm. 5 p.
Association of Florida Native Nurseries. 2001. West, E. and L.E. Arnold. 1952. The native trees
Florida gardening with native plants—Legume. of Florida. University of Florida Press,
http://www.nsis.org/garden/family/legume.html. Gainesville, FL. 212 p.
4 p.
Winrock International. 2001. Nitrogen fixing trees
The Champion Tree Project. 2001. National and shrubs. http.//www.winrock.org
champion trees, Florida. /forestry/factpub/nftlist /htm. 25 p.
http://www.championtrees.org/database/champi
onsFL.htm. 8 p. Workman, R.W. 1980. Growing Native. The
Sanibel-Captiva Conservation Foundation, Inc.
Francis, J.K. and A. Rodríguez. 1993. Seeds of 137 p.
Puerto Rican trees and shrubs: second
576
Pluchea carolinensis (Jacq.) G. Don cure-for-all
ASTERACEAE
Synonyms: Conyza symphytofolia sensu Gillis

Pluchea odorata authors, non P. odorata (L.) Cass.
Conyza carolinensis Jacq.
Conyza coresii Kunth
Pluchea cortesii (Kunth) DC.
John K. Francis
purple to white. The achenes are brown, 0.6 to 0.8

mm long, and have a pappus of 10 to 15
yellowish-white bristles. Chromosome number is
2n = 20 (author’s observation, Howard 1989,
Liogier 1997, Long and Lakela 1976, Stevens and
others 2001)
Range.—Cure-for-all is native to Florida,

Bermuda, the Bahamas, the West Indies, Mexico,
Central America, and Colombia, Venezuela, and
Ecuador in South America (Stevens and others
2001, Natural Resources Conservation Service
2002). The species has naturalized in Hawaii,
Guam and other Pacific Islands, Taiwan, West
Africa, and probably in many other places in the
tropics (Peng and others 1998, Pacific Islands
Ecosystems at Risk 2002, Stevens and others
2001).
Ecology.—Cure-for-all is adapted to a wide

variety of soils and sites. It tolerates excessively-
well to poorly-drained soils, the full range of soil
textures, acid and alkaline reactions, salt and salt
spray, and compaction. A minimum of about
1000 mm of mean annual precipitation is required
General Description.—Cure-for-all, also known in upland sites, but it grows in much drier climates
as sourbush, sweetscent, wild tabacco, cattle along streams and near mangroves and marshes.
tongue, salvia, sauge rouge, guerit-tout, tabak Cure-for-all may be found from near sea level to
djab, zówèy mouten, is a shrub 1 to 3 m in height 1,000 m in elevation in Hawaii (University of
and up to 6 cm in basal stem diameter. Stem wood Hawaii Botany 2002). The species is intolerant
is moderately soft and brittle. Older plants are and cannot endure overhead shade or severe
supported by many flexible, lateral roots and may competition from brush or grass. It is common in
be sparsely branched to somewhat branchy. The disturbed areas such as construction sites,
twigs are stout. The foliage is concentrated on the riverbanks, the margins of hammocks, road cuts
branch ends. Leaves have petioles 1 to 3 cm long, and fill, vacant lots, eroded sites, landslides,
and ovate to elliptic blades 6 to 20 cm long with burned areas, and abandoned fields. In Hawaii,
rounded bases, pointed ends, usually entire edges, cure-for-all quickly invades burned areas, but
upper surfaces glabrate to densely pubescent, and being early successional, is soon replaced by other
lower surfaces velvet-pubescent. The terminal species (Smith and Tunison 1992). Plants sprout
inflorescences are broadly rounded corymbose after fires if they are not too intense (University of
panicles of heads of over 500 flowers each. The Hawaii Botany 2002).
corollas are usually pink, but vary from light
Reproduction.—Cure-for-all blooms in spring
577
and summer in Florida (Long and Lakela 1976) P. Lohr, and D. Atha. 2000. Medical plants used
and produces seeds prolifically. A collection of by Latino healers for woman’s health conditions
seeds from Puerto Rico weighed an average of in New York City. Economic Botany 54(3):
0.000025 g/seed or 40 million seeds/kg. Because 344-357.
the seeds failed to germinate on moist filter paper,
it is not known whether this represents a Gavilán-Yodú, R. and P.N. Hechavarría. 2002.
reasonable estimate for the species (author’s Efecto analgésico y antiinflamatorio de la tintura
observation). The seeds are wind-dispersed and de salvia (Pluchea carolinensis) al 30 % en ratas
probably require wet, bare soil to germinate and Wistar. Instituto Superior de Ciencias Médicas,
establish themselves. La Habana, Cuba. http://fcmfajardo.sld.cu/
jornada/trabajos/salvia/resumen/htm. 5 p.
Growth and Management.—In Puerto Rico,
most cure-for-all live for 2 to 4 years before dying Howard, R. A. 1989. Flora of the Lesser Antilles,
or dying back to the root and resprouting. Sprouts Leeward and Windward Islands. Vol. 6. Arnold
grow about 1.5 m in the first year. Establishment Arboretum, Harvard University, Jamaica Plain,
of new plants can probably be assured by MA. 658 p.
scarifying the soil before the wet season in the
presence of a seed source. Where it is necessary to Liogier, H.A. 1990. Plantas medicinales de Puerto
eliminate cure-for-all, grubbing out the plants or Rico y del Caribe. Iberoamericana de Ediciones,
spraying with broadleaf herbicides is Inc., San Juan, PR. 566 p.
recommended until tested treatments are available.
A seed insect, Acinia picturata (Diptera: Liogier, H.A. 1997. Descriptive flora of Puerto
Tephritidae), was introduced in Hawaii and Rico and adjacent islands. Vol. 5. Editorial de la
although now well established, has had no Universidad de Puerto Rico, Río Piedras, PR.
significant effect on the shrub (Alyokhin and 436 p.
others 2001).
Benefits.—Cure-for-all has an excellent ability to Tropical Florida. Banyan Books, Miami, FL.
colonize and stabilize disturbed areas and act as a 962 p.
nurse crop for later-successional species. Although
not specifically reported, as other members of the Natural Resources Conservation Service. 2002.
family, it probably provides a source of nectar and Plant profile: Pluchea carolinensis (Jacq.) G.
pollen for honeybees and other insects. It has Don. http://plants.usda.gov/cgi_bin/plant_profile
several herbal applications including aromatic .cgi?symbol=PLCA10. 3 p.
baths, control of fever, treatment of uterine
fibroids, relief of sore throat and stomach pain, Nelson, G. 1996. The shrubs and woody vines of
poultices for wounds and skin ulcers, as an Florida. Pineapple Press, Inc., Sarasota, FL.
analgesic, and for the treatment of malaria (Balick 391 p.
and others 2000, Liogier 1990, Vélez and van
Overbeek 1950). Analgesic and antiinflammatory Pacific Island Ecosystems at Risk. 2002. Invasive
effects have been demonstrated in laboratory trials plant species: Pluchea carolinensis (Jacq.) G.
with rats (Gavilán-Yodú and Hechavarría 2002). Don, Asteraceae. http://hear.org/pier3/plcar.htm.
2 p.
References
Peng, C.-I., C.-H. Chen, W.-P. Leu, and H.-F.
Alyokhin, A.V., R.H. Messing, and J.J. Duan. Yen. 1998. Pluchea Cass. (Asteraceae: Inuleae)
2001. Utilization of the exotic weed Pluchea in Taiwan. Botanical Bulletin of Academia
odorata (Asteraceae) and related plants by the Sinica 39(4): 287-297.
introduced biological control agent Acinia
picturata (Diptera: Tephritidae) in Hawaii. Smith, C.W. and J.T. Tunison. 1992. Fire and alien
Biocontrol Science and Technology 11: 711- plants in Hawai’i: Research and management for
718. native ecosystems. In: C.P. Stone, C.W. Smith,
and J.T. Tunison, eds. Alien plant invasions in
Balick, M.J., F. Kronenberg, A.L. Ososki, M. native ecosystems of Hawai’i: Management and
Reiff, A. Fugh-Berman, B. O’Connor, M. Roble, Research. University of Hawaii Cooperative
578
National Park Resources Studies Unit, University of Hawaii Botany. 2002. Alien plants
Honolulu, HI. p. 394-409. of Hawaii: Pluchea symphytifolia (Mill.) Gillis.
http://www.botany.hawaii.edu/faculty/cw_smith
Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. /plu_sym.htm. 1 p.
Monographs in Systematic Botany. Vol. 85, No. Vélez, I. and J. van Overbeek. 1950. Plantas
1. Missouri Botanical Garden Press, St. Louis, indeseables en los cultivos tropicales. Editorial
MO. 943 p. Universitaria, Río Piedras, PR. 497 p.
579
Prosopis glandulosa Torr. honey mesquite
FABACEAE
Synonyms: Prosopis chilensis (Molina) Stuntz

Prosopis odorata Torr. & Frém.

mesquites is common and identification of the
intermediate forms may be difficult (Uchytil
1990). The mesquite introduced into Hawaii is also
called “algarroba” but is a different species
(Sohmer and Gustafson 1987).
Range.—Honey mesquite is a common shrub in

Oklahoma, southern Kansas, Texas, New Mexico,
extreme southwestern Utah, Arizona, southeastern
California, and southern Nevada, and in Nuevo
Leòn, Tamaulipas, and Coahuila in Mexico
(Barneby 1989, Powell 1998).
Ecology.—Honey mesquite grows at elevations

below 1,800 m in a variety of soils and habitats. It
is an aggressive plant and will occupy disturbed
soils. It has a very deep (to 20 m) and wide root
system that confers drought tolerance (Dayton
1931, Kearney and others 1960). It grows in areas
Hunt Institute of Death Valley that receive only 5 cm annual
rainfall (Bainbridge and others 1990). It is
General Description.—Prosopis glandulosa is moderately frost tolerant. It has always had a wide
commonly known as honey mesquite, mesquite, geographical range but has been reported as being
glandular mesquite, prairie mesquite, algarobo, historically restricted to moist, rather narrow
and la péchita. The Spanish conquistadores named habitats such as arroyos, floodplains, and canyons
it “algaroba” as it reminded them of the algarroba, (Dick-Peddie 1993, Powell 1998). However,
or carob tree of Spain (USDAFS 1988). Mesquite contrary to these reports mesquite was not totally
are deciduous shrubs, or small trees, armed with restricted to these more moist habitats but was a
one or two, rarely absent, straight, stout, stipular component of upland communities prior to
spines and pinnately twice-compound leaves. European settlement and may have been over-
Flowers are in fragrant, globose heads and the looked by early explorers as it tended to be a low,
fruits are straight, slightly curved, long small shrub obscured in thick grasslands (Smiens
(approximately 20 cm) pods with partitioned and and others 1992). This breadth of habitat is
embedded seeds (Barneby 1989, Vines 1986). consistent with its hardiness to adverse
Typically the plants are 1- to 2-m high shrubs but environmental conditions.
may develop into 10 m tall trees (Bowers 1993).
An unusually large individual in Texas was Reproduction.—Honey mesquite flowers in
reported to be 12.8 m tall with a 15.8 m crown March to May, and the fruits mature before the
spread and 2.4 m girth (Powell 1998). In some summer rains in June through September (Bowers
habitats the plants remain less than 0.5 m tall 1993, Vines 1986). However, depending upon
(USDAFS 1988). Three varieties of P. glandulosa environmental conditions, flowering may extend
are recognized: var. torreyana, var. prostrata, and into September (Everitt and Drawe 1993, Vines
var. glandulosa (Kartesz 1994). Prosopis 1986). The fragrant, creamy-yellow to yellowish-
glandulosa has been known erroneously as P. green flowers are insect pollinated (Everitt and
juliflora (Kartesz 1994) that grows in the West Drawe 1993, Uchytil 1990). The fruit is a many-
Indies. Inter- and intraspecific hybridization within seeded, brown, shiny, dehiscent pod (legume).
580
Scarified seeds germinate readily and can remain as Peniocereus greggii Britt. and Rose (Tull 1987,
undigested and viable as they pass through animal author’s personal observation). It is not clear
digestive tracts (Wright 1982). The germination whether this is due to a favorable microclimate or
rate typically ranges from 75 to 90 percent (Vines a response to grazing pressures. Seeds of mesquite
1986). Seeds frequently contain weevil larvae are nutritionally rich and are important food for a
(Vines 1986). The pollen is reported responsible large number of wildlife species (Graham 1941,
for some hay fever (Kearney and others 1960) Tull 1987). Honey mesquite seeds may constitute
implying that it is also disseminated by wind. 5 to 37 percent of scaled quails' diets year round
(Davis and others 1975). Deer, javelina, and
Growth and Management.—Since livestock smaller wildlife species such as jackrabbits, feed
grazing was introduced in the Southwestern United on both pods and vegetation. It is important
States and Northern Mexico, honey mesquite has livestock feed, especially in times of drought.
increased to such an extent as to be considered a However, if livestock eat too large amounts,
pest. However, mesquite is a valuable soil binder rumen stasis, impaction, and death may result
and slows erosion of otherwise denuded land. The (Stubbendieck and others 1993). It is an important
increase in honey mesquite has been attributed to a “honey plant” and bees that forage its flowers
variety of factors. The two most often cited are a produce excellent quality honey (Dayton 1931). It
reduction in perennial grasses, which when healthy provides a good source of nectar and food for
and dense reduces mesquite seedling butterfly adults and larvae (Taylor and others
establishment, and the reduced incidence of fire 1997). The wood is resistant to termites and decay,
due to livestock over-grazing and fire suppression and is used for building. It is also prized as bar-b-
(Bahre 1995, Humphrey 1958, Fisher 1977). Other cue fuel as it is a hard, close-grained wood that
reasons include: reduced effectiveness of burns hot and slowly (Barneby 1989, CWAR
invertebrate seed predators, increased 2002). Honey mesquite was essential to Native
dissemination of seed by livestock and/or Americans who made breads from the pods, fuel
kangaroo rats, land clearing and cultivation, soil and utensils from the wood, medicines from the
compaction by livestock that hinders grass stems and leaves, rope from the roots, and dye
establishment, decreased activity of jackrabbits from the black pitch of the trunks (Moerman 1998,
and wood rats, and climate change (Bahre 1995, Tull 1987). The gum exudates of the bark may be
Glendening 1952). Depending upon the situation, a substitute for gum arabic and have been used to
herbicide and mechanical treatments are often both mend pottery (Powell 1998). An intoxicating
economical and effective in controlling scattered beverage is made from fermented meal (Bowers
and dense stands of mesquite (Schmutz and others 1993).
1992). Chaining and burning the trees destroy
above-round tissue, but old stumps of honey References
mesquite readily resprout. Burning is effective
only on small mesquite plants and where fuel is Bahre, C.J. 1995. Human impacts on the
adequate to carry a fire (Schmutz and others 1992). grasslands of southeastern Arizona In: M.P.
McClaran and T.R. Van Devender, eds. The
Benefits.—Honey mesquite plants bind soils, and Desert Grassland. The University of Arizona
plantings are recommended for erosion control Press, Tucson, AZ. p. 230-264.
(Graham 1941, Rorabaugh 1995). Mesquite,
particularly var. torreyana, is tolerant of saline Bainbridge, D.A., R.A. Virginia, and W.M. Jarrell.
soils and is a particularly useful cover in degraded 1990. Honey Mesquite. Forest, Farm, and
areas (Rorabaugh 1995, Schmutz and others 1992). Community Tree Network (FACT Net)
It has been used, with limited success, in http://www.winrock.org/forestry/factnet.htm.
revegetating copper mine wastes (Norem and
others 1982). Mesquite has been used in riparian Barneby, R.C. 1989. Fabales. Intermountain Flora,
restoration projects and provides excellent habitat Vascular Plants of the Intermountain West,
for many species of wildlife and birds (Rorabaugh U.S.A. Vol. 3, Part B. New York Botanical
1995). In one study, the number of quail using an Garden, Bronx, NY. 279 p.
area decreased as a consequence of mesquite
removal (Goodwin and Hungerford 1977). Honey Bowers, J.E. 1993. Shrubs and trees of the
mesquite also provides shelter for the germination Southwest Deserts. Southwest Parks and
and development of forbs, grasses, and cacti such Monuments Assoc. Tucson, AZ. 140 p.
581
Center for Wood Anatomy Research. 2002. U.S. Kearney, T.H., R.H. Peebles, and collaborators.
Department of Agriculture. Forest Service. 1960. Arizona flora. 2nd ed. University of
Tech. Fact Sheet. Prosopis. http://www2.fpl. California Press, Berkeley, CA. 1085 p.
fs.fed.us/TechSheets/HardwoodNA/htmlDocs/
prosop1.html. Moerman, D.E. 1998. Native American Ethno-
Davis, C.A., R.C. Barkley, and W.C. Haussamen.
1975. Scaled quail foods in southeastern New Norem, M.A., A.D. Day, and KL. Ludeke. 1982.
Mexico. Journal of Wildlife Management An evaluation of shrub and tree species used in
39(3): 496-502. revegetating copper mine wastes in the United
States. Journal of Arid Environments 5: 299-
Dayton, W.A. 1931. Important western browse 304.
plants. U.S. Department of Agriculture. Misc.
Publ. 101. Washington, DC. 214 p. Powell, A.M. 1998. Trees and shrubs of Trans-
Pecos and adjacent areas. University of Texas
Dick-Peddie, W.A. 1993. New Mexico Vegetation Press, Austin, TX. 498 p.
– past, present, and future. University of New
Mexico Press, Albuquerque, NM. 244 p. Rorabaugh, J.C. 1995. A superior accession of
western honey mesquite (var. torreyana) for
Everitt, J.H. and D.L. Drawe. 1993. Trees, Shrubs riparian restoration projects. Desert Plants.
and Cacti of South Texas. Texas Tech 11(4) 32-40
University Press, Lubbock, TX. 213 p.
Schmutz, E.M., E.L. Smith, P.R. Ogden, M.L.
Fisher, CE. 1977. Mesquite and modern man in Cox, J.O. Klemmedson, J.J. Norris, and L.C.
southwestern North America. In: B.B. Simpson, Fierro. 1992. Desert Grassland. In: R.T.
ed. Mesquite, its biology in two desert shrub Coupland ed. Ecosystems of the World. Vol.
ecosystems. Downden, Hutchinson, and Ross, 8A. Elsevier, London and New York. 469 p.
Stroudsburg, PA. p 17-88.
Smiens, F.E., D.D. Diamond, and C.W. Hanselka.
Glendening, G.E. 1952. Some quantitative data on 1992. Coastal Prairie. In: R.T. Coupland, ed.
the increase of mesquite and cactus on a desert Ecosystems of the World. Vol. 8AElsevier,
grassland range in southern Arizona. Ecology. London and New York. 469 p.
33 (3): 319-328.
Sohmer, S.H. and R. Gustafson. 1987. Plants and
Goodwin, J.G. and C.R. Hungerford. 1977. Habitat flowers of Hawaii. University of Hawaii Press,
use by native Gambel’s and scaled quail and Honolulu, HI. 160 p.
released masked bobwhite quail in southern
Arizona. Research Paper RM-197. USDA Stubbendieck, J., S.L. Hatch, and C.H. Butterfield.
Forest Service, Rocky Mountain Research 1993. North American Range Plants. 4th ed.
Station, Fort Collins, CO. 8 p. Univ. of Nebraska Press, Lincoln, NB. 493 p.
Graham, E.H. 1941. Legumes for erosion control Taylor, R.B., J. Zrutledge, and J.G. Herrera. 1997.
and wildlife. U.S. Department of Agriculture A field guide to common south Texas shrubs.
Misc. Pub. 412. Washington D.C. 153 p. Texas Parks and Wildlife Press, Austin, TX.
p. 106.
Humphrey, R.R. 1958. The desert grassland: a
history of vegetational changes and an analysis Tull, D. 1987. Edible and Useful Plants of Texas
of causes. Botanical Review 24: 193-252. and the Southwest. University of Texas Press.
Austin, TX. 518 p.
vascular flora of the United States, Canada and Uchytil, R.J. 1990. Prosopis glandulosa var.
Greenland. Vol.1 - Checklist. 2nd ed. Timber glandulosa. U.S.D.A. Forest Service, Rocky
Press, Portland, OR. 622 p. Mountain Research Station, Fire Sciences
Laboratory Fire Effects Information System,
http:// www.fs.fed. us/databse/feis/.
582
United States Department of Agriculture Forest Wright, R.A. 1982. Aspects of desertification in
Service. 1988. Range Plant Handbook. Dover Prosopis dunelands of southern New Mexico.
Publications, Inc. New York. 837 p. Journal of Arid Environments 5: 277-284.

of the Southwest. University of Texas Press.
583
Proustia vanillosma C. Wright sweet yellowcrown
ASTERACEAE
Synonyms: Proustia krugiana Urban

Perezia vanillosma Molinet & Gómez
John K. Francis
1800 mm of mean annual precipitation from a few

meters above sea level to elevations of about 550
m. Sweet yellowcrown grows on a wide variety of
well-drained soils derived from sedimentary
(including ultramaphic) rocks. It is moderately
tolerant to shade, growing under open to
moderately dense forest stands, as well as in
openings and at forest edges. The species is most
common in remnant primary stands and secondary
forests that have never been completely converted
to agriculture. Sweet yellowcrown slowly invades
abandoned fields and pastures after they become
forested. Although occasionally common and
forming small thickets, plants are usually scattered
(author’s observations).
General Description.—Sweet yellowcrown, a
name assigned by the Natural Resources Reproduction.—Sweet yellowcrown in Puerto
Conservation Service (2003), is a clambering Rico flowers near the end of the wet season
woody shrub reaching 1 to 3 m in height and 5 m (November and December) and matures seed at the
of extension. The species usually has multiple beginning of the dry season (January and
stems that sometimes reach 2 cm in diameter. February). A group of seeds collected in Puerto
They are covered with a lightly striated, gray- Rico averaged 0.000654 g/seed or 1,529 seeds/g.
brown bark and a green inner bark, and have a Sown on moist blotter paper, 88 percent of the
brittle, greenish-white wood with a 2-mm pith. seeds germinated within 13 days. Germination is
The stems layer (root) whenever they come in epigeal. Seeds are dispersed by wind. Seedlings
contact with the soil. The roots are brown and occur infrequently in forests. Once established,
flexible. Wood of branches is flexible, strong, and plants spread locally by layering (author’s
hard. The alternate, light-green leaves are thin to observations).
leathery with a rough surface, elliptic to narrowly
ovate, pointed to rounded at the tips, have margins Growth and Management.—Sweet yellowcrown
rough to dentate, and are up to 9 cm long. The has a moderate growth rate (about 0.5 m of stem
fragrant inflorescences are leafy, terminal panicles extension/year) and can live for at least a decade.
of heads that have three to seven florets. The Apparently plants (clones) survive longer than
corolla is yellow, 8 to 10 mm long. The seeds have individual stems. No planting or natural stand
an 8-mm straw-colored pappus (Liogier 1997). management experience is published or known to
Turner (1993) proposed a new genus, the author.
Berylsimpsonia, to replace Proustia in the Greater
Antilles. Benefits.—Sweet yellowcrown helps protect the
soil, furnishes cover for wildlife, and adds to the
Range.—Sweet yellowcrown is native to Puerto aesthetics of the forest.
Rico, Hispaniola, and Cuba (Liogier 1997). It is
not known to have been planted or naturalized References
elsewhere.
Ecology.—The species occurs in dry and moist Rico and adjacent islands. Vol. 5. Editorial de la
areas of Puerto Rico that receive from about 750 to
584
Universidad de Puerto Rico, San Juan, PR. cgi_bin/plant_profile.cgi?symbol=PRVA. [not
436 p. paged].
Natural Resources Conservation Service. 2003. Turner, B.L. 1993. Berylsimpsonia (Compositae:
Plants profile: Proustia vanillosma C. Wright, Mutiseae), a new genus for the Greater Antilles.
sweet yellowcrown. http://plants.usda.gov/ Phytologia 74: 349-355.
585
Prunus americana Marsh. American plum
ROSACEAE
Synonyms: Prunus americana Marsh. var. floridana Sarg.

Prunus domestica L. var. americana (Marsh.) Castigl.
John K. Francis
The fruits have a thick skin and sweet-tart, yellow

flesh. The brown stone forms a flattened oval,
ridged on one edge and grooved on the other
(Britton and Shafer 1908, Sargent 1923).
Range.—American plum is native to southeastern

Canada and the conterminous states of the United
States except Texas, Washington, Idaho, Oregon,
Nevada, and California where it is planted and
probably naturalized (Natural Resources
Conservation Service 2003, Sargent 1923). It has
been suggested that the current broad range is, in
part, the result of planting by Native Americans
and white settlers (Treeguide 2003). The species is
widely planted in temperate areas of the World and
probably has naturalized in some of them.
Ecology.—Across its vast range, American plum

grows in a wide variety of sites. In the eastern and
mid portions of the continent, it grows on
General Description.—American plum, also roadsides, old fields, vacant lots, fencerows,
known as Pottawattami plum, wild plum, yellow gullies, clearings, edges of woodlots, and open
plum, red plum, river plum, goose plum, hog prairies. In the West, it grows in riparian areas and
plum, and ciruela, is a shrub or small gnarled tree ditch banks, at the edges of fields, and in moist
generally 1 to 8 m in height (rarely to 11 m) with foothills. American plum grows at elevations up to
stem diameters up to 30 cm. Maximum size 2,300 m in Utah. The species grows on sand
depends on the environment. It usually produces a through clay soils on moist to somewhat dry sites.
single stem and branches low to the ground but It tolerates a moderate amount of salt in the soil
forms thickets by suckering from the roots. Stem (Forest Service 2003). American plum is intolerant
bark is about 1.25 cm thick, gray or dark brown of shade and will not grow in the understory of
tinged with red, with the outer layer separating forest or persist if overtopped by trees (Treeguide
into thin, persistent plates. The twigs are green at 2003). Other members of the genus and
first, becoming orange-brown and darker as they presumably American plum recover rapidly from
age. The plants become branchy and somewhat fires by sprouting (Forest Service 2003).
thorny, especially in older individuals. The wood
is hard, heavy, and close-grained, with reddish- Reproduction.—Flowering occurs from March
brown heartwood and light tan sapwood. Leaves through May and fruit ripens from late June
are dark green, elliptic to obovate, 4 to 10 cm long, through October, depending on habitat (Forest
with an elongated point at the tip and a rounded Service 2003, Oklahoma Biological Survey 1999).
base, margin serrate or doubly serrate, and having The flowers are pollinated principally by
usually glandless petioles 1.5 to 2 cm long. honeybees (Apis mellifera L.). Good fruit and seed
Flowers are white, five-petaled, about 2.5 cm crops are borne every 1 to 2 years (Grisez and
across, growing in two- to five-flowered lateral others 2003). Fresh fruits collected by the author
umbels. They are unpleasantly aromatic. Fruits in Utah averaged 6.572 + 0.139 g/fruit. Air-dried
(globose to oblong drupes) are 1.8 to 2.5 cm long seeds separated from them averaged 0.5336 +
and red to yellow (usually orange-red) when ripe. 0.0095 g/seed or 1,874 seeds/kg. Grisez (1974)
586
reported 1,918 cleaned seeds/kg, and that 60 of the plants to domestic ruminants and wild game
percent germinated after cold stratification. animals varies from good to poor depending on the
Germination is hypogeal (Young and Young animal species and the location (Forest Service
1992). Seeds are dispersed by mammals and birds. 2003). The cover provided by American plum
Animals that ingest the seeds, such as black bear thickets is important to many wild animals. The
(Ursus americanus), are more effective dispersers fruits are eaten by a number of species including
than those that do not (Forest Service 2003). Seeds bluejays (Cyanocitta cristata), brown thrashers
ingested by coyotes (Canis latrans) had a (Toxostoma rufum), mockingbirds (Mimus
significantly lower germination rate than uneaten polyglottos), red-headed woodpeckers
seeds (Cypher and Cypher 1999). Seeds may (Melanerpes erythrocephalus), bobwhite quail
remain in the soil seed bank for many years until (Colinus virginianus), white-tailed deer
disturbance creates conditions for growth (Odocoileus virginianus), raccoons (Procyon
(Treeguide 2003). After dispersed plants become lotor), squirrels (Sciurus spp.), and coyotes (Canis
well established, they begin forming clonal latrans) (Cowley 2003, Cypher and Cypher 1999,
thickets. Suckers may appear as much as 3 m away Kaiser 2001). The flowers furnish nectar food for
from the parent plants (Colorado Springs Utilities great purple hairstreak [Atlides halesus (Cramer)]
2003). and Sweadner’s Jupiter hairstreak [Callophrys
gryreus (Hübner)] butterflies (Cowley 2003).
Growth and Management.—American plum is Fruits are eaten raw or cooked, made into jams and
only capable of a moderate growth rate, usually jellies, and were dried for winter food in former
less than 30 cm in height per year (Michigan State times by Native Americans and pioneers. Several
University Extension 2003). Minimum seed- horticultural varieties of plum have been derived
bearing age is 4 years (Grisez and others 2003). from American plum. It has also been used in
Individual stems rarely live longer than 20 years hybrid crosses with other species of plums
(Treeguide 2003). However, clones may last much (Bircher and Bircher 2000). The wood, which has
longer. American plum fruits should be collected a specific gravity of 0.73 (Britton and Shafer
when fully mature for best quality seed. This can 1908), makes excellent firewood except for being
be done by hand stripping or shaking or beating crooked and limby. American plum is used in
onto a tarp spread under the shrub or tree. The amenity planting for wildlife and in revegetation
seeds should be cleaned of all pulp by macerating projects.
and washing. The seeds are usually 96 to 100
percent filled. Seeds of this species can be stored References
at room temperature for up to 30 months without
loss of viability. Seeds to be stored for longer Bircher, A.B. and W.H. Bircher. 2000.
should be surface dried and placed in a sealed Encyclopedia of fruit trees and edible flowering
container at 1 to 5 °C. Stratification of 90 to 150 plants in Egypt and the Subtropics. The
days at 2 to 5 °C before planting is recommended American University in Cairo Press, Cairo,
for after-ripening of the seed. Alternately, seed Egypt. 568 p.
may be sown in nursery beds 2.5 to 5 cm deep in
the fall. Seedlings are ready for lifting (bare-root) Britton, N.L. and J.A. Shafer. 1908. North
at the end of one growing season (Grisez and American trees. Henry Holt and Company, New
others 2003). Plantings may be made with bare- York. 964 p.
root stock and potted nursery seedlings. Direct
seeding in the fall with unstratified seed or spring Colorado Springs Utilities. 2003. Plant details.
with stratified seed in prepared seedspots will http://www.csu.org.cgi-bin/xeri/Xeriinclude?
normally yield new seedlings. The species can be Xeridetail?PIS-pra. 2 p.
grafted or used as root stock, and it has been
successfully propagated from stem cuttings (Forest Cowley, M. 2003. Rose family (Rosaceae).
Service 2003). http://www.nsis.org/garden/family/rose.html.
4 p.
Benefits.—American plum is an early spring
bloomer and with its white flowers beautifies the Cypher, B.L. and E.A. Cypher. 1999. Germination
forests and prairies for a week or more each year. rates of tree seeds ingested by coyotes and
The species also helps protect the soil and provides raccoons. American Midland Naturalist 142(1):
benefits to wildlife and humans. The browse value 71-76.
587
Forest Service. 2003. Fire effects information Natural Resources Conservation Service. 2003.
system: species: Prunus americana. http://fs. Prunus americana Marsh. http://plants.usda.
fed.us/database/feis/plants/tree/pruame/all.html. gov/cgi_bin/plant_profile.chi?symbol=PRAM.
14 p. 4 p.
Grisez, T.J. 1974. Prunus L. cherry, peach, and Oklahoma Biological Survey. 1999. Prunus
plum. In: C.S. Schopmeyer, tech. coord. Seeds americana Marsh. http://www.biosurvey.ou.
of woody plants in the United States. U.S. edu/shrub/prun-ame.htm. 2 p.
Washington, DC. p. 658-673. Sargent, C.S. 1923. Manual of the trees of North
America (exclusive of Mexico). Houghton
Grisez, T.J., J.R. Barbour, and R.P. Karrfalt. 2003. Mifflin, Boston, MA. 910 p.
Prunus L. cherry, peach, and plum. In: F.T.
Bonner and R.G. Nisley, eds. Woody plant seed Treeguide. 2003. American plum, Prunus
manual. http://wpsm.net/Prunus.pdf. [not americana Marsh. http://www.treeguide.com/
paged]. Species.asp?SpeciesID=770. 3 p.
Kaiser, J. 2001. 2001 featured plant, American Young, J.A. and C.G. Young. 1992. Seeds of
plum, Prunus americana. In: Plants for woody plants in North America. Dioscorides
conservation, Vol. 4, No. 1. Elsberry Plant Press, Portland, OR. 407 p.
Materials Center, Elsberry, MO. p. 1.
Michigan State University Extension. 2003.

Prunus americana—American plum. http://
www.msue.msu.edu/msue/imp/modzz/00002025
.html. 1 p.
588
Prunus emarginata (Dougl. ex Hook.) D. Dietr. bitter cherry
ROSACEAE
Synonyms: None
Christopher Ross
persists for a long time but does not occur until

late in the post-disturbance seral progression.
Seed dispersal is by animals (Halperin 1989,
personal observation).
Reproduction.—Three to 12 five-petaled flowers

on 3- to 12-mm pedicels occur in sometimes flat-
topped racemes. The sepals and hypanthium are
glabrous to puberulent. White petals are 4 to 8 mm
long. Fruits are obvoid to round about 7 to 14 mm
long, smooth, with a red to purple fleshy pulp
(Wilken 1993). Seeds may at times undergo heavy
predation and mortality while green, apparently
from sawflies of the genus Hoplocampa (Weaks
2001). Rodents, including chipmunks, eat the
seeds (John Francis and Maurice Beck, personal
communication).
Fire Effects.—Although bitter cherry is a prolific

sprouter after fire, Leege (1979) found that
repeated fires eventually result in heavy plant
mortality. However, in those cases, seedlings
developed to replace mature plants lost to fire.
General Description.—Bitter cherry ranges from Growth and Management.—Bitter cherry has
a shrub to a tree of 10 m. It often forms dense been effectively controlled on conifer planting
thickets. Clustered, deciduous elliptic to obovate, sites by application of the herbicide 2,4,5-T (now
crenate-serrate leaves 20 to 65 mm in length are on banned in the United States) (Bock and others
3 to 12 mm petioles (Wilken 1993, Mozingo 1978). It may be heavily attacked by tent
1987). The leaves, stems, and flowers all have a caterpillars and other insects at times.
very strong cyanide odor (bitter almond) when
crushed. Benefits.—Blue grouse and other birds are fond of
the fruits of bitter cherry. They are also a preferred
Range.—Bitter cherry is found in cooler parts of summer and fall food of black bears (Unsworth
Arizona, California, far western Nevada, Idaho, and others 1989). Leaves are browsed by sheep
Utah, New Mexico and Montana and north to and cattle, despite their cyanogenic properties,
British Columbia (Mozingo 1987, Esser 1995). which have caused reported stock poisonings
(Mozingo 1987). In the Pacific Northwest, bitter
Ecology.—Bitter cherry site dominance may not cherry is eaten by deer and elk (Crouch 1968),
peak until three decades after disturbance, although it is not a preferred forage (Leege 1979,
suggesting that it is not an obligate seed bank Klebenow 1965). Native Americans used long
species (Oakley and Franklin 2001). However, in fibrous strips of bark for twine and basketry, and
some cases following fire it may quickly dominate used the roots for a variety of medicinal purposes
a site (personal. observation.). However, others but made little use of the bitter fruit (Turner and
(Morgan and Neuenschwander 1988) have Bell 1973). Bitter cherry essence is marked by
considered it an obligate seed bank species. It may herbalists, who claim that it aids patience and
achieve major dominance of disturbed sites, which short attention spans. Bitter cherry has also been
589
used for mine and highway reclamation (Everett Morgan, P. and L.F. Neuenschwander. 1988.
and others 1980 and personal observation). Seed-bank contributions to regeneration of shrub
species after clear-cutting and burning.
References Canadian Journal of Botany 66:169-72.
Bock, J.H., M. Raphael, and C.E. Bock. 1978. A Mozingo, H. N. 1987. Shrubs of the Great Basin.
comparison of planting and natural succession University of Nevada Press, Reno Nevada. 342
after a forest fire in the northern Sierra Nevada. p.
Journal of Applied Ecology 15:597-602.
Oakley, B.B. and J. F. Franklin. 2001. Bitter
Crouch, G.L. 1968. Forage availability in relation cherry (Prunus emarginata) distribution,
to browsing of Douglas-fir seedlings by black- successional dynamics, and implications for the
tailed deer. Journal of Wildlife Management 32: role of the seed bank.
542-553. http://www.cisti.nrc.ca/cisti/journals/cjb/b98-
162.html. p. 1725-1732.
Esser, L.L. 1995. U.S. Department of Agriculture,
Forest Service, Rocky Mountain Research Turner, N.C. and M.A.M. Bell. 1973. The
Station, Fire Sciences Laboratory (2001, Ethnobotany of the Southern Kwakiutl Indians
October). Fire Effects Information System, of British Columbia. Economic Botany 27: 257-
http://www.fs.fed.ss/database/feis/. [not paged]. 310.
Everett, R.L., R.O. Meeuwig, and R. I. Butterfield. Unsworth, J.W., J.J. Beecham, and L.R. Irby.
1980. Revegetation of untreated acid spoils at 1989. Female black bear habitat use in West-
Leviathan mine, Alpine County, California. central Idaho. Journal of Wildlife Management
California Geology 32: 8-10. 53: 668-673.
Weaks, A. 2001. Bitter cherry sawfly.

Halperin, C.B. 1989. Early successional patterns of http://tardigrade.org/natives/cherrysawfly.html.
forest species: Interactions of life history traits 3 p.
and disturbance. Ecology 70: 704-720.
Wilken, D.H. 1993. Prunus In: Hickman, J.C., ed.
Klebenow, D.A. 1965. A montane forest winter 1993. The Jepson Manual: higher plants of
deer habitat in western Montana. Journal of California. University of California Press,
Wildlife Management 29: 27-33. Berkeley and Los Angeles. 1,400 p.
Leege, T.A. 1979. Effects of repeated prescribed

burns on Northern Idaho elk browse. Northwest
Science 53:107-113.
590
Prunus pumila L. sand cherry
ROSACEAE
Synonyms: Cerasus pumila (L.) Michx.

Prunus susquehanae Willd.
Prunus cuneata Raf.
Prunus depressa Pursh.
Prunus besseyi Bailey.
Don C. Bragg
are erect and diffusely branched, with some

decumbent branches on active dunes.
P. pumila L. var. cuneata (Raf.) Bailey,
the Appalachian sand cherry, has oblong to
oblong-obovate leaves with acute bases, often 20
to 30 mm wide at maturity, pale green above and
glaucous below. Stems are erect or diffusely
branched. This variety is most common on dry or
rocky sites.
P. pumila L. var. depressa (Pursh)
var. pumila Gleason, the flat sand cherry, has leaves that are
narrow (10 to 20 mm wide), oblanceolate, often
with obtuse, long-tapering bases, pale green above
and whitish below. Stems are prostrate, forming
low mats up to 2 m wide. New shoots are reddish,
highly lustrous, and often freely rooted.
P. pumila L. var. besseyi (Bailey)
Gleason, the western sand cherry, has leaves that
are somewhat glaucous underneath, oblanceolate,
up to 18 mm wide, with acute to acuminate tips
and long-cuneate bases.
var. cuneata
Range.—Sand cherry is widely distributed in the
Illustration source: (used with permission) Cranbrook northern half of the United States and eastern
Institute of Science Canada, from New Brunswick down the Atlantic
Seaboard to North Carolina, westward to Utah,
General Description.—Sand cherry, also called Wyoming, and Montana, and as far north as
beach plum, “cerise de sable” (Fernald 1923), or northern Ontario and Quebec (BONAP 1999,
dwarf American cherry, is a diffusely-branched, Cusik 1985). The varieties of sand cherry are
low growing (0.5 to 3 m tall, depending on variety generally geographically distinct (BONAP 1999,
and habitat) to sometimes decumbent or prostrate Gleason 1952). Prunus pumila var. pumila is most
shrub (Fernald 1923, Gleason 1952, Lamson- common in the Great Lakes region, especially in
Scribner 1891). Older stems develop a grayish, Michigan, Minnesota, and Wisconsin. Prunus
glabrous bark, while younger twigs are often pumila var. depressa has been reported from New
tannish- to reddish-brown or brown. Its alternate Brunswick to Pennsylvania and New Jersey, with
leaves have glandular petioles up to half the blade isolated populations in Wisconsin, Tennessee, and
length. While Fernald (1923) listed three different possibly Kentucky. Prunus pumila var. cuneata
species of sand cherry (P. pumila, P. susquehanae, ranges from Maine to Minnesota (and possibly
and P. depressa), Gleason (1952) recognized four Arkansas), southeast to Indiana and North
varieties of P. pumila:P. pumila L. var. pumila, the Carolina. Prunus pumila var. besseyi is primarily a
Great Lakes sand cherry, has narrow (10 to 18 mm western subspecies, extending from Minnesota to
wide) oblanceolate leaves, narrowly cuneate at Kansas westward to Utah and Montana, with
their base, acute or acuminate at their apex, isolated subpopulations noted as far east as
lustrous on their top surface and pale below. Stems
591
Michigan. The widespread distribution of sand mammals. Sand cherry can also vegetatively
cherry has helped to conserve the species, although propagate (Olson 1958).
it is considered locally threatened by some States
(e.g., Arkansas, Ohio) because of the rarity of the Growth and Management.—Sand cherry growth
sometimes specialized habitats it occupies is best under open canopy conditions. As with
(Arkansas Department of Planning 1974, Cusik many other species that specialize on poor sites,
1985, Emmitt and Cusik 1983). rates of growth are usually better on higher quality
locations (i.e., those with abundant moisture and
Ecology.—As its name suggests, sand cherry is nutrients). However, these conditions also promote
abundant in sandy areas, although var. cuneata is the growth of competitors that can exclude this
most common on rocky sites (Billington 1943, low-growing, shade-intolerant species. Sand
Gleason 1952). In the Great Lakes region, sand cherry’s inconspicuous stature and spreading root
cherry is often found on deep, excessively drained system leave it vulnerable to overshading,
glacial sand plains or sand dunes bordering major trampling, soil compaction, erosion, and other
bodies of water (Cowles 1899, McAtee 1920, surface disturbances (Cusick 1985, Emmitt and
Walp 1935). Sand cherry also grows along gravel Cusick 1983). Road grading, for example, may
bars or shorelines, cliff faces, rocky slopes, or have destroyed the two known pockets of sand
even on calcareous, saline, or serpentine soils cherry in Arkansas (Arkansas Department of
(Fernald 1923, Gleason 1952). The habitat Planning 1974). Habitat protection coupled with
preferences of sand cherry have restricted its the restoration of openings should help conserve
abundance in many parts of its range. For example, this species.
the sand cherry in Arkansas appears to have been
limited to two locations on remnants of the Grand Benefits.—The small stature of sand cherry has
Prairie in Prairie County (Smith 1988). Sand minimized its economic benefits, as it does not
cherry is opportunistic in its distribution, produce merchantable wood and fruit production is
frequently colonizing road cuts, gravel pits, or generally limited. At least one cultivated hybrid is
railroad beds (Cusik 1985, Fernald 1923, Stevens commercially available. Popular for its colorful
1961). Preferred habitat is typically open, with few foliage, the purpleleaf sand cherry [P. x cistena
trees, other shrubs, or herbs to compete for light, (N.E. Hansen) Koehne] is a cross between P.
nutrients, and water on the harsh sites it occupies. pumila and P. cerasifera Ehrh. Fernald (1923)
However, some have reported sand cherry as fondly described the fruit of sand cherry along the
abundant in closed forests (Gysel 1966). Human rivers of New England and southeastern Canada:
activities and habitat alteration from changing “...its juicy black ‘plums’ are highly prized either
natural disturbance regimes have been blamed for raw, cooked or as the source of a rich syrup-like
localized extinctions of sand cherry (Arkansas jelly,” suggesting that specialty food markets may
Department of Planning 1974, Cusick 1985, be possible. While the ecological benefits of sand
Drayton and Primack 1996). cherry are poorly understood, the vanishing
habitats occupied by this species often have
Reproduction.—Depending on geographic considerable value. For example, var. cuneata was
location, from April to June a sand cherry may one of the primary floristic components of the
produce two to four white insect-pollinated Albany Pine Bush in northern New York, which is
flowers in umbels scattered amongst its leaves. an important local refuge for numerous
Voss (1954) reported that butterflies pollinated amphibians and reptiles (Stewart and Rossi 1981).
sand cherry flowers in northern Michigan. Sand Sand cherry may also play a critical pioneering
cherry fruits ripen by late July or August and are role in the ecosystems where it is abundant. Its
typically reddish- or purplish-black to nearly deep root network and dense thickets help to
black, without bloom, subglobose to globose, and stabilize shifting sand, allowing for the invasion of
10 to 15 mm in diameter (Billington 1943, Fernald other plant species and colonization by important
1950, Gleason 1952). Prunus pumila var. pumila is invertebrates such as ants (Olson 1958, Talbot
virtually inedible for humans (Rehder 1958), 1934). Soil stabilization and organic matter
though Fernald (1923, 1950) mentions the production by sand cherry also contribute to
palatability of var. cuneata and var. besseyi. Given nitrogen biogeochemistry in dune habitats
their size, fleshy and edible fruit, and large stone, (Robertson and Vitousek 1981).
seed dispersal is primarily through birds and small
592
References Gysel, L.W. 1966. Ecology of a red pine (Pinus
resinosa) plantation in Michigan. Ecology 47:
Arkansas Department of Planning. 1974. Arkansas 465-472.
natural areas plan. Arkansas Department of
Planning, Little Rock, AR. 248 p. Lamson-Scribner, F. 1891. A sketch of the flora of
Orono, ME. Botanical Gazette 16: 228-234.
Billington, C. 1943. Shrubs of Michigan. Bulletin
20. Cranbrook Institute of Science, Bloomfield McAtee, W.L. 1920. Notes on the jack pine plains
Hills, MI. 249 p. of Michigan. Bulletin of the Torrey Botanical
Club 47: 187-190.
BONAP. 1999. Synthesis of the North American
flora, version 1.0 (CD). North Carolina Olson, J.S. 1958. Rates of succession and soil
Botanical Garden, University of North changes on southern Lake Michigan sand
Carolina, Chapel Hill, NC. dunes. Botanical Gazette 119: 125-170.
Cowles, H.C. 1899. The ecological relations of the Rehder, A. 1958. Manual of cultivated trees and
vegetation on the sand dunes of Lake shrubs, 2nd ed. The Macmillan Co., New York.
Michigan. Botanical Gazette 27: 167-202. 996 p.
Cusick, A.W. 1985. Prunus pumila var. pumila L. Robertson, G.P. and P.M. Vitousek. 1981.
(Great Lakes Sand Cherry). Abstract. Ohio Nitrification potentials in primary and
Dept. of Natural Resources Division of Natural secondary succession. Ecology 62: 376-386.
Areas and Preserves. http://www.dnr.
state.oh.us/ODNR/dnap/Abstracts/P/prupumi. Smith, E.B. 1988. An atlas and annotated list of
htm. 2 p. the vascular plants of Arkansas, 2nd ed.
Department of Botany and Microbiology,
Drayton, B. and R.B. Primack. 1996. Plant species University of Arkansas, Fayetteville, AR.
lost in an isolated conservation area in 489 p.
metropolitan Boston from 1894 to 1993.
Conservation Biology 10: 30-39. Stevens, O.A. 1961. Plants of Fargo, North
Dakota. American Midland Naturalist 66: 171-
Emmitt, D.P. and A.W. Cusick. 1983. Prunus 177.
pumila var. cuneata Willd. (Sand Cherry).
Abstract. Ohio Dept. of Natural Resources Stewart, M.M. and J. Rossi. 1981. The Albany
Division of Natural Areas and Preserves. Pine Bush: a northern outpost for southern
http://www.dnr.state.oh.us/ODNR/dnap/ species of amphibians and reptiles in New
Abstracts/P/prupumi2.htm. 2 p. York. American Midland Naturalist 106: 282-
292.
Fernald, M.L. 1923. The identities of the sand
cherries of eastern America. Rhodora 25: 69- Talbot, M. 1934. Distribution of ant species in the
74. Chicago region with reference to ecological
factors and physiological toleration. Ecology
Fernald, M.L. 1950. Gray’s manual of botany, 8th 15: 416-439.
ed. American Book Co., New York. 1,632 p.
Voss, E.G. 1954. The butterflies of Emmet and
Gleason, H.A. 1952. Illustrated flora of the Cheboygan Counties, Michigan with other
Northeastern United States and adjacent notes on northern Michigan butterflies.
Canada, Vol. 2. Lancaster Press, Inc., American Midland Naturalist 51: 87-104.
Lancaster, PA. 655 p.
Walp, R.L. 1935. Shrubs of Cheboygan and
Emmet Counties, Michigan. American
Midland Naturalist 16: 230-247.
593
Prunus virginiana L. chokecherry
ROSACEAE
Synonyms: Cerasus demissa Nutt.

Cerasus virginiana L.
Padus demissa Nutt.
Padus melanocarpa A. Nelson
Padus virginiana L.
Prunus demissa Nutt.
Bruce L. Welch
Hypanthium and pedicels are also glabrous. The

mature fruit 6 to 8 mm thick, known as a drupe, is
dark red to black in color (Cronquist and others
1997, Welsh and other 1987). The diploid
chromosome number of chokecherry is 16, 26, or
32 (Cronquist and others 1997).
Taxonomy.—Two varieties of chokecherry are

recognized: var. melanocarpa and var. demissa
(Cronquist and others 1997). They are in the words
of Cronquist and others 1997: “weakly separated
as follows: 1. Leaves glabrous or rarely with a
few tufts of hairs in the axils of the lateral leaf
veins beneath; drupe blackish.........var.
Melanocarpa (A. Nelson) Sarg. 2. Leaves
pubescent; drupe dark red; California, in White
Mountains and Sierra Nevada, westward
......var. demissa (Nutt.) Torr.”
Range.—Chokecherry occupies a variety of sites

General Description.—Chokecherry is a across Canada and the United States (Little 1976).
thornless, deciduous shrub or small tree that grows It grows in 20 ecosystems and occurs in 47 of
to 8 m tall (Cronquist and others 1997). Young Kuchler’s plant associations (Pacific Southwest
stems are finely puberulent, greenish at first then Experiment Station. 2002). The Society of
become glabrous and reddish in color. Older stems American Foresters list chokecherry as occurring
are ashy-gray with a reddish-brown undertone in 51 of its cover types, and the Society for Range
(Cronquist and others 1997, Welsh and other Management finds chokecherry growing in 40 of
1987). The alternate leaves are simple. A pair of its rangeland cover types (Pacific Southwest
reddish glands appears on the petioles near the Experiment Station 2002). It ranges from
base of the leaves. Leaves are 2 to 10 cm in length Newfoundland in eastern Canada to British
and 1.5 to 7 cm in width. Leaves are elliptic to Columbia in the west (Little 1976). Scatted stands
oblong-ovate, finely serrated and abruptly can be found as far north as the southern portion of
acuminate apically. At the base, the leaves are the Northwest Territories of Canada and as far
acute to rounded. Leaves are dark green above and south as western Texas, Southwest, and southern
pale beneath. The midrib is impressed above and California (Little 1976). In the United States
prominent beneath. (Cronquist and others 1997, chokecherry occurs in the Northeast, Midwest,
Welsh and others 1987). Inflorescence is a raceme Central Plains, Pacific Northwest, and
that is 4 to 20 cm long with leafy peduncles 2 to 8 Intermountain regions (Little 1976). Isolated
cm long. The white perfect flowers are 10 to 20 patches occur in Virginia, West Virginia,
mm wide and numerous with 4 to 17 mm long Tennessee, Kentucky, North Carolina and
pedicels. Petals are white and are 4 to 6 mm long Oklahoma (Little 1976).
and suborbicular. Sepals are fringed and glabrous.
594
Ecology.—Usually chokecherry can be found excessive livestock grazing has damaged some
growing in canyon bottoms, sheltered slopes, populations in many areas of the northern Great
along streams and roads. It grows in a wide range Plains. Hydrogen glycoside prunasin is a toxin
of soils, ranging from Entisols to Mollisols that produced by chokecherry that could be poisonous
have textures ranging from sandy loams to clays. It to grazing livestock. Highest levels (5 percent) are
can be found from 177 (Michigan) to 3,100 (Utah) found in new stems and leaves. These levels
m in elevation where the combinations of soil and diminish over the growing season to 1.2 to 2.2
topography permit greater than average percent.
accumulation of moisture (Pacific Southwest
Experiment Station. 2002). Weakly salty soils are Benefits.—Chokecherry adds to the biodiversity
tolerated by chokecherry, but it cannot tolerant of a multitude of ecosystems. It provides habitat
soils that are poorly drained or suffer from and food for a number of wildlife species, and
prolonged flooding. It grows in soil pH ranges watershed protection. Fruits, leaves, or twigs are
from 3.5 to 7.6. It is found in numerous habitat and eaten by bears, moose, bighorn sheep, pronghorn,
plant associations that range from post disturbance elk and deer. A number of small of mammals also
invaders to early successional to climax or stable consume chokecherry including coyotes,
(Pacific Southwest Experiment Station 2002). snowshoe hares, red foxes, bobcats, raccoons, and
Chokecherry is intolerant to intermediate in porcupines. Chokecherry fruits are eaten by many
tolerance of shade and resprouts from root crowns birds including robins, western, eastern, and
and rhizomes readily, thus giving it the ability to mountain bluebirds, European starlings, and
persists under open or forest canopies of moderate Columbian sharp-tailed grouse. Livestock also eat
densities. It is well adapted to disturbance by fire. chokecherry. Winter twigs contain 38.9 percent
Chokecherry is a primary host of the eastern tent total digestible nutrients, which is lower than a
caterpillar and a fungus Plowrightia stansburiana, number of winter shrubs (Dietz 1972, Welch
which causes black knot-like tumors on stems 1981.) Winter protein, calcium, and phosphorus
(Pacific Southwest Experiment Station 2002). levels are above average. Fruits are harvested to
Western X virus can kill entire stands of make wines, syrups, jellies, and jams. Chokecherry
chokecherry. plants are planted as ornamentals, for enhancing
backyard wildlife habitats, and as windbreaks.
Reproduction.—Chokecherry is among the first
of the deciduous woody shrubs to leaf out and to References
flower in the spring. It can reproduce sexually
from seeds and asexually from root crowns and Auger, J., S.E. Meyer, and H.L. Black. 2002. Are
rhizomes. Seeds are encased in a stony endocarp American black bears (Ursus americanus)
and have an after-ripening requirement for legitimate seed dispersers of fleshy-fruited
germination. Heat treatment improves shrubs?American Midland Naturalist. 147:352-
germination. Seeds weigh about 0.095 grams 367.
(Grisez 1974). Passing through digestive systems
of mammals and birds may enhance germination Cronquist, A., N.H. Holmgren, and P.K. Holmgren
(Auger and others 2002). Also a number of birds 1997. Intermountain flora: vascular plants of the
and mammals may be responsible for long Intermountain west, U.S.A. Vol. 3. Part A;
distance dispersal (Auger and others 2002). Subclass Rosidae (except Fabales). The New
Chokecherry seeds persists in the soil seedbank . York Botanical Garden, New York. 446 p.
Growth and Management.—Heights of this

Dietz, D.R. 1972. Nutritive value of shrubs. In:
species vary greatly as to variety and site quality.
C. M. McKell, J. P. Blaisdell, J. R. Goodin, tech.
Chokecherry may grow as high as 12 m with a
eds. Wildland shrubs–their biology
truck diameter 20 cm on some sites in the Great
andutilization, an international symposium;
Basin. It has a deep root system that can grow to
Proceedings; 1971 July; Logan, UT. General
depths exceeding 1.8 m with lateral roots more
Technical Report INT-1. U.S. Department of
than 10.6 m in length. Rhizomes ranging from 1 to
Agriculture, Forest Service, Intermountain
2 cm in diameter are produced by this species.
Chokecherry rhizomes sprout at a faster rate with a
UT. p. 289-302.
higher percentage of sprouts than Gambel oak. It is
moderately tolerant of browsing. However,
595
Grisez, T.J. 1974. Prunus L. Cherry, peach, and Pacific Southwest Experiment Station. 2002. Fire
plum. In: C.S. Schopmeyer, tech. coord. Seeds effects information system. http://www.
of woody plants in the United States. fs.fed.us/ database/feis/plant. 33 p.
Agriculture Handbook 654. U.S. Department of
Agriculture, Forest Service, Washington, DC.
Welsh, S. L., N. D. Atwood, S. Goodrich, L.
p. 658-673.
C.Higgins. 1987. A Brigham Young
University, Utah flora. Great Basin Naturalist
Little, E.L., Jr. 1976. Atlas of United States trees.
Memoirs 9. Provo, UT. 894 p.
Vol. 3. Minor Western hardwoods.
Miscellaneous Publication 1,314. U.S.
596
Psidium guajava L. guava
MYRTACEAE
Synonyms: Psidium pumilum Vahl

Psidium guajava L. var. pumilum (Vahl) Griseb.
Psidium guajava L. var. cujavillum King & Urban
John K. Francis
flowers are usually solitary with four or five white

petals. Fruits (berries) are globose or pear-shaped,
with a prominent, persistent calyx. They are at
first hard and green, becoming softer, and yellow
at maturity and 3 to 5 cm long. Inside the thin rind
is a sweet-tart flavored pink or yellow pulp
containing many hard, angular, yellow seeds about
2 mm long. The species has 2n = 21, 22, 30, or 33
chromosomes (Howard 1989, Liogier 1994, Long
and Lakela 1976, Pennington and Sarukhan 1968,
Range.—Guava originated somewhere in the

Neotropics and was spread nearly to the extent of
its adapted climate before the arrival of Europeans
(Morton 1987). Seeds found in Peruvian
archeological sites seem to indicate that it
originated in that area (Rain-tree 2002). Guava has
been planted in nearly every tropical and frost-free
subtropical country and has naturalized in most of
them (Howard 1989).
Ecology.—Guava grows on soils of all textures

General Description.—Guava, also known as derived from most parent materials. Well-drained
guayaba, guayabo, arazá-puitá, goyavier, and and poorly drained soils, soils with pH’s from 4.5
gobiabiera, is an evergreen shrub or small tree 2 to to 9.4, mildly salty soils, and soils both rich and
8 m in height and up to 40 cm in diameter at breast poor in basic cations are tolerated. Guava grows at
height. Plants may have a single stem, especially if near sea level in coastal environments up to 2,300
crowded in secondary forest, but individuals m in elevation in Ecuador (Morton 1987). It grows
receiving ample light usually develop secondary naturally in areas of Puerto Rico that receive from
stems arising from the main stem near the ground. about 1000 to 3000 mm of mean annual
The branches and stems are usually crooked and precipitation. Guava withstands drought very well.
have a smoothish, cream to reddish-brown bark The species is moderately intolerant of shade. It
between thin, irregular scales that peel off. Guava develops a broad, low crown if open grown, grows
bark is 5 to 8 mm thick (Pennington and Sarukhan a more vertical crown with side shade, and
1968). The sapwood is light brown and the becomes tall and spindly in intermediate crown
heartwood is reddish brown, hard, heavy (specific positions. Saplings can endure a few years in the
gravity of 0.8), and strong (Little and Wadsworth understory of low basal-area secondary forests.
1964). The roots are slender. The young twigs are Guava survives the competition of weeds, grass,
four-angled, slightly winged, and green, turning and brush well. Growth is benefited by root
brown with age. The leathery and light green or association with arbuscular mycorrhizal fungi
yellow green opposite leaves have petioles 4 to 7 (Samarao and Martins 1999). Although they rarely
mm long, and elliptic or oblong blades, 8 to 14 cm kill the plants, a number of insects and diseases
long that are short or round pointed at both ends. affect the species. As many as 80 percent of
The foliage is aromatic when crushed. The axillary unprotected fruits may be attacked by
597
Mediterranean fruit flies (Ceratitis capitata Wied.) The fruit rinds are candied or stewed in syrup.
(Popenoe 1948). This thin-barked species is easily Nearly all the commercial production comes from
top-killed by fire and is sensitive to frost (von improved varieties with large fruits and few seeds.
Carlowitz 1991). The ripe fruits contain the following components:
water 84 percent, ash 0.7 percent, protein 0.8
Reproduction.—Guava flowers and fruits nearly percent, fiber 5.6 percent, total sugars 5.4 percent,
throughout the year (Little and Wadsworth 1964). starch 2.5 percent, and fat 1.0 percent (Popenoe
Many individual plants bear just once per year but 1948). The fruit pulp is rich in vitamins A and C
not necessarily synchronized with other guava (Morton 1987). Guava wood is used for tool
plants in the area. Honey bees (Apis mellifera) are handles, carving, and fuel (Advisory Committee
the chief pollinators (Morton 1987). Time from on Technology Innovation 1983). In folk
flowers to ripe fruits ranges from 102 to 124 days medicine, extracts of roots, bark, and leaves are
(Samson 1986). Thirty-one ripe fruits collected used to treat gastroenteritis, vomiting, diarrhea,
from wild plants in Puerto Rico averaged 32.8 g dysentery, wounds, ulcers, toothache, coughs, sore
and ranged from 13.5 to 61.8 g. Air-dried seeds throat, inflamed gums, and a number of other
averaged 0.0079 + 0.0001 g/seed or 127,000 conditions (Morton 1987). Guava leaf tea is
seeds/kg. Sown on moist peat, 63 percent widely used to control blood sugar of diabetics in
germinated between 15 and 60 days of sowing. Japan and elsewhere. It has been shown to be
Germination was epigeal (author’s observation). effective in vitro, in mice, and in human
Birds and mammals disperse the seeds (Invasive volunteers (Deguchi and others 1998). The basis
Species Specialist Group 2002). Root cuttings are for herbal treatment of diarrhea was established by
often used for commercial propagation of demonstrating inhibition of eight bacteria species
improved varieties (Morton 1987). Branches and and amoebas, and antispasmodic activity (Tona
stems layer (root) when they come in contact with and others 1999).
moist soil (author’s observation), and suckers also
arise from roots near the trunk. Established plants References
coppice readily and withstand repeated cutting
(Advisory Committee on Technology Innovation Advisory Committee on Technology Innovation
1983). 1983). Firewood crops. Vol. 2. National
Academy of Sciences, National Academy Press,
Growth and Management.—Guava seedlings Washington, D.C. 92 p.
grow at a moderate rate, but older plants grow
more slowly. Under good conditions, guava begins Deguchi, Y., K. Osada, K. Uchida, H. Kimura, M.
bearing fruits in 3 or 4 years. Plants live 40 years Yoshikawa, T. Kudo, H. Yasui, and M.
or more (Popenoe 1948). It is probably unwise and Watanuki. 1998. Effects of extract of guava
unnecessary to plant wild guava because the leaves on the development of diabetes in the
species is tough and aggressive and often invades db/db mouse and on the postprandial blood
agricultural (especially cattle pasture) and forest glucose of human subjects. Nippon
lands. Control is sometimes attempted, especially Nogeikagaku Kaishi 72(8): 923-931.
in cattle pastures and plantations. Sheep and goats
graze the leaves and strip the bark and have been Howard, R.A. 1989. Flora of the Lesser Antilles,
used to control it. Several herbicides are effective Leeward and Windward Islands. Vol. 5. Arnold
in controlling infestations (Pacific Island Arboretum, Harvard University, Jamaica Plain,
Ecosystems at Risk 2002). Repeated heavy MA. 604 p.
plowing or repeated burning are also effective
measures (Mune and Parham 1956). Invasive Species Specialist Group. 2002. Notes
from Jim Space on his survey of invasive plant
Benefits.—Guava helps protect the soil and can be species in Tonga. The World Conservation
a major participant in reforestation of disturbed Union. http://www.issg.org/features/
areas and abandoned pastures. It furnishes food invasives_on_tonga.html. 4 p.
and cover for wildlife. Domestic animals eagerly
consume the fruits. Guava is one of the most Liogier, H.A. 1994. Descriptive flora of Puerto
important fruits in the tropics and is exported to Rico and adjacent Islands. Vol. 3. Editorial de la
temperate areas. It is consumed fresh and made Universidad de Puerto Rico. Río Piedras, PR.
into juice, jams and jellies, and paste or “cheese.” 461 p.
598
Little, E.L., Jr. and F.L. Wadsworth. 1964. Rain-tree. 2002. Guava. Raintree Nutrition, Inc.,
Common trees of Puerto Rico and the Virgin Austin, Texas. http://www.rain-tree.com/guava.
Islands. Agriculture Handbook 249. U.S. htm. 5 p.
548 p. Samarao, S.S. and M.A. Martins. 1999. Influence
of arbuscular mycorrhizal fungi, associated with
Long, R.W. and O. Lakela. 1976. A flora of addition of rutin, on the growth of guava
Tropical Florida. Banyon Books, Miami, FL. (Psidium guajava L.). Revista Brasileira de
962 p. Fruticultura 21(2): 196-199.
Morton, J. 1987. Guava. In: J.F. Morton. Fruits of Samson, J.A. 1986. Tropical fruits. 2nd ed.
warm climates. Julia F. Morton, Maimi, FL. Longman Scientific & Technical, Harlow, UK.
p. 356-363. 336 p.
Mune, T.L. and J.W. Parham. 1956. Weed control: Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
Guava and its control in Fiji. Agricultural Montiel, eds. 2001. Flora de Nicaragua.
Journal, Fiji 27(3/4): 103-108. Monographs in Systematic Botany Vol. 85, No.
Pacific Island Ecosystems at Risk. 2002. Psidium p. 945-1,910.
guajava L., Myrtaceae. http://hear.org/pier_v3.3/
psgua.htm. 4 p. Tona, L., K. Kambu, K. Mesia, K. Cimanga, S.
Apers, T. de Bruyne, L. Pieters, J. Totte, A.J.
Pennington, T.D. and J. Sarukhan. 1968. Arboles Vlietinck, and T. de Bruyne. 1999. Biological
tropicales de México. Instituto Nacional de screening of traditional preparations from some
Investigación Forestales, Secretaría de medicinal plants used as antidiarrhoeal in
Agricultura y Ganadaría. Ciudad de México, Kinshasa, Congo. Phytomedicine 6(1); 59-66.
México. 413 p.
von Carlowitz, P.G. 1991. Multipurpose trees and
Popenoe, W. 1948. Manual of tropical and shrubs: Sources of seeds and inoculants.
subtropical fruits. Hafner Press, New York. International Council for Research in
474 p. Agroforestry, Nairobi, Kenya. 328 p.
599
Psorothamnus scoparius (Gray) Rydb. broom dalea
FABACEAE
Synonyms: Dalea scoparia Gray

Parosela scoparia (A. Gray) Heller
James E. Nellessen
Ecology.—This species is common in Plains-Mesa

Sand Scrub habitats and can be one of the
dominant shrubs in this vegetation type. It is a
deep-sand tolerant or deep-sand adapted shrub. It
may occur on gypsum sands as well as quartz
sands. More specifically it occurs in a mixed shrub
series in association with Artemisia filifolia Torr.,
Atriplex canescens (Pursh) Nutt., Oryzopsis
hymenoides (Roem. & Schult.) Ricker ex Piper,
Sporobolus spp., and other mixed grasses and
forbs (Dick-Peddie 1993).
Reproduction.—Broom dalea often appears

relatively dormant, with little new growth, until
summer monsoon rains begin, whereupon it
rapidly enters the blooming stage. The dark blue
flowers are grouped in semi-spherical spikes, of a
few to several flowers, the clusters 1 to 2 cm in
diameter (Carter 1997, Martin and Hutchins 1980-
General Description.—Broom dalea is also 81). The fragrant blooms attract numerous
known as broom indigobush and purple sage pollinating bees. Fruits are short, egg shaped pods
(Dick-Peddie 1993, Ivey 1995). This is a highly with hairs and glands. Under proper moisture
branched shrub with slender twigs and small conditions seeds may germinate the following
leaves. Its overall growth form gives it a broom- year. New plants may reach reproductive status in
like appearance. It may reach heights of 1.5 m or as little as 3 to 6 years depending on local
more, but is typically about 1 m when mature. environmental conditions.
Horizontal spread is likewise about 1 m but in
older plants may be as much as 2 m or more. Pollination.—Numerous species of bees are
Leaves are alternate, linear to linear-spatulate, attracted to the blue flowers and collect pollen
from 1 to 2 mm in width and 0.5 to 1.9 cm in from broom dalea. The plant emits a noticeable
length, sometimes trifoliate, and gland-dotted sweet, aromatic scent when in bloom. During the
(Carter 1997, Martin and Hutchins 1980-81). peak of summer blooming, these shrubs are often
Populations in New Mexico are actually much focal points for bee foraging activity. Some bees
more frequently trifoliate than some floras would visiting broom dalea include crepuscular (twilight
indicate. Flowers are pealike, dark blue, grouped active) bees of the family Colletidae (yellow-faced
in semi-spherical clusters, and fragrant. and plasterer bees), subfamily Diphaglossinae,
species Caupolicana ocellata Michener; and
Range.—Broom dalea occurs from far western family Anthophoridae (cuckoo, digger, and
Texas through central and southern New Mexico carpenter bees), subfamily Anthophorinae, Tribe
into eastern Arizona and northern Mexico Eucerini, species Martinapis luteicornis
(Kearney and others 1951, Martin and Hutchins (Cockerell) (Rozen and Rozen 1986). These bee
1980-81, Warnock 1974). It occurs at elevations species were observed in southeastern Arizona.
from about 915 m to 1,830 m in upland desert Both groups of bees nest in the ground, in crevices
scrub habitat, usually in sandy soils. in rocks or walls, in plant stalks or other cavities.
Both groups use the pollen, or pollen mixed with
nectar as a food source.
600
Growth and Management.—This species grows Dick-Peddie, W.A. 1993. New Mexico
relatively quickly and can reach 1 m in height or Vegetation: Past, Present and Future. University
diameter within 4 to 7 years depending on local of New Mexico Press, Albuquerque, NM. 244 p.
environmental conditions. The sandy Plains-Mesa
Sand Scrub habitats in which this shrub may be a Ivey, R.D. 1995. Flowering Plants of New Mexico,
dominant are by nature poor rangelands for 3rd Ed. Published by the author, NM. 504 p.
livestock.
Kearney, T.H., R. Peebles, and Collaborators.
Benefits.—Broom dalea is an important natural 1951 with 1960 supplement. Arizona Flora.
stabilizer of dune and other sandy soils. Its University of California Press, Berkeley, CA.
relatively rapid growth rate allows for reasonably 1,085 p.
quick establishment in situations of shifting and
disturbed soils. Like many members of the legume Lucero, M.E., R.E. Estell, and E.L. Frederickson.
family, it is a nitrogen fixer and forms rhizobial 2002. The essential oil composition of
root nodules (Allen and Allen 1981). In fact, this Psorothamnus scoparius (A. Gray) Rydb.
species has been documented to transmit the United States Department of Agriculture,
rhizobial bacteria to another genus within the Agricultural Research Service, Jornada
legume family, Crotalaria (Wilson 1939). Broom Experimental Range http://www.nal.
dalea is not an important range or browse plant for usda.gov/ttic/tektran. In press, Journal of
livestock and may increase under grazing. It is not Essential Oils.
substantially browsed, likely due to oils contained
within the glands (Warnock 1974). Very little, Martin, W.C. and C.E. Hutchins. 1980-1981
until recently, has been known about the volatile (reprinted 2001). A Flora of New Mexico. Vol.
oil composition of this species. Sixty-four volatile 1. Bishen Singh Mahendra Pal Singh, India and
compounds have been identified (Lucero and Koeltz Scientific Books, Germany. p. 1-1,276.
others 2002). The three most abundant compounds
were gamma-terpinene (22.3 percent), p-cymene Rozen, Jr., J.G. and Rozen B.L. 1986. Bionomics
(14.0 percent), and alpha-pinene (9.0 percent). of crepuscular bees associated with the plant
Some of these are known deterrents to herbivore Psorothamnus scoparius: Hymenoptera,
browsing in other plant species and may serve the Apoidea. Journal of the New York
same function in broom dalea. Entomological Society. 94(4): 472-479.
References Warnock, B.H. 1974. Wildflowers of the

Guadalupe Mountains and the Sand Dune
Allen, O.N. and E.K. Allen. 1981. The Country, Texas. Sul Ross State University,
Leguminosae: A Source Book of Alpine, TX. 176 p.
Characteristics, Uses, and Nodulation.
University of Wisconsin Press, WI. 812 p. Wilson, J.K. 1939. Symbiotic promiscuity of two
species of Crotalaria. Journal of the American
Carter, J.L. 1997. Trees and Shrubs of New Society of Agronomy 31: 934-939.
601
Psychotria brachiata Sw. palo de cachimbo
RUBIACEAE
Synonyms: Psychotria neurotricha DC.

Cephaelis polycephala Schltdl.
Palicourea caerulea (Ruiz & Pav.) Roem. & Schult.
John K. Francis
others 2001).
Range.—Palo de cachimbo is native to the

Greater Antilles, Trinidad, and Guatemala to Peru
(Liogier 1997). The Missouri Botanical Garden
(2002) lists some 73 herbarium specimens from
South America. However, some researchers (Croat
1978, Stevens and others 2001) contend that the
South American range is based on
missidentification of the similar P. caerullea Ruiz
& Pav. Palo de cachimbo is not known to have
been planted or naturalized elsewhere.
Ecology.—Palo de cachimbo grows mostly in wet

forest areas that receive from 2000 to 3000 mm of
mean annual precipitation. It occurs at elevations
from near sea level to 800 m in Nicaragua
(Stevens and others 2001). Palo de cachimbo is
intolerant of shade. Plants compete well with
herbs and other shrubs in openings but do not
survive long after a forest canopy closes over
them. The species is common to uncommon in
General Description.—Palo de cachimbo is an roadsides, secondary forest, and openings in
evergreen shrub occasionally reaching 5 m in primary forest. Plants sampled in secondary forest
height but usually 2 to 3 m in height and 3 to 6 cm in Puerto Rico had the following nutrient levels in
in basal diameter. The shrub usually has several their leaves: 2.36 + 0.03 percent N, 0.12 + 0.00
stems formed by suckers from the lateral roots a percent P, 2.08 + 0.25 percent K, and 12.66 + 0.71
few cm out from the central stem and by branches percent ash. Lower levels of those nutrients were
low on the principal stems. The wood has a reported for twigs and stem (Lugo 1992).
moderately low density, is moderately strong, and
has faint annual rings. Palo de cachimbo plants are Reproduction.—Palo de cachimbo flowers from
supported by relatively shallow lateral root February to August and fruits from July to April in
systems, at least in soils that have poorly aerated Nicaragua (Stevens and others 2001). The species
subsoil. The roots are tan colored and flexible. The is a good producer of fruits and seeds. A collection
branches and twigs are green, slender, and usually of fresh fruits from Puerto Rico weighed an
paired. Shiny, dark-green leaves are ovate to average of 0.1569 + 0.0125 g/fruit. Air-dried seeds
lanceolate, entire and pointed at both ends, 7 to 20 separated from them averaged 0.0048 + 0.0002
cm long and 3 to 8 cm broad, with a 1- to 3-cm g/seed or 208,000 seeds/kg. Placed in commercial
petiole. Small yellow to greenish-white flowers potting mix, 48 percent germinated between 34
are tightly clustered in open terminal panicles. The and 57 days following sowing. Seeds are
4-mm ellipsoidal berries are tightly clustered. presumably dispersed by birds. Seedlings are
They are dark purple or dark blue at maturity and common in the vicinity of fruit-producing plants.
usually contain two hemispherical seeds having
the inside face concave and the outside face with Growth and Management.—Palo de cachimbo
five ridges (Croat 1978, Liogier 1997, Stevens and grows slowly in the early seedling stage. It grows
602
0.5 to 0.8 m/year for 2 or 3 years from sprouts. Liogier, H.A. 1997. Descriptive flora of Puerto
Individual stems probably live from 10 to 20 Rico and adjacent islands. Vol. 5. Editorial de la
years, but plants may persist longer by sprouting. Universidad de Puerto Rico, Río Piedras, PR.
Natural regeneration probably can be encouraged 436 p.
by disturbance designed to create forest openings.
Planting and management experience have not Lugo, A.E. 1992. Comparison of tropical tree
been published. plantations with secondary forests of similar
age. Ecological Monographs 62: 1-41.
Benefits.—Palo de cachimbo contributes to the
biodiversity of forests, helps protect the soil, and Missouri Botancial Garden. 2002. W3Tropicos:
furnishes food and cover for wildlife. nomenclature data base. http://mobot.mobot.
org/cgi-bin/search_vast#samer. [not paged].
References
Croat, T.B. 1978. Flora of Barro Colorado Island. Montiel, eds. 2001. Flora de Nicaragua.
Stanford University Press, Stanford, CA. 943 p. Monographs in Systematic Botany Vol. 85, No.
p. 1,911-2,666.
603
Psychotria deflexa DC. garricillo
RUBIACEAE
Synonyms: Psychotria patens of authors, not of Sw.

Psychotria flexulosa of authors, not of Willd.
Palicourea patens (Sw.) Urban
John K. Francis
surface (Croat 1978, Liogier 1997, Stevens and

other 2001).
Range.—Garricillo is native to the Greater

Antilles, Trinidad, southern Mexico, Central
America, and South America to Bolivia (Croat
1978, Liogier 1997, Stevens and others 2001). It is
not known to have naturalized outside the native
range.
Ecology.—Garricillo is a shrub of moist and wet

forests, areas receiving from about 1600 mm to
over 3000 mm of mean annual precipitation.
Although the species may occur from a few meters
above sea level to over 1,000 m in elevation
(Instituto Nacional de Biodiversidad 2002), it
generally grows at medium to high elevations
(Liogier 1997) because there is greater available
moisture. The species grows on poorly drained to
well drained soils generally with loamy to clayey
texture and pH’s from about 4.5 to 6.5. Garricillo
is intermediate in tolerance to shade and grows in
the forest understory (if not too dark), at forest
edges, in overgrown fencerows (Molano and
others 2002), and in openings. Most fruit and seed
production takes place in openings or situations
with increased to full sunlight. The species
General Description.—Garricillo (a Panamanian competes well with herbs and low brush.
name), sometimes referred to by the general name
“cachimbo,” is an evergreen shrub usually 1.5 to Reproduction.—Garricillo flowers from May to
2.0 m and 1.0 to 2.0 cm in basal diameter, July and matures fruits mostly from August to
sometimes reaching 3 m in height. Multiple stems December in Panama (Croat 1978). The flowers
are common, but not thick clumps. The wood is are both insect-pollinated and self-pollinated
moderately hard with faint annual rings. The (Faivre 2002). Fresh fruits collected in Puerto
angular twigs and the foliage are glabrous. Rico weighed an average of 0.0758 + 0.0006
Persistent, bilobed, long-pointed stipules are g/fruit. Air-dried seeds separated from them
characteristic of the species. Paired, dark-green, weighed an average of 0.0057 + 0.0001 g/seed, or
elliptic to oblong lanceolate leaves are 5 to 18 cm 175,000 seeds/kg. There was an average of 1.7
long with 3- to 10-mm petioles. Inflorescences are seeds/fruit in this sample. Sown without
few-flowered terminal panicles. The small, sessile pretreatment in moist potting mix, 69 percent
flowers are white. Fruits are subglobose, 3 to 5 germinated between 68 and 117 days after sowing.
mm in diameter, white or pale blue. Normally, Fruit and seed production is good and consistent.
there are two hemispherical seeds per fruit that are Birds disperse the seeds (Molano and others
striated on the convex surface and have a medial 2002). Seedlings are common but suffer high
groove on the opposite, flat or slightly concave mortality. Damaged plants resprout. Prostrate
604
stems of young plants layer (root) readily when in Allentown, PA. http://botany2002.org.section3/
contact with the soil. abstracts/21.shtml. 1 p.
Growth and Management.—Seedlings of Instituto Nacional de Biodiversidad. 2002. Lista de

garricillo grow slowly at first. Older plants and especímenes de Psychotria deflexa. Costa Rica.
sprouts add from 20 to 40 cm to their height each http://www.inbio.ac.cr/bims/k03/p13/c045/o014
year. Individual stems live about 10 years; plants 2/f01359/g008913/s028252.htm. 4 p.
may last longer by producing new sprouts.
Although the species is more or less common, it Liogier, H.A. 1997. Descriptive flora of Puerto
does not appear to cause any difficulties for forest Rico and adjacent islands. Vol. 5. Editorial de la
users. Universidad de Puerto Rico, Río Piedras, PR.
436 p.
Benefits.—Garricillo contributes to the
biodiversity and biomass of forest communities, Molano, J.G., M.P. Quiceno, and C. Roa. 2002. El
helps protect the soil, and furnishes food and papel de las cercas vivas en un sistema de
cover for wildlife. producción agropecuaria en el Piédemonte
Llanero. http://www.lead.virtualcentre.org/
References es/ele/conferencia2/vbconfe3.htm. 18 p.
Croat, T.B. 1978. Flora of Barro Colorado Island.

Stanford University Press, Stanford, CA. 943 p.
Monographs in Systematic Botany Vol.85, No.
Faivre, A.E. 2002. Reproductive biology of seven
understory Psychotria species on Barro
MO. p. 1,911-2,666.
Colorado Island, Panama. Cedar Crest College,
605
Psychotria microdon (DC.) Urban thicket wild coffee
RUBIACEAE
Synonyms: Psychotria pinularis Sessé & Moc.

Rondeletia microdon DC.
John K. Francis
others 2001). It is not known to have been planted

or naturalized elsewhere.
Ecology.—Thicket wild coffee is occasional to
relatively common in secondary and remnant
forests from near sea level to 840 m (Stevens and
others 2001). Mean annual precipitation for the
species in Puerto Rico ranges from 750 to about
1700 mm. It grows on a variety of well-drained
soils that developed from alluvium, and igneous
and sedimentary (including limestone) rocks.
Thicket wild coffee is moderately intolerant to
intermediate in tolerance to shade. It will grow and
fruit in openings and low basal-area stands and
survives in forest stands with moderate basal areas.
The species endures light grazing but disappears
under heavy grazing pressure.
General Description.—Thicket wild coffee, also
Reproduction.—In Nicaragua, thicket wild coffee
known as café bâtard, and café marron, is a
flowers May through September and fruits August
semideciduous arching or scrambling shrub 1 to 3
through September (Stevens and others 2001).
m in height with a basal diameter of up to 4 cm.
Fruit and seed production is moderate. A
There is usually a single stem from the ground in
collection of fresh fruits from Puerto Rico weighed
undisturbed plants with multiple branches low on
an average of 0.259 + 0.005 g/fruit. Air-dried
the stem. The arching stems sometimes extend 2 or
seeds separated from them averaged 0.0187 +
3 m horizontally and tend to be very branchy. The
0.0004 g/seed or 53,000 seeds/kg. Sown without
wood is of medium density, moderately weak and
pretreatment on moist blotter paper, 78 percent
brittle. Stem and branch bark is smooth and tan
outside and green within. The plant is supported
Germination is epigeal. The seeds are apparently
by a tap-and-lateral root system of dark tan and
dispersed primarily by birds. Tapirs eat the fruits,
flexible roots. The foliar display is medium to thin.
but the seeds are destroyed in the process (Olmos
Membranous, glabrous, obovate to oblanceolate,
1997). Seedlings are common near seed sources in
light-green leaves are 4 to 11 cm long with 10- to
Puerto Rico. Survival to adulthood is relatively
18-mm petioles. The inflorescences are terminal
rare, however.
corymbiform cymes with three to five rays. The
flowers have a five-toothed white corolla tube 5 to
Growth and Management.—Thicket wild coffee
10 mm long. Fruits are red or red-orange
is slow growing as a new seedling and grows at a
ellipsoidal berries about 7 mm long. They usually
moderate rate as a sapling and adult. Plants survive
contain two tan seeds that are flattened on one side
at least 5 years and perhaps much longer. Planting
and rounded on the other (Howard 1989, Liogier
and management experience has not been
1997, Stevens and others 2001).
published. The species is not common enough or
aggressive enough to warrant control.
Range.—Thicket wild coffee is native to the
Greater Antilles (except Jamaica), the Lesser
Benefits.—Thicket wild coffee contributes to the
Antilles, from Central Mexico south through
biodiversity of forests, helps protect the soil, and
Central America to Bolivia and eastwards to
furnishes food and cover for wildlife. The fruits
Guiana (Howard 1989, Liogier 1997, Stevens and
are slightly sweet and edible but not highly
606
desirable. The species is a larval host for the Olmos, F. 1997. Tapirs as seed dispersers and
Xylophanes pluto (Fabricius) moth in Puerto Rico predators. IUCN/SSC Tapir Specialist Group.
(Torres-Bauzá 2000). http://www.tapirback.com/tapirgal/iucn-ssc/tsg/
action97/ap97-05.htm. 12 p.
References
Howard, R.A. 1989. Flora of the Lesser Antilles, Montiel, eds. 2001. Flora de Nicaragua,
Leeward and Windward Islands. Angiospermas. Monographs in Systematic
Dicotyledoneae. Part 3. Vol. 6. Arnold Botany Vol. 85, No. 3. Missouri Botanical
Arboretum, Harvard University, Jamaica Plain, Garden Press, St. Louis, MO. p. 1,911-2,666.
MA. 658 p.
Torres-Bauzá, J.A. 2000. Ciclo de vida y aspectos
Liogier, H.A. 1997. Descriptive flora of Puerto de la biología de Xylophanes pluto (Fabricius)
Rico and adjacent islands. Vol. 5. Editorial de la en Puerto Rico (Lepidoptera: Sphingidae).
Universidad de Puerto Rico, Río Piedras, PR. Caribbean Journal of Science 36(3-4): 227-232.
436 p.
607
Psychotria nervosa Sw. shiny-leafed wild coffee
RUBIACEAE
Synonyms: Psychotria undata Jacq.

Psychotria chimarrhoides DC.
Psychotria lanceolata Nutt.
Psychotria hirsuta Spreng.
Psychotria oligotricha DC.
Psychotria portoricensis DC.
John K. Francis
lanceolate, with sunken veins and elongated tips.
A large, flattened bud is almost always visible
between the youngest pair of leaves. The flower
clusters, which are terminal or axillary, are nearly
globose and contain a number of small, white
flowers. The clusters become more diffuse as the
fruits mature. The fruits (drupes) are bright red,
elliptic or globose at maturity and 6 to 7 mm long
(Howard 1989, Liogier 1997, Nelson 1996,
Stevens and other 2001). They are fleshy and juicy
with little flavor. Each contains two seeds that
have a rounded side with shallow striations and a
flattened side with a division down the middle like
a coffee bean (author’s observation).
Range.—Shiny-leafed wild coffee is native to

Florida, the West Indies, Southern Mexico,
Central America, and Colombia, Ecuador, and
Venezuela in South America (Liogier 1997,
Ecology.—Shiny-leafed wild coffee grows in dry

and moist forests that receive from about 700 to
2000 mm of mean annual precipitation. At the
moist end of the precipitation range, it tends to be
found only on excessively drained sites. It is cold-
sensitive, tolerating temperatures only to about
freezing (Dolan 2002). It grows well above the
General Description.—Shiny-leafed wild coffee,
frost line in Florida; presumably shiny-leafed wild
also known as wild coffee, Bahaman wild coffee,
coffee freezes to the ground and resprouts later.
wood balsam, Seminole balsamo, café marrón,
Well-drained soils with the range of textures and
palo moro, ti café marron, ti kafé mawon, and
both igneous and sedimentary (especially
remaroogu, is a 1- to 3-m erect shrub. Stem-like
limestone) parent materials are colonized. Wild
low branches are common, and multiple stems are
coffee is moderately tolerant of shade, usually
formed in response to damage to the trunk. The
growing in the understories of low basal area
stems, sometimes up to 7 cm in diameter, are
forests. The shrub may also be found in natural
covered with a bone-colored, smooth bark that is
and artificial openings. In Florida, it is usually
hairy in some variants. The plant is supported by
found in shell ridges, hammocks, and pinelands
stiff tap and lateral roots and abundant fine roots.
(School of Forest Resources and Conservation
Relatively few branches and twigs are formed
2002). Shiny-leafed wild coffee is occasionally
with foliage maintained only near the tips. The
found in pure clumps, but more often it grows as
leaves are shiny light or dark green, opposite,
well-distributed plants.
simple, glabrous or nearly so, elliptic to
608
Reproduction.—Shiny-leafed wild coffee blooms References
in spring and summer in Florida (Nelson 1996)
and irregularly through the year in Nicaragua Dave’s Garden Network. 2002. Wild coffee
(Stevens and others 2001). The flowers are visited (Psychotria nervosa). The Plants Database.
by honeybees and butterflies (Dave’s Garden http://plantsdatabase.com/go/2326.html. 2 p.
Network 2002) that presumably pollinate them.
A collection of fruits from Puerto Rico weighed an Dolan, S. 2002. Native shrubs.
average of 0.256 + 0.009 g/fruit. Air-dried seeds http://www.indialantic.com/contents/ecology/
from those fruits weighed an average of 0.0175 + shurbs.htm. 3 p.
0.0004 g/seed or 57,000 seeds/kg. About 85
percent of fruits from the collection contained two Howard, R.A. 1989. Flora of the Lesser Antilles,
seeds; the reminder contained one filled seed and Leeward and Windward Islands.
one aborted seed. Sown in commercial potting Dicotyledoneae. Part 3. Vol. 6. Arnold
mix, 95 percent of the filled seeds germinated Arboretum, Harvard University, Jamaica Plain,
between 64 and 164 days after sowing (author’s MA. 658 p.
observation).
Growth and Management.—Shiny-leafed wild Rico and adjacent islands. Vol. 5. Editorial de la
coffee has a moderate growth rate. Small wildlings Universidad de Puerto Rico, Río Piedras, PR.
and pruned larger plants transplant well. They also 436 p.
can be grown easily from seed. Once established
in a garden or natural area, they will reproduce Nelson, G. 1996. The shrubs and woody vines of
and eventually fill the area (Workman 1980). Florida. Pineapple Press, Inc., Sarasota, FL.
Where a seed source is available, probably the 391 p.
best strategy to promoting natural establishment is
to manipulate the stand for an open understory. Olmos, F. 1997. Tapirs as seed dispersers and
predators. IUCN/SSC Tapir Specialist Group.
Benefits.—Shiny-leafed wild coffee is a http://www.tapirback.com/tapirgal/iucn-ssc/tsg/
widespread and important understory plant in dry action97/ap97-05.htm. 12 p.
and moist forest and as such contributes to the
biodiversity and biomass accumulation, and School of Forest Resources and Conservation.
protects against soil erosion. It is frequently 2002. Florida forest plants: wild coffee
planted in Florida as a native landscape plant in (Psychotria nervosa). University of Florida,
both formal and natural gardens (Workman 1980). Gainesville, FL. http://www.plantatlas.usf.edu/
It is appreciated for its attractive foliage and bright maps.asp?plantID=2811. 2 p.
red berries and because it attracts birds and
butterflies (Dave’s Garden Network 2002). The Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
fruits are food for tapirs (Olmos 1997), a number Montiel, eds. 2001. Flora de Nicaragua,
of bird species (School of Natural Resources and Angiospermas. Monographs in Systematic
Conservation 2002), and probably many other Botany Vol. 85, No. 3. Missouri Botanical
animals. Shiny-leafed wild coffee contains the Garden Press, St. Louis, MO. p. 1,911-2,666.
alkaloid dimethyltryptamine, which is
hallucinogenic (Ted 2002) and may eventually Ted, M. 2002. Smokable dimethyltryptamine from
prove to have medicinal applications. The seeds organic sources. http://www.deoxy.org/
were once used as a coffee substitute (Dave’s smokedmt.htm. 11 p.
Garden Network 2002).
Sanibel, FL. 137 p.
609
Purshia tridentata (Pursh) DC. antelope bitterbrush
ROSACEAE
Synonyms: Kunzia tridentata Spreng.

Tigarea tridentata Pursh
Nancy L. Shaw and Stephen B. Monsen

eastward into western Montana, the Black Hills of
South Dakota, and northwestern New Mexico
(Cronquist and others 1997, Welsh and others
1987).
Ecology.—Antelope bitterbrush grows on well-

drained, slightly basic to slightly acidic soils at
elevations from 60 to 3,510 m (Nord 1965). It
often grows as the dominant shrub with bluebunch
wheatgrass and a variety of forbs. It is also a
common associated species in many big sagebrush
(Artemisia tridentata L.), mountain brush, pinyon–
juniper (Pinyon L.-Juniperus L.), ponderosa pine
(Pinus ponderosa P. & C. Lawson), and
occasionally lodgepole pine (Pinus contorta
Dougl. ex Loud.) communities (McArthur and
others 1983, Nord 1965). It is a pioneering species
on steep, rocky, unstable disturbances. Erect forms
Illustration source: USDA Forest Service 1937 generally do not survive wildfires, but layering
forms may resprout. Antelope bitterbrush is
General Description.—Antelope bitterbrush, also sometimes associated with the nitrogen-fixing
known as antelope brush, antelope buckbrush, actinomycete Frankia, but root nodulation varies
deerbrush, or quininebrush, is an intricately with factors such as moisture availability and soil
branched, deciduous shrub varying widely in chemistry (Righetti and others 1983). Emerging
growth habit from low, decumbent, spreading seedlings are susceptible to a number of damping-
forms to upright arborescent plants over 4 m in off organisms, while seedlings and mature plants
height (Blauer and others 1975, Welsh and others may be damaged by grasshoppers, Great Basin tent
1987). Leaves are alternate, simple, pinnatifid or caterpillars (Malacosoma fragile Stetch), and other
apically three-toothed and sometimes glandular. insects (Nord 1965, Shaw and Monsen 1983).
Flowers are insect-pollinated and borne on short
spurs of previous year’s growth. They are Reproduction.—Plants flower in April to June,
numerous and white to yellow with petals 5 to 9 and fruits ripen in late June to August depending
mm long (Welsh and others 1987). The fruit is a upon elevation and latitude (Nord 1965). Seed
cartilaginous achene with a persistent tapering production is related to precipitation and leader
style and a dark pinkish gray pyriform seed growth in the previous year, but yields are often
(Blauer and others 1975). Purshia DC. ex Poir. drastically reduced by insect predators (Shaw and
likely derived from Cowania (D. Don) (Jabbes Monsen 1983). Seeds dehisce rapidly at maturity.
2000, McArthur and others 1983) and hybrids High percentages of the achenes are cached by
between Cowania and Purshia are common. rodents, and most successfully establishing natural
Purshia glandulosa Curran is considered a recruitment is from these caches (Evans and others
stabilized hybrid between Cowania and Purshia 1983, Nord 1965, Van der Wall 1994). Large
tridentata. quantities of seed are hand harvested from
wildland stands each year. Dry seed may be stored
Range.—Antelope bitterbrush is widely in a warehouse for up to 15 years (Stevens and
distributed from British Columbia southward on others 1981). A 28-day moist prechill is required
the east side of the Cascade and Sierra Nevada to release seed dormancy (AOSA 1993, Booth
Mountains to California and northern Arizona and
610
1999, Meyer 1989). Site-adapted seed sources 1993. Rules for testing seeds. Journal of Seed
should be selected for plantings. Antelope Technology 16: 1–113.
bitterbrush should be drill seeded in late fall or
winter to provide overwinter prechilling. It may be Blauer, A.C., A.P. Plummer, E.D. McArthur, R.
seeded with other native species that are not highly Stevens, and B.C. Giunta. 1975. Characteristics
competitive. Seeding failures resulting from rodent and hybridization of important Intermountain
predation of seed can be avoided by seeding in late shrubs. I. Rose family. U.S. Department of
fall on fairly large disturbances that do not provide Agriculture, Forest Service, Intermountain
cover for the rodents (Evans and others 1983). Forest and Range Research Station, Ogden, UT.
Containerized or 1-0 bareroot seedlings are easily 35 p.
grown, but fungicide treatments may be required
to reduce losses to damping-off organisms. Booth, D.T. 1999. Imbibition temperatures affect
Nursery stock can be planted on steep, rocky bitterbrush seed dormancy and seedling vigor.
slopes that are inaccessible to seeding equipment. Journal of Arid Environments 48: 35-39.
It establishes best if planted in early spring when
antelope bitterbrush in the surrounding area is yet Cronquist, A., N.H. Holmgren, and P.K.
dormant. Herbaceous vegetation must be cleared Holmgren. 1997. Intermountain flora: vascular
around each seedling to reduce competition. plants of the Intermountain West, U.S.A. Vol. 3,
part 1. Subclass Rosidae: (except Fabales). The
Growth and Management.—Antelope New York Botanical Garden, The Bronx, NY.
bitterbrush seedings, plantations, and wild stands 446 p.
are vulnerable to heavy use by wild or domestic
ungulates and predation by gophers, mice, birds, Evans, R.A., J.A. Young, G.J. Cluff, and J.K.
and insects. Seedlings are not competitive with McAdoo. 1983. Dynamics of antelope
weedy annuals or most introduced perennial bitterbrush seed caches. In: A.R. Tiedemann and
grasses. Many mature stands are damaged by K.L. Johnson, comps. Proceedings: Research
continuous and close browsing by deer and and management of bitterbrush and cliffrose in
livestock. western North America. GTR-INT-152. U.S.
Benefits.—Next to sagebrush, antelope bitterbrush Intermountain Forest and Range Experiment
is probably one of the most widespread shrubs in Station, Ogden, UT. p. 195–202.
the Great Basin region, although its prevalence has
been reduced by range fires and exotic plant Jabbes, M. 2000. Hybridization and its
introductions (Mozingo 1987). It is important for a evolutionary consequences in Purshia and
host of wildlife and provides nutritious browse and Cowania. Ph.D. dissertation. University of
essential cover on fall, winter, and summer mule Idaho, Moscow, ID. 185 p.
deer ranges (Plummer and others 1968). Protein
content varies among populations depending upon McArthur, E.D., H.C. Stutz, and S.C. Sanderson.
over-winter leaf retention. New growth may also 1983. Taxonomy, distribution, and cytogenetics
be heavily used in spring and summer. Because of of Purshia, Cowania, and Fallugia (Rosoideae,
the varied growth forms, attractive foliage, and Rosaceae). In: A.R. Tiedemann and K.L.
showy flowers, antelope bitterbrush has potential Johnson, comps. Proceedings, Research and
as an ornamental in low-maintenance landscapes. management of bitterbrush and cliffrose in
Populations of antelope bitterbrush with distinctive western North America. GTR-INT-152. U.S.
attributes have been recognized and are Department of Agriculture, Intermountain Forest
commercially harvested and sold. Native and Range Experiment Station. p. 4–24.
Americans reportedly used bitterbrush as a remedy
for smallpox, measles, tuberculosis, and Meyer, S.E. 1989. Warm pretreatment effects on
pneumonia, and as an antiseptic for rashes and antelope bitterbrush (Purshia tridentata)
insect bites (Mozingo 1987). germination response to chilling. Northwest
Science 63: 146–153.
References
Mozingo, H.N. 1987. Shrubs of the Great Basin: a
AOSA [Association of Official Seed Analysts]. natural history. University of Nevada Press,
Reno, NV. 342 p.
611
Nord, E.C. 1965. Autecology of bitterbrush in Research and management of bitterbrush and
California. Ecological Monographs 35: 307– cliffrose in western North America. GTR-INT-
334. 152. U.S. Department of Agriculture, Forest
Plummer, A.P., D.R. Christensen, and S.B. Experiment Station, Ogden, UT. p. 55–69.
Monsen. 1968. Restoring big game range in
Utah. Publication 68-3. Utah Division of Fish Stevens, R., K.R. Jorgensen, and J.N. Davis. 1981.
and Game, Salt Lake City, UT. 183 p. Viability of seed from thirty-two shrub and forb
species through fifteen years of warehouse
Righetti, T.L., C.H. Chard, and D.N. Munns. 1983. storage. Great Basin Naturalist 41: 274–277.
Opportunities and approaches for enhancing
nitrogen fixation in Purshia, Cowania and USDA Forest Service. 1937. Range plant
Fallugia. In: A.R. Tiedemann and K.L. Johnson, handbook. U.S. Government Printing
comps. Proceedings: Research and management
of bitterbrush and cliffrose in western North
Office, Washington, D.C. 512 p.
America. GTR-INT-152. U.S. Department of
Agriculture, Forest Service, Intermountain Van der Wall, S.B. 1994. Seed fate pathways of
Forest and Range Experiment Station, Ogden, antelope bitterbrush: dispersal by seed-caching
UT. p. 214–224. yellow pine chipmunks. Ecology 75: 1911-1926.
Shaw, N.L. and S.B. Monsen. 1983. Phenology Welsh, S.L., N.D. Atwood, L.C. Higgins, and S.
and growth habits of nine antelope bitterbrush, Goodrich. 1987. A Utah flora. Great Basin
desert bitterbrush, Stansbury cliffrose, and Nturalist Memoirs 9. Brigham Young University
Apache plume accessions. In: A.R. Tiedemann Press. 894 p.
and K.L. Johnson, comps. Proceedings:
612
Quercus havardii Rydb. Havard shin oak
FAGACEAE
Synonyms: None
James E. Nellessen
Hutchins 1980). Havard shin oak is a member of

the white oak group, (i.e. it does not bear bristle
tips on the leaves and the fruits mature in the first
year). There are two varieties: Q. havardii var.
havardii and Q. havardii var. tuckeri Welsh
(Kartesz 1994).
Range.—This species occurs from western

Oklahoma, and western Texas, into the
southeastern portion of New Mexico, with
outlying populations elsewhere in these states and
as far west as Utah and Arizona (Great Plains
Flora Association 1986, Peterson and Boyd 1998).
Havard shin oak has recently been documented
from Kansas for the first time (Freeman 2000).
General Description.—Havard shin oak is also The elevational range of habitats in which shin oak
may occur is from 500 m to 1,525 m. Variety
known as just Havard oak, shin oak or shinnery
tuckeri occurs in the Four Corners area (Utah and
oak, but the names sand shinnery oak, sand scrub
oak, panhandle oak, and midget oak have also Arizona populations); these populations appear to
be hybrids between shin oak and Q. turbinella
been applied to it (McCoy 1981). Peterson and
Greene (scrub live oak) and Q. gambelii Nutt.
Boyd (1998) supply a fairly comprehensive review
of this species. Havard shin oak grows to 1 m tall (Gambel oak) (Flora of North America 1997).
at maturity, but hybrids with other white oak
species may attain greater height. It forms dense Community Ecology.—Havard shin oak
historically covered an estimated 5 to 7 million
thickets because of an extensive underground
acres within the southern Great Plains (Peterson
system of horizontal rhizomes. The rhizomes are
usually 3 to 10 cm in diameter but may be as thick and Boyd 1998). Of this, 1 million acres occurred
in Oklahoma, 1.5 million acres in New Mexico,
as 15 cm. Most of the rhizomes and roots occur in
and 3.5 million acres in Texas. About 30 percent
the upper 60 cm of soil, but the plant is known to
root to depths of 9 m. The bark is gray becoming of the acreage in Texas has been converted to
cropland and grassland, and about 10 percent has
papery scaly to rough with age. The twigs are
been converted in Oklahoma and New Mexico.
brown to grayish, smooth to densely hairy with
short yellowish or grayish hairs. The buds are dark Shin oak occurs on deep sandy soils and plains,
including sand dunes, and is the major shrub
red-brown, semi-globose, and lightly hairy. The
species comprising Plains-Mesa Sand Scrub
leaves are simple, alternate, leathery, somewhat
variable in shape, oblong, ovate, or lanceolate in vegetation in southeastern New Mexico and
similar sandy soil habitats in western Texas
outline, with rounded shallow lobes, although
including portions of the Llano Estacado ("staked"
sometimes they may be more coarsely toothed
than lobed (the teeth are not bristle-tipped), plains) (Dick-Peddie 1993). The shin oak or
shinnery vegetation series variously places Havard
lustrous green and often smooth above, sometimes
shin oak in dominance with Schizachyrium
with hairy patches, densely yellowish to brownish
green felty-hairy below, the veins prominent. The scoparium (Michx.) Nash (little bluestem),
Artemisia filifolia Torr. (threadleaf sagebrush),
leaves range in length from 1.5 to 10 cm, and in
width from 1.2 to 5.5 cm, with occasionally curled Andropogon hallii Hack. (sand bluestem), Aristida
purpurea Nutt. (purple three-awn), Bothriochloa
or wavy margins (Carter 1997, Correll and
saccharoides (Sw.) Rydb. (silver beardgrass),
Johnston 1970, Flora of North America 1997,
Great Plains Flora Association 1986, Martin and Yucca campestris McKelvey (plains yucca),
613
Atriplex canescens (Pursh) Nutt. (four-wing considerable amounts of water during wet weather
saltbush), Sporobolus airoides (Torr.) Torr. (alkali with up to 50 percent of their mass being water.
sacaton), and Eriogonum annuum Nutt. (annual This gives the plant a great ability to survive
buckwheat). But there are a wide variety of other drought. During spring drought it may not leaf out
shrubs, forbs and grasses comprising numerous or may defoliate and leaf out later once water
other vegetation series within the greater shinnery becomes available. It generates more negative leaf
community. The tall grasses such as sand water potentials, -25 to -32 bars, than many other
bluestem, big bluestem (Andropogon gerardii species in the surrounding community. Soils under
Vitmann), and Indiangrass [Sorghastrum nutans shin oak, in addition to being sandy, are low in
(L.) Nash] have been reduced in abundance nitrogen (0.017 percent), low in organic matter
through overgrazing of Havard shin oak (0.36 percent) neutral to slightly alkaline in pH,
communities. Havard shin oak itself may remain and low in clay and carbonates (a caliche layer is
stable or also decrease with overgrazing, the cover inhibitory). Havard shin oak has been reported to
of grass and Havard shin oak often being have allelopathic effects on other plant species.
positively correlated. Stands of Havard shin oak on
sites with shallow water tables, such as in Chavez Reproduction.—Shin oak flowers in April to
County New Mexico, may contain cottonwoods. May. As with other oaks, male flowers are born in
Hybrids with other oak species, such as Q. stellata pendulous catkins, the female flowers solitary or a
Wang (post oak), occur and may be found within few clustered within leaf axils. The male catkins
typical Havard shin oak stands, the hybrids often are 2 to 3 cm long and densely flowered. The
attaining greater height (2 to 3 m), and stand out as female catkins are 3 to 7 mm long with one to five
readily visible clumps from the surrounding flowers. For a small shrubby oak, the acorns are
shorter plants. These hybrids tend to have quite large, 12 to 25 mm long, up to about 20 mm
intermediate characteristics such as larger leaves wide, brown, ovoid to oblong in shape. They are
and smaller acorns. These hybrids tend to be more born solitary or in pairs and the cup, 10 to 12 mm
common within the eastern portion of Havard shin deep, covers about a third to a half of the fruit. The
oak’s range (Correll and Johnston 1970, Flora of fruits mature from mid-July into September during
North America 1997, Peterson and Boyd 1998). the first year of development on the current
Havard shin oak also hybridizes with Q. mohriana season’s branches (Carter 1997, Correll and
Buckley ex Rydb. (Mohr shin oak) that occurs Johnston 1970, Flora of North America 1997,
within the rocky “breaks” area of the Texas Great Plains Flora Association 1986, Martin and
panhandle. Hybrids are intermediate in Hutchins 1980). Acorns of the white oak group,
characteristics and habitat location (Muller 1952). show little or no dormancy, and will germinate
immediately (Young and Young 1992). Acorn
Autoecology.—The plant primarily spreads by its moisture content for this group cannot drop below
underground rhizomes to form large clonal clumps 30 to 50 percent otherwise seed viability will be
that may range in size from as small as 2 to 15 m lost. Within a particular clone, acorn crops
in diameter to as large as 0.8 ha. Isozyme generally only occur every 3 to 10 years. Within a
electrophoresis identified 37 clones/ha, larger larger landscape scale area containing many
clones being free of interclonal growth, and the different clones, at least one or a few clones will
sample population was outcrossing based on have acorns. Most acorns are attacked by weevils
Hardy-Weinberg expected genotype ratios (Mayes and will decompose quickly. There is little
and others 1998). There are no data on horizontal reproduction by seed because of the short lifespan
growth rates but they are believed to be slow. The of the acorn, the need for good environmental
plants are long-lived, estimated at hundreds to conditions for germination and establishment at
thousands of years (Peterson and Boyd 1998). The the time of acorn drop, attack by weevils, and
rootstocks are massive, comprising the bulk of the decomposers.
plant mass (73 percent); the above-ground shoots
are merely small side shoots to the below ground Growth and Management.—For purposes of
parts. Above-ground biomass has been measured agriculture and livestock, shin oak has generally
at 1,821 kg/ha (Sears and others 1986) and annual been managed as an undesirable species. Havard
production has been estimated to be anywhere shin oak has sometimes been classified as an
from 550 to 4,500 kg/ha depending on the site increaser under livestock grazing, but reports do
(reviewed in Peterson and Boyd 1998). The roots not substantiate this (reviewed in Peterson and
(which are mycorrhizal) and rhizomes absorb Boyd 1998). Its poor reproduction by seed restricts
614
its spread primarily to the slow growing rhizome catkins and buds were higher in livestock grazed
system. An estimated 100,000 acres of land in plots than ungrazed plots. Crude protein and other
New Mexico have been treated with the herbicide nutritional qualities did not differ between
tebuthiuron for brush control. Tebuthiuron treatments (Boyd and others 2001). The acorns are
gradually kills the plant over a 3 to 4 year period. eaten by a variety of wildlife and are a staple food
Other chemicals that have historically been used for lesser prairie chickens (Tympanuchus
on Havard shin oak are 2,4-D, benzoic acids, pallidicinctus Ridgeway) in the fall and winter
2,4,5-T, and picloram. Tebuthiruron should not be seasons. Havard shin oak communities are
used in drought years, and the treated land should important breeding and nesting habitats for the
not be summer grazed for 2 years. Various lesser prairie chicken. Prescribed fire in shin oak
techniques of rotational and continuous grazing in habitats can improve grass and forb cover and
shin oak communities have been tested with mixed associated insect abundance for prairie chickens
results for livestock productivity. But rotational but at least a 2 to 3-year recovery period, no
grazing is best for overall ecosystem management burning in spring, and burning in patches are
including maintenance of proper native grass necessary (Boyd and Bidwell 2001). The rare sand
covers, and management of the lesser prairie dune lizard (Sceloporus arenicolus Degenhardt
chicken and perhaps other wildlife. Goats have and Jones), a threatened species in New Mexico, is
been used to control shin oak with reasonable restricted to Havard shin oak communities. Both
success, although goats eating it continuously will Native Americans and early Spanish colonists
lose weight. In addition to livestock use and direct would eat the acorns raw or baked. A flour can be
agricultural conversion of shin oak habitat, another made from the large acorns by first boiling them,
reason for shin oak management concerns the boll then roasting, chopping into pieces, drying for 30
weevil (Anthonomus grandis Boheman). This minutes, then passing them through a grinder a
cotton pest is known to overwinter in oak leaf litter couple of times (Warnock 1974).
and to reinfest cotton fields for up to a half mile
away (reviewed in Peterson and Boyd 1998). References
Above-ground shoots may live 11 to 15 years.
Shoot regeneration is rapid, 30 to 60 cm within the Boyd, C.S. and T.G. Bidwell. 2002. Effects of
first year, when above ground tissues are removed prescribed fire on shinnery oak (Quercus
mechanically or through fire. The response to fire havardii) plant communities in western
is variable and dependent on timing of the fire. Oklahoma. Restoration Ecology 10(2): 324-
Burning reduced shin oak cover by at least 25 333.
percent, with the greatest reduction, 50 percent,
following spring burns (Boyd and Bidwell 2002). Boyd, C.S. and T.G. Bidwell. 2001. Influence of
Winter and annual fires increased rhizomatous prescribed fire on lesser prairie chicken habitat
grass cover, and although oak cover was reduced, in shinnery oak communities in western
regrowth resulted in greater stem densities (e.g. Oklahoma. Wildlife Society Bulletin 29(3):
increasing from 20 stems/ha to 40 stems/ha). 938-947.
Benefits.—The sandy soils that Havard shin oak Boyd, C.S., L.T. Vermeire, T.G. Bidwell, and R.L.
grows in are susceptible to wind erosion. The Lochmiller. 2001. Nutritional quality of
extensive underground rootstock system and dense shinnery oak buds and catkins in response to
above ground shoots make it an important soil burning or herbivory. Southwestern Naturalist
stabilizer. Oak leaves can be good forage for sheep 46(3): 295-301.
and goats, but are less utilized by cattle.
Consumption of too much, especially young spring Carter, J.L. 1997. Trees and Shrubs of New
shoots, can be toxic and result in illness or even Mexico. Johnson Books, Boulder, CO. 534 p.
death for cattle and lambs (Gay and Dwyer 1998,
Warnock 1974). As with many oak species, Correll, D.S. and M.C. Johnston. 1970. Manual
toxicity is due to phenolics and tannins. Later in of the Vascular Plants of Texas. Texas Research
the growing season cattle will utilize it in small Foundation, Renner, TX. 1,881 p.
and varying degrees. The crude protein content of
foliage is about 9 percent. The plants are browsed Dick-Peddie, W.A. 1993. New Mexico
by deer, pronghorn antelope, and southern plains Vegetation: Past, Present and Future.
woodrats. Phenolic content (15 to 20 percent) of
615
University of New Mexico Press, Albuquerque, Mayes, S.G., M.A. McGinley, and C.R. Werth.
NM. 244 p. 1998. Clonal population structure and genetic
variation in sand-shinnery oak, Quercus
Flora of North America Editorial Committee. havardii (Fagaceae). American Journal of
1997. Flora of North America, Vol 3. Botany 85(11): 1609-1617.
Magnoliophyta: Magnoliidae and Hamamelidae.
Flora of North America Editorial Committee, McCoy, D. 1981. Roadside Trees and Shrubs of
Oxford University Press, New York. 590 p. Oklahoma. University of Oklahoma Press,
Norman, OK. 116 p.
Freeman, C.C. 2000. Vascular plants new to three
states in the central United States. Transactions Muller, C.H. 1952. Ecological control of
of the Kansas Academy of Science 103(1-2): 51- hybridization in Quercus; A factor in the
54. mechanism of evolution. Evolution 6: 147-161.
Gay, C.W., Jr. and D.D. Dwyer. 1998 Reprint. Peterson, R.S. and C.S. Boyd. 1998. Ecology and
New Mexico Range Plants. Cooperative Management of Sand Shinnery Communities: A
Extension Service Circular 374. Revisions by: Literature Review. General Technical Report
C. Allison, S. Hatch, and J. Schickedanz. New RMRS-GTR-16. United States Department of
Mexico State University, Las Cruces, NM. Agriculture Forest Service, Rocky Mountain
84 p. Research Station, Fort Collins, CO. 44 p.
Great Plains Flora Association. 1986. Flora of the Sears, W.E., C.M. Britton, D.B. Wester, and R.D.
Great Plains. University Press of Kansas, KS. Pettit. 1986. Herbicide conversion of a sand
1,392 p. shinnery oak (Quercus havardii) community:
effects on biomass. Journal of Range
Kartesz, J.T. 1994. A Synonymized Checklist of Management 39: 399-403.
the Vascular Flora of the United States, Canada,
and Greenland, Vol. 1, 2nd Edition. Biota of Warnock, B.H. 1974. Wildflowers of the
North America Program of the North Carolina Guadalupe Mountains and the Sand Dune
Botanical Garden. Timber Press, Portland, OR. Country, Texas. Sul Ross State University,
622 p. Alpine, TX. 176 p.
Martin, W.C. and C.E. Hutchins. 1980 (reprinted Young, J.A. and C.G. Young. 1992. Seeds of
2001). A Flora of New Mexico, Vol. 1. Bishen Woody Plants in North America. Dioscorides
Singh Mahendra Pal Singh, India and Koeltz Press, Portland, OR. 407 p.
Scientific Books, Germany. 1,276 p.
616
Randia aculeata L. white indigo berry
RUBIACEAE
Synonyms: Randia mitis L.

Gardenia randia Sw.
Randia latifolia Lam.
Randia aculeata L. var. mitis (L.) Griseb.
John K. Francis
white fruits (or ivory with a salmon blush), with a

brittle shell-like skin, have a blue or black pulp
and five to 10 round, flattened seeds (Howard
1989, Liogier 1997, Little and others 1974). There
are populations and transition populations with
and without spines, with large leaves and small
leaves, and with large and small fruits. These gave
rise to variety designations that are no longer
recognized (Howard 1989). White indigo berry is
also known as box-brier, inkberry, prickle bush,
fishing rod, five fingers, goat horn, tintillo, tintero,
palo de cotorra, árbol de navidad, sota-caballo,
resuelesuele, café cimmarrón, espino cruz,
crucete, crucito, maíz tostado, raboe, peetsch-
kitam, cabai nache, croc-à-chien, bois-lance, ti
coco, and petit coco (Howard 1989, Little and
others 1974).
Range.—White indigo berry is native to southern

Florida, Bermuda, the Bahamas, the Caribbean
Islands, and from Mexico through Central
America and South America as far south as
Colombia (Little and others 1974, Smithsonian
Institution 2001). It is not known to have
Ecology.—White indigo berry grows in most

General Description.—White indigo berry is a types of soils including excessively drained and
shrub or small tree from 2 to 6 m in height. It is well drained, but not poorly drained soils. It
spiny and may have one to several stems. White inhabits areas that receive from about 700 to 2200
indigo berry has smooth to slightly fissured gray mm of annual precipitation from near sea level to
bark, opposite, often horizontal, branches, a thin over 600 m of elevation in Puerto Rico. In Florida,
crown, and rough appearance. The plant is white indigo berry grows in unburned pinelands
supported by an extensive tap and lateral root and along the margins of coastal hammocks
system. The roots have a corky, ivory colored bark (Nelson 1996). In Puerto Rico, the species grows
and are stiff and woody. The spines are paired and in dry and moist forests, more commonly over
the foliage is crowded on the ends of short lateral limestone rocks, but also over igneous rocks and
twigs. The leaves are almost stalkless and entire, ultramafics (serpentine). Because white indigo
ovate, obovate, or orbicular. They are 1 to 6 cm berry is moderately intolerant of shade, it must
long and 0.5 to 3 cm broad, slightly thickened, grow in openings or under low-density forest. The
shiny green above and light green below. The species invades abandoned land, but not quickly,
solitary flowers are small, white, funnel-shaped, and because of slow growth, is not common in
with a five-lobed corolla, and located on very early secondary forest following cultivation.
short lateral spurs. The 6- to 13-mm, globose, However, white indigo berry does hang on
617
tenaciously after disturbance and is prominent in References
secondary forests arising after logging or partial
clearing for pasture. Plants are usually well Dade County. 2001. Some trees and shrubs native
disbursed and seldom, if ever, form thickets. to South Florida. http://www.fnps.org/dade/
pubs/SomeTrees.html. 6 p.
Reproduction.—White indigo berry blooms and
fruits irregularly throughout the year. Fruits (n = Howard, R.A. 1989. Flora of the Lesser Antilles,
73) collected in Puerto Rico weighed an average Leeward and Windward Islands.
of 0.472 + 0.014 g/fruit. Fourteen of the fruits Dicotyledoneae. Part 3. Vol. 6. Arnold
contained from three to 14 seeds and averaged 8.2 Arboretum, Harvard University, Jamaica Plain,
seeds. The air dry seeds from these fruits averaged MA. 658 p.
0.0264 + 0.0003 g/seed or 38,000 seeds/kg. Sown
in commercial potting mix, 25 percent germinated Liogier, H.A. 1990. Plantas medicinales de Puerto
between 46 and 168 days after sowing. Rico y del Caribe. Iberoamericana de Ediciones,
Growth and Management.—Weaver (1990)
measured 12 plants over a 5-year period in closed Liogier, H.A. 1997. Descriptive flora of Puerto
low forest in St. John, U.S. Virgin Islands and Rico and adjacent islands. Vol. 5. Editorial de la
found the vary slow diameter growth rate of 0.06 Universidad de Puerto Rico, Río Piedras, PR.
+ 0.02 cm/yr. A 1-m tall sapling in a moist area of 436 p.
Puerto Rico with a basal diameter of 2 cm had 10
growth rings (author’s observation). Ornamental Little, E.L., Jr., R.O. Woodbury, and F.H.
plants also grow slowly (Workman 1980). Seeds Wadsworth. 1974. Trees of Puerto Rico and the
for nursery production should be cleaned by Virgin Islands. Vol. 2. U.S. Department of
maceration and wet sieving before sowing. Agriculture, Agriculture Handbook 449.
Production of seedlings in containers is Washington, DC. 1,024 p.
recommended. Because growth is slow, a year or
more in the nursery is required followed by Nelson, G. 1996. The shrubs and woody vines of
protection from weeds in the field. Florida. Pineapple Press, Inc. Sarasota, FL.
391 p.
Benefits.—White indigo berry has been used as an
ornamental to a limited extent in Florida and, Smithsonian Institution. 2001. Centers of plant
because it is native and resists drought and diversity: Columbian Central Massif.
hurricanes, it is currently being recommended for http://www.nmnh.si.edu/botany/projects/cpd/sa/
planting (Dade County 2001). It is available as sa29.htm. 14 p.
nursery-grown plants, and plants from the wild
can be successfully dug up and transplanted. The Vélez, I. and J. van Overbeek. 1950. Plantas
fruits are edible although of poor flavor indeseables en los cultivos tropicales. Editorial
(Workman 1980). White indigo berry provides Universitaria, Río Piedras, PR. 497 p.
food and nesting sites for birds (Vélez and van
Overbeek 1950) and nectar for butterflies (Dade Weaver, P.L. 1990. Tree diameter growth rates in
County 2001). The wood is hard and heavy and Cinnamon Bay Watershed, St. John, U.S. Virgin
useful for fuel. The shrub is sometimes decorated Islands. Caribbean Journal of Science 26(1-2):
for a Christmas tree, hence the Spanish name, 1-6.
árbol de navidad. Other names (tintillo and
inkberry) arose from the former use of the berries Workman, R.W. 1980. Growing native. The
for dye and for ink (Little and others 1974). The Sanibel-Captiva Conservation Foundation, Inc.,
plant is also frequently formed into bonsai plants. Sanibel, FL. 137 p.
The fruits are used in herbal medicine to control
dysentery. An unspecified part of the plant is used
to control fever, and the latex is reported to
effectively stop bleeding (Liogier 1990).
618
Rauvolfia viridis Willd. ex Roemer & J.A. Schultes bitter bush
APOCYNACEAE
Synonyms: Rauvolfia lamarckii A. DC. in DC.

Rauvolfia latifolia A. DC in DC.
Rauvolfia lanceolata sensu Griseb.
John K. Francis
(Departamento de Botánica 1999), and Colombia

(Missouri Botanical Garden 2001).
Ecology.—In Puerto Rico, bitter bush grows in

dry and moist forests that receive from 750 to
about 1700 mm of annual precipitation. Usually
rocky or otherwise poor sites are colonized in
areas of both igneous and sedimentary (including
limestone) rocks. The species grows from near sea
level to about 400 m in elevation. Bitter bush
requires full or partial sunlight. It grows in open,
disturbed areas and low basal area, secondary and
remnant forest. It withstands competition from
herbs and shrubs well but eventually succumbs to
competition from tall grass swards and closed
forests. Although never abundant, the species
becomes more common in overgrazed areas.
General Description.—Bitter bush, also known Reproduction.—A sample of bitter bush fruits
as bellyache bush, balsam, sassafras, milk tree, collected in Puerto Rico weighed an average of
and ti bois-lait, is a branchy shrub to 2.5 m in 0.124 + 0.005 g/fruit. Air-dried seeds separated
height (Howard 1989, Liogier 1995). The species from these fruits averaged 0.0247 + 0.0004 g/seed
rarely becomes a small tree to 4 m in height or 40,500 seeds/kg. Sown in commercial potting
(Liogier 1995) and 7 cm in diameter at breast mix without any pretreatment, 5 percent
height (Weaver 1990). Usually, there are multiple germinated after 11 months. Birds that eat the
stems, as suckers arising from lateral roots, fruits disburse the seeds.
especially after fire or other disturbance. The root
system is dominated by lateral roots, which Growth and Management.—Growth of bitter
support sinker roots and a moderate amount of bush is slow. The author estimated height growth
fine roots. The ovate or ovate-elliptic leaves are 5 of young shrubs from sprouts at 0.5 m/year.
to 15 cm long by 3 to 5 cm broad, and grow in Weaver (1990) measured the diameter growth rate
whorls of four or rarely three leaves. The several- of two old small trees and found an average of
to many-flowered inflorescences are usually only 0.01 cm/yr. No planting or management
terminal. The tiny flowers are tubular and white. experience has been reported.
The fruits are spherical or slightly flattened and 5
to 8 mm in diameter. Each contains two seeds Benefits.—The roots of bitter bush were analyzed
(Howard 1989, Liogier 1995). for alkaloids. Ajmalidine, a-yohimina, reserpinine,
quebrachidine, and yohimbina were isolated.
Range.—According to Howard (1989) and Reserpine, a sedative present in other Rauvolfias,
Liogier (1995) bitter bush is native to Hispaniola, was not found (Martínez Pérez and others 1997).
Puerto Rico, the Virgin Islands, the Lesser Bitter bush furnishes minor food and cover for
Antilles, Trinidad, Margarita, and Venezuela. birds and other wildlife. With other dry forest
There are also records for Cuba (Martínez-Pérez vegetation, it protects the soil from erosion.
and others 1997), Navassa Island (New York
Botanical Garden 2001), Costa Rica
619
References Tápanes, and V. Fuente Fiallo. 1997. Bases
débiles minoritarias de la raíz de Rauvolfia
Departamento de Botánica. 1999. Especies de viridis Roem. et Schult. Revista Cubana Farm
plantas nuevas para Costa Rica. Departamento 31(3): 188-191.
de Botánica, Inventario Biótico.
http://www.minae.go.cr/estrategia/estudio/planta Missouri Botanical Garden. 2001. Current
s1.htm. 4 p. specimen list for Rauvolfia viridis.
http://mobot.mobbot.org/cgi-bin/search_vast.
Howard, R.A. 1989. Flora of the Lesser Antilles, 1 p.
Dicotyledoneae. Part 3. Vol. 6. Arnold The New York Botanical Garden. 2001. Flora of
Arboretum, Harvard University, Jamaica Plain, Navassa Island: vascular plant checklist.
MA. 658 p. http://www.nybg.org/bsci/hcol/navassa/checklist
vasc.html. 9 p.
Rico and adjacent islands. Vol. 4. Editorial de la Weaver, P.L. 1990. Tree diameter growth rates in
Universidad de Puerto Rico, Río Piedras, PR. Cinnamon Bay Watershed, St. John, U.S. Virgin
617 p. Islands. Caribbean Journal of Science 26(1-2):
1-6.
Martínez Pérez, J., A. Bello Alarcío, O. Montero
620
Rhododendron macrophyllum D. Don ex G. Don Pacific rhododendron
ERICACEAE
Synonyms: Hymenanthyes macrophyllum (D. Don ex G. Don) H.F. Copel

Rhododendron californicum Hook.
Christopher Ross
livestock losses (Kingsbury 1964). About 30

diterpine, polyhydroxylated cyclic hydrocarbons
are found in rhododendrons and related species
(Harborne and Baxter 1993). Sucking nectar from
flowers has caused illness in humans, as has
consumption of rhododendron honey (Alberta
Government 2002). The species has recently been
identified as a host of the algae Phytophthora
ramorum responsible for sudden oak death in
California, but the effects are not yet known
(University of California Cooperative Extension
2002). A variety of insect and fungi may attack
rhododendrons, including several caterpillars such
as cutworms, loopers, and leafrollers (Foss and
Antonelli 1999).
Reproduction.—Pacific rhododendron flowers are

insect pollinated. Fruits are oblong capsules that
split along the sides to release minute seeds (4400
to 12,500/g) after ripening. Germination occurs
with no pretreatment, but requires light (Young
and Young 1992). Vegetative reproduction by
General Description.—Pacific rhododendron, crown sprouting is common after fire, logging, and
also known as coast rhododendron and California other disturbance. Propagation is by seeds or
rosebay, is a coarse-branched evergreen shrub to 4 cuttings.
m tall. The alternate leathery ovate to obovate
leaves are 7 to 15 cm long and 3 to 6 cm wide. Fire Effects.—Rhododendron has been classed as
Flowers are showy and bell shaped, about 4 cm a residual shrub (rather than a colonizer) after fire
long, with rose pink to purple corolla, five petals, (Dyrness 1973). It recovers well but slowly after
five sepals, and 10 stamens (Wallace 1993). fire, primarily from vegetative sprouting from root
crowns and stem bases. Fire dramatically reduces
Range.—Pacific rhododendron occurs near the it initially, and peak abundance is not reached until
Pacific coast from British Columbia south to late in the fire recovery process (Halpern 1989). It
Monterey, California, and in the coastal and is scarce after severe burns (Dyrness 1973).
Cascade mountain ranges of Oregon, Washington,
and California (Hitchcock and others 1959, Growth and Management.—Pacific
Wallace 1993). It occurs in redwood, Douglas fir, rhododendron typically occupies drier sites within
yellow pine, and mixed evergreen forests fron sea its range (Dyrness 1973). Like some other
level to over 1,000 m elevation (CalFlora 2000). rhododendrons, it may have a mycorrhizal
relationship that enables it to thrive on low-
Ecology.—Pacific rhododendron grows in acidic nutrient soils (USDA USFS 2002). The tendency
(pH 5.5) to neutral fine textured soils (USDA to resprout from stem crowns has caused difficulty
2002). Pacific rhododendron, like many others of for conifer reforestation (Halverson and others
the genus, is highly toxic to humans and animals. 1986), so that spring spraying with triclopyr has
The poison is a neurotoxin, andromedotoxin, or been used for control in some areas (Burrill and
grayanotoxin, which has occasionally caused large others 1989). At California’s Kruse Rhododendron
621
State Reserve, Pacific rhododendron is considered for the western hemlock zone: Mt. Hood
to be an early seral stage after fire or logging, and National Forest. R6-ECOL-232A. U.S.
before tanoak, which is being thinned in order to Department of Agriculture, Forest Service,
set back the succession process and maintain the Pacific Northwest Region, Portland, OR.
rhododendrons (California State Parks 2002). 111 p.
Benefits.—Pacific rhododendron is widely Harborne, J.B. and H. Baxter. 1993.

available from commercial nurseries and is valued Phytochemical Dictionary. Taylor and Francis,
for native landscaping purposes. It has served as a London and Washington, D.C. 791 p.
source of genetic material for many ornamental
cultural selections. It is also used in erosion Hitchcock, C.L., R. Cronquist, and M. Ownbey.
control (Olson 1974). It has no other common uses 1959. Vascular plants of the Pacific northwest.
(USDA 2002). Part 4: Ericaceae through Campanulaceae.
University of Washington Press. Seattle, WA.
References 510 p.
Alberta Government. 2002. Rhododendron Kingsbury, J.M. 1964. Poisonous plants of the
toxicity. http://www.Agric.gov.ab.ca/crops. United States and Canada. Prentice Hall, Inc.
[not paged]. Englewood, NJ. 626 p.
Burrill, L.C., W.S. Braunworth Jr., and R.D. Olson, D.F., Jr. 1974. Rhododendron L.
William, comps. 1989. Pacific Northwest weed rhododendron. In: C.S. Schopmeyer, ed. Seeds
control handbook. Oregon State University, of woody plants in the United States.
Extension Service, Agricultural Agricultural Handbook 450. U.S. Department
Communications. Corvallis, OR. 276 p. of Agriculture, Forest Service, Washington,
D.C. p. 709-712.
CalFlora. 2000. Information on California plants
for education, research and conservation. University of California Cooperative Extension.
Berkeley, California: The CalFlora Database. 2002. Sudden oak death. http://cemarin.
http://www. Calflora.org/ [not paged]. ucdavis.edu. 2 p.
California State Parks–Kruse Rhododendron State USDA NRCS. 2002. Plants Source and Reference.
Reserve. 2002. http://cal-parks.ca.gov. http://plants.usda.gov. [not paged].
Dyrness, C.T. 1973. Early stages of plant U.S. Department of Agriculture, Forest Service,
succession following logging and burning in the Rocky Mountain Research Station, Fire
western Cascades of Oregon. Ecology 54: 57- Sciences Laboratory. 2002. Fire Effects
69. Information System, http://www.fs.fed.us/ [not
paged].
Foss, C. and A. Antonelli. 1999. Hortsense.
http://pep.wsu.edu/hortsense. [not paged]. Wallace, G. 1993. Rhododendron. In: J.C.
Hickman, ed. 1993. The Jepson Manual:
Halpern, C.B. 1989. Early successional patterns of higher plants of California. U.C. Press,
forest species: Interactions of life history traits Berkeley and Los Angeles, CA. 1,400 p.
and disturbance. Ecology 70: 704-720.
Young, J.A. and C.G. Young. 1992. Seeds of
Halverson, N.M., C. Topik, and R. Van Vickle. woody plants of North America. Disceroides
1986. Plant association and management guide Press, Portland, OR. 407 p.
622
Rhododendron maximum L. rosebay rhododendron
ERICACEAE
Synonyms: none
Barton D. Clinton
optimum carbon gain, and has a tremendous
capacity for avoiding cavitation during freeze-thaw
cycles (Lipp and Nilsen 1997). Where extensive
overstory mortality has eliminated most of the
overstory, this species forms a thick and
continuous subcanopy known locally as ‘laurel
slicks’ or ‘laurel hells’. Rosebay rhododendron is
an important structural and functional component
of southern Appalachian forest ecosystems. What
isn’t clear is whether or not we are in a period of
advancement or retreat for this species. For
example, on poorly drained sites on ridge or upper
slope positions, large areas of rosebay
rhododendron, particularly at the high elevations,
have recently died out presumably due to the
General Description.—Rosebay rhododendron, phytophthora fungus, or due to recent prolonged
sometimes known as ‘great laurel’, is an periods of below-average precipitation. Yet,
understory evergreen shrub. It ranges in height rosebay rhododendron now occupies sites that
from 2 to 8 m, and can occasionally reach historically were free of evergreen understory.
diameters of 20 to 25 cm. Leaves are There are still important questions to be answered
schlerophyllous, simple, alternate, and oblong (10 regarding this species to completely understand its
to 30 cm long, 5 to 8 cm wide). It retains its waxy, role in forest understories.
deep-green leaves for up to 8 years, but once shed
are slow to decompose. It produces large, showy, Reproduction.—Rosebay rhododendron is clonal.
white to purple flowers each June. It is capable, however, of reproducing both
vegetatively and sexually. It reproduces
Range.—Worldwide, there are over 800 species in vegetatively through a process called ‘layering’
the genus Rhododendron. Most of the 26 North where it produces roots from aboveground woody
American species occur in the East along the parts when in contact with the forest floor. The
mountain chain stretching from Nova Scotia in the fruit is produced from showy flowers from March
north to Georgia and Alabama in the south, but are to August. The fruit is an oblong capsule that
concentrated in the southern Appalachians (Dobbs ripens in the fall, and splits along the sides soon
1998). Rosebay rhododendron is the most after ripening to release large numbers of minute
frequently occurring and dominant species of this seed (approx. 400 per capsule) (Schopmeyer
genus in the southern Appalachian region 1974). Microsite requirements for seed
(Swanson 1994), and occurs occasionally on mesic germination are relatively specific (e.g., high in
hill-slopes throughout the upper Piedmont organic matter such as rotting logs); hence, the
Crescent of the Southeastern United States. majority of reproduction is vegetative resulting in
a clonal distribution.
Ecology.—Approximately 1.2 million ha in the
southern Appalachians are occupied by this Growth and Management.—Seeds from rosebay
species (Dobbs 1995) where it dominates the rhododendron are minute and it is estimated that
understory. This species has historically been approximately 11 million are contained in 1 kg.
confined to riparian areas and other mesic sites but Commercial seed production is generally from
takes advantage of disturbed areas where it is cultivated hybrids. Seeds from wild sources are not
present to advance onto sub-mesic sites. It prefers commonly sold commercially. Rosebay
deep well-drained acid soils high in organic matter rhododendron is a slow-growing shrub and has a
where it produces a thick, peat-like humus. It very high sprout potential. If mechanical removal
prefers low to medium light conditions for is attempted in the case of forest management,
623
extremely high densities are attained by this Clinton. B.D. and J.M. Vose. 1996. Effects of
species in a manner of a few years. Prescribed fire Rhododendron maximum L. on Acer rubrum L.
has also been used to control this species but with seedling establishment. Castanea 61(1):38-45.
limited success (Clinton and Vose 2000).
Clinton, B.D. and J.M. Vose. 2000. Plant
Benefits.—Rosebay rhododendron is a striking succession and community restoration
and aesthetically pleasing feature of mesic following felling and burning in the southern
southern Appalachian forests. It is one of the Appalachian Mountains. In: W. K. Moser and
largest and hardiest rhododendrons grown C.F. Moser, eds. Fire and forest ecology:
commercially. Several cultivars with white to innovative silviculture and vegetation
purple flowers have been selected for the management. Tall Timbers Fire Ecology
horticultural trade (Brown and Kirkman 1990). Conference Proceedings, No. 21. Tall Timbers
Where it occurs naturally, it produces a showy, Research Station, Tallahassee, FL. p . 22-29.
white, pink, or light purple flower primarily in
June, but occurs from March into August. Rosebay Dobbs, M.M. 1995. Spatial and temporal
rhododendron maintains deep-green foliage year distribution of the evergreen understory in the
round. This species affords protection to steep southern Appalachians. Master’s Thesis,
watersheds and shelter for wildlife. The wood is University of Georgia, Athens, GA. 100 p.
very hard and is occasionally used for specialty
wood products. Dobbs, M.M. 1998. Dynamics of the evergreen
understory at Coweeta Hydrologic Laboratory,
Detrimental Effects.—For all its prized qualities North Carolina. PhD Dissertation, University
as a naturally occurring component of the of Georgia, Athens GA. 179 p.
landscape or as plantings in residential and
commercial landscaping, rosebay rhododendron Lipp, C.C. and E.T. Nilsen. 1997. The impact of
can have an inhibitory effect on regeneration of subcanopy light environment on the hydraulic
other plant species. There is some evidence to vulnerability of Rhododendron maximum to
suggest that due to fire suppression and the freeze-thaw cycles and drought. Plant, Cell
absence of other cultural activities (i.e., mountain- and Environment 20:1,264-1,272.
land grazing), this species has advanced beyond
the mesic forest sites into sub-mesic understories Nilsen, E.T., B.D. Clinton, T.T. Lei, O.K. Miller,
(Dobbs 1998). The significance of this movement S.W. Semones, and J.F. Walker. 2001. Does
onto previously unoccupied sites centers around Rhododendron maximum L. (Ericaceae) reduce
the impacts of rosebay rhododendron on plant the availability of resources above and
succession (Clinton and Vose 1996) and resource belowground for canopy tree seedlings?
availability (Nilsen and others 2001). Rosebay American Midland Naturalist 145:325-343.
rhododendron is associated with reduced woody
and herbaceous seedling abundance throughout its Nilsen, E.T., J.F. Walker, O.K. Miller, S.W.
range, and hence poses a serious impediment to Semones, T.T. Lei, and B.D. Clinton. 1999.
the production of wood products. The Inhibition of seedling survival under
mechanism(s) by which rosebay rhododendron Rhododendron maximum (Ericaceae): could
reduces seedling survival has been the subject of allelopathy be a cause? American Journal of
much debate. Possible sources of inhibition Botany 86(11):1,597-1,605.
include allelopathy, competition for resources
including light, physical and chemical attributes of Schopmeyer, C.S. 1974. Seeds of woody plants in
the forest floor and soil, and interactions between the United States. Agricultural Handbook 450.
some or all sources (Nilsen and others 1999, U.S. Department of Agriculture, Forest
Nilsen and others 2001). Service, Washington, DC. 883 p.
References Swanson, R.E. 1994. A Field Guide to the Trees

and Shrubs of the Southern Appalachians. The
Brown, C.L. and L.K. Kirkman. 1990. Trees of Johns Hopkins University Press, Baltimore,
Georgia and Adjacent States. Timber Press, MD. 399 p.
Portland, OR. 292 p.
624
Rhus copallinum L. shining sumac
ANACARDIACEAE
Synonyms: None
Kristina Connor
glossy, dark green leaflets (Brickell and Zuk 1996,

Odenwald and others
1996, Seiler and Peterson 2001). Leaflets are

paired, olbong-lanceolate in shape, and from 3 to
10 cm long (Bailey and Bailey 1976, Krussmann
1976, Johnson and Hoagland 1999). Buds are
glabrous, and terminal buds are absent (Synor and
Cowen [no date]). Sap is sticky and yellow
(Johnson and Hoagland 1999).
Range.—Shining sumac grows from southern

Ontario south along the Coastal Plain to Florida. It
extends westward into eastern Texas, and inland
from central Michigan and Wisconsin to Kansas
and Oklahoma (USDA-ARS [no date], Coladonato
1992). It is also found in Cuba. Recognized
varieties are R. copallinum var. latifolia Engl., R.
copallinum var. lanceolata Gray, and R.
copallinum var. leucantha (Jacq.) DC (USDA-
ARS [no date], USDA-NRCS 2001).
Ecology.—Shining sumac is a deciduous, clonal

shrub. It is an early pioneer species and grows best
on well drained soils in full sunlight (Coladonato
1992, Odenwald and others 1996, Seiler and
Peterson 2001). It can form thickets in abandoned
fields, along roadsides, in glades and open woods,
and in waste areas (Tenaglia 2002, Oplin 2001,
Arborquest 2001). It is difficult to get rid of, once
established, due to its ability to spread from
rhizomes (Arborquest 2001). This characteristic
also makes it well adapted to fire (Coladonato
Illustration source: Grelen and Duvall 1966 1992). Shining sumac will tolerate compacted soil,
drought, pollution, heavy pruning, and
General Description.—Shining sumac, also transplanting, but the shallow roots make it
known as black sumac, dwarf sumac, flameleaf susceptible to uprooting and the stems can break in
sumac, mountain sumac, shiny sumac, and winged strong wind storms (Arborquest 2001). It can be a
sumac, is an upright, deciduous shrub or (rarely) serious competitor with young pines and
small tree from 3 to 6 m tall (Bailey and Bailey hardwoods (Coladonato 1992).
1976, Odenwald and others 1996). Bark ranges in
color from light brown to gray to reddish-brown. Reproduction.—Shining sumac is a clonal species
Shoots and twigs are hairy and reddish in color. which sprouts from roots and the root crown. It
Twigs have conspicuous lenticels. The sparsely also regenerates sexually (Coladonato 1992). From
branched, flat crown is composed of alternate, May to August, plants produce panicles of small
pinnately compound leaves approximately 15 to 30 yellow-green dioecious flowers that have five
cm long, with wings between each of the 7 to 27 petals and five sepals (Tenaglia 2002). The
625
panicles are pubescent, about 12 to 15 cm long,
and can be either terminal or axillary (Brickell and Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third:
Zuk 1996, Johnson and Hoagland 1999, A Concise Dictionary of Plants Cultivated in the
Krussmann 1976). Brown and Kirkman (1990) United States and Canada. McMillan Publishing
report the presence of polygamous flowers but Co., Inc., New York. 1,312 p.
state that shining sumac is functionally dioecious
since the pistils of the polygamous flowers abort. Brickell, C. and J.D.. Zuk. 1996. The American
Thus, both male and female plants are needed for Horticultural Society A-Z Encyclopedia of
good fruit set to occur (Arborquest 2001) The 3- to Garden Plants. DK Publishing, Inc., New York.
5- mm pubescent fruits are small drupes that 1,092 p.
contain a single nutlet (Coladonato 1992). Fruits
ripen from August to October. They turn dark red Brinkman, K.A. 1974. Rhus L. Sumac. In: C.S.
when ripe and often remain on the plant Schopmeyere, tech. coord. Seeds of woody
throughout the winter. The hard-coated seeds plants in the United States. Agriculture
germinate poorly without pretreatment, but a 1 hr Handbkook 450. U.S. Department of
soak in sulfuric acid at room temperature can Agriculture, Forest Service Washington, D.C.
result in a 75 percent germinative capacity 883 p.
(Brinkman 1974).
Brown, C.L. and L.K. Kirkman. 1990. Trees of
Growth and Management.—Shining sumac is a Georgia and adjacent states. Timber Press,
fast growing but short lived clonal shrub, first Portland, OR. 292 p.
cultivated in 1688 (Brinkman 1974). It is planted
as an ornamental and used in mass plantings Coladonato, M. 1992. Rhus copallinum. In: U.S.
because of its brilliant red fall foliage but it can Department of Agriculture, Forest Service,
spread well beyond desired borders because of its Rocky Mountain Research Station Fire Sciences
ability to sprout from the roots. Laboratory, Fire Effects Information System
http://www.fs.fed.us/database/feis. 9 p.
Benefits.—Shining sumac is nonpoisonous to
humans (USDA-ARS [no date], Seiler and Grelen, H.E. and V.L. Duvall. 1966. Common
Peterson 2001, Synor and Cowen [no date]), and plants of longleaf pine-bluestem range. Research
the fresh fruits can be used to make a lemon- Paper SO-23. U.S. Department of Agriculture,
tasting beverage. It was also used by Native Forest Service, Southern Forest Experiment
Americans to treat dysentery and mouth sores Station, New Orleans, LA. 96 p.
(Tenaglia 2002). Dense thickets of shining sumac
serve as cover for birds and mammals. The seeds Johnson, F.L. and B.W. Hoagland. 1999. Rhus
are eaten by a variety of birds, while the flowers copallinum L. Catalog of the Woody Plants of
attract butterflies. Deer and rabbits commonly Oklahoma, Oklahoma Biological Survey.
browse the twigs in winter; rabbits also eat the http://www.biosurvey.ou.edu. 1 p.
bark (Grelen and Duvall 1966, Oplin 2001,
Arborquest 2001, Coladonato 1992). It is not, Krüssmann, G. 1976. Manual of Cultivated Broad-
however, a preferred food and is considered a poor Leaved Trees and Shrubs. Volume I, A-D.
to moderately important browse (Coladonato Timber Press, Beaverton, OR. 448 p.
1992). However, the seeds, bark and leaves, while
low in nutritive value, have a high tannin content Odenwald, N.G., C.F. Fryling, Jr., and T.E. Pope.
and have been used by the leather industry (Bailey 1996. Plants for American Landscapes.
and Bailey 1976, Brown and Kirkman 1990, Louisiana State University Press, Baton Rouge,
Coladonato 1992). Its ability to sprout from the LA. 266 p.
roots, coupled with its rapid growth rate, make
shining sumac a good species to plant for erosion Oplin. 2001. Dwarf Sumac--Rhus copallina. Ohio
control (Arborquest 2001). Public Library Information Network (OPLIN)
and the Ohio Historical Society (OHS).
References http://www. oplin.lib.oh.us/products/tree. 2 p.
Arborquest. 2001. Rhus copallina. Seiler, J.R. and J.A. Peterson. 2001.Shining
http://www.arborquest.com. 2 p. sumac. Virginia Tech. Dendrology Webpage
626
http://www.fw.vt.edu/dendro/dendrology/syllab USDA, ARS National Genetic Resources Program.
us/ivirginica.htm. 1 p. Germplasm Resources Information Network-
(GRIN). [no date]. Online Database. National
Synor, T.D. and W.F. Cowen. [no date]. Ohio Germplasm Resources Laboratory, Beltsville,
trees. Rhus--Sumac. The Ohio State University MD. http://www.ars-grin.gov/npgs/
College of Food, Agricultural, and searchgrin.html.
Environmental Sciences, Extension Bulletin
700-00. http://www.ag.ohio- USDA, NRCS. 2001. The PLANTS database.
state.edu/~ohioline/b700. 3 p. National Plant Data Center, Baton Rouge, LA.
http://plants.usda.gov. [not paged].
Tenaglia, D. 2002. Rhus copallinum. The Missouri
Flora Website. http://www.missouriplants.com.
4 p.
627
Rhus glabra L. smooth sumac
ANACARDIACEAE
Synonyms: Rhus borealis Greene

Rhus calophylla Greene
Rhus canadensis Mill.
Rhus carolineana Mill.
Rhus cismontana Greene
Rhus occidentalis (Torr.) Blank. (additional synonyms: Institute of
Systematic Botany 2003).
John K. Francis
gray-brown stem but often occurs in clonal
thickets formed by suckers from horizontal roots.
The relatively few branches and twigs are thick,
have a soft pith, and large, horseshoe-shaped leaf
scars. Stems and branches ooze a milky-resinous
sap when cut. The alternate leaves are uneven,
pinnately compound, up to 46 cm long, and
support seven to 29 leaflets. Leaflets are almost
stalkless and have serrate edges. (Clark 1973). The
foliage turns a brilliant red or scarlet during the
fall. Small, greenish-yellow flowers are grouped in
tight, 15- to 25-cm terminal panicles. The fruits are
dry, hairy, red to scarlet drupes that contain one
hard ovoid, dark-gray nutlet. The fruits have a
tangy, sour taste (Johnson 1995). Smooth sumac is
dioecious (male and female plants) (Wyman
1969). Total above-ground dry biomass may be
predicted by the equation: dry weight = 1.5130 +
0.6292 D2H, where D = diameter at 1 cm above
groundline, H = total height and r2 = 0.974
(Reeves and Lenhart 1988).
Range.—Smooth sumac is native to every state of

the United States (except Alaska and Hawaii) and
from Nova Scotia to British Colombia and south
into northern Mexico (Abrams 1951, Fire Sciences
Laboratory 2003, Natural Resources Conservation
Service 2003). However, it is most common in the
eastern half of United States north of Florida
(Hutchison 2003). It has been planted widely as an
ornamental but has not been reported to have
Ecology.—Smooth sumac prefers rich, well-
Hunt Institute
drained soil but will grow on excessively drained,
sandy soils and eroded sites without topsoil. In
General Description.—Smooth sumac, also
much of the Eastern range, it grows as a seral
known as common sumac, Rocky Mountain
species on disturbed sites such as logged areas,
sumac, red sumac, scarlet sumac, lemonade sumac,
abandoned fields, neglected pastures, old
western sumac, and white sumac, is a deciduous
construction sites, strip mines, stream overflow
shrub or a small tree from 0.5 to 6 m in height and
areas, roadsides, and fence rows (Hutchison 2003).
1 to 18 cm in stem diameter. It usually has a single
628
In Western areas it grows in stable communities on requires more than ordinary maintenance to keep
mountains, hillsides, and canyons. The species is smooth sumac from spreading vegetatively. One
used as a climax indicator in a number of shrub- method of keeping plants in check is to establish
grassland communities (Fire Sciences Laboratory them in soil space limited by rock or concrete
2003). Smooth sumac usually grows with full sun barriers (Clark 1973). The tops are killed by fires,
but tolerates partial shade. It grows from low but underground rhizomes (7.5 to 30 cm deep) are
elevations in many areas and up to 2,100 m in not damaged and sprout vigorously, increasing the
Arizona (Kearney and Peebles 1951). density and extent of smooth sumac stands. Seeds
from the soil seed bank are also stimulated to
Reproduction.—Smooth sumac flowers from germinate. Spring fires are more beneficial than
May to late July, depending on location. Fruits late summer fires (Fire Sciences Laboratory 2003).
ripen in September and October and persist When it is necessary to control the plant, it is
through the fall and winter (Fire Sciences recommended that they be lopped during or
Laboratory 2003). The flowers are pollinated by shortly after flowering and the sprouts spot-
bees (Rowe and Blazich 2003). Smooth sumac sprayed with glyphosate (Hutchison 2003).
produces at least some fruits and seeds nearly
every year (Fire Sciences Laboratory 2003). There Benefits and Detriments.—Smooth sumac helps
are about 2,000 seeds per well-formed panicle protect the soil in disturbed areas, adds to the
(Browse-Shrub and Forb Committee 1985), much aesthetics of wildlands, and furnishes food and
less in some areas and during dry years. Rowe and cover for wildlife. The fruits are eaten by a wide
Blazich (2003) report 50,600 to 105,600 fruits/kg variety of birds (Johnson 1995). Smooth sumac
and 52,800 to 277,200 seeds/kg. Plummer and foriage is rated poor in both energy and protein
others (1968) found 137,700 seeds/kg in samples value (Fire Sciences Laboratory 2003). White tail
from Utah. Air-dried fruits collected by the author deer (Odocoileus virginianus) browse it during fall
in Utah averaged 0.0202 + 0.0004 g/fruit. Seeds and winter but do not weaken the plants (Strauss
separated from them averaged 0.0103 + 0.0002 1991). It is also browsed by mule deer (Odocoileus
g/seed or 97,000 seeds/kg. The seeds are dispersed hemionus) (Fire Sciences Laboratory 2003).
principally by birds and form a soil seed bank Smooth sumac is valued as an ornamental for its
(Fire Sciences Laboratory 2003). Once individual brilliant red and scarlet fall colors and because it
plants are established, horizontal roots sucker to can be planted almost anywhere in the Continental
form clonal thickets. United States. It is particularly useful for
Nursery plants are usually produced from vegetating sandy banks and for low screens (Clark
seed, although production from cuttings is also 1973) and has been recommended for establishing
done to preserve favorable traits of select strains “living snow fences.” The species has been planted
(such as assuring only female plants). Rootstalks to revegetate roadsides and cuts, landfills, and strip
pulled up from wild stands in the early spring mines (Fire Sciences Laboratory 2003, Plummer
when the ground is wet will survive almost 100 and others 1968). In addition to using dried leaves
percent if planted properly (Plummer and others as an adulterant for tobacco and making drinks
1968). Seed can be collected in quantity by hand in from the fruits, Native Americans employed
fall or early winter. Seeds may be separated from smooth sumac to treat a large number of ailments,
the fruits by rubbing or beating followed by particularly mouth and throat sores, burns, to
screening (Rowe and Blazich 2003) and can be control diarrhea, and to promote urination (Fire
stored for as long as 5 years without loss of Sciences Laboratory 2003, Giese 1996, Moerman
viability (Fire Sciences Laboratory 2003). 1986). Basis for medicinal uses may be found in
Scarification to break physical (endocarp) the fact that methanol extracts of ground branches
dormancy may be done by a 3 to 4 hr soak in were effective (although not as effective as
concentrated sulfuric acid (Rowe and Blazich commercial antibiotics) against total of 11 species
2003) or emersion for less than 1 min. in boiling of bacteria. The antibacterial compounds were
water (Li and others 1999). identified as methyl gallic acid, gallic acid, and the
methyl ester of 3,4,5-trihydroxybenzoic acid
Growth and Management.—Smooth sumac has a (Saxena and others 1994). Some people develop
moderate growth rate of 30 to 46 cm in height per allergic dermatitis after contacting the sap, but the
year (Michigan State University Extension 1999). species is not as dangerous as many other Rhus
Although clones may last much longer, individual species (Clark 1973). Thickets of smooth sumac
stems are not long-lived, perhaps 2 to 10 years. It can prevent the development of tree seedlings
629
through shade and root competition and arrest in seeds of two North American Rhus species
succession (Putz and Canham 1992). (Anacardiaceae). American Journal of Botany
86(11): 1,505-1,511.
References
Michigan State University Extension. 1999. Rhus
Abrams, L. 1951. Illustrated flora of the Pacific glabra—smooth sumac. http://www.msue.msu.
States. Vol. 3. Stanford University Press, edu/msue/imp/modzz/00001245.html. 1 p.
Browse-Shrub and Forb Committee. 1985. America. Research Reports in Ethnobotany Vol.
Handbook on seeds of browse-shrubs and forbs. 1. University of Michigan Museum of
Tech. Pub. R8-TP8. Association of Official Seed Anthropology, Ann Arbor, MI. 534 p.
Analysts and U.S. Department of Agriculture,
Forest Service, Southern Region, Atlanta, GA. Natural Resources Conservation Service. 2003.
246 p. Plants profile: Rhus glabra L., smooth sumac.
Clark, L.J. 1973. Wild flowers of British symbol=RHGL. 5 p.
Columbia. Gray’s Publishing Limited, Sidney,
BC, Canada. 591 p. Plummer, A.P., D.R. Christensen, and S.B.
Monsen. 1968. Restoring big-game range in
Fire Sciences Laboratory. 2003. Fire effects Utah. Pub. 68-3. Utah Division of Fish and
information system: Rhus glabra. U.S. Game, Salt Lake City, UT. 183 p.
Rocky Mountain Research Station, Fire Sciences Putz, F.E. and C.D. Canham. 1992. Mechanism of
Laboratory, Missoula, MT. http://www.fs. arrested succession in shrublands: root and shoot
fed.us/database/feis/plants/shrub/rhugla/all.html. competition between shrubs and tree seedlings.
28 p. Forest Ecology and Management 49(3-4): 267-
275.
Giese, P. 1996. Red sumac: multi-tribal use.
http://www.kstrom.net/isk/food/rhususes.html. Reeves, H.C. and J.D. Lenhart. 1988. Fuel weight
4 p. prediction equations for understory woody
plants in eastern Texas. Texas Journal of
Hutchison, M. 2003. Vegetation management Science 40(1): 49-53.
guideline: smooth sumac (Rhus glabra). Illinois
nature Preserves Commission, Belknap IL. Rowe, D.B. and F.A. Blazich. 2003. Rhus L.,
http://www.inhs.uiuc.edu/VMG/smsumac.html. sumac. In: F.T. Bonner and R.G. Nisley, eds.
5 p. Woody plant seed manual. U.S. Department of
Agriculture, Forest Service, Washington, DC.
Institute of Systematic Botany. 2003. Atlas of http://wpsm.net.Rhus.pdf. 16 p.
Florida plants: Rhus glabra. http://www.
plantatlas.usf.edu/main.asp?plantID=3941. 2 p. Saxena, G., A.R. McCutcheon, S. Farmer, G.H.N.
Towers, and R.E.W. Hancock. 1994.
Johnson, F.D. 1995. Wild trees of Idaho. Antimicrobial constituents of Rhus glabra.
University of Idaho Press, Moscow, ID. 212 p. Journal of Ethnopharmacology 42(2): 95-99.
Kearney, T.H. and R.H. Peebles. 1951. Arizona Strauss, S.Y. 1991. Direct, indirect, and
flora. University of California Press, Berkeley cumulative effects of three native herbivores on
and Los Angeles, CA. 1,032 P. a shared host plant. Ecology 72(2): 543-558.
Li, X., J.M. Baskin, and C.C. Baskin. 1999. Wyman, D. 1969. Shrubs and vines for American
Anatomy of two mechanisms of breaking gardens. The Macmillan Company, London.
physical dormancy by experimental treatments 613 p.
630
Rhus microphylla Engelm. ex Gray little-leaf sumac
ANACARDIACEAE
Synonym: Rhoeidium microphyllum (Engelm.) Greene
James E. Nellessen
1993, Whitford 2002). It demonstrated a 46

percent nitrogen and phosphorus resorption
efficiency from leaves prior to leaf drop in the
autumn (Whitford 2002). This was lower than the
obligate riparian shrubs, desert willow [Chilopsis
linearis (Cavanilles) Sweet] and cutleaf brickellia
(Brickellia laciniata A. Gray), but higher than the
nitrogen fixing mesquite (Prosopis glandulosa
Torrey). Hence, the greater resorption efficiencies
of these shrubs may be due to their biology and
ecology as it relates to plant nitrogen demand and
nitrogen availability within their local habitats
(Killingbeck and Whitford 2001). Littleleaf sumac
General Description.—Little-leaf sumac is as well as Torrey yucca (Yucca torreyi Shafer) and
sometimes referred to as desert sumac because of javelina bush (Condalia warnockii MC Johnston)
its occurrence in habitats that are drier than the were the most frequently used species for nesting
habitats of other sumacs. Another name is birds in a Chihuahuan Desert location in south-
correosa. This highly branched shrub may reach central New Mexico (Kozma and Mathews 1997).
heights of up to 4 to 5 m and is frequently as broad This observation was made despite the fact that
as it is tall. Some twigs may be somewhat spiny- little-leaf sumac was lower in density (209
tipped. Leaves are odd-pinnately compound, stems/ha in arroyo sites and six stems/ha in upland
alternate, with five to nine elliptic to oblong sites) than other shrub species. Twenty-seven bird
leaflets, which are sessile, 8 to 18 mm long, 2 to 6 species were monitored from 1993-1995. Large
mm wide, and generally not leathery or shiny. junipers may function as nurse plants, aiding the
Leaflet margins are entire and may be somewhat establishment of certain shrub species, including
revolute, and the rachis is winged (Carter 1997, little-leaf sumac, in western Texas (McPherson
Correll and Johnston 1970, Epple 1995, Great and others 1988).
Plains Flora Association 1986, Kearney and others
1951, Martin and Hutchins 1980-81). Leaves Reproduction.—Flowers are small, greenish
persist on the plant until the first hard frost. white with five petals and five stamens. Flowers
appear before the leaves in both axillary and
Range.—Little-leaf sumac occurs from western terminal clusters. Blooming may occur from
Texas, southwestern Oklahoma, through the March through May depending on local
southern half of New Mexico into southeastern environmental conditions and availability of
Arizona, and south into Mexico. It occurs at moisture. Fruits are red to orange drupes, from 4.5
elevations from about 1,060 m to 1,830 m. to 6 mm in diameter, with a covering of glandular
hairs, and maturing in July to August.
Ecology.—This species is a common member of
Chihuahuan Desert grassland vegetation although Seed Dormancy.—Seed dormancy appears to be
its distribution is not uniform. It may occur in caused by a water impermeable endocarp (Li and
canyons, along river banks, arroyos, rocky others 1999). After incubation on moist substrate
hillsides, even alkali flats. It grows on both for 4 weeks, 29 to 34 percent of all seeds had
limestone and sandy soils and can be a dominant imbibed water, and after 1 year imbibition had
in some sandy-dune soils forming extensive climbed to 93 to 100 percent among all of the
coppice conditions. Littleleaf sumac will often seeds. This type of physical dormancy occurs in
grow adjacent to arroyos but is considered a other members of the genus Rhus (Young and
facultative or semiriparian species (Dick-Peddie Young 1992), and some members of the genus
631
also have physiological dormancy (Baskin and University of New Mexico Press, Albuquerque.
Baskin 2001). There is a high proportion of 244 p.
species with physical dormancy in savannas, hot
deserts, and similar climates with distinct dry and Emmerich, W.E., J.D. Helmer, K.G. Renard, and
wet seasons, the wet season being the season L.J. Lane. 1984. Fate and effectiveness of
favorable for germination. The impermeable tebuthiuron applied to a rangeland watershed. J.
endocarp in the genus Rhus may have evolved as of Environmental Quality 13(3): 382-386
long ago as the Middle Eocene epoch (Baskin and
Baskin 2001). Epple, A.O. 1995. A Field Guide to the Plants of
Arizona. Falcon Publishing Inc., Helena, MT.
Growth and Management.—Little-leaf sumac is 347 p.
not one of the faster growing shrubs. It produces
numerous stems and resprouts readily. Great Plains Flora Association. 1986. Flora of the
Tebuthiuron, an herbicide often used for brush Great Plains. University Press of Kansas.
control on rangelands, proved effective in little- 1,392 p.
leaf sumac control (Emmerich and others 1984).
Tebuthiuron was applied at 0.84 kg/ha to Kearney, T.H., R. Peebles, and Collaborators.
rangeland in both pelleted and dissolved forms. 1951 with 1960 supplement. Arizona Flora.
After 21 months there was a 100 percent kill of University of California Press, Berkeley, CA.
little-leaf sumac, tarbush (Flourensia cernua DC.), 1,085 p.
and four-wing saltbush [Atriplex canescens
(Pursh) Nutt.], all dominant species on the test site. Killingbeck, K.T. and W.G. Whitford. 2001.
Tebuthiuron moved to a soil depth of 15 cm within Nutrient resorption in shrubs growing by design,
8 months, and after 21 months 38 percent of the and by default in Chihuahuan Desert arroyos.
application was remaining, while only 0.47 percent Oecologia, Berlin 128(3): 351-359.
was detected in runoff water.
Kozma, J.M. and N.E. Mathews 1997. Breeding
Benefits.—The fruits are readily eaten by many bird communities and nest plant selection in
birds. Little-leaf sumacs are also important nesting Chihuahuan desert habitats in south-central New
sites for birds. The fruits may be steeped in warm Mexico. Wilson Bulletin 109(3): 424-436.
or cold water to make soothing tea-like drinks that
can sooth sore throats (Bowers and Wignall 1993, Li, Xiaojie, J.M. Baskin, and C.C. Baskin. 1999.
Warnock 1974). Seed morphology and physical dormancy of
several North American Rhus species
References (Anacardiaceae). Seed Science Research 9(3):
247-258.
Baskin, C.C. and J.M. Baskin. 2001. Seeds:
Ecology, Biogeography, and Evolution of Martin, W.C. and C.E. Hutchins. 1980-1981
Dormancy, and Germination. Academic Press, (reprinted 2001). A Flora of New Mexico. 2
San Diego. 666 p. volumes. Bishen Singh Mahendra Pal Singh,
India and Koeltz Scientific Books, Germany.
Bowers, J.E. and B. Wignall. 1993. Shrubs and 2,591 p.
Trees of the Southwest Deserts. Southwest
Parks and Monuments Association, Tucson, AZ. McPherson, G.R., H.A. Wright, and D.B. Wester.
140 p. 1988. Patterns of shrub invasion in semiarid
Texas USA grasslands. American Midland
Carter, J.L. 1997. Trees and Shrubs of New Naturalist 120(2): 391-397.
Warnock, B.H. 1974. Wildflowers of the
Correll, D.S. and M.C. Johnston. 1970. Manual of Guadalupe Mountains and the Sand Dune
the Vascular Plants of Texas. Texas Research Country, Texas. Sul Ross State University,
Foundation, Renner, TX. 1,881 p. Alpine, TX. 176 p.

Vegetation: Past, Present and Future.
632
Whitford, Walter G. 2002. Ecology of Desert Young, J.A. and C.G. Young. 1992. Seeds of
Systems. Academic Press, San Diego, CA. Woody Plants in North America. Dioscorides
343 p. Press, T. R. Dudley, general editor, Portland,
OR. 407 p.
633
Rhus trilobata Nutt. oakleaf sumac
ANACARDIACEAE
Synonyms: Rhus aromatica Ait. var. trilobata (Nutt.) Gray

Toxicodendron trilobatum Kuntze
Schmaltzia oxyacanthoides Greene
Schmaltzia pubescens Osterh.
Nancy L. Shaw and Ann M. DeBolt
often treated as a variety of that more eastern

species.
Range.—Oakleaf sumac is a widespread and

highly variable taxon, ranging from Alberta to
Iowa and south to California, Mexico, and Texas
(Harrington 1964). It grows at elevations from
1,050 to 2,400 m on sites that receive 250 to 500
mm of precipitation (Hitchcock and others 1961,
USDA Forest Service 1937, Wasser 1982).
Ecology.—In the Intermountain Region, oakleaf

sumac is common in mountain brush and pinyon-
juniper communities (Hitchcock and others 1961,
Welch and others 1987) and also grows in
openings in communities of oakbrush, ponderosa
pine, and other conifers (Vories 1981). At lower
Illustration credit: E.G. Hurd
elevations it may be found in the blackbrush, basin
big sagebrush, or Wyoming big sagebrush zones.
General Description.—Also known as skunkbush In these areas it occurs on streambanks and
sumac or lemonade sumac, oakleaf sumac is a terraces, floodplains, seep, and spring margins,
morphologically variable deciduous species with a mesic slopes, talus slopes, swales, drainageways,
spreading, mound-like, or erect growth form. It and on other sites receiving extra moisture (Welsh
frequently forms clumps or thickets 0.6 to 2.1 m in and others 1987). It is frequently associated with
height (Hitchcock and others 1961, Thornburg riparian communities, growing intermixed with
1982, Welch and others 1987). Young branches other shrubs or as scattered thickets in
are brownish and puberulent, turning gray with communities dominated by herbaceous species.
age. The alternate, trifoliate leaves are glossy, dark Adapted to a wide range of soil conditions, oakleaf
green, and 3 to 7 cm long. Leaflet lobes are sumac grows on basic to acidic soils with textures
shallowly to deeply incised and puberulent below. ranging from clayey to sandy.
The terminal leaflet is fan-shaped; the lower two
are about half as large and shallowly crenate-lobed Reproduction.—Flowers appear in early spring,
(Hitchcock and others 1961). Plants are polygamo- usually March or April (Brinkman 1974). Fruits
dioecious (nearly dioecious, but with some ripen from June to September (Plummer and
male/female flowers). Inflorescences consist of others 1968, Shaw 1984, van Dersal 1938) but
clusters of spikes located near branch tips many remain on the shrub well into winter. Seeds
(Goodrich and Neese 1986). Flowers are are dispersed primarily by birds but also by
inconspicuous, pale-yellow and mostly imperfect, rodents and other animals (Brinkman 1974). There
developing before the leaves. Fruits are sticky, are approximately 44,000 cleaned seeds/kg
subglobose, orange-red drupes 6 to 8 mm in (Brinkman 1974, Plummer and others 1968).
diameter with a lemony flavor when fresh. This Germination is restricted by the presence of a hard
species closely resembles R. aromatica Ait. and is seedcoat and embryo dormancy, with the degree of
dormancy varying considerably among
populations. Seedcoat permeability of most
634
populations is improved by a 0.3 to 2 hour in the United States. Agriculture Handbook 450.
sulphuric acid scarification (Babb 1959, Brinkman U.S. Department of Agriculture, Forest Service,
1974, Glazebrook 1941, Heit 1968), while 30 to Washington, DC. p. 715-719.
120 days of moist prechilling is required to release
embryo dormancy (Babb 1959, Swingle 1939). Clark, J.W. and J.E. Depuit. 1981. Analysis of
Mechanically rupturing the seedcoat improves direct seeding methods for establishment of
germination (McKeever 1938), but techniques selected native shrub species on minesoils in
have not been developed for treating large seed southeastern Montana. In: L.H Stelter, E.J.
lots. Following fire, oakleaf sumac generally DePuit, and S.A. Mikol, tech. coords. Shrub
sprouts vigorously from the root crown and establishment on disturbed arid and semi-arid
rhizomes (Wasser 1982). lands. Wyoming Game and Fish Department,
Cheyenne, WY. p. 89-103.
Growth and Management.—Local seed sources
should be selected for range seedings or Glazebrook, T.B. 1941. Overcoming delayed
transplanting projects as the range of adaptability germination in the seed of plants valuable for
for most populations is not known (Swenson erosion control and wildlife utilization. Master’s
1957). Scarified seed may be fall planted to thesis, University of Idaho, Moscow, ID. 97 p.
provide overwinter moist prechilling. Seed should
be planted approximately 1.3 cm deep or slightly Goodrich, S. and E. Neese. 1986. Uinta Basin
deeper in dry, coarse textured soils. Seedlings are flora. U.S. Department of Agriculture, Forest
considered to be drought and cold hardy but may Service, Intermountain Region, Ogden, UT. 319
succumb to competition from herbaceous species p.
when young (Clark and Depuit 1981, Monsen
1987). Range or wildland seedings normally Harrington, H.D. 1964. Manual of the plants of
require 2 to 5 years for establishment (Plummer Colorado. The Swallow Press, Inc., Chicago, IL.
and others 1968). 666 p.
Benefits.—Oakleaf sumac can be used to stabilize Heit, C.E. 1968. Thirty-five years’ testing of tree
disturbed riparian areas. It provides excellent and shrub seed. Journal of Forestry 66: 632-634.
ground cover, and the spreading root system
reduces soil erosion. It is particularly useful for Hitchcock, C.L., A. Cronquist, M. Ownbey, and
restoration of meadows and other riparian sites J.W. Thomson. 1961. Vascular plants of the
where extensive erosion and gully cutting has Pacific Northwest. Part 3: Saxifragaceae to
lowered water tables. Oakleaf sumac thickets Ericaceae. University of Washington Press,
provide excellent cover for many small birds and Seattle, WA. 614 p.
mammals, and winter browse where shrubs are
limited. It is increasingly being used by LHBH [Liberty Hyde Bailey Hortorium]. 1976.
homeowners seeking drought-tolerant native Hortus third: a concise dictionary of plants
species since it is commercially available, easy to cultivated in the United States and Canada.
grow, long-lived, responds well to pruning, and Third Edition, Macmillan Publishing, New
has attractive red, orange to yellow fall color. York. 1,290 p.
Native Americans have used oakleaf sumac for
food, tobacco substitute, basketry, and as a McKeever, D.G. 1938. The effects of various
mordant in dyeing (LHBH 1976). methods of treatment on germination of seeds of
some plants valuable for game and erosion
References purposes. Master’s thesis, University of Idaho,
Moscow, ID. 128 p.
Babb, M.F. 1959. Propagation of woody plants by
seed. Bulletin 26. Alaska Agricultural Monsen, S.B. 1987. Shrub selections for pinyon-
Experiment Station. [Place of publication juniper plantings. In: R.L. Everett (comp.).
unknown]. 12 p. Pinyon-juniper conference. INT-GTR-215. U.S.
Brinkman, K.A. 1974. Rhus L. Sumac. In: C.S. Intermountain Research Station, Ogden, UT. p.
Schopmeyer, tech. coord. Seeds of woody plants 316-334.
635
Plummer, A.P., D.R. Christensen, and S.B. USDA Forest Service. 1937. Range plant
Monsen. 1968. Restoring big game range in Handbook. U.S. Department of Agriculture,
Utah. Publication 68-3. Utah Division of Fish Forest Service, Washington, DC. Republished
and Game, Salt Lake City, UT. 183 p. by Dover Publications, London. 816 p.
Shaw, N. 1984. Producing bareroot seedlings of van Dersal, W.R. 1938. Native woody plants of the
native shrubs. In: P.M. Murphy, comp. The United States: Their erosion-control and wildlife
challenge of producing native plants for the values. Miscellaneous Publication 303. U.S.
Intermountain area. INT-GTR-168. U.S. Department of Agriculture, Washington, DC.
Department of Agriculture, Forest Service, 302 p.
Intermountain Forest and Range Experiment
Station, Ogden, UT. p. 6-15. Vories, K.C. 1981. Growing Colorado plants from
seed: a state of the art. Vol. 1: Shrubs. INT-
Swenson, W.S. 1957. Squawbush in windbreaks in GTR-103. U.S. Department of Agriculture,
eastern Colorado. Journal of Soil and Water Forest Service, Intermountain Forest and Range
Conservation 12: 184-185. Experiment Station, Ogden, UT. 80 p.
Swingle, C.F. 1939. Seed propagation of trees, Wasser, C.H. 1982. Ecology and culture of
shrubs, and forbs for conservation planting. selected species useful in revegetating disturbed
SCS-TP-12. U.S. Department of Agriculture, lands in the West. FWS/OBS-82/56. U.S.
Soil Conservation Service, Washington, DC. Department of the Interior, Fish and Wildlife
198 p. Service, Office of Biological Services, Western
Energy and Land Use Team, Washington, DC.
Thornburg, A.A. 1982. Plant materials for use on 347 p.
surface mined land in arid regions. T.P. 157,
EPA 600-7-79-133. U.S. Department of Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
Agriculture, Soil Conservation Service, Higgins, eds. 1987. A Utah flora. Great Basin
Washington, DC. 88 p. Naturalist Memoir 9. Brigham Young
636
Ribes aureum Pursh golden currant
GROSSULARIACEAE
Synonyms: Chrysobotrya aurea (Pursh.) Rydb.
five petals. They are solitary or in erect clusters, or

racemes, of up to 15 flowers. The smooth, hairless,
spherical fruit is a juicy several-seeded berry
approximately 0.5 to 1.5 cm across and variously
colored from yellow, red, brown to black. This
description is after Carter 1997, Cronquist and
others 1997, Hitchcock and Cronquist 1973, and
Vines 1986. Kartesz (1994) and the U.S.
Department of Agriculture (2003) database
PLANTS recognize three varieties: the type
variety aureum, variety gracillimum (Coville and
Britt.) Jepson that is synonymous with Ribes
gracillimum, and variety villosum DC that is
synonymous with Ribes odoratum H. Wendl.
Despite some confusing references to the contrary,
Ribes aureum var. aureum is not a synonym of
Ribes odoratum. The likely source of the
taxonomic confusion is that Ribes odoratum H.
Wendl is synonymous with Ribes aureum Gray, as
opposed to the more familiar and taxonomically
accepted, Ribes aureum Pursh.
Illustration credit: J. Shoemaker in Wasser 1982
Range.—Golden currant, Ribes aureum var.
General Description.—Ribes aureum Pursh. is aureum, is a wide-ranging species across Western
commonly known as golden currant, but local North America (Cronquist and others 1997). It is
names also include buffalo, flowering, fragrant, common from Alberta, Canada, to western Texas
and clove currant (Elmore 1976). The species (Kershaw and others 1998, Powell 1998). Ribes
name “aureum” is derived from the Latin word for aureum var, odoratum is native to mid-America,
“golden-yellow” and refers to the rich-yellow west of the Great Lakes (Cronquist and others
colored flowers. In some areas flowers may also 1997). Both varieties have become naturalized in
be a cream or reddish color (Derig and Fuller Europe and possibly in other parts of the world
2001). The derivation of “Ribes” is apparently (Stace 1997, Cronquist and others 1997, Komarov
debatable. Gledhill (1992) suggests it is derived 1939).
from the Persian word for “acid-tasting” whereas
Vines (1986) indicates “Ribes” may be derived Ecology.—Golden currant grows along water
from an ancient Danish name. Another likely courses and in moist meadows in a variety of
derivation is a link to historical cultural use. In the vegetation communities from grasslands, shrub
Middle East, a popular syrup was made from an lands, piñon juniper woodland, western hardwood,
Oriental species of rhubarb (Rheum ribes) called pine, and fir-spruce forests (Marshall 1995, Dick-
“Ribas” (USDA FS 1988, ISTA 1968). In Western Peddie 1992, Welsh and others 1993). Common
Europe, currants were substituted for the rhubarb associates include rabbitbrush (Chrysothamnus
in this prized juice, and apparently this use species), maples (Acer species), serviceberry
influenced Linnaeus to adopt “Ribes” as the (Amelanchier species), chokecherry (Prunus
generic name for currants (USDA FS 1988). virginiana L.), juniper (Juniperus species),
Golden currant is a deciduous, unarmed (spineless) ponderosa pine (Pinus ponderosa P. & C.
shrub. It is usually from 60 cm to 3 m tall but may Lawson), willows (Salix species), and
grow to a little over 4 m. The light green, cottonwoods (Populus species). The elevation at
semiglossy leaves are usually three-lobed. The which it occurs tends to depend somewhat upon
flowers are large (up to 9 mm long) and each has latitude, but generally plants are found between
637
approximately 400 to 2,800 m (Cronquist and and plants can sprout after fire (Tykač 1990,
others 1997, Carter 1997, Powell 1998). Golden Marshal 1995). Tolerance to fire in the dormant
currant grows in full sunlight but also does well in state is good (Wasser 1982). However, the
partial shade (Tykač 1990). It is tolerant to a range severity, or heat, of the fire is likely to be
of weakly acid (pH 6) to weakly basic soils important to survival, and very hot fires can kill
(Wasser 1982). Bacterial diseases have been noted shrubs completely (Marshall 1995).
to infect canes of some genotypes, but resistant
varieties are available (Wasser 1982). Ribes Benefits.—Golden currant rates from “poor” and
species are intermittent hosts of the white pine “fair” to “medium” palatability for cattle (Dayton
blister rust (Cronartium ribicola), a fungal 1931, USDA FS 1988). It has good palatability for
pathogen that can be ruinous to most native white sheep (Dayton 1931). In Montana, twigs and
pine species. Because of this, Ribes species have foliage can make up to 2 to 5 percent of a sheep’s
been subject to specific eradication programs in diet (Martin and others 1951). The fruits are an
many white pine (Pinus monticola Dougl. ex D. important food source for a variety of wildlife
Don) regions (Martin and others 1951, Van such as songbirds, chipmunks, mice, and ground
Arsdel and others 1998). However, the squirrels (Martin and others 1951, Marshall 1995,
significance of specifically golden currant in the Tykač 1990). The fruits may contribute up to 10
life cycle and pathology of this rust is uncertain. percent of a chipmunk and ground squirrels’ diet
Throughout the Sacramento Mountains of New (Martin and others 1951). Both fruit and foliage
Mexico, R. pinetorum Greene was a major are used by a range of species that include grouse,
contributor to the spread of rust to nearby pines, coyote, and beaver (Martin and others 1951). Elk
but blister rust was not found on golden currant have been reported to eat the foliage in Colorado,
that generally grew below the elevation of the and one report indicated that mule deer browsed it
pines (Van Arsdel and others 1998). infrequently, but cattle not at all, during a 1-year
study in Montana (Dusek 1975). Golden currant
Reproduction.—Golden currant generally flowers has been planted for erosion control and in
in June through August, although flowers may be restoration projects (Vines 1986, Wasser 1982). It
found from March until the first frost (Epple 1995, is particularly used for restoration of game habitat
McGregor and others 1986, USDA FS 1988, Vines (Wasser 1982). When golden currant is used in
1986). The fragrant flowers are insect pollinated. vegetation restoration projects it is usually a small
Some insects have adapted to the long narrow component of the seed mix and used at
flower tube by boring a hole at its base to “steal” approximately 461 to 922 grams per hectare
the nectar (Derig and Fuller 2001). Golden currant (Wasser 1982). The berries, both fresh, dried, and
readily crosses with other Ribes species (Wasser made into jams and cakes, have been used for food
1982). The fruits mature in summer and fall. The by many Native American tribes as well as by
seed is a densely hairy achene. The cleaned seed European settlers (Elmore 1976, Moerman 1998,
averages approximately 478,398 to 513,672 Welsh and others 1993). It has also been used as a
individuals per kg with a germination rate of 60 to pharmaceutical by some of the tribes of the Great
98 percent (Vines 1986, Wasser 1982). The seeds Plains. The dried, pulverized inner bark was
undergo a physiological dormancy that is broken sprinkled on sores, used with other ingredients in a
by 60 days of cold stratification (Baskin and poultice, and a decoction was taken for leg
Baskin 2001). Germination is epigeal. The plant swelling (Moerman 1998). The flowers are edible,
also spreads by growing shoots from rhizomes apparently flavorful, and eaten by gourmets
(Wasser 1982). (Elmore 1976). Golden currant is a popular
ornamental plant in North America, Europe, and
Growth and Management.—Golden currant is parts of the former Soviet Union (Bailey and
easily propagated from hardwood cuttings in June others 1976, Komarov 1939, Tykač 1990).
or September and bare rooted plants are best
planted when leafless in the spring or fall (Tykač References
1990). When starting plants from seed, fall
seeding is preferred, but stratified seed can be Bailey, L.H., E.Z. Bailey, and the staff of the
planted in the spring (Wasser 1982). Golden Liberty Hyde Bailey Hortorium. 1976. Hortus
currant is used as a rootstock for grafting on other Third–a concise dictionary of plants cultivated
species of currant that root poorly (Tykač 1990). in the United States and Canada. Macmillian
Cutting back old shoots stimulates new growth, Publishing Company, New York. 1,290 p.
638
Baskin, C.C. and J.M. Baskin. 2001. Seeds, Kartesz, J.T. 1994. A synonymized checklist of the
ecology, biogeography, and evolution of vascular flora of the United States, Canada and
dormancy and germination. Academic Press, Greenland. Vol.1 - Checklist. 2nd Ed. Timber
New York. 666 p. Press, Portland, OR. 622 p.
Carter, J.L. 1997. Trees and shrubs of New Kershaw, L., A. MacKinnon, and J. Pojar. 1998.
Mexico. Johnson Books, Boulder, CO. 534 p. Plants of the Rocky Mountains. Lone Pine
Publishing, Edmonton, Canada. 383 p.
Cronquist, A., N.H. Holmgren, and P.K.
Holmgren. 1997. Intermountain flora, vascular Marshall, K.A. 1995. Ribes aureum. In:
plants of the Intermountain West, U.S.A. Vol. Department of Agriculture, Forest Service,
3, Part A. New York Botanical Garden, Bronx, Rocky Mountain Research Station, Fire
NY. 446 p. Sciences Laboratory. Fire Effects Information
System, http://www.fs.fed.us/ database/feis/.
Dayton, W. A. 1931. Important western browse 13 p.
plants. Miscellaneous Publication 101. U.S.
Department of Agriculture, Washington DC. Martin, A C., Zim, H. S., and Nelson, A.L. 1951.
214 p American wildlife and plants. 2nd printing.
McGraw-Hill Book Company, Inc., New York.
Derig, B.B. and M.C. Fuller. 2001. Wild berries of 500 p.
the West. Mountain Press Publishing Company,
Missoula, MT. 235 p. McGregor, R. L., T.M. Barkley, R.E. Brooks, and
E.K. Schofield. 1986. Flora of the Great Plains.
Dick-Peddie, W.A. 1992. New Mexico University Press of Kansas, Lawrence, KS.
Vegetation– past, present, and future. 1,402 p.
University of New Mexico Press, Albuquerque,
NM. 244 p. Moerman, D.E. 1998. Native American ethno-
Dusek, G.L. 1975. Range relations of mule deer
and cattle in prairie habitat. Journal of Range Powell, A.M. 1998. Trees and shrubs of Trans-
Management 39(3): 605-615. Pecos and adjacent areas. University of Texas
University Press, Austin, TX. 498 p.
Elmore. F.H. 1976. Shrubs and Trees of the
Southwest Uplands. Southwest Parks and Stace, C. 1997. New Flora of the British Isles. 2nd
Monuments Assoc. Tucson, AZ. 214 p. Ed. Cambridge University Press, New York.
1,130 p.
Epple, A.O. 1995. A field guide to the plants of
Arizona. Falcon Press Publishing Co., Helena, Komarov. B.K. 1939. Flora SSSR. Vol. 9. Flora of
MT. 347 p. the USSR translated 1964 by the Israel Program
for Scientific Translations, Jerusalem, Israel. p.
Gledhill, D. 1992. The names of plants. 2nd 205
edition, reprinted 1992. Cambridge University
Press, Cambridge, UK. 202 p. Tykač, J. 1990. The illustrated guide to ornamental
shrubs. Treasure Press, London, UK. 224 p.
Hitchcock, C.L. and A. Cronquist. 1973. Flora of
the Pacific Northwest. University of U.S. Department of Agriculture. 2003. Plants
Washington Press, Seattle, WA. 730 p. database. http://plants.usda.gov/cgi _bin/plant_
profile.cgi?symbol=FAPA# [not paged].
International Seed Testing Association (ISTA).
1968. A multilingual glossary of common United States Department of Agriculture Forest
plant-names 1. Field crops, grasses and Service (USDA FS). 1988. Range Plant
vegetables. International Seed Testing Handbook. Dover Publications, Inc. New York.
Association, Wageningen, The Netherlands. 838 p.
371 p.
639
Van Arsdel, E.P., D.A. Conklin, J.B. Popp, and Wasser, C.H. 1982. Ecology and culture of
B.W. Geils. 1998. The distribution of white selected species useful in revegetating disturbed
pine blister rust in the Sacramento Mountains of lands in the West. FWS?OBS-82/56. Fish and
New Mexico. Finnish Forest Research Institute Wildlife Service, U.S. Department of the
Research Papers 712: 275-283. Interior, Washington, DC. 347 p.
Vines, R.A. 1986. Trees, shrubs, and woody vines Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
of the Southwest. University of Texas Press. Higgins. 1993. A Utah flora. 2nd Edition,
Austin, TX. 1,104 p. Revised. Brigham Young University, Provo,
UT. 986 p.
640
Ribes cereum Dougl. wax current
GROSSULARIACEAE
Synonyms: Ribes inebrians Lindl.

Ribes cereum Dougl. var. inebrians (Lindl.) C.L. Hitchc.
John K. Francis
Range.—Wax current is native to the states of

Washington, Oregon, California, Nevada, Utah,
Arizona, New Mexico, Oklahoma, Idaho,
Montana, Wyoming, North Dakota, South Dakota,
and Nebraska, as well as the provinces of British
Columbia and Alberta. There are three recognized
varieties. Variety cerium is apparently found in all
the range except Oklahoma. Variety colubrinum
C.L. Hitchc. occurs in Washington, Oregon, Idaho,
(and probably British Columbia). Variety
pedicellare Brewer & S. Wats., known as whisky
current, occupies the range east and south of
Washington and Oregon (Natural Resources
Conservation Service 2003).
Ecology.—Wax current is most common and

grows best in the open but it can grow under open
conifer forest canopies. It therefore classifies as
intolerant of shade. New seedlings appear after
fires (that may scarify seeds) and when the soil is
disturbed and opened to sunlight (Marshall 1995).
The species grows on well-drained soils of the full
range of textures that have developed from a wide
Drawing source: Britton and Brown 1913 variety of parent materials. These soils are often
rocky or gravelly. Wax current grows at elevations
General Description.—Wax current, also known of 1,500 to 4,000 m (Marshall 1995) in areas with
as squaw current, is a deciduous shrub 0.5 to 1.5 m warm, dry to moist summers and cool to cold
in height. Plants support one to several stems and winters. Minimum annual precipitation is around
are unarmed, dark gray to brown, sticky, and hairy 330 mm/year. Wax current occurs in many
when young, becoming glabrous with age. The vegetation types including open and broken
yellow-green to grayish-green, simple leaves are coniferous forests, exposed subalpine slopes and
circular to kidney-shaped with three to seven ridges, and sagebrush (Artemisia spp.) steps
shallow lobes and a fine toothed or rounded- (author’s observation).
toothed margin. Petioles are 0.4 to 2.2 cm long and
blades are 0.5 to 2.5 long and 0.7 to 3.0 cm broad. Reproduction.—Wax current flowers between
Inflorescences are two- to four-flowered racemes April and June and ripens fruit in August (Pfister
that are shorter than the leaves. The 4- to 7-mm 1974). Flowers are pollinated by insects (Plants
long, glandular, tubular, greenish-white to pale For a Future 2003). There was an average of
pink flowers have five tiny petals. Fruits (berries) 553,000 seeds/kg from fruits collected in
are red, shiny, and 6 to 8 mm in diameter. Each California (Pfister 1974). Seeds are dispersed by
fruit contains several rough-surfaced, elliptical birds and small mammals, as well as by gravity
seeds about 2.5 mm long. There are 2n = 16 (Marshall 1995). At 21 °C, air-dried seeds retained
chromosomes (Abrams 1944, Brayshaw 1996, some viability for 27 years (Pfister 1974).
Mesler and Sawyer 2003, Pfister 1974, Welsh and Reproduction is mainly by seed; the species has
others 1987). only a “weak” ability to sprout from the root
crown (Marshall 1995).
641
Growth and Management.—Fruits must Brayshaw, T.C. 1996. Trees and shrubs of British
generally be picked by hand. Seeds are cleaned of Columbia. UBC Press, Vancouver, B.C.,
fruit by maceration, and air dried. They can be Canada. 373 p.
stored in sealed containers at ambient temperatures
or refrigerated (Pfister 1974). Seeds require 4 to 5 Britton, N.L., and A. Brown. 1913. Illustrated flora
months of cold stratification (at temperatures near of the northern states, Canada, and the British
freezing) after which they can be planted in the possessions. Vol. 2, 2nd Ed. Scribner, New York.
spring (Marshall 1995). Alternately, they can be 735 p.
sown in the fall and allowed to stratify naturally.
Plants can also be started from cuttings of semi- Marshall, K.A. 1995. Ribes cereum. In: U.S.
hardened wood (Plants For a Future 2003). Adult Department of Agriculture, Forest Service,
plants can be eliminated from stands by fire, Rocky Mountain Research Station, Fire Sciences
grubbing, and probably herbicides. However, in at Laboratory, Fire Effects Information System.
least one study, because many new seedlings http://www.fs.fed.us/database/feis/plants/shurb/
appeared, the number of plants increased in sites ribcer/all.html. 14 p.
disturbed by treatments (Marshall 1995). Because
wax current does poorly in shade, tree planting is Mesler, M.R. and J.O. Sawyer, Jr. 2003.
suggested as a long-term solution. Grossulariaceae, gooseberry family. http://
ucjeps.berkeley.edu/chi-bin/get_JM_treatment.
Benefits.—Wax current contributes to the beauty pl?4451,4452,4463. 3 p.
of forest lands where it grows and helps protect the
soil from erosion. Shade from their crowns shelters Natural Resources Conservation Service. 2003.
conifer seedlings and thus aids forest succession Plants profile: Ribes cereum Doul.
(Marshall 1995). The species is used to a limited http://plants.usda.gov/cgi_bin/plant_profile.chi?
extent as a foundation plant in formal landscaping symbol=RICE. 3 p.
(Wildland Nursery 2003). Wax current is fair to
poor browse for domestic livestock and wild Oehlke, B. 2003. Agapema homogena moth, Dyar,
ungulates and is mainly eaten when little else is 1908. http://www3.islandtelecom.com/
available. New annual growth contains about 5.6 ~oehlkew/sahomoge.htm. 4 p.
percent protein. Several species of birds and
rodents consume the fruits (Marshall 1995). The Pfister, R.D. 1974. Ribes L., current, gooseberry.
larva of the Rocky Mountain Agepema moth In: C.S. Schopmeyer, tech. coord. Seeds of
(Agapema homogena Dyar) feed on the foliage woody plants in the United States. Agriculture
(Oehlke 2003). Wax current berries are edible but Handbook 450. U.S. Department of Agriculture,
have little flavor. They are picked for making Forest Service, Washington, DC. p. 720-727.
jams, jellies, and pies and were formerly used by
Native Americans for making pemmican (Marshall Plants For a Future. 2003. Database search results:
1995). People of the Secwepemc tribe of British Ribes cereum. http://www.ibiblio.org/pfaf/cgi-
Columbia ate them to relieve diarrhea (Secwepemc bin/arr_html?Ribes+cereum&CAN=LATIND.
Nation 2003). An infusion of inner bark was used 5 p.
as a wash for sore eyes (Plants For a Future 2003).
Wax current and other Ribes species are alternate Secwepemc Nation. 2003. Secwepemc
hosts for white pine blister rust (Cronartium ethnobotanical garden: other plants. Sccwepemc
ribicola), which can be lethal to five-needle pines. Nation, Kamloops, BC, Canada. 4 p.
Efforts to eradicate Ribes from several forest areas
failed and did not decrease the incidence of blister Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
rust (Marshall 1995). Higgins. 1987. A flora of Utah. Great Basin
References University Press, Provo, UT. 894 p.
Abrams, L. 1944. Illustrated flora of the Pacific Wildland Nursery. 2003. Wildland nursery:
States. Vol. 2. Stanford University Press, shrubs. Wildland Nursery, Joseph, UT.
Stanford, CA. 635 p. http://www.wildlandnursery.com/shrubs/shrubs
M-Z.html. 6 p.
642
Ricinus communis L. castor bean
EUPHORBIACEAE
Synonyms: none
John K. Francis
found growing naturally and in cultivation as an
ornamental throughout the tropical and subtropical
areas of the United States and its territories.
Ecology.—Disturbance is required for successful

natural stands of castor bean. The species is an
intolerant pioneer. If disturbance is not repeated, it
will be succeeded in a few years by grass, vines, or
trees. Castor bean is competitive and most
frequently seen in flood zones, on neglected
farmland, and roadsides. In Puerto Rico, wild
stands rarely succeed on exposed subsoils and
highly weathered soils such as Ultisols. In
cultivation, where excessive vegetative growth is a
detriment, castor beans are often grown on
marginal agricultural soils (CSIR 1972). Castor
bean is not frost hardy. Many pests and diseases
are known worldwide, but they rarely present a
serious threat to cultivated or wild stands.
Reproduction.—Castor bean flower clusters grow

erect at the ends of twigs. Male flowers form on the
lower part and female flowers on the upper part of
each raceme. The seed capsules are covered with
weak spines (spineless types exist). The capsules
General Description.The castor bean, also
split into three parts, each containing one seed. The
known as the castor-oil plant, higüerito, higüerillo,
seeds are shiny brown with darker streaks or spots
palma Christi, carrapateira, ricin commún, and
and resemble a bloated tick (Little and others
many other common names, is grown as an annual
1974). Two seed sizes are reported in India: 1,164
herb in temperate countries but grows to treelike
seeds/kg and 2,610 seeds/kg (Kadambi and Dabral
proportions on fertile ground in tropical climates.
1955). Stored seeds should be kept cool. Seeds
The plant usually has only one stem arising from
stored at ambient temperature lose 75 percent of
the soil but develops a number of low robust
their viability within 3 months (CISR 1972).
branches when open grown. Castor bean is
Soaking the seeds for 12 hours prior to sowing is
herbaceous when young but becomes woody with
reported to improve germination (CISR 1972).
age. The wood is soft and light with a thick central
Germination starts in 7 to 10 days and is complete
pith. Occasionally, irregular brown heartwood
in 25 to 45 days (Kadambi and Dabral 1955).
develops (Kadambi and Dabral 1955). The bark is
There are hundreds of natural forms and cultivated
light brown, smooth, and exhibits rings at the
varieties of the species, including annual and
nodes and raised lenticels. There are a moderate
perennial types.
number of large, star-shaped leaves with 7 to 9
long pointed lobes (Little and others 1974). Castor
bean has been cultivated in Africa and Asia from Growth and Management.—Castor beans are
ancient times and used medicinally (CISR 1972). planted in rows spaced from 1 to 2 m apart with
spacing within the rows of about 0.5 m. When
Range.—Castor bean is a native of tropical Africa grown as an annual crop, it takes 5 to 9 months
but has naturalized in moist tropical and from planting to harvest (CSIR 1972). Castor beans
subtropical regions throughout the world. It may be are reported to survive for 8 to 10 years in India
643
and may reach 6 m in height (Kadambi and Dabral and horses are especially vulnerable. Fatal doses
1955). Annual varieties reach 1 to 2 m in height. are from 2.5 to 6 seeds for humans and about 6
Seed yields under cultivation vary from 200 to seeds for horses (CISR 1972). Symptoms are
1,700 kg/ha, depending on variety and site quality stomach irritation, diarrhea, abdominal pain,
(CISR 1972). increased heart rate, profuse sweating, collapse,
and convulsions. Broken seeds can cause skin
Benefits.—Castor beans contain from 31 to 61 irritation. The foliage is only slightly toxic
percent oil (CISR 1972). After decorticating, they (Anonymous 2000). It is advisable to completely
are subject to a series of hot or cold presses eliminate castor bean from pastures, especially
followed by solvent extraction, each step yielding a horse pastures, and pinch off flowers of ornamental
different grade of oil (Kirschenbauer 1960). The plants to prevent possible poisoning of children.
largest producers are Brazil and India. The
principal consumer is the United States References
(Encyclopedia Brittanica 2000), but an annual
demand for 100,000 tons of castor oil is reported Anonymous. 2000. Castorbean, castor oil plant.
for Europe (NF-2000 Database 2000). The seed http://www.vet.perdue.edu/depts/addl/toxic/
cake remaining after extracting the oil is used as plant11. htm. 1 p.
fertilizer or cooked to destroy the toxin and
incorporated into animal feeds. Not as popular as it CSIR. 1972. The wealth of India. Raw materials.
once was, castor oil is still widely used in Vol. 9. Publications & Information Directorate,
traditional and herbal medicine, especially in less Council for Scientific and Industrial Research,
developed countries. Its principal use in medicine New Delhi. 472 p.
is as a purgative and laxative. Castor oil is also
used as a lubricant, lamp fuel, a component of Encyclopedia Britanica. 2000. Castor oil.
cosmetics, and in the manufacture of soaps, http://www.Britanica.com/bcom/eb/article/4/0,57
printer’s ink, plastics, fibers, hydraulic fluid, break 16,2105+1+20724,00htm?query=castoroil%20
fluid, varnishes, paints, embalming fluid, textile oil. [not paged].
dyes, leather finishes, adhesives, waxes, and
fungicides (Encyclopedia Brittanica 2000, CISR Kadambi, K. and S.N. Dabral. 1955. The
1972). It is gradually being replaced as a raw silviculture of Ricinus communis Linn. Indian
material for some of the uses by petroleum-based Forester 81(1): 53-58.
products. In India, the leaves are used as food for
eri silk worms (Kadambi and Dabral 1955). The Kirschenbauer, H.G. 1960. Fats and oils, an
stalks from fields are burned for fuel in India and outline of their chemistry and technology.
have been shown to be suitable for short-fiber pulp Reinhold Publishing Corporation, New York.
(Kadambi and Dabral 1955). The species has been 240 p.
planted for dune stabilization (CISR 1972). Castor
bean is widely planted as an ornamental. Its large, Little, E.L., Jr., R.O. Woodbury, and F.H.
star-shaped leaves make it a bold foliage plant. Wadsworth. 1974. Trees of Puerto Rico and the
Some varieties have red- or purplish-colored leaves Virgin Islands. Vol. 2. Agriculture Handbook.
and stems. 449. U.S. Department of Agriculture, Forest
Service, Washington, DC. 1,024 p.
Detrimental Effects.—Castor bean may become a
weed in neglected cropland and pasture. It is not NF-2000 Online Database Information Items.
difficult to control through cultivation and mowing. 2000. Crops: castor oil (Ricinus communis).
Of greater concern than its weedy potential is the http://www.nf-2000.org/secure/Crops/S600.htm
high toxicity of its seeds, which contain ricin, a [not paged].
water-soluble protein. Even a small amount of
masticated seed is likely to cause death. Humans
644
Rondeletia inermis (Spreng.) Krug & Urban cordobancillo
RUBIACEAE
Synonyms: Catesbaea inermis Spreng.

Rondeletia inermis angustifolia Krug & Urban
Rondeletia inermis oblongifolia Krug & Urban
Rondeletia inermis intermedia Krug & Urban
Rondeletia inermis latifolia Krug & Urban
John K. Francis
small, yellow (or white to purple or brown)

flowers with four corolla lobes are borne on
peduncles 0.5 to 3.0 cm long. The fruit is a two-
celled, 3- to 4-mm, globose capsule containing
tiny, wrinkled, brown seeds (author’s observations,
Liogier 1997, Little and others 1974).
Range.—Cordobancillo is endemic to Puerto Rico

and its offshore island, Muertos (Little and others
1974). There are several races differing in size and
shape of leaves and amount of pubescence (Liogier
1997). The species is not known to have been
Ecology.—The species is intermediate in tolerance

to shade. It usually grows in the understory of
open to moderately dense forests, or in small
openings or edges. Cordobancillo may be found in
both primary remnants and secondary forests. It
invades disturbed areas but takes many years to do
so. Cordobancillo grows in a wide range of well-
drained, neutral to mildly acidic soils derived from
metamorphic (including ultramafic) rocks. It
grows from a few meters above sea level to
General Description.—Cordobancillo is an elevations of 1,000 m (Little and others 1974) in
upright evergreen shrub or occasionally a small areas that receive from about 750 to 2500 mm of
tree sometimes reaching 4.5 m in height and 7.5 mean annual precipitation.
cm in stem diameter. The plant occasionally takes
a vine-like form. Older plants usually have Reproduction.—Cordobancillo flowers and fruits
multiple stems arising at or near ground level. throughout the year. A collection of air-dried seeds
Stems and branches are slender, mostly straight. from the moist limestone hills near Bayamon,
The bark is gray or dark brown and nearly smooth. Puerto Rico averaged 1,863,000 seeds/kg. Seed
Inner bark is green. The wood is hard or dispersal is probably by gravity and wind.
moderately hard and stiff; sapwood is yellowish Collecting seeds is difficult because the capsules
and heartwood is light brown. Cordobancillo ripen, dry, and open a few at a time and
plants are supported by a tap and lateral root immediately release their tiny seeds.
system of stiff, brown roots. Foliage is mostly
grouped at the ends of twigs. The opposite leaves Growth and Management.—Cordobancillo
are variable: with or without hairs, cartaceous to appears to have a slow to moderate growth rate
coriaceous, linear to elliptic, and 1 to 10 cm long. and be relatively long-lived. By sprouting from the
There are deltoid stipules about 2 mm long and base and root crown, plants live longer than
petioles 3 to 6 mm long. Groups of one to five individual stems. There is no published experience
645
on plantation establishment or management of References
natural stands.
Benefits.—Cordobancillo helps protect the soil, Rico and adjacent islands. Vol. 5. Editorial de la
furnishes cover for wildlife, and adds to the Universidad de Puerto Rico, Río Piedras, PR.
aesthetics of Puerto Rican forests. Too small for 436 p.
lumber or poles, the wood is occasionally used for
fuel. Little, E.L., Jr., R.O. Woodbury, and F.H.
646
Rondeletia pilosa Sw. cordobancillo peludo
RUBIACEAE
Synonyms: Rondeletia triflora Vahl

Oldenlandia longiflora Lam.
Hedyotis longiflora (Lam.) Spreng.
John K. Francis
capsules produce many minute brown seeds

(Liogier 1997, Little and others 1974).
Range.—Cordobancillo peludo is native to Puerto

Rico, the Virgin Islands, and Monsarrat (Liogier
Ecology.—Cordobancillo peludo occurs in areas

that receive from about 750 to about 1000 mm of
mean annual precipitation from near sea level to
about 450 m in elevation. Soil texture does not
appear to be a limiting factor. All are well- to
excessively-well drained. Soil pH’s range from
about 5.5 to 8.0. Parent materials may be
metamorphic (including ultramaphic) rocks. It is
moderately intolerant and grows in open areas as
well as under low basal-area low forest. Because
the species is not highly competitive,
cordobancillo peludo is more likely to survive
where growth otherwise is limited—such as very
rocky terrain. It is scattered to rare in coastal
General Description.—Cordobancillo peludo, thickets, and remnant and middle to late secondary
also known as cachimbo peludo and lechecillo, is forests in upland areas.
an evergreen shrub or small tree commonly 1.5 to
2 m in height and 2 cm in basal diameter but Reproduction.—Cordobancillo peludo appears to
sometimes reaching 5 m in height. It may have flower and fruit throughout the year. In terms of
several stems from the base. The form is usually numbers, production of seeds is huge. The seeds
upright but occasionally vine-like. Stem bark is are dispersed by the wind. However, seedlings are
gray and smooth. The plants examined by the uncommon. The seeds number a few million/kg.
author had a tan colored, robust, shallow lateral Of a group of seeds sown on moist filter paper, 69
root system with sparse secondary and tertiary percent germinated between 15 and 70 days after
roots. The twigs are long, slender, forking, and sowing. Germination is epigeal.
ringed at the nodes. Soft hairs cover the twigs and
lower surfaces of the leaves. The leaves are Growth and Management.—Growth of
opposite, with entire, elliptic blades 4 to 10 cm cordobancillo peludo is slow. Basal sprouts in one
long and 2 to 3.5 cm broad, and pointed at both stand grew about 30 cm/year. One plant with a
ends, and have a petiole 3 to 6 mm long. 2.5-cm basal diameter had 10 growth rings.
Inflorescences are axillary and usually have three Individual stems probably last 10 to 20 years, and
flowers with peduncles up to 7 cm long. The plants can prolong their lives by coppicing. No
flowers have four-lobbed hairy calyxes and planting or management experience is known to
corollas. The corollas are white with red-tipped the author. Probably the best management strategy
lobes. The two-celled capsules are rounded, 3 mm is to protect remnant stands and late-secondary
in diameter, with calyx lobes persistent. The forests containing the species.
647
Benefits.—Cordobancillo peludo contributes to Universidad de Puerto Rico, Río Piedras, PR.
the aesthetics of the forest, it helps protect the soil, 436 p.
and furnishes cover for wildlife. The wood is
useful for small-diameter fuel. Little, E.L., Jr., R.O. Woodbury, and F.H.
Liogier, H.A. 1997. Descriptive flora of Puerto Washington, DC. 1,024 p.
648
Rosa woodsii Lindl. Wood’s rose
ROSACEAE
Synonyms: Rosa fendleri Crep
Rosa californica Watson.
Rosa macounii Greene
Rosa chrysocarpa Rydb.
Bruce L. Welch
constriction (Cronquist and others 1997). The

sepals are erect or spreading in the fruit (Blauer
and others 1975). Petals, five in all, are 1 to 2.5 cm
long (Cronquist and others 1997, Welsh and others
1987). Hips (fruits) are red to purple or nearly
black in color and ellipsoid to subglobose in shape
(Cronquist and others 1997, Welsh and others
1987). Hip size varies from 6 to 15 mm wide and
up to 2.0 cm in length and contains from 15 to 30
achenes (Blauer and others 1975). Each achene is
3 to 4 mm long with stiff hairs along one side.
Achenes weigh from 0.007 to 0.013 g (Gill and
Pogge 1974). The diploid chromosome number of
Wood’s rose is 14 (Blauer and others 1975).
Taxonomy.—Two varieties of Wood’s rose are

recognized: var. ultramontana and var. woodsii
(Blauer and others 1975, Cronquist and other
1997, Welsh and others 1987). Variety
ultramontana is the most prevalent in the
General Description.—Wood’s rose is a prostrate Intermountain area of Western United States.
to upright shrub from 0.5 to 3.0 m tall (Blauer and Woodsii is found on the prairies and plains of
others 1975). It is a strongly armed to nearly central North America and is smaller and stiffer
unarmed shrub with well-developed, straight, or than var. utlramontana with smaller leaflets that
less often, somewhat curved infrastipular prickles, often have gland-tipped teeth (Blauer and others
and often with other stout or weak prickles 1975, Cronquist and others 1997, Welsh and
(Cronquist and others 1997). The opposite others 1987).
compound leaves are composed of five to 11
leaflets, which are oval, elliptic, or obovate in Range.—Wood’s rose occurs from Minnesota
shape, with serrate margins (Blauer and others west and northwest to Alaska and British
1975). Leaflets are 2 to 5 cm long and 1 to 2.5 cm Columbia, south to Arizona, northern Mexico and
wide; the teeth are not gland-tipped except in var. western Texas and north to western Kansas and
woodsii. Flowers are light-pink to deep-rose color North Dakota. It is the most widespread native
and are borne on lateral shoots from old wood, rose in Alberta (Blauer and others 1975, Cronquist
which are usually over 10 cm long (Blauer and and others 1997, Pacific Southwest Experiment
others 1975). Flowers are solitary or two to several Station 2002). It grows in 24 ecosystems and
in small corymbiform cymes that terminate the occurs in 51 of Kuchler’s plant associations
lateral branches of the season and are relatively (Pacific Southwest Experiment Station. 2002).
small (Cronquist and others 1997). The The Society of American Foresters list Wood’s
hypanthium is glabrous and 3 to 5 mm thick at rose as occurring in 55 of its cover types (Pacific
anthesis (Cronquist and others 1997). Pedicels are Southwest Experiment Station 2002).
glabrous (Cronquist and others 1997). The five
persistent sepals are from 1 to 2 cm long and 2 to Ecology.—Wood’s rose can grow at elevations
3.5 mm wide at the base. The tip of each sepal is reaching 3,549 m (Pacific Southwest Experiment
usually a little expanded above the median Station 2002). It occurs on bluffs, dry grassy
649
slopes, and on sandhills throughout the prairies and food for a number of wildlife species, and
and on riverbanks and clearings in boreal and watershed protection. Fruits, leaves, or twigs are
subalpine forests. It can be found on the eastern eaten by a number of birds and mammals. These
slopes of the Rocky Mountains and throughout the include pronghorn, elk, mule deer, white-tailed
Great Basin. It is found along roadsides and deer, porcupines, beavers, squirrels, coyotes, bear,
south-facing cutbanks (Pacific Southwest small nongame birds and upland game birds
Experiment Station. 2002). In the Great Basin, (Pacific Southwest Experiment Station 2002).
Wood’s rose prefers moist sites along streams, in Livestock also eat Wood’s rose. Crude protein
seepage areas along fences, irrigation canals, content of Wood’s rose leaves vary from 5.7 (fall)
marsh lands, lake shores, and hillsides in to 16.4 (spring) percent of dry matter and stems
palustrine and lacustrine habitats (Welsh and from 5.4 (winter) to 12.0 (spring) percent (Dietz
others 1987). When conditions are favorable, 1972). Phosphorus content of leaves varied from
nearly impenetrable thickets of Wood’s rose are 0.29 (fall) to 0.48 (spring) percent of dry matter
formed along some mountain streams (Pacific with stems varying from 0.12 (winter) to 0.32
Southwest Experiment Station 2002). It flourishes (spring) percent (Dietz 1972). In vitro digestible
in moderate shade to full sunlight and so makes for Wood’s rose was 65.9 percent of dry matter
good growth on all aspects (Blauer and other digested for spring tissues and 54.5 percent for
1975). Wood’s rose is adapted to a wide range of winter tissues (Dietz 1972). These in vitro
soil types and textures (Pacific Southwest digestible values would rank Wood’s rose as a
Experiment Station 2002). Growth is generally highly digestible spring and winter forage (Welch
best on moderately fertile, well-drained clay loam, 1989, Welch and Andrus 1977). The dense
sandy loam, or sandy soils. It is tolerant of thickets formed by Wood’s rose are used for
moderately acid to weakly basic soils. (Pacific nesting and escape cover by many birds and small
Southwest Experiment Station 2002). It is seldom mammals (Pacific Southwest Experiment Station
found where the average annual precipitation is 2002). The extensive rhizomes and good
less than 260 mm (Blauer and others 1975). survivability and revegetation characteristics
makes this species an effective tool in erosion
Reproduction.—Seeds are produced on 2 to 5 control (Pacific Southwest Experiment Station
year old plants. Seeds of Wood’s rose are ingested 2002). Good results have been obtained with
with the hip and dispersed in the dropping of birds rooting of hardwood cuttings, direct seeding, and
and mammals. The seeds have a seed coat transplanting trials (Pacific Southwest Experiment
dormancy and require warm or cold stratification. Station 2002). Native Americans made extensive
Seed viability ranged from 44 to 64 percent for use of Wood’s rose roots, stems, leaves, flowers,
seed collected in northern Alberta. The seeds and hips for foods and therapeutic purposes. The
remain viable for 2 to 5 years. Seeds will hips are a source of vitamin C and are dried for
germinate within 30 to 40 days (Blauer and others use in flavoring teas, jellies, fruitcakes, and
1975, Gill and Pogge 1974, Pacific Southwest puddings. Wood’s rose is used as an ornamental
Experiment Station 2002). Wood’s rose spreads near homes to attract birds and other wildlife
vegetatively through underground rhizomes, (Pacific Southwest Experiment Station 2002).
sprouting from the root crown, and by layering
(Blauer and other 1975, Pacific Southwest References
Experiment Station 2002).
Blauer, A.C., A.P. Plummer, E.D. McArthur, R.
Management.—Wood’s rose has a fairly high Stevens, and B.C. Giunta. 1975. Characteristics
tolerance to browsing (Pacific Southwest and hybridization of important intermountain
Experiment Station 2002). It is susceptible to shrubs. I. Rose family. Research Paper INT-
various leaf spots, as well as to leaf rusts, gray 169. U.S. Department of Agriculture, Forest
mold, powdery mildew, common gall, and stem Service, Intermountain Forest and Range
cankers (Pacific Southwest Experiment Station Experiment Station, Ogden, UT. 36 p.
2002). The principal forest insect pests are tent
caterpillar, rose leaf hopper, and a subspecies of Cronquist, A., N.H. Holmgren, and P.K.
western tussock moth (Pacific Southwest Holmgren. 1997. Intermountain flora: vascular
Experiment Station 2002). plants of the Intermountain west, U.S.A. Vol. 3.
Part A; Subclass Rosidae (except Fabales). The
Benefits.—Wood’s rose adds to the biodiversity New York Botanical Garden., New York. 446 p.
of a multitude of ecosystems. It provides habitat
650
Dietz, D. R. 1972. Nutritive value of shrubs. In: Welch, B.L. 1989. Nutritive value of shrubs. In: C.
C. M. McKell, J. P. Blaisdell, and J. R. Goodin, M. McKell, ed. The biology and utilization of
tech. eds. Wildland shrubs–their biology and shrubs. Academic Press, Inc., San Diego, CA.
utilization, An international symposium; p. 405-424.
Proceedings; 1971 July; Logan, UT. General
Technical Report INT-1. U.S. Department of Welch, B.L. and D. Andrus. 1977. Rose hips-a
Agriculture, Forest Service, Intermountain possible high-energy food for wintering mule
Forest and Range Experiment Station, Ogden, deer? Research Note INT-221. Intermountain
UT. p. 289-302. Forest Experiment Station, U.S. Department of
Agriculture, Forest Service, Ogden, UT. 6 p.
Gill, J.D. and F.L. Pogge. 1974. Rosa L. Rose In:
C.S. Schopmeyer, tech. coord. Seeds of woody Welsh, S.L, N.D. Atwood, S. Goodrich, and L.C.
plants in the United States. Agric. Handbook Higgins. 1987. A Utah flora. Great Basin
654. U.S. Department of Agriculture, Forest Naturalist Memoirs. No. 9. Brigham Young
Service, Washington, DC. p. 732-737. University, Provo, UT. 894 p.
Pacific Southwest Experiment Station. 2002. Fire

effects information system.
www.fs.fed.us/database/feis.plant. 33p.
651
Rubus discolor Weihe & Nees Himalayan blackberry
ROSACEAE
Synonyms: Rubus procerus auct. non P. Muell. ex Genev

Rubus fruticosus L.
Rubus macrostemon (Focke) Sampaio
Rubus thyrsanthus (Focke) Foerster.
Rubus praecox Bertol.
Rubus grabowskii Weihe ex Gunther et al.
John K. Francis
After its release to gardeners, it quickly spread to

wildlands on the West Coast and later to other
parts of the United States (Jacobson 2001).
Taxonomists disagree on its identity. Some
maintain that it is in fact Rubus armeniacus Focke
(Ceska 1999). Himalayan blackberry hybridizes
with R. thyrsiger Banning & Focke, R. calvatus
Lees ex Bloxam, and R. schlechtendalii Weihe ex
Link (Tirmenstein 1989).
Range.—Himalayan blackberry originated in the

Old World, probably Europe. It has been under
semicultivation for a long time and has escaped
into the wild in Western and East-Central United
States, Hawaii, parts of Europe, Australia, New
General Description.—Himalayan blackberry, Zealand, South Africa, and probably southern
also known as Armenian blackberry or just South America (Ceska 1999, Natural Resources
blackberry, is a deciduous to evergreen, spiny, Conservation Service 2003, Starr and others 2003).
arching woody shrub that may reach heights of 4 In many populations, it is not always clear from
m with stem (cane) lengths up to 10 m. The the literature whether it is Himalayan blackberry or
multiple stems are fluted with stout, red, straight or other closely related taxa.
recurved prickles along ridges. The canes grow
straight upward at first but arch as they become Ecology.—Himalayan blackberry grows in vacant
long. The root crown may reach 20 cm in lots, logging sites, burned areas, along rivers,
diameter; the root may penetrate up to 1.5 m into roads, fences, and railroad tracks. It grows best in
the soil, and lateral roots extend 30 to 60 cm. The full sunlight but does well under a light forest
bark is green to purplish-red. The 7- to 20-cm-long cover. However, it will not grow under a dense
leaves are usually penta-foliate except on fruiting canopy. Himalayan blackberry colonizes most soil
branches where they can be trifoliate or unifoliate. types in moist areas but is confined to riparian
The petioles, rachises, and lower surfaces of the areas and the edges of irrigated fields in interior
midveins are also armed. The blades are 3 to 12 dry climates. It grows at elevations from near sea
cm long, ovate to orbicular, dark green, and have level on the West Coast to 1,500 m in the inland
biserrate margins. The white to pink, five-peteled West (Welsh and others 1987). Himalayan
flowers 2- to 2.5-cm broad are borne three to 20 in blackberry is confined at the northern extent of its
terminal panicles. Fruits are aggregates 12 to 20 range to mild coastal areas and low-elevation,
mm in diameter of black, juicy, sweet druplets on sheltered inland sites. It tolerates periodic flooding
a torus (central core). Each drupe contains one of fresh or even brackish water. The species
hard, flattened seed that is covered with a fine vigorously sprouts from root systems after fires
network of pits and ridges (Brayshaw 1996, (Tirmenstein 1989).
Bruzzese 1998, Starr and others 2003, Welsh and
others 1987). Luther Burbank introduced the fruit
cultivar he called Himalayan blackberry in 1885.
652
Reproduction.—Himalayan blackberry ecosystems in places such as Hawaii (Starr and
commonly flowers from June to August. The others 2003). Plants can be eliminated by grubbing
flowers are intensely visited by honey bees, but with follow-up treatment a year later, repeated
insect pollination is not necessary for seed mowing, and by pasturing with goats (Cox 2003).
production. In fact, sexual reproduction may be Several herbicides are effective using foliar spray,
rare; 17 samples from Himalayan blackberry stem injection, treatment of cut stumps, and basal
plants collected throughout Australia proved to spray methods (Starr and others 2003, Tirmenstein
have no genetic variation (Evans and others 1998). 1989). Dead canes accumulate in older stands and
Good seed crops are produced nearly every year are as much a nuisance to remove as the plants
(Tirmenstein 1989). There are 324,000 cleaned themselves. Because of dormant seeds in the soil,
seeks/kg. The seeds are dispersed by birds and checking and retreatment for many years may be
mammals as well as by gravity. Passing through required to completely eliminate the species.
birds and mammals is reported to increase
germination by 30 percent in the first year Benefits.—Himalayan blackberry provides food
(Bruzzese 1998). Seeds in the soil seed bank and cover for many mammals and birds. It is little
remain viable for several years (Brinkman 1974). used by domestic livestock except for goats.
Stands in Victoria, Australia have been estimated However, deer, elk, rabbits, porcupines, beavers,
to produce 7,000 to 13,000 seeds/m2/year and mountain beavers consume leaves, buds,
(Bruzzese 1998). Himalayan blackberry seedlings twigs, and cambium, especially during the winter
commonly appear after fires or disturbance that months. A large number of species consume the
exposes the soil and allows sunlight to reach the fruits and a number rely on the thickets of stems
surface. Once seedlings become established, most for escape and reproductive cover (Tirmenstein
subsequent reproduction is vegetative. Plants 1989). Himalayan blackberry fruits are among the
reproduce by sprouts from rhizomes and by most delicious of wild fruits but are difficult to
layering (rooting) at the nodes when stems come in pick because of the spines. They are eaten fresh,
contact with the ground. canned, used to flavor ice cream, and made into
pies, jams, jellies, juices, and wines. Some people
Growth and Management.—Canes do not bloom object to the seeds getting caught in one’s teeth
during their first year in which they make most of when eaten whole, but it is a minor inconvenience
their growth (2 to 8 m of elongation). Canes bloom that can be avoided by juicing the fruit. In spring,
and fruit in their second and sometimes third years the succulent canes can be pealed and eaten, fresh
and die at the end of their second or third years. or cooked (Jacobson 2001). The brambles are
Individual root crowns live a maximum of 7.5 allowed to grow over fences and trellises to create
years in Australia (Bruzzese 1998). By suckering barriers almost impenetrable to people and
from rhizomes and layering, plants (clones) can livestock. While the species does protect the soil
endure almost indefinitely. Live biomass of a stand from erosion and helps revegetate disturbed sites,
in Victoria, Australia totaled 3.1 tons/ha (59 the thickets and mounds of brambles restrict
percent above-ground and 41 percent below- movement of hikers and woods workers, and
ground) and 27 tons of dead canes and litter suppress other vegetation and slows succession.
(Bruzzese 1998). Scarification is required for
prompt germination. Although several methods References
will work, soaking in concentrated H2SO4 for 50 to
60 minutes followed by 90 days of cold Brayshaw, T.C. 1996. Trees and shrubs of British
stratification is recommended. About 33 percent Columbia. UBC Press, Vancouver, BC, Canada.
germination in about 70 days can be expected. 373 p.
Sowing in late summer for spring germination can
substitute for artificial cold stratification. Sowing Brinkman, K.A. 1974. Rubus L., blackberry,
depth is 3 to 9 mm. (Brinkman 1974). The species raspberry. In: C.S. Schopmeyer, tech. coord.
can be propagated by digging up and replanting Seeds of woody plants in the United States.
suckers. However, many more plants can be Agriculture Handbook 450. U.S. Department of
started by using root cuttings. Root pieces between Agriculture, Forest Service, Washington, DC.
3 and 13 mm in thickness and 10 to 18 cm long are p. 738-743.
recommended. They are grown for 1 year in the
nursery bed before outplanting (Shoemaker 1978). Bruzzese, E. 1998. The biology of blackberry in
Himalayan blackberry can be troublesome in many south-eastern Australia. Plant Protection
habitats and is a potential threat to native Quarterly 13(4): 160-162.
653
Ceska, A. 1999. Rubus armeniacus—a correct e&keywordquery=Rubus+discolor&earl=plant_
name for Himalayan blackberries. Botanical search.cgi. 5 p.
Electronic News ISSN 1188-603X. http://www.
ou.edu.cas/botany-micro/ben/ben230.html. 3 p. Shoemaker, J.S. 1978. Small fruit culture. The
AVT Publishing Company, Inc., Westport, CN.
Cox, C. 2003. Nonchemical methods for removing 357 p.
unwanted blackberry plants. Journal of Pesticide
Reform 23(1): 10-11. Starr, F., K. Starr, and L. Loope. 2003. Plants of
Hawai’i: Rubus discolor. http://www/jear/
Evans, K.J., D.E. Symon, and R.T. Roush. 1998. prg/starr/hiplants/reports/html/rubus_discolor.ht
Taxonomy and genotypes of the Rubus m. 6 p.
fruticosus L. aggregate in Australia. Plant
Protection Quarterly 13(4): 152-156. Tirmenstein, D. 1989. Rubus discolor. In: U.S.
Jacobson, A.L. 2001. Himalaya blackberry: Rubus Rocky Mountain Research Station, Fire Sciences
armeniacus Focke. http://www.arthurleej.com/ Laboratory, Fire Effects Information System.
a-himalayabb.html. 4 p. http://www.fs.fed.us/database/feis/plants/shurb/
rubdis/all.html. 15 p.
Plants profile: Rubus discolor Weihe & Nees, Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
Himalayan blackberry. http://plants.usda.gov/ Higgins. 1987. A Utah flora. Great Basin
cgi_bin/plant_search.cgi?mode=Scientific+Nam Naturalist Memoirs 9. Brigham Young
University Press, Provo, UT. 894 p.
654
Rubus parviflorus Nutt. thimbleberry
ROSACEAE
Synonyms: Rubus nutkanus Moc. ex Ser.

Rubus lacer Kuntze
Rubacer parviflorum Rydb.
Bossekia parviflora Greene
Rubus nutkanus var. parviflorus Focke
John K. Francis
sorrel, and western thimble raspberry, is a

spineless, deciduous shrub 0.5 to 2.5 m in height.
Twigs and new stems are green and glandular; old
stems are covered with tan, papery, shredding
bark. Stems support relatively few branches.
Clonal groups of plants are usually represented by
several to many somewhat dispersed stems that
have arisen from rhizomes (underground stems).
Leaves, which have a maple-leaf appearance, are
simple, alternate, and palmately veined with three
to seven lobes and have entire to serrate margins.
The yellow-green to dark-green blades are 4.5 to
15 cm long and 5.5 to 20 cm broad and are
supported by long petioles. Perfect, white flowers
2 to 4 cm broad are grouped in terminal corymbs
of 2 to 7 flowers. The fruits, which are aggregates
of tiny drupes, are hemispherical, thin, somewhat
dry, fragile, and dark pink to bright red, and detach
easily from the central core (torus), which has five
sepals at its base. The hard seeds are 1.5 to 2.5 mm
long (Abrams 1944, Rook 1998, Welsh 1974)
Range.—Thimbleberry occurs from Alaska to

western Ontario, from Michigan westward, and
south through California, Arizona, New Mexico,
and northern Mexico (Natural Resources
Conservation Service 2003, Welsh 1974, Welsh
and others 1987). It is also reported in
Massachusetts, probably as a naturalized
population. There are two recognized varieties of
thimbleburry, var. parviflorus, accounting for
virtually all of the range, and var. velutinus (Hook.
& Arn.) Greene, which is densely hairy on the
underside of leaves, found in California (Abrams
1944, Natural Resources Conservation Service
2003).
Ecology.—Thimbleberry grows on most types of

Illustration source: USDA-Forest Service collection, moist soils, including skeletal soils. The species is
Hunt Institute most common in cool, moist sites, particularly on
north slopes. Thimbleberry also occurs on dry,
General Description.—Thimbleberry, also known exposed ridges in high-elevation areas. It grows
as western thimbleberry, salmonberry, mountain from near sea level to 2,900 m in elevation in
655
Wyoming (Tirmenstein 1989). Thimbleberry is spring sowing. Also, scarifying with H2SO4 prior
moderately shade tolerant. Although it is most to the cold stratification may enhance germination.
abundant and grows and fruits best in 60 to 100 Tests have given maximum germination estimates
percent sunlight, the species persists in the of 62 percent (Tirmenstein 1989). Hardwood
understory of closed stands (Tirmenstein 1989). It cuttings can be easily rooted. New plants can also
occurs both as scattered individual plants and as be started from rhizome cuttings and plant
large continuous stands. Habitat includes burned divisions. Wildlings survive well but are slow to
sites, logged areas, avalanche tracks, roadsides, become established (Washington State University
natural shrublands, hardwood forest understories, Cooperative Extension 2003). Occasionally, it is
understories of low to moderate basal area conifer desirable to release conifer seedlings in and under
forests, and river overflow terraces and shorelines. thimbleberry thickets or to make way for other,
It is a part of a large number of different plant more palatable forage species. The species is
communities. Thimbleberry is resistant to fire, moderately susceptible to herbicides. Picloram,
readily resprouting from rhizomes. In fact, the 2,4-D, and glyphosate have been effective in
species generally benefits from all kinds of appropriate applications (Tirmenstein 1989).
disturbance. It usually appears during the first year
after disturbance and often dominates the Benefits.—Thimbleberry is an important shrub of
understories of logged areas within 5 years. the understory and forest openings. It helps protect
Moderate to light thimbleberry stands can serve as the soil and adds to the beauty of the forest. The
a good protective cover for conifer seedlings. In species is occasionally planted as an ornamental
recreational sites, the species showed relatively for its fragrant flowers and brilliant orange to
low resistance to trampling (Tirmenstein 1989). maroon fall foliage (Borialforest.org 2003, Rook
1998). It is also occasionally planted in
Reproduction.—Thimbleberry generally blooms conservation plantings in disturbed areas.
in June and July. Its fruits mature in July and Thimbleberry is relatively low in energy and
August (Borialforest.org 2003). However, at the protein (4 to 8 percent), and is little used by cattle
extremes of its range, it may bloom anywhere and horses and is only fair forage for sheep. It is
from May to September and fruit between late sometimes important for deer, elk, and other wild
June through September (Tirmenstein 1989). The ungulates during the summer while the leaves are
flowers are insect pollinated. Good fruit and seed still present. Rodents consume bark, buds, and
crops are usually produced every year except in foliage to a limited extent. On the other hand, the
high-elevation areas where production may be fruits are an important food item for numerous
unreliable (Tirmenstein 1989). The seeds are wild mammals and birds. Thickets of thimbleberry
dispersed by birds and mammals that eat the fruits, are also important escape, resting, and
and by gravity. The seeds accumulate in the soil reproductive cover for many species of wildlife
and duff to form a seed bank and germinate in (Tirmenstein 1989). Fruits are certainly edible to
great numbers following fire and other humans but reports vary greatly on their
disturbances that remove the forest canopy (Rook palatability (Clark 1976, Welsh 1974). The author
1998). After establishment by seeds, stands has eaten them for years and finds them inferior to
thicken through sprouts from rhizomes (Rook cultivated raspberries (Rubus idaeus L.) but well
1998). worth the trouble of picking while hiking in the
forest. The fruits are made into an excellent jelly
Growth and Management.—The stems (or and were once dried for later use by Native
canes) live for 2 or 3 years. In the first year, canes Americans. The tender young shoots are juicy and
grow in height and develop only leaves. During sweet and can be boiled or eaten fresh. The leaves
the second and third years height growth continues are sometimes made into herb teas
and flowers and fruits are produced. The canes (Borialforest.org 2003, Washington State
then die and are replaced by others that sprout Department of Transportation 2003). Native
from rhizomes. Maximum height of current canes Americans applied poultices of leaves to burns and
is reached within 10 years after establishment wounds and took decoctions of roots as a tonic, for
(Tirmenstein 1989). Seeds must be collected by vomiting, and certain internal disorders (Moerman
hand and should be cleaned of adhering fruit. 1986).
Seeds may be sown in the fall for spring
germination or may be warm (20 to 30 °C for 90
days) and cold stratified (2 to 5 °C for 90 days) for
656
References Tirmenstein, D. 1989. Rubus parviflorus. In: U.S.
Abrams, L. 1944. Illustrated flora of the Pacific Rocky Mountain Research Station, Fire Sciences
States. Vol. 2. Stanford University Press, Laboratory, Fire Effects Information System.
Stanford, CA. 635 p. http://www.fs.fed.us/database/feis/plants/shurb/
rubpar/all.html. 37 p.
Borialforest.org. 2003. Rubus parviflorus,
thimbleberry. http://www.borialforest.org/ Washington State Department of Transportation.
shrubs/shrub41.htm. 2 p. 2003. Environmental affairs: Rubus parviflorus.
http://www.wsdot.wa.gov/environment/eao/
Clark, L.J. 1976. Wild flowers of the Pacific culres/ethbot/q-s/RubusPar.htm. 2 p.
Northwest. Gray’s Publishing Limited, Sidney,
BC, Canada. 604 p. Washington State University Cooperative
Extension 2003. Native plants: Rubus
Moerman, D.E. 1986. Medicinal plants of Native parviflorus var. parviflorus. http://cahedb.
Americans. Technical Report 19. Museum of wsu.edu/nativePlant/scripts/webDisplayText.asp
Anthropology, University of Michigan, Ann ?ID=nv044. 1 p.
Arbor, MI. 534 p.
Welsh, S.L. 1974. Anderson’s flora of Alaska.
Natural Resources Conservation Service. 2003. Brigham Young University Press, Provo, UT.
Plants profile: Rubus parviflorus Nutt. 724 p.
http://plants.usda.gov/cgi_bin/topics.cgi. 4 p.
Rook, E.J.S. 1998. Rubus parviflorus, Higgins. 1987. A flora of Utah. Great Basin
thimbleberry. http://www.rook.org.earl/bwca/ Naturalist Memoirs 9. Brigham Young
hature/shurbs/rubuspar.html. 7 p. University Press, Provo, UT. 894 p.
657
Rubus rosifolius Sm. roseleaf raspberry
ROSACEAE
Synonyms: Rubus coronaries (Sims) Sweet
John K. Francis
Puerto Rico) and Brazil and to many of the islands

of the Pacific (including Hawaii) (Liogier 1985,
Lu and Boufford 2002, Pacific Island Ecosystems
at Risk 2002, Peixoto 2002).
Ecology.—Although roseleaf raspberry invades

forest understories in some habitats (Pacific Island
Ecosystems at Risk 2002), it is moderately shade
intolerant, requiring partial sun to flower and fruit;
it does best in full sun. Preferred habitats include
natural and artificial openings in primary and
secondary forest, such as tree-fall gaps, landslides,
abandoned fields, neglected pastures, and
roadsides. Roseleaf raspberry has a high moisture
General Description.—Roseleaf raspberry, also requirement, which usually means more than 1800
known as wild raspberry, Mauritius raspberry, mm of mean annual precipitation and no severe
Queensland raspberry, fresa de montaña, annual droughts. The species usually grows at
framboisier, and akalakala, is an evergreen moderate to high elevations. It is reported at 2,200
scrambling shrub, freestanding to 0.75 m, taller m in Tahiti and 1,730 m in Hawaii (Pacific Island
when supported on other vegetation, and extending Ecosystems at Risk 2002). A variety of well-
laterally 2 m or more. Individual plants usually drained to poorly-drained soils are colonized.
have several arching stems (canes) with basal
diameters of 0.5 to 1 cm and have a 3-mm pith. Reproduction.—In China, roseleaf raspberry
Stems are green with reddish coloration at the flowers in March through May and fruits in June
nodes and lower portions. Wood of the lower and July (Lu and Boufford 2002). The species
stems is moderately hard and brittle. The stems, blooms and fruits throughout the year in
twigs, and leaf petioles and rachises are armed continually moist areas of Puerto Rico and
with short, straight or curved prickles. Adult plants Equatorial Africa (Burkill 1997). Rubus species
have a fibrous lateral root system. Leaves are odd- are pollinated by insects, particularly honey bees
pennate with mostly seven ovate to linear, doubly (Apis mellifera L.) (Dalaplane and Mayer 2000).
serrate leaflets. The terminal or axile Fresh fruits collected in Puerto Rico averaged
inflorescences contain one to four, 1.5 to 3 cm 1.516 + 0.102 g/fruit. Air-dried seeds separated
diameter flowers with five white petals. The from them averaged 0.00045 g/seed or 2.2 million
globose to narrowly oblong aggregate fruit is seeds/kg. The seeds are dispersed by birds and
bright red and juicy when ripe. The pitted rodents (Pacific Island Ecosystems at Risk 2002).
(foveolate), yellow seeds (pyrenes) are about 1.5 Stems root (layer) whenever they come in contact
mm long (author’s observation, Howard 1988, with the soil, which helps it fill in newly colonized
Liogier 1985, Lu and Boufford 2002). habitat.
Range.—Roseleaf raspberry is apparently native Growth and Management.—Stems (canes) of

from China and Taiwan south through Indochina roseleaf raspberry originating from sprouts grow 1
and Indonesia along the eastern coast of Australia, m or more per year and live about 1.5 years. New
Mauritius, New Caledonia, the Solomon Islands, sprouts then arise to replace them and the plant
and Vanuatu, and eastward as far as northeastern lives on (potentially) for several years. In Puerto
India. However, it is difficult to tell the native Rico, clumps and thickets of roseleaf raspberry
from naturalized range that extends from Japan to usually disappear in 2 or 3 years due to mounting
Africa and through the Caribbean (including competition and encroaching shade from trees,
658
shrubs, and herbs. The species is occasionally Howard R.A. 1988. Flora of the Lesser Antilles,
planted from wildlings and rooted layers for Leeward and Windward Islands.
personal use in gardens. It is controlled along with Dicotyledoneae, Part 1. Vol. 4. Arnold
other vegetation by slashing or mowing, and by Arboretum, Harvard University, Jamaica Plain,
tillage to prepare ground for crops. MA. 673 p.
Benefits.—Roseleaf raspberry helps return Liogier H.A. 1985. Descriptive flora of Puerto
disturbed areas to forest vegetation, protects the Rico and adjacent islands, Spermatophyta. Vol.
soil, and furnishes food and cover for wildlife. 1. Editorial de la Universidad de Puerto Rico,
Leaves and tender shoot are browsed by livestock. Río Piedras, PR. 352 p.
The berries are eaten out of hand and made into
drinks and preserves throughout its range. Fresh Liogier, H.A. 1990. Plantas medicinales de Puerto
fruits are sold nearly year-round in Ugandan Rico y del Caribe. Iberoamericana de Ediciones,
markets (Burkill 1997). The fruits contain Inc. 566 p.
relatively high (1.58 and 12.93 mg/100g fresh
weight respectively) of vitamins E and C (Wei and Lu, L. and D.E. Boufford. 2002. Rubus Linnaeus
Payne 2002). Roseleaf raspberry has been grown [Draft]. In: A.R. Brach and Hong S., eds. Flora
as a ground cover in Cameroon plantations of China, Harvard University Herbaria,
(Burkill 1997). A double petaled variety (R. Cambridge, MA. http://hua.huh.harvard.edu/
rosifolius var. coronaries (Sims) Focke) is grown china/mss/volume09/Rosaceae-AGH_Rubus_
as an ornamental (Garden Plant Conservation coauthoring.htm. 200 p.
2002). Infusions of flower petals are used to
control diarrhea, vomiting, and other flu Nortan, A. 2002. Bush tucker: Roseleaf raspberry
symptoms, and as a tonic (Liogier 1990). An (Rubus rosifolius). Rumbalara Environmental
infusion of the leaves was brewed in former times Ecucation Centre, Sydney, NSW, Australia.
to relieve menstrual cramps, morning sickness, and http://rumbalara-e.schools.nsw.edu.au/
labor pains (Nortan 2002). bushtucker/ Rubus_rosifolius.htm. [not paged].
References Pacific Island Ecosystems at Risk. 2002. Rubus

rosifolius Sm., Rosaceae. http://hear.org/
Burkill, H.M. 1997. The useful plants of West pier_v3.3/ruros.htm. 2 p.
Tropical Africa. Ed. 2. Vol. 4. Royal Botanical
Garden, Kew, UK. 960 p. Peixoto, M. 2002. Plantas Brasileiras: Rosaceae,
Rubus rosifolius. http://sites.uol.com.br/
Dalaplane, K.S. and D.F. Mayer. 2000. Crop mpeixoto/outras/rosifolius.html. 2 p.
pollination by bees. CABI Publishing,
Wallinford, UK. 352 p. Wei, C. and T.J. Payne. 2002. Raspberry (Rubus
L.). Washington Red Raspberry Commission,
Garden Plant Conservation. 2002. Rubus rosifolius Lyndon, WA. http://www.red-raspberry.org/
‘Coronarius.’ http://www.manntaylor.com/ china.pdf. 7 p.
plantweek9c.html. 7 p.
659
Salix arctica Pallas arctic willow
SALICACEAE
Synonyms: Salix anglorum Cham.

Salix brownie (Anderss.) Bebb
Salix caespitosa Kennedy
Salix pallasii Anderss.
Salix petrophila Rydb.
Dallwitz (1980). Identification may be difficult.

Environment can affect the morphology to such an
extent it appears a unique species is present (Argus
and others 1999). Although many subspecies have
been described, they have generally been discarded
in favor of the type species. In addition, hybrids
are reported, and introgression with other species,
such as S. cascadensis in Utah and Salix reticulata
in Russia, is well documented (Hultén 1968,
Komarov 1936, Pojar and MacKinnon 1994,
Range.—Arctic willow is a circumpolar species

occurring in North America, Europe, and Asia
(Britton 1901, Fertig and Markow 2001, Hultén
1968, Komarov 1936). In North America it grows
in the sub-alpine and alpine tundra zones from
Quebec to Alaska and south, through the Rocky
Mountains, into New Mexico (Kershaw and others
Hunt Institute
1998). Occurrences are also reported from Utah
(Welsh and others 1993) and the Ruby Mountains
General Description.—Salix arctica is commonly of Nevada (Charlet 1994). It has disjunct
known as arctic willow. This deciduous, prostrate, occurrences on the west coast of the U.S.A.,
trailing shrub is usually less than 10 cm tall and occurring in California and in the northern
commonly forms mats. However, its growth Cascade Mountains in Washington and in the
habitat is variable, and in the Pacific Northwest it Wallowa Mountains in Oregon (Hitchcock and
grows up to 50 cm tall (Pojar and MacKinnon Cronquist 1973).
1994). The broadly oval, alternate leaves are 2 to 6
cm long and sparsely hairy when mature. They are Ecology.—Arctic willow grows in dry meadows,
a glossy green on the upper side and pale green- rocky tundra, fellfields, and in relatively moist
grey below (Kershaw and others 1998, Fertig and sites in fen lands and at the edge of snowfields.
Markow 2001). Plants are dioecious, possessing Habitat tends to be related to latitude. In the
male and female flowers on separate, “unisexual” mountains of New Mexico it is an obligate wetland
plants. The flowers of both sexes are arranged in species, whereas at higher latitudes it grows in
dense, erect clusters or catkins. The female catkins both dry and moist habitats (Cooper and others
are densely hairy, 1 to 8 cm (usually 2 to 4 cm) 1997, Dick-Peddie 1993, Fertig and Markow 2001,
long and are borne on lateral, sometimes appearing Hultén 1968). It may be locally dominant in some
terminal, branchlets (Kershaw and others 1998, cushion plant communities and co-occur with
Fertig and Markow 2001). The male, staminate, other species of dwarf willow such as Salix
catkins are up to 4 cm long. Flowers appear with reticulata L. (snow willow). In New Mexico
the leaves. There are several more synonyms than tundra, arctic willow and snow willow are often
those listed above and may be referred to in found associated with lichens and other vascular
Kartesz (1994), Argus and others (1999), and plants in snowbank and rivulet communities
(Dick-Peddie 1993). Similarly in Montana and
660
elsewhere in the Rocky Mountains, a common S. chambers (Jones and others 1999). Overall net
arctica/Polygonum bistortoides Pursh community assimilation was higher in the dry habitat than in
was identified in “water-receiving positions” the wet habitat, and higher in females than in
(Cooper and others 1997). However, the latter sites males, although there appeared to also be some
were also reported to have a good probability of habitat-sex interactions (Jones and others 1999).
experiencing wind scouring, and some of the Elevated temperature enhanced development and
associated vegetation suggested the area could be growth of both male and female catkins, but the
relatively dry (Cooper and others 1997). The response of the leaves to elevated temperatures
elevation at which it occurs is also associated with was more variable particularly among male
latitude. Plants are only found above 3,350 m in individuals (Jones and others 1999). These
New Mexico, whereas they occur at sea level and findings indicate the importance of defining both
up to 700 m in northern parts of its range, such as sex and habitat conditions when making
Greenland (Carter 1997, Argus and others 1999). observations on this dioecious species, and they
It grows in a range of soils, from acidic to may also have significance in predicting the
calcareous (Argus and others 1999, Cooper and response of Arctic willow to global climate
others 1997, Pojar and MacKinnon 1994). A change.
mutualistic association exists between arctic
willow and ectomycorrhizal fungi that are Benefits.—The majority of Salix species provide
primarily agarics, or gilled mushrooms (Cripps important browse for wildlife and livestock within
and Horak 2002). their range (Dayton 1931). Arctic willow,
particularly the young leaves stems and buds,
Reproduction.—Arctic willow reproduces by provide valuable browse for ptarmigan and other
seed and also vegetatively by rooting at the nodes arctic animals including muskox and reindeer
of stems (Fertig and Markow 2001). Flowering is (Kershaw and others 1998, Komarov 1936, Tolven
from June into August. On calm, sunny days the and others 2001). Arctic willow is tolerant of
female catkins can be up to 8.5 oC warmer and the varied conditions, and cuttings have been rooted
male catkins up to 7 oC warmer than the and used in restoration projects (Bittman 1997).
surrounding air temperature (Kershaw and others Willow leaves and bark are commonly used for
1998). The warmer temperatures speed pollen and medicinal purposes and the roots, twigs, and bark
seed development and also attract insects are used for a multitude of fiber work, such as
(Kershaw and others 1998). basketry and clothing by Native American peoples
(Moerman 1998). Arctic willow has been used for
Growth and Management.—Seeds have a similar purposes by peoples native to Asian Arctic
physiological dormancy and require a 30-day regions (Moerman 1998). The Yakuts have used
stratification period for germination (Baskin and an infusion as a substitute for tea called “chai-
Baskin 2001). Seeds germinate at 25 oC although talak” (Komarov 1936). It is used as fuel in some
minimum temperatures for germination are not regions where it is the only source of wood
defined (Densmore and Zasada 1983). Thompson (Kershaw and others 1998). Arctic willow is
(1992) reported that seeds of Salix species are cultivated, and it is especially popular for rock
“normally absent” from the seed bank in alpine gardens (Bailey and others 1976, Pojar and
and arctic communities. Individual shrubs live for MacKinnon 1994).
60 to 85 years although one specimen was reported
to be 180 and another 236 years old (Savile 1979, References.
Raup 1959). Habitat appears to influence the sex
that is most abundant. Dawson (1987) reported Argus, G.W., C.L. McJannet, and M.J. Dallwitz.
female plants were significantly more numerous 1999 onwards. ‘Salicaceae of the Canadian
than male plants in mesic-wet, more fertile, low Arctic Archipelago: Descriptions, Illustrations,
soil-temperature sites, whereas male plants were Identification, and Information. http://www.
most prevalent in drier, less fertile sites. In mun.ca/biology /delta/arcticf/.
addition, both habitat and sex may influence a
plant’s response to environmental conditions such Bailey, L.H., E.Z. Bailey, and the staff of the
as temperature. In a 3-year gas exchange field Liberty Hyde Bailey Hortorium. 1976. Hortus
study, male and female willows from dry and wet Third. Macmillian Publishing Company, New
habitats were subjected to passively enhanced York. 1,290 p.
summer temperature using small open-top
661
Baskin, C.C. and J.M. Baskin. 2001. Seeds, Dick-Peddie, W.A. 1993. New Mexico
ecology, biogeography, and evolution of vegetation. University of New Mexico Press,
dormancy and germination. Academic Press, Albuquerque, NM. 244 p.
New York. 666 p.
Fertig, W. and S. Markow. 2001. Guide to the
Bittman, K.K. 1997. High elevation native species willows of Shoshone National Forest. General
island model for mine reclamation. Quintette Technical Report RMRS-GTR-83. U.S.
Operating Corporation, Tumbler Ridge, BC. Department of Agriculture, Forest Service
http://www.teckcominco.com/presentations/kb- Rocky Mountain Research Station, Fort Collins,
highelv-1997.pdf. CO. 80 p.
Britton, N.L. 1901. Manual of the Flora of the Hitchcock, C.L., and A. Cronquist. 1973. Flora of
northern states and Canada. Henry Holt and the Pacific Northwest. University of Washington
Company, New York. 1,080 p. Press, Seattle, WA. 730 p.
Hultén, E. 1968. Flora of Alaska and neighboring

Carter, J.L. 1997. Trees and shrubs of New
territories Stanford University Press, Stanford,
CA. p. 340-341.
Charlet, D. 1994. Plant Species in the Ruby
Jones M.H., S.E. Macdonald, and G.H.R. Henry.
Mountains, Nevada.
1999. Sex- and habitat-specific responses of a
http://www.brrc.unr.edu/data/plants/ ruby.html.
high arctic willow, Salix arctica, to experimental
climate change. Oikos 87(1): 129-138.
Cooper, S.V., P. Lesica, and D.Page-Dumrose.
1997. Plant community classification for alpine
vegetation on the Beaverhead National Forest,
vascular flora of the United States, Canada and
Montana. General Technical Report INT-362.
Greenland. Vol.1 - Checklist. 2nd Ed. Timber
USDA Forest Service Intermountain Research
Press, Portland, OR. 622 p.
Station, Ogden, UT. 61 p.
Kershaw, L., A. MacKinnon, and J. Pojar. 1998.
Cripps, C.L., and E. Horak. 2002. A survey Plants of the Rocky Mountains. Lone Pine
of alpine agaricales in the Rocky Mountains. Publishing, Edmonton, Canada. 383 p.
http://plantsciences.montana.edu/alpinemushroo
ms/ rocky_mountain_ alpine_field_sites.htm Komarov, V.L. 1936. Flora of the U.S.S.R.
Volume V. Izdatel’stvo Akademii Nauk SSSR,
Dallwitz, M.J. 1980. A general system for coding Moscow-Leningrad, U.S.S.R. Translated from
taxonomic descriptions. Taxon, 29: 41–46. Russian by N. Landau 1970. Israel Program for
scientific translations, Jerusalem, Israel and The
Dawson, T. E. 1987. Comparative Smithsonian Institution Washington, D.C. p.
ecophysiological adaptations in arctic and alpine 36-37.
populations of a dioecious shrub, Salix arctica
Pall. Ph.D. dissertation, University of Moerman, D.E. 1998. Native American ethno-
Washington, Seattle, WA. 208 p. botany. Timber Press, Portland, OR. 927 p.
Dayton, W. A. 1931. Important western browse Pojar, J. and A. MacKinnon. 1994. Plants of the
plants. Miscellaneous Publication 101. U.S. Pacific Northwest coast–Washington, Oregon,
Department of Agriculture, Washington DC. British Columbia and Alaska. Lone Pine
214 p. Publishing, Edmonton, Canada. 526 p.
Densmore, R. and Zasada, J.C. 1983. Seed Raup, H.M. 1959. The willows of boreal western
dispersal and dormancy patterns in northern America. Contributions from the Gray
willows: Ecological and evolutionary Herbarium of Harvard University 185: 3–95.
significance. Canadian Journal of Botany 61:
3207-3216
662
Savile, D.B.O. 1979. Ring counts in Salix arctica Tolven, A. J., Schroderus, and G.H.R. Henry.
from northern Ellesmere Island. Canadian Field- 2001. Age- and stage-based bud demography of
Naturalist 93: 81-82. Salix arctica under contrasting muskox grazing
pressure in the High Arctic. Evolutionary
Thompson, K. 1992. The functional ecology of Ecology 15: 4-6
seed banks. In: M. Fenner, ed., Seeds, the
ecology of regeneration in plant communities. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
CAB International, Wallingford, Oxon, U.K. Higgins. 1993. A Utah flora. Second Edition,
373 p. revised. Brigham Young University, Provo, UT.
986 p.
663
Salix exigua Nutt. coyote willow
SALICACEAE
Synonyms: Salix interior Rowlee
James E. Nellessen
times longer than wide, 2 to 16 cm long, 4 to 15
mm wide, the margins are smooth or irregularly
toothed, with one to five teeth per cm, and the
petioles are short, 2 to 7 mm long (Argus 1986,
Britton and Brown 1913, Carter 1997, Gleason and
Cronquist 1963, Great Plains Flora Association
1986, Martin and Hutchins 1980-81, Newsholme
1992). Upon emergence the young twigs and
leaves are densely silvery hairy, but with time and
aging, the upper leaf surface reduces to a
moderately gray hairy or smooth surface,
sometimes slightly shiny. Plants in moist sites at
higher elevations in the mountains may be
noticeably less hairy (Mozingo 1987). The lower
leaf surface generally remains whitish and much
hairier. Coyote willow is one of few willow
species with stomata on both leaf surfaces (Argus
1986). Stipules are small and sometimes absent.
The chromosome number is 2n = 38 (Dorn 1998).
Taxonomy.—Coyote willow has a complicated

taxonomy. It is a member of Salix section
Longifoliae. Several botanists now consider S.
interior as a much less hairier to smooth form of S.
exigua that occurs in the Great Plains and Eastern
United States, while variety exigua is the hairier
Western form (Argus 1986, Carter 1988, Dorn
1998, Gleason and Cronquist 1963, Great Plains
Flora Association 1986, Weber and Wittman
2001a, b). The Eastern interior is more distinctly
General Description.—Coyote willow is also toothed, leaf veins are more prominent, and the
known as sandbar willow, basket willow, fruits (capsules) slightly longer. But characteristics
narrowleaf willow, slender willow, riverbank are highly variable across the entire range of this
willow, acequia willow, long-leaved willow, and plant. Under this taxonomic organization and
gray willow (Elmore 1976). It generally ranges in using the Dorn (1998) treatment, there would be
height from 0.5 to 3 m, is occasionally a small tree two subspecies: S. exigua subsp. exigua (coyote
to 6 m and trunk diameter to 13 cm, with one willow) and S. exigua subsp. interior (Rowlee)
author (Dorn 1998) reporting up to 17 m in height. Cronq. (sandbar willow). The Western subspecies
Coyote willow is a clonal species that spreads by is divided into four varieties: var. exigua, var.
creeping rootstocks that generate new shoots hindsiana (Benth.) Dorn, var. sessilifolia (Nutt.)
forming multistemmed dense thickets. Coyote Dorn, and var. columbiana Dorn. The Eastern
willow is one of few willow species that reproduce subspecies, interior, contains no varieties under
this way. As with other willows the species is Dorn’s treatment. There are various synonyms
dioecious. The bark is greenish to reddish on (Argus 1986, Britton and Brown 1913, Dorn 1998,
young twigs and ashy gray to grayish brown on Gleason and Cronquist 1963, Great Plains Flora
older stems. Lenticels are elliptic and raised. The Association 1986, Kearney and others 1951,
buds are reddish brown. The leaves are alternate, Newsholme 1992). A close to complete list of 34
simple, long and narrow, linear to lanceolate in synonyms can be found in Kartesz (1994).
shape, tapering at both ends, generally six or more Hybridization is likely an important factor
664
complicating the taxonomy for this species, the (Tamarix ramosissima Ledebour), with 52 percent
entire section, likely a syngameon undergoing of the leaf mass remaining after 142 days
geographic speciation (Brunsfield and others (Pomeroy and others 2000). A number of insects
1992). attack coyote willow, including leaf mining
beetles, lepidopterans, sawflies (Euura spp.), and
Range.—This is a widely distributed species of galling insects. A bud galling midge (Rabdophaga
plains and lower elevations to lower montane sp.) affects both terminal and lateral shoots,
habitats in the Western United States, as well as arresting growth of the immediate shoot (Declerck
throughout Eastern deciduous forests if S. interior and Price 1994). Coyote willow responds by
is considered a subspecies. The Western form producing new lateral buds and shoots, but many
(exigua) occurs from Saskatchewan to Eastern do not survive.
British Columbia in Canada, south through the
United States to Eastern and Southern California, Reproduction.—The male flowers are in densely
Arizona, New Mexico, and into Northern Mexico flowered catkins from 1 to 6 cm in length, each
and the Western Great Plains. The Eastern form or flower with two stamens, the filaments hairy. The
subspecies (interior) occurs from New Brunswick female catkins are loosely flowered, from 2 to 10
in Eastern Canada, westward throughout the Great cm long, the ovaries are smooth to slightly silky
Plains, as far west as Idaho, north to Alaska, south with a short or almost nonexistent style. Each
to Eastern New Mexico, and extends into the ovary bears nine to 15 ovules (Argus 1986).
Southeast along a line from Northern Virginia to Catkin scales are yellow and deciduous. The
Tennessee and Louisiana. The two subspecies catkins appear with the leaves or shortly after the
overlap in the Western Great Plains. In the leaves in spring (March to May) on short leafy
Western mountains, coyote willow can be found as stalks. The flowers are bee pollinated. There may
high as 2,900 m in elevation. even be a second blooming period from June to
August (Argus 1986). The fruits are smooth
Ecology.—Coyote willow commonly forms capsules from 4 to 8 mm long, green, turning
thickets along streams and rivers and is an yellow to reddish brown upon ripening. The fruits
important riparian species stabilizing streambanks. open by two valves releasing tiny seeds (22,000/g
It will commonly colonize sand bars along rivers, [Brinkman 1974]) with long hairs that aid in wind
hence the frequently used alternate common name dispersal. The seeds are short-lived, do not require
sandbar willow. It also occurs in ditches, and on stratification, must fall on a moist substrate, and
the edges of swamps, lakeshores, and other can germinate within 12 to 24 hours under proper
wetland habitats. This species will grow anywhere conditions (Young and Young 1992). Successful
groundwater is close to the surface. This includes germination is reported at 22 C for the Western
sites where human activities result in surface water subspecies exigua (Brinkman 1974) and 5 to 25
availability on a reasonably regular and reliable C for the Eastern subspecies interior (Densmore
basis (exclusive of saline conditions). Coyote and Zasada 1983, Baskin and Baskin 2001).
willow is also one of the few willow species that Flower-and-fruit-bearing shoots have been
can be found in lower elevation, hot desert areas in observed to sprout new leafy shoots and become
the Western United States. But because if its wide new side branches (Argus 1986).
elevational range it is both a floodplain-plains and
montane riparian species (Dick-Peddie 1993). Due Growth and Management.—Along the Rio
to the extensive rootstock and clonal nature of the Grande in New Mexico, clearing riparian areas of
species, cut stands of coyote willow will the invasive, nonnative salt cedar prior to peak
regenerate relatively quickly. The presence of an river flows in the spring can facilitate the
aerenchyma-containing wetland plant, Typha reestablishment of natives such as coyote willow
latifolia L., may facilitate a nonaerenchyma (Taylor and others 1999). In Alberta, Canada,
bearing species like coyote willow to survive in successful seedling establishment of several
low oxygen wetland soils (Callaway and King riparian species, including coyote willow, occurred
1996). Leaf water potentials as low as -2.7 MPa best on deposits 80 to 120 cm above the late
have been measured for high elevation populations summer stream flow (Rood and Mahoney 2000).
in Wyoming (Foster and Smith 1991). Coyote In Manitoba, Canada, average annual height
willow leaves within the Colorado River (Arizona) increases were 30 cm, stem diameter increases
decomposed substantially more slowly than those were 2.6 mm, flowering began in 2 to 3 year old
of Populus fremontii S. Watson and salt cedar stems, stem mortality was greatest in 3 to 6 year
665
old clones, entire plant senescence first occurred at Baskin, C.C. and J.M. Baskin. 2001. Seeds:
12 years, and the oldest plant recorded was 31 Ecology, Biogeography, and Evolution of
years on a point bar in the Assiniboine River Dormancy, and Germination. Academic Press,
(Ottenbreit and Staniforth 1992). San Diego, CA. 666 p.
Benefits.—Because coyote willow forms dense Brinkman, K.A. 1974. Salix L. Willow. In: C.S.
thickets and spreads clonally, it is an important Schopmeyer, tech. coord. Seeds of Woody
stabilizer of streambanks. It is frequently and Plants in the United States. U.S. Department of
successfully used in riparian and wetland Agriculture Forest Service, Handbook 450.
revegetation and restoration. It is an important Washington, DC. p. 746-750.
food for wildlife such as deer and beaver. It is
considered good browse for sheep and fair for Britton, N.L. and A. Brown. 1913 (1970 Dover
cattle, but cattle generally find it more palatable edition). An Illustrated Flora of the Northern
later in the growing season (Mozingo 1987, U.S. and Canada, Vol. 1. Dover Publications
Stubbendieck and others 1997, USDA 1937). Inc., New York. 680 p.
Coyote willow is used in intensive culture coppice
plantations (Aravanopoulos and others 1999), was Brunsfield, S.J., D.E. Soltis, and P.S. Soltis. 1992.
one of 32 woody plants tested for biomass Evolutionary patterns and processes in Salix:
gasification, and was one of three species that evidence from chloroplast DNA. Systematic
exceeded 0.30 liter methane per gram of volatile Botany 17(2): 239-256.
solids (Turick and others 1991). Coyote willow
has been successfully grown in tissue culture with Callaway, R.M. and L. King. 1996. Temperature
roots developing from shoots longer than 1 cm driven variation in substrate oxygenation and the
(Stoehr and others 1989). Native Americans have balance of competition and facilitation. Ecology
used the slender stems in making baskets and have 77(4): 1,189-1,195.
made a tea from the bark to treat fever and
headache. Willow bark, as with most willows, is Carter, J.L. 1988. Trees and Shrubs of Colorado.
bitter and acts as an astringent for diarrhea, has Johnson Books, Boulder, CO. 165 p.
been used for fevers, pain, arthritis, rheumatism,
and a poultice for cuts, ulcers, and rashes. Salicylic Carter, J.L. 1997. Trees and Shrubs of New
acid, commonly used in asperin as acetylsalicylic Mexico. Johnson Books, Boulder, CO. 534 p.
acid, was originally derived from salicin contained
in the bark of willows (Foster and Duke 1990). Declerck, F.R. and P.W. Price. 1994. Impact of a
Salicin is a member of the chemical class of bud-galling midge on bud populations of Salix
phenols and affords some protection against exigua. Oikos 70(2): 253-260.
browsing mammals such as rabbits. It tends to
attain a greater concentration in low growing Densmore, R. and J.C. Zasada. 1983. Seed
willows but becomes lost in larger, taller willows dispersal and dormancy patterns in northern
(Harborne 1988). Oestriol (an oestrogen), a human willows: ecological and evolutionary
sex hormone, has also been found in willow significance. Canadian Journal of Botany 61:
flowers. 3,207-3,216.
References Dick-Peddie, W.A. 1993. New Mexico

Vegetation: Past, Present and Future. University
Aravanopoulos, F.A., K.H. Kim, and L. Zsuffa. of New Mexico Press, Albuquerque, NM. 244 p.
1999. Genetic diversity of superior Salix clones
selected for intensive forestry plantations. Dorn, R.D. 1998. A taxonomic study of Salix
Biomass and Bioenergy 16(4): 249-255. section Longifoliae (Salicaceae). Brittonia
50(2): 193-210.
Argus, G.W. 1986. The Genus Salix (Salicaceae)
in the Southeastern United States. Systematic Elmore, F.H. 1976. Shrubs and Trees of the
Botany Monographs, Vol. 9. The American Southwest Uplands. Southwest Parks and
Society of Plant Taxonomists, USA. 170 p. Monuments Association, Tucson, AZ. 214 p.
666
Foster, S.A. and J.A. Duke. 1990. A Field Guide to willow). Canadian Journal of Botany 70(6):
Medicinal Plants (Eastern/Central). Peterson 1,141-1,146.
Field Guide Series, Houghton Mifflin Co.,
Boston, MA. 366 p. Pomeroy, K.E., J.P. Shannon, and D.W. Blinn.
2000. Leaf breakdown in a regulated desert
Foster, J.R. and W.K. Smith. 1991. Stomatal river: Colorado River, Arizona, U.S.A.
conductance patterns and environment in high Hydrobiologia 434: 193-199.
elevation phreatophytes of Wyoming, USA.
Canadian Journal of Botany 69(3): 647-655. Rood, S.B. and J.M. Mahoney. 2000. Revised
instream flow regulation enables cottonwood
Gleason, H.A. and A.R. Cronquist. 1963. Manual recruitment along the St. Mary River, Alberta,
of the Vascular Plants of Northeastern U.S. and Canada. Rivers 7(2): 109-125.
Adjacent Canada. D. Van Nostrand Co., New
York. 810 p. Stoehr, M.U., M. Cai, and L. Zsuffa. 1989. In-vitro
plant regeneration via callus culture of mature
Great Plains Flora Association. 1986. Flora of the Salix exigua. Canadian Journal of Forest
Great Plains. University Press of Kansas, KS. Research 19(12): 1,634-1,638.
1,392 p.
Stubbendieck, J., S.L. Hatch, and C.H. Butterfield.
Harborne, J.B. 1988. Introduction to Ecological 1997. North American Range Plants, 5th Edition.
Biochemistry, 3rd Ed. Academic Press, London. University of Nebraska Press, Lincoln, NE.
356 p. 501 p.
Kartesz, J.T. 1994. A Synonymized Checklist of Taylor, J.P., D.B. Wester, and L.M. Smith. 1999.
the Vascular Flora of the United States, Canada, Soil disturbance, flood management, and
and Greenland, Vol. 1, 2nd Edition. Biota of riparian woody plant establishment in the Rio
North America Program of the North Carolina Grande floodplain. Wetlands 19(2): 372-382.
Botanical Garden. Timber Press, Portland, OR.
622 p. Turick, C.E., M.W. Peck, D.P. Chynoweth, D.E.
Jerger, E.H. White, L. Zsuffa, and W.A.
Kearney, T.H., R. Peebles, and Collaborators. Kenney. 1991. Methane fermentation of woody
1951 with 1960 supplement. Arizona Flora. biomass. Bioresource Technology 37(2): 141-
University of California Press, Berkeley, CA. 147.
1,085 p.
U.S. Department of Agriculture Forest Service.
Martin, W.C. and C.E. Hutchins. 1980-1981 1937 (1988 Dover edition). Range Plant
(reprinted 2001). A Flora of New Mexico, Vol. Handbook. Dover Publications Inc., New York.
1. Bishen Singh Mahendra Pal Singh (India) and 816 p.
Koeltz Scientific Books (Germany). 1,276 p.
Weber, W.A. and R.C. Wittman. 2001a. Colorado
Mozingo, H.N. 1987. Shrubs of the Great Basin: Flora: Eastern Slope. University Press of
A Natural History. University of Nevada Press, Colorado, Boulder CO. 521 p.
Reno, NV. 342 p.
Weber, W.A. and R.C. Wittman. 2001b. Colorado
Newsholme, C. 1992. Willows: The Genus Salix. Flora: Western Slope. University Press of
Timber Press, Portland, OR. 224 p. Colorado, Boulder CO. 488 p.
Ottenbreit, K.A. and R.J. Staniforth. 1992. Life Young, J.A. and C.G. Young. 1992. Seeds of
cycle and age structure of ramets in an Woody Plants in North America. Dioscorides
expanding population of Salix exigua (sandbar Press, Portland, Oregon. 407 p.
667
Salix scouleriana Barratt ex Hook. Scouler’s willow
SALICACEAE
Synonyms: Salix brachystachys Benth.

Salix capreoides Anderss.
Salix flavescens Nutt.
Salix nuttallii Sarg.
Salix stagnalis Nutt.
John K. Francis
female flowers on different trees. Tiny flowers are

grouped in “pussy willow”-like catkins. The
anthers, two per flower, are yellow, sometimes
tipped with red; pistols are red. Fruits are light
reddish-brown, long-pointed capsules about 0.75
cm long. At maturity, they open to release a white
fluff with imbedded tiny seeds. The species has 2n
= 76 or 114 chromosomes (Anonymous 2003,
Department of Ecology 2003, Harrington 1964,
Peattie 1953, Sargent 1923, Viereck and Little
1972).
Range.—Scouler’s willow occurs from south

central Alaska to central Manitoba, in the Black
Hills of South Dakota, in the Rocky Mountains
south to New Mexico and Arizona, and along the
coast through British Colombia, Washington,
Oregon, and California (Natural Resources
Conservation Service 2003, Peattie 1953,
Treeguide.com 2003).
Illustration source: USDA-Forest Service Collection,
Hunt Institute Ecology.—Scouler’s willow is the most common
upland willow through most of its range. It invades
General Description.—Scouler’s willow, also quickly and abundantly after fires and logging
known as fire willow, Nuttall willow, mountain (Viereck and Little 1972). Mineral soil seedbeds
willow, and black willow, is a deciduous shrub or are required for seedling establishment (Forest
small tree, depending on the environment. It Practices Branch 1997). In northern areas,
usually has multiple stems that reach 2 to 7 m in Scouler’s willow occurs in muskegs, willow
height in dry, cold, high elevation, and other thickets, disturbed areas, and forests (Welsh 1974).
difficult environments, and 10 to 20 m in favorable At lower latitudes, the species grows in former
sites. The stems are straight and support few clearcuts, burned areas, thinned forests, and areas
branches generally resulting in narrow crowns. of natural disturbance such as avalanche areas and
The root system is fibrous, deep, and widespread. river flood zones. These are all moist, well-drained
The thick sapwood is nearly white, and heartwood to poorly drained sites. Although this willow
is light brown tinged with red. Stem bark is thin, tolerates drier conditions than most other willows,
gray or dark brown with broad, flat ridges. Twigs it does not tolerate xeric conditions. Scouler’s
are stout and whitish green. Leaves are willow is a component in a large number of
oblanceolate to elliptic, 5 to 12.5 cm long, mostly vegetation types throughout its range (Anderson
short-pointed at the apex and tapered toward the 2001). With few exceptions, it is the only willow
base with entire to sparsely wavy-toothed margins. found growing with other trees in upland Western
The leaves are dark-green and nearly hairless forests (Johnson 1995). Soils of all textures,
above and white- or grayish-hairy below. including skeletal soils and soils derived from
Scouler’s willow is dioecious, having male and most parent materials are colonized. Sites may
668
vary from near sea level to about 3,000 m in temperatures. It is an important browse species for
elevation (Peattie 1953). Scouler’s willow is top- domestic livestock and wild animals. Cattle, sheep,
killed by all but gentle fires, but usually greater and goats all like it as browse. It is sometimes the
than 65 percent of the plants sprout quickly most preferred food species for white tailed, black
afterwards (Anderson 2001). The species is tailed and mule deer, elk, moose, and bighorn
intolerant of shade, and when overtopped by sheep. Small mammals, bears, upland game birds,
conifers and other hardwoods, it begins to decline. and waterfowl feed to a lesser extent on leaves,
buds, and seeds. Fresh browse (twigs and leaves)
Reproduction.—Scouler’s willow flowers from contain 41 percent dry matter, 4 percent protein, 2
April through June, flowers appearing before percent fat, 20.8 percent nitrogen-free extract, 11.2
leaves, often while snow is still on the ground, and percent crude fiber, and good quantities of mineral
fruiting occurs from May through July, depending nutrients (Anderson 2001). The cover provided by
on area. The flowers are insect pollinated. There Scouler’s willow is important for mammals and
are about 14,300 cleaned seeds/g. Germination, birds. The flowers provide pollen and nectar to
which is epigeal, begins to occur in 12 to 24 hours honey bees in early spring (Anderson 2001). The
after seeds alight on wet ground. Germination wood, which is soft and close-grained, is not sawn
usually reaches 95 percent in 1 or 2 days into lumber but is used to a limited extent for
(Brinkman 1974). The seeds are dispersed by the firewood and wood carving (Viereck and Little
wind. Plants sprout from the root collar when cut 1972). The Secwepemc people of British
or top-killed. Pieces of stem and root will root and Colombia used Scouler’s willow wood for
grow if partially buried in moist soil (Forest smoking fish, drying meat, and constructing
Practices Branch 1997). fishing weirs, the inner bark for lashing, sowing,
cordage, and headbands, and decoctions of twigs
Growth and Management.—Annual height for treating pimples, body odor, and diaper rash
growth of sprouts from cut stems varies from 1 to (Secwepemc Cultural and Education Society
3 m/year. Up to 60 sprouts are produced per stem 2003).
(Forest Practices Branch 1997). Maximum height
at 20 years is about 9 m. At higher elevations, References
shrubs reach 4 to 5 m in 15 years after which
growth slows until a maximum height of 10 m is Anderson, M.D. 2001. Salix scouleriana. In: U.S.
reached (Natural Resources Conservation Service Department of Agriculture, Forest Service,
2003). Fruits should be collected by hand or with Rocky Mountain Research Station, Fire Sciences
pruning poles as soon as they turn from green to Laboratory, Missoula, MT. http://www.fs.
yellow. The capsules are air-dried until opening. fed.us/database/feis/plants/tree/salsco/all.html.
Generally, the seeds should be sown as soon as 57 p.
possible because they remain viable for only a few
days. Seed can be stored in sealed containers under Anonymous. 2003. Treatment from the Jepson
refrigeration for 4 to 6 weeks, but germination manual: Salicaceae. University of California,
begins to drop rapidly after 10 days. Seeds are Berkeley, CA. http://ucjeps.berkeley.edu/chi-
broadcast on well-prepared beds that are kept bin/get_JM_treatment.pl?7043,7050,7079. 3 p.
continually moist until germination and seedling
emergence. Light is required for successful Brinkman, K.A. 1974. Salix L., willow. In: C.S.
germination (Brinkman 1974). Recommended Schopmeyer, tech. coord. Seeds of woody plants
spacing using rooted cuttings for erosion control is in the United States. Agriculture Handbook 450.
1.8 m by 1.8 m; for unrooted whips or shorter U.S. Department of Agriculture, Forest Service,
cuttings, 0.6 m. Rooted cuttings can be grown to 3 Washington, DC. p. 746-750.
m tall in containers. Cuttings should be 45 to 60
cm long, and whips (not recommended) should be Department of Ecology. 2003. Plant selection
1.2 m long (Department of Ecology 2003). guide. State of Washington, Department of
Ecology, Olympia, WA. http://www.ecy.wa.
Benefits.—Scouler’s willow protects the soil and gov/programs/sea/pubs/93-30/table3.html. 6 p.
helps return sites to forest cover following
disturbance. When growing along streams, it helps Forest Practices Branch. 1997. Operational
protect the stream banks from erosion and shade summary for vegetation management: willow
the watercourse, thus maintaining cooler water complex. ISBN 0-7726-3166-2. Forest Practices
669
Branch, Ministry of Forests, Victoria, British Secwepemc Cultural and Education Society. 2003.
Colombia, Canada. 11 p. Flood plain garden. Secwepemc Nation,
Kamloops, BC, Canada. http://www.secwepemc.
Harrington, H.D. 1964. Manual of the plants of org/SECethnogardens2/flood_plain_garden.htm.
Colorado. Sage Books, Denver, CO. 666 p. 4 p.
Johnson, F.D. 1995. Wild trees of Idaho. Treeguide.com. 2003. Scouler willow, Salix
University of Idaho Press, Moscow, ID. 212 p. scouleriana Barratt ex Hook. http://www.
treeguide.com/Species.asp?Region=NorthAmeri
Natural Resources Conservation Service. 2003. can&SpeciesID=996. 2 p.
Plants profile: Salix scouleriana Barratt ex
Hook. http://plants.usda.gov/chi_bin/ Viereck, L.A. and E.L. Little, Jr. 1972. Alaska
plant_search.cgi?mode=Scientific+Name&keyw trees and shrubs. Agriculture Handbook 410.
ordquery=Salix+scouleriana&earl=plant_search. Forest Service, U.S. Department of Agriculture,
4 p. Washington, DC. 265 p.
Peattie, D.C. 1953. A natural history of western Welsh, S.L. 1974. Anderson’s flora of Alaska and
trees. Houghton Mifflin Company, Boston, MA. adjacent parts of Canada. Brigham Young
751 p. University Press, Provo, UT. 724 p.
Sargent, C.S. 1923. Manual of the trees of North

America (exclusive of Mexico). Houghton
Mifflin, Boston, MA. 910 p.
670
Salvia apiana Jepson white sage
LAMIACEAE
Synonyms: Audibertia polystachya Benth.

Ramona polystachya Briq.
Audibertiella polystachya Briq.
Salvia californica Jepson not Bdg.
Arlee M. Montalvo
side, convex on the other side, and often grey-

brown to light brown.
Range.—White sage ranges from Santa Barbara

County, California south to the middle of Baja
California and east to the western edge of the
Colorado desert and is an important component of
inland and coastal sage scrub vegetation, with peak
abundance in Riverside, Orange, and San Diego
Counties, California. It is found on dry slopes and
benches in many plant communities below 1500 m
(occasionally higher), including chaparral, coastal
sage scrub, yellow-pine forest, and the upper edges
of desert scrub (Munz and Keck 1968).
General Description.—White sage is a rounded, 1
to 2.5 m tall shrub of low elevations that has Ecology.—Plants often thrive in droughty soils
highly aromatic, 3 to 9 cm long, whitish to pale and in habitats with high summer temperatures.
gray-green leaves that persist throughout the year. Westman (1981a) reports occurrence on coarse to
The thick, petioled, lance-oblong leaves have loamy soils derived from sandstone, conglomerate
crenulate margins and are covered with short and sandstone, shale, granitic/dioritic rocks, and
appressed hairs and oil glands that give the leaves volcanic rocks. In western Riverside County,
a silvery sheen. The attractive white to pale plants also occur on serpentine outcrops (author’s
lavender flowers with short greenish calyx are observation). White sage tends to occur with
clustered in compact cymose branches of a tall Artemisia californica Less. and Eriogonum
graceful, often pink-tinged inflorescence (thyrsoid fasciculatum Benth. Plants are notably drought
panicle) that extends from 0.5 to 1.5 m above the tolerant even though roots are thought to be less
foliage (Munz and Keck 1968). The lower lip of than 1.5 m deep (Hellmers and others 1955). The
the two-lipped corolla is long and ruffled and crowded, whitish leaves are often held vertically
obstructs the corolla tube, whereas the upper lip is and become folded during drought, likely
reduced to a small lobe. Anthers release most adaptations for keeping leaves cool. In addition
pollen prior to maturity of stigmas. The two many leaves dehisce late in the dry season. In a
stamens are exerted to about 15 mm, diverging up desert wash in Baja California during summer
and outward from their insertion on the lower lip. drought, leaf conductance, transpiration rates, and
The exerted stigma protrudes forward about 15 turgor potentials were high, and plants did not
mm beyond the floral tube. There is a north to show signs of drought stress (Schmitt and others
south and east to west geographic gradient in floral 1993). In chaparral, white sage had higher water
morphology (Meyn and Emboden 1987). For potentials throughout the day well into the summer
example, longer lips occur toward the coast. On drought than did most of the evergreen shrubs
the desert edge, there is a form with more compact measured (Poole and Miller 1975). Water
inflorescences called S. apiana var. compacta potentials decreased late in the dry season and
Munz (Munz and Keck 1968). Plants have n = 15 recovered after November at the end of the
chromosomes (Epling and others 1962). The four seasonal drought. The roots of white sage form
nutlets are about 2 to 3 mm long, somewhat associations with arbuscular mycorrhizal fungi that
rectangular, narrow in cross-section, keeled on one may also assist plants in low nutrient and droughty
671
habitats (personal communication, James Beaver, year (Storms 1999). Seedling survival is expected
Indiana University, Bloomington). to be much lower in dry years and weedy areas.
Fire is a normal disturbance factor of most Most growth occurs after winter rains begin and
plant communities within which white sage continues until the soil becomes dry in summer.
occurs. Following fire, this shrub frequently Leaf growth over the season results in a seasonal
sprouts from distinct basal burls (Keeley 1998), dimorphism in leaf size (Westman 1981b). Smaller
but year-old seedlings are killed by fire, and leaves are produced on short shoots that develop
repeated burning kills mature plants (Zedler and from leaf axils along the main elongated stems and
others 1983). Seedlings can emerge from the are often retained when the older, longer leaves
seedbank in the first season after fire, but reported dehisce during drought. With the return of
densities are low (Zedler and others 1983, Keeley seasonal rains, the short shoots elongate into long
1998), and high intensity fires can kill seeds shoots, and the retained short leaves continue to
(Keeley and Fotheringham 1998). expand and subtend the next generation of short
shoots and leaves.
Reproduction.—Inflorescences arise from the
upper nodes in spring and produce flowers from Hybridization.—Epling (1938) reports that white
late May to July. Based on the floral morphology, sage hybridizes with other Salvias with the same
only some of the larger bees are capable of chromosome number, including S. mellifera
transferring pollen to the highly exerted stigmas. Greene, S. munzii Epl., S. leucophylla Greene, S.
Grant and Grant (1964) report three species of clevelandii (Gray) Greene, S. eremostachya Jeps.,
Xylocopa and a species of Bombus as effective S. pachyphylla Epl. Ex Munz., and S. vaseyi
pollinators. Hummingbirds, honeybees, bombyliid (Porter) Parish. Hybrid zones between S. mellifera
flies, and small species of native bees (primarily (black sage) and white sage are especially
Anthophora, Diadasia, and Osmia) sometimes common. Their hybrids are fully viable but, on
visit flowers (Grant and Grant 1964) but are average, suffer reduced fertility relative to parental
thought to be ineffective as pollinators. The species (Epling 1947a, Meyn and Emboden 1987),
protandrous flowers and presence of interspecific and there are fewer seeds/flower in F1 backcrosses
hybrids are consistent with an outcrossing to both parental species (Grant and Grant 1964).
breeding system. Inflorescences are large and Hybrids exhibit a range of intermediate floral and
many-flowered, making some self-pollination leaf traits (Epling 1947a, b, Anderson and
likely. The small nutlets fall from the dried calyx Anderson 1954) and differ in leaf anatomy (Webb
during mid to late summer but are secondarily and Carlquist 1964). Most hybrids are thought to
dispersed by harvester ants and seed caching be F1 crosses or backcrosses to black sage and
rodents. Seed traps revealed that seeds dispersed they tend to grow close to the parental types.
up to 3 m away and that there was a higher Although black and white sage overlap in range,
secondary removal rate of seeds in coastal sage they have different habitat affinities. Their ranges
scrub (80 percent) than in grassland (0 percent) or overlap in most of coastal southern California, but
the ecotone of the two habitats (about 40 percent) white sage is not found north of Santa Barbara
(DeSimone and Zedler 2001). County and it ranges farther south into Baja
California and eastward into the edge of the desert
Establishment and Growth.—Seedlings emerge than does black sage. Both species can occur
in the cool wet season in late winter to early spring intermingled, but black sage tends to be found in
and can take 2 years or more to reach maturity, flatter and wetter areas, while white sage is usually
depending on rain patterns and growing on drier slopes (Epling 1947a, Anderson and
conditions. Seedling establishment in shrublands is Anderson 1954, Grant and Grant 1964, Gill and
associated with gaps in mature vegetation and Hanlon 1998). As such, most sites contain a single
within grasslands in areas where herbs have been species, but hybrids are present at most sites where
removed by gophers (DeSimone and Zedler 1999, the species co-occur (Meyn and Emboden 1987).
2001). Seedlings show a relatively high growth Authors have speculated about factors that may
rate, little herbivore damage and they fold their limit the success of hybrids and maintain black and
leaves in response to drought (DeSimone and white sage as distinct species. Epling (1947a) first
Zedler 2001). In plots that were hand weeded and suggested that hybrids were less adapted to wet
seeded in a twice-normal rainfall year, about a sites than black sage and less adapted to dry sites
quarter of the seedlings survived to the second than white sage, but that they may be better
672
adapted to recently disturbed conditions. Hybrids trimming back the dried inflorescence stalks to
usually occur on recently disturbed areas, encourage compact growth. Plants can be easily
suggesting that hybrids are better adapted to started from seeds in the fall or planted from
intermediate habitats (Anderson and Anderson containers.
1954, Meyn and Emboden 1987). Within a natural
hybrid zone, Gill and Hanlon (1998) found that Growth and Management.—Because of its
xylem pressure potential was significantly higher historical abundance, fast growth, attractive nature,
in white than in black sage, that hybrids were and importance to the biodiversity of sensitive
intermediate to the two parental species, and that sage scrub habitat, white sage is used in many
white sage drops fewer leaves during drought. restoration and roadside revegetation projects,
These data support that white sage is more drought especially in endangered sage scrub vegetation.
adapted than black sage. It is possible that Both container plants and seeds are used
selection will maintain the integrity of the hybrid successfully in dry-land restoration in southern
zones if the intermediate hybrids do better in California. Seeds are collected from the dry,
intermediate habitats relative to their performance fruiting inflorescences, usually in July and August,
in the main habitats of the parental species. depending on site, and processed to remove chaff
The potential for reproductive isolation of the and insects. S&S Seeds (Carpenteria, CA) and
species within areas of overlap has been examined Mirov and Krabel (1939) report 715,000 to
most thoroughly by Grant and Grant (1964), 750,200 bulk seeds/kg, with often 70 percent
augmenting earlier observations (Epling 1947a, purity and 50 percent germination. Seeds should
Anderson and Anderson 1954). Black sage be planted before the onset of the cool rainy
generally flowers earlier than white sage and the season. If seed germination tests show low
short overlap in flowering time of one to several germination but high seed viability, seeds can be
weeks limits hybridization. In addition, white sage treated with dry or liquid smoke before sowing. If
has large, highly modified flowers that are liquid smoke is used, seeds must be air-dried prior
pollinated almost exclusively by large carpenter to dry broadcasting. Dormant seeds can result in a
and bumblebees (Xylocopa and Bombus), while beneficial seed bank and establishment over
black sage has smaller, unmodified flowers that several years. Hydroseeding can be successful on
are pollinated primarily by honeybees and small steep slopes and road cuts. Shallow planting
native solitary bees. methods such as hydroseeding and imprinting are
likely to produce higher germination than drilling
Seed germination.—Keeley (1987) showed that because seed germination is improved with
exposure to 70 oC for 1 hour followed by exposure to light. Based on seedling survival data
incubation in light substantially increased (Storms 1999, DeSimone and Zedler 2001), even
germination over controls, that heat shock for 5 under well-weeded and rainy conditions, fewer
minutes at 100 oC had no effect, and that exposure than 25 percent of seedlings should be expected to
for 5 min at 120 oC killed seeds. Leachate from survive through their first year. This species has
charred wood did not increase germination decreased dramatically in percent cover in western
(Keeley 1987), but cool smoke treatment increased Riverside County since the early 1930’s (Minnich
germination (Keeley and Fotheringham 1998). and Dezzani 1998). Successful restoration will
Liquid smoke can also be used to significantly depend on controlling the factors associated with
increase germination (author’s observation). its decrease, especially invasive plants and fire
Horticulture.—The generally rounded form with frequency, so that shrubs can establish, mature,
attractive whitish leaves and long graceful, pink- and buildup a seed bank.
tinged inflorescence stalks make this an attractive
shrub for background areas of dry landscape Benefits.—White sage leaves are an important
gardens (Perry 1992, Keator 1994, Clebsch 1997). browse of mountain sheep in the winter and spring
Plants require full sun, well-drained soil, do not (Perry and others 1987). Leaves contain essential
tolerate constantly damp conditions, and are oils and a variety of diterpines and triterpenes,
sensitive to conditions below –7 oC (Keator 1994, including carnosic acid, oleaolic acid, and ursolic
Clebsch 1997), although populations in desert acid, some of which have been linked to the
scrub are likely to be more cold tolerant than those medicinal use of white sage by native Tribes.
from coastal locations. Plants should be kept dry Some may also deter many herbivores (Dentali
all summer long. Clebsch (1997) recommends and Hoffman 1992). A tincture of the leaves has
673
been used internally as a diaphoretic and diuretic, Epling, C., H. Lewis, and P.H. Raven. 1962.
and externally to wash skin. The tincture may have Chromosomes of Salvia: section Audibertia.
antimicrobial properties. Antibacterial and Aliso 5: 217-221.
antifungal activity of several compounds have
been verified in vitro (Dentali and Hoffman 1992). Gill, D.S. and B.J. Hanlon. 1998. Water potentials
In addition to medicinal use, the dried leaves have of Salvia apiana, S. mellifera (Lamiaceae), and
been used as a smudge and crushed as a soapless their hybrids in the coastal sage scrub of southern
shampoo (Bean and Saubel 1972, Moore 1989). California. Madroño 45: 141-145.
Seeds have been used as food and leaves for
flavoring. Grant, K.A. and V. Grant. 1964. Mechanical
isolation of Salvia apiana and Salvia mellifera
References (Labiatae). Evolution 18: 196-212.
Anderson, E. and B.R. Anderson. 1954. Hellmers, H., J.S. Horton, G. Juhren, and J.
Introgression of Salvia apiana and Salvia O'Keefe. 1955. Root systems of some chaparral
mellifera. Annals of the Missouri Botanical plants in southern California. Ecology 36: 667-
Garden 41: 329-338. 678.
Bean, J.L. and K.S. Saubel. 1972. Temalpakh: Keator, G. 1994. Complete Garden Guide to the
Cahuilla Indian Knowledge and Usage of Plants. Native Shrubs of California. Chronicle Books,
Malki Museum Press, Morongo Indian San Francisco, CA. 314 p.
Reservation, CA. 225 p.
Keeley, J.E. 1987. Role of fire in seed germination
Clebsch, B. 1997. A Book of Salvias: Sages for of woody taxa in California chaparral. Ecology
Every Garden. Timber Press Inc., Portland, OR. 68: 434-443.
221 p.
Keeley, J.E. 1998. Postfire ecosystem recovery and
Dentali, S.J. and J.J. Hoffmann. 1992. Potential management: The October 1993 large fire
antiinfective agents from Eriodictyon episode in California. In: J. M. Moreno (ed.)
angustifolium and Salvia apiana. International Large Forest Fires. Backbuys Publishers, Leiden,
Journal of Pharmacognosy 30: 223-231. Netherlands. p. 69-90.
DeSimone, S.A. and P.H. Zedler. 1999. Shrub Keeley, J.E., and C.J. Fotheringham. 1998. Smoke-
seedling recruitment in unburned Californian induced seed germination in California
coastal sage scrub and adjacent grassland. chaparral. Ecology 79: 2,320-2,336.
Ecology 80: 2,018-2,032.
Meyn, O. and W.A. Emboden. 1987. Parameters
DeSimone, S.A. and P.H. Zedler. 2001. Do shrub and consequences of introgression in Salvia
colonizers of southern Californian grassland fit apiana X S. mellifera (Lamiaceae). Systematic
generalities for other woody colonizers? Botany 12: 390-399.
Ecological Applications 11: 1,101-1,111.
Minnich, R.A. and R.J. Dezzani. 1998. Historical
Epling, C. 1938. The California salvias. Annals of decline of coastal sage scrub in the Riverside-
the Missouri Botanical Garden 25: 95-188. Perris plane, California. Western Birds 29: 366-
391.
Epling, C. 1947a. The genetic aspects of natural
populations: actual and potential gene flow in Mirov, N.T. and C. J. Kraebel. 1939. Collecting and
natural populations. The American Naturalist 81: handling seeds of wild plants. Civilian
104-113. Conservation Corps, Forestry Publication No. 5,
United States Government Printing Office,
Epling, C. 1947b. Natural hybridization of Salvia Washington, D.C. 42 p.
apiana and S. mellifera. Evolution 1: 69-78.
674
Moore, M. 1989. Medicinal Plants of the Desert wash in Baja California, Mexico. Journal of Arid
and Canyon West. Museum of New Mexico Environments 24: 155-164.
Press, Santa Fe, NM. 184 p.
Storms, N. 1999. Restoration of a Native Shrubland
Munz, P.A. and D.D. Keck. 1968. A California in an Area of Frequent Disturbance and High
Flora with Supplement. University of California Nitrogen Deposition. M.S. Thesis. University of
Press, Berkeley, CA. 1681 + 224 p. California, Riverside. 60 p.
Perry, B. 1992. Landscape Plants for Western Webb, A.-A. and S. Carlquist. 1964. Leaf anatomy
Regions: an Illustrated Guide to Plants for Water as an indicator of Salvia apiana-mellifera
Conservation. Land Design Publishing, introgression. Aliso 5: 437-449.
Claremont, CA. 318 p.
Westman, W.E. 1981a. Factors influencing the
Perry, W.M., J.W. Dole, and S.A. Holl. 1987. distribution of species of California coastal sage
Analysis of the diets of mountain sheep from the scrub. Ecology 62: 439-455.
San Gabriel Mountains, California. California
Fish and Game 73: 156-162. Westman, W.E. 1981b. Seasonal dimorphism of
foliage in Californian coastal sage scrub.
Poole, D.K. and P.C. Miller. 1975. Water relations Oecologia 51: 385-388.
of selected species of chaparral and coastal sage
communities. Ecology 56: 1,118-1,128. Zedler, P.H., C.R. Gautier, and G.S. McMaster.
1983. Vegetation change in response to extreme
Schmitt, A.K., C.E. Martin, V.S. Loeschen, and A. events: the effect of a short interval between fires
Schmitt. 1993. Mid-summer gas exchange and in California chaparral and coastal scrub.
water relations of seven C3 species in a desert Ecology 64: 809-818.
675
Salvia mellifera Greene black sage
LAMIACEAE
Synonyms: Audibertia stachyoides Benth.

Ramona stachyoides Briq.
Audibertiella stachyoides Briq.
Arlee M. Montalvo. and Paul A. McMillan
dark brown and are about 1 mm wide and 2 mm

long.
Range.—Black sage occurs on dry slopes and

alluvial fans within interior and coastal sage scrub
and lower montane chaparral from sea level to
1,200 m, ranging from Contra Costa County, CA
(near San Francisco) south to northern Baja
California (Munz and Keck 1968, Hickman 1993).
On leeward slopes of the Coast Ranges it extends
into desert scrub (Keeley 1986). It is one of the
dominant shrubs of sage scrub (Westman 1983),
and can form dense, nearly monospecific stands on
steep slopes.
Ecology.—Black sage occurs on a variety of soils

derived from sandstone, shale, granite, and
especially serpentinite, and gabbro basalt
(Westman 1981). Plants produce numerous
volatile compounds, including camphor, cineole,
terpenes, and sesquiterpenes (Neisess and others
General Description.—Black sage has a dark 1987, Gonzalez and others 1992, Arey and others
appearance, especially during drought. Salvia is 1995), which may be important in herbivore or
derived from the Latin word for safe and refers to pathogen defense.
the medicinal properties of sages. The specific
Black sage is semideciduous, depending on
epithet “mellifera” means “honey producing” and
site and severity of drought, and shallow rooted.
refers to its use by nectar foraging bees. Shrubs are
Plants are drought tolerant by leaf curling rather
open, 1 to 2 m tall with twisted, furrowed woody
than drought avoiding through leaf drop (Gill and
stems at the base, and herbaceous greenish branch
Hanlon 1998). Leaf drop is drought induced
tips that are square in cross section. Twigs and
possibly due to embolisms that occur in xylem
leaves are glandular, highly aromatic, and
tissues (Kolb and Davis 1994). At Stanford, CA,
opposite. The oblong-elliptical blades are 2 to 6
leaves began to senesce and drop in October in
cm long, dark green and rugulose above (convex
both irrigated and unirrigated treatments, leaf
between obvious veins) with crenulate (finely
stomata did not control transpiration when xylem
scalloped) margins, and taper to a few to 12 mm
water potentials were low, and rates of
long petiole (Munz and Keck 1968). Many small,
photosynthesis did not decline until after leaf drop
6- to 12-mm long, two-lipped flowers occur in
(Gigon 1979). In Santa Barbara County, CA, most
compact clusters subtended by green bracts and are
leaves senesced between May and July, 32 percent
spaced 2 to 6 cm apart along the often branched
persisted into the following growing season, leaf
inflorescence. Corollas vary from pure white to
curling and uncurling was related to water stress,
pale blue or lavender. The style and stamens
and leaf drop was more related to photoperiod
extend just beyond the corolla tube. Flowers bear
than to low xylem potentials (Gill and Mahal
four oblong nutlets (single–seeded fruits) within a
1986).
persistent calyx. Nutlets vary from mottled gray to
676
Reproduction.—The timing of growth and occurred (E. Allen, L. Egerton-Warburton, A.
flowering is closely adapted to the Mediterranean Montalvo, unpublished report).
climate of southern California, with cool wet
winters and hot dry summers (Grant and Grant Seed germination.—Seeds of black sage often
1964, Gill and Mahall 1986). Common garden have low germination rates unless exposed to light
studies with 12 populations revealed that date of or components of fire such as charred wood,
first flowering varies with source population and smoke, or KNO3 (Keeley 1986, Thanos and
ranged from early February to early May Rundel 1995, Keeley and Fotheringham 1998).
(Montalvo, unpublished data). Also, shifts in Exposure to dry cool smoke for 5 min (Keeley and
flowering time occur in years with contrasting Fotheringham 1998) yields higher germination
rainfall patterns (Meyn and Emboden 1987). than charred wood. A 12 to 15 hour soak in a 1:25
The self-compatible flowers are dilution of Regen 2000 Smokemaster, a liquid
hermaphroditic with stamens dispersing pollen smoke product, also breaks dormancy (author’s
before stigmas are receptive. Flowers are observation). Heat of fire may reduce germination
pollinated by small to meduim-sized solitary bees of seeds from sage scrub populations (Keeley and
in families Andrenidae, Anthophoridae, Fotheringham 1998). Oddly, seeds from desert
Halictidae, Megachilidae, and Xylocopinae as well populations, where fires are less frequent, were
as introduced honey bees (Grant and Grant 1964). stimulated to germinate by heat and charred wood,
Rarer visitors include large-bodied Bombus and but not by light (Keeley 1986).
Xylocopa bees, syrphid and bombyliid flies, and
Anna’s hummingbirds. Each flower produces up to Genetics, Geographic Variation, and Fitness.—
four seeds that are gravity dispersed in June and Black sage is genetically variable over its
July from the dry, persistent calyces, and can be geographic range. In a survey of 12 populations
secondarily dispersed by ants. Seeds accumulate in (Montalvo, Clegg, and Ellstrand unpublished, 14
a dormant seed bank (Keeley 1986). allozyme loci), expected heterozygosity was high
(He = 0.23) and alleles averaged 2.81 per locus.
Establishment and Growth.—Black sage occurs Within sites, a low inbreeding coefficient (f =
in plant communities adapted to fire and drought. 0.072) suggests the protandrous flowers and
Seedlings emerge in clearings between adult pollinators combine to promote outcrossing. There
shrubs, but most emerge in the first 2 years is significant but low structure both among and
following fire, primarily from February to April within populations, and results are consistent with
(Keeley 1986, Westman and O’Leary 1986). a relatively high level of historical gene flow
Plants take two or more growing seasons to reach (Theta-p (~FST, sites relative to total population) =
maturity. After fire, plants sometimes resprout 0.041; Theta-s (plots relative to sites) = 0.057; and
from the base (Went and others 1952, Keeley F (FIT) = 0.125).
1986, 1998). However, where fire intensity is high, Populations cluster by genetic similarity, and
shrubs tend to be killed, and recruitment is geographic floristic associations and this is
primarily by seed. Across a range of conditions, somewhat mirrored by distribution patterns of
resprouting success averaged 15.8 percent, and morphological variation and timing of flowering.
shrubs that resprouted had significantly smaller Flowers from populations in the Santa Monica
basal diameters, suggesting resprouting decreases Mountains and Simi Hills can be strikingly bluish.
with age (Keeley 1998). The seasonal pattern of In the drier, more interior hills and valleys of
growth and dormancy varies both geographically Riverside Co., flowers tend to be white to pale
and between years. Generally, seasonal growth blue or lavender, and plants flower earlier. These
begins after the start of fall rains, most new leaves differences are retained in common gardens.
are produced by mid March, and flowering occurs Differences in timing of flowering could affect
from late winter to late spring. seed production of translocated populations if
Roots are colonized by arbuscular pollinator activity, seed predation, or important
mycorrhizal (AM) fungi. After 4 months of growth physical environmental factors differ from home
in a greenhouse, root and shoot mass was higher in sites. Plants from contrasting populations also
seedlings inoculated with AM fungi than in smell different, likely due to differences in
controls, but in younger plants the opposite composition of volatile chemicals. Such
differences among populations may be adaptive
677
and contribute to their success under particular Within a hybrid zone, Gill and Hanlon (1998)
environments. Montalvo and Ellstrand found that xylem pressure potential was
(unpublished) evaluated seedling survival, growth significantly higher in white than in black sage and
and reproduction of 12 populations in a common that putative hybrids were intermediate to the two
garden. A combination of climatic data and soil parental species. These data support that S. apiana
traits was used to calculate environmental is more drought adapted than S. mellifera, and that
similarity among garden and source sites. After the hybrids are intermediate. In addition,
three growing seasons, populations from sources reproductive isolation is favored by flowering
most similar to the garden site were the most differences. S. apiana flowers later than black
successful (measured as survival x flower sage, limiting opportunities to hybridize (Grant
production). This effect was small (r2 = 0.18) but and Grant 1964). In addition, white sage has
is consistent with the presence of weak adaptive larger, highly modified flowers that are pollinated
differences among source populations. almost exclusively by large bees in Xylocopa and
Bombus. Black sage has smaller, two-lipped
Hybridization.—Black sage hybridizes with the flowers that are rarely visited by these large-
shrubs S. apiana Jeps. (n = 15), S. leucophylla bodied bees.
Greene (2n = 30), and S. clevelandii (Gray) Greene
(2n = 30), and rarely with annuals S. columbariae Growth and Management.—Seeds are collected
Benth (n = 13) and S. carduacea Benth (n = 16) from June-August and should be cleaned to
(Epling 1938, 1947a, Epling and others 1962, remove seed predators. Air separation can be used
Munz and Keck 1968). Most hybridization is with to remove lighter unfilled seeds that are sometimes
S. apiana. These hybrids are fully viable but suffer abundant. There are about 1,375,000 seeds/bulk kg
reduced pollen fertility (Epling 1947a, Meyn and (personal communication with Victor Schaff, S&S
Emboden 1987) and numbers of seeds/flower in Seeds, Carpenteria, CA). Treatment of seeds with
F1 back crosses to parental species (Grant and dry or liquid smoke increases germination
Grant 1964). Hybrids differ from both parental substantially. Seeds should be air-dried before
species by a range of intermediate floral, leaf, and handling. Plants can be sown in flats for
anatomical traits (Epling 1947a, b, Anderson and subsequent transplanting or directly sown into the
Anderson 1954, Webb and Carlquist 1964). Most ground in the fall. Plants can be started from
are thought to be F1s or backcrosses to S. mellifera cuttings, but this should be avoided for restoration
and tend to grow near parental types. Hybrid zones because it reduces genetic variation. Prostrate
can be stable in position but may shift in cultivars (e.g., Clebsch 1997) should not be used
morphology (Meyn and Emboden 1985). This in restoration. Black sage grows in well-drained
suggests differences in the relative success of soil and can tolerate some summer water in a
hybrids and backcrossed progeny over time. The garden.
influence of gene exchange on floral form of black
sage does not appear to extend beyond hybrid Benefits.—Black sage is important for restoration,
zones (Epling 1947a, b). erosion control, and native landscape gardening in
Several authors have speculated on the California. Plants provide cover and seeds for
factors promoting hybridization while also wildlife, and in western Riverside Co. the listed
maintaining black and white sage as distinct California gnatcather frequently nests in
species. While ranges overlap in most of southern vegetation containing black sage (Weaver 1998).
California, S. apiana is not found north of Santa The seeds were eaten and the leaves used for
Barbara County and it ranges farther south into flavoring by native California tribes (Bean and
Baja California and eastward. In areas of contact, Saubel 1972). In addition, the leaves have a long
S. mellifera tends to occur in flatter and wetter history of medicinal use by native tribes (Bocek
microsites, while S. apiana occurs on drier slopes 1984). Their diterpene compounds have
(Epling 1947a, Anderson and Anderson 1954, antimicrobial activity against gram (+) bacteria
Grant and Grant 1964, Meyn and Emboden 1987, (Moujir and others 1996).
Gill and Hanlon 1998). Hybrids also occur in
recently disturbed areas, suggesting that hybrids Decreasing Populations.—Black sage
are adapted to intermediate habitats (Anderson and populations are declining. Increased ignition
Anderson 1954, Meyn and Emboden 1987). sources and invasion of shrublands by annual
grasses have increased fire frequency, jeopardizing
678
recovery of seed banks (Westman and O’Leary their hybrids in the coastal sage scrub of
1986, Haidinger and Keeley 1993, Minnich and southern California. Madroño 45: 141-145.
Dezzani 1998). Black sage is also inhibited by
exotic annual grasses, black mustard (Went and Gill, D.S. and B.E. Mahall. 1986. Quantitative
others 1952, Storms 1999), and air pollution phenology and water relations of an evergreen
(Westman 1985, Preston 1988). and deciduous chaparral shrub. Ecological
Monographs 56: 127-143.
References
Anderson, E. and B. R. Anderson. 1954. González, A.G., L.S. Andrés, Z.E. Aguiar, and
Introgression of Salvia apiana and Salvia J.G. Luis. 1992. Diterpenes from Salvia
mellifera. Annals of the Missouri Botanical mellifera and their biogenetic significance.
Garden 41: 329-338. Phytochemistry 31(4): 1,297-1,305.
Arey, J., D.E. Crowley, M. Crowley, M. Resketo, Grant, K.A. and V. Grant. 1964. Mechanical
and J. Lester. 1995. Hydrocarbon emissions isolation of Salvia apiana and Salvia mellifera
from natural vegetation in California's south (Labiatae). Evolution 18: 196-212.
coast air basin. Atmospheric Environment 29:
2,977-2,988. Haidinger, T.L. and J.E. Keeley. 1993. Role of
high fire frequency in destruction of mixed
Bean, J. and K. Saubel. 1972. Temalpakh: Cahuilla chaparral. Madroño 40: 141-147.
Indian knowledge and usage of plants. Malki
Museum Press, Morongo Indian Reservation, Hickman, J.C., editor. 1993. The Jepson Manual:
CA. 225 p. Higher Plants of California. University of
California Press, Ltd., Los Angeles, CA. 1,400
Bocek, B. 1984. Ethnobotany of Costanoan p.
Indians, California, based on collections by
John P. Harrington. Economic Botany 38: 240- Keeley, J.E. 1986. Seed germination patterns of
255. Salvia mellifera in fire-prone environments.
Oecologia 71: 1-5.
Clebsch, B. 1997. A Book of Salvias: Sages for
Every Garden. Timber Press, Portland, OR. 221 Keeley, J.E. 1998. Postfire ecosystem recovery
p. and management: The October 1993 large fire
episode in California. In: J. M. Moreno, ed.
Epling, C. 1938. The California salvias. Annals of Large Forest Fires. Backbuys, Leiden, The
the Missouri Botanical Garden 25: 95-188. Netherlands. p. 69-90.
Epling, C. 1947a. The genetic aspects of natural
populations: actual and potential gene flow in Keeley, J.E. and C.J. Fotheringham. 1998. Smoke-
natural populations. The American Naturalist induced seed germination in California
81: 104-113. chaparral. Ecology 79: 2,320-2,336.
Epling, C. 1947b. Natural hybridization of Salvia Kolb, K.J. and S.D. Davis. 1994. Drought
apiana and S. mellifera. Evolution 1: 69-78. tolerance and xylem embolism in co-occurring
Epling, C.,H. Lewis, and P.H. Raven. 1962. species of coastal sage and chaparral. Ecology
Chromosomes of Salvia: section Audibertia. 75: 648-659.
Aliso 5: 217-221.
Meyn, O. and W.A. Emboden. 1987. Parameters
Gigon, A. 1979. CO2-gas exchange, water and consequences of introgression in Salvia
relations and convergence of Mediterranean apiana X S. mellifera (Lamiaceae). Systematic
shrub-types from California and Chile. Botany 12: 390-399.
Oecologia Plantarum 14: 129-150.
Minnich, R.A. and R.J. Dezzani. 1998. Historical
Gill, D.S. and B.J. Hanlon. 1998. Water potentials decline of coastal sage scrub in the Riverside-
of Salvia apiana, S. mellifera (Lamiaceae), and Perris plane, California. Western Birds 29: 366-
391.
679
Moujir, L., A.M. Gutiérrez-Navarro, L. San Weaver, K.L. 1998. Coastal sage scrub variations
Andres, and J.G. Luis. 1996. Bioactive of San Diego County and their influence on the
diterpenoids isolated from Salvia mellifera. distribution of the California gnatcatcher.
Phytotherapy Research 10: 172-174. Western Birds 29: 392-405.
Munz, P.A. and D.D. Keck. 1968. A California Webb, A.-A. and S. Carlquist. 1964. Leaf anatomy
Flora with Supplement. University of California as an indicator of Salvia apiana-mellifera
Press, Berkeley, CA. 1,681 + 224 p. introgression. Aliso 5: 437-449.
Neisess, K.R., R.W. Scora, and J. Kumamoto. Went, F.W., G. Juhren, and M.C. Juhren. 1952.
1987. Volatile leaf oils of California Salvias. Fire and biotic factors affecting germination.
Journal of Natural Products 50: 515-517. Ecology 33: 351-364.
Preston, K.P. 1988. Effects of sulphur dioxide Westman, W.E. 1981. Factors influencing the
pollution on a Californian coastal sage scrub distribution of species of California coastal sage
community. Environmental Pollution 51: 179- scrub. Ecology 62: 439-455.
195.
Westman, W.E. 1983. Xeric Mediterranean-type
Storms, N. 1999. Restoration of a native shrubland shrubland associations of Alta and Baja
in an area of frequent disturbance and high California and the community/continuum
nitrogen deposition. Master of Science in debate. Vegetatio 52: 3-19.
Botany Thesis. University of California,
Riverside. 60 p. Westman, W.E. 1985. Air pollution injury to
coastal sage scrub in the Santa Monica
Thanos, C.A. and P.W. Rundel. 1995. Fire- Mountains, southern California. Water, Air, and
followers in chaparral: nitrogenous compounds Soil Pollution 26: 19-41.
trigger seed germination. Journal of Ecology
83: 207-216. Westman, W.E. and J.F. O'Leary. 1986. Measures
of resilience: the response of coastal sage scrub
to fire. Vegetatio 65: 179-189.
680
Sassafras albidum (Nutt.) Nees sassafras
LAURACEAE
Synonyms: Sassafras sassafras (L.) Karst.

Laurus albidus Nutt.
Laurus sassafras L.
Sassafras officinale Nees & Eberm.
Sassafras variifolium (Salsb.) Kuntze
Sassafras albidum (Nutt.) Nees var. molle (Raf.) Fern.
John K. Francis
side lobe (“mitten”), and trilobate leaves with the

center lobe being symmetrical and somewhat
larger than the side lobes. Leaves are 8 to 15 cm
long. Small yellowish-green flowers grow in
axillary racemes. The fruit is a fleshy, oblong, dark
blue or black drupe partially set in a red or orange
thickened base (pedicel and calyx). The seeds are
oblong, pointed, and brown (Griggs 1990, Sargent
1923, Vines 1982).
Range.—Sassafras is native to all of the States

east of a line running between Lake Superior and
western Texas, and in extreme southern Ontario
(Natural Resources Conservation Service 2003,
Sullivan 1993). It is planted as an ornamental in at
least the United Kingdom, Germany, and Spain
(Oberle 2003, Plants for a Future 2003, Sanchez de
Lorenzo 2003). There are no reports of it
naturalizing outside its native range. The genus
contains two other species, natives of China and
Taiwan (Center for Wood Anatomy Research
2003)
Ecology.—Sassafras is intolerant of shade, and

reproduction from seed is rare under closed-
General Description.—Sassafras is also known as canopy forest. However, it is a pioneer in old
white sassafras, common sassafras, ague tree, fields and an invader of disturbed and early
cinnamon wood, saloop, smelling stick, gumbo secondary forests. It grows as scattered individual
file, and mitten tree. It is a shrub or small tree up plants or as dense thickets, often of sucker origin.
to 15 m in height and 35 cm in diameter. The Removal of the forest overstory by almost any
species grows as a medium to large shrub in means (i.e. gypsy moth, logging, fire) results in
Florida, in the northern part of its range, and on increased density of the species. On the other
sand or poor, rocky sites. It may have single or hand, as competition increases and forest canopies
multiple stems. Older stems have deeply furrowed, close, sassafras decreases in importance. Although
red-brown bark. Slender, brittle, twigs with thick, uncommon, it persists in old-growth stands. The
white, mucilaginous pith are covered by green species grows from a few meters above sea level
bark. The species depends primarily on a shallow, to 1,220 m (Sullivan 1993). Mean annual
lateral root system. Long lateral roots develop precipitation ranges from 760 to 1400 mm/year
suckers that in turn develop their own root system. and the average frost-free period ranges from 160
The alternate dark green leaves occur in three to 300 days. Average January temperatures range
forms: ovate or elliptical leaves, leaves with one from -7 to 13 °C and average July temperatures
681
range from 21 to 27 °C. (Griggs 1990). Sassafras Benefits.—Sassafras adds beauty in both summer
grows in a wide variety of soil types and grows and winter to forest lands where it grows. It has
best on well-drained sandy loams with a pH been used to help restore depleted soils of old
between 6.0 and 7.0 (Griggs 1990). Seedlings and fields and is considered superior to black locust
saplings are easily top-killed by fire but readily (Robinia pseudoacacia L.) and pines in this
sprout. Thick-barked trees are injured by fire application (Grigg 1990). Sassafras is browsed
(Sullivan 1993). Sassafras is sometimes attacked year-round by white tailed deer. The plants are
by leaf blight caused by Actinopelte dryina, by leaf also utilized to some extent by woodchucks,
spots caused by Mycosphaerella sassafras, and by rabbits, beavers, and bears. Many species of birds
a Nectria canker. A number of insects also attack and some small mammals consume the fruits.
the species (Griggs 1990). Crude protein ranges from 21.0 percent (leaves in
April) to 6.1 percent (twigs in January). The fruits
Reproduction.—Flowers open in early spring have a high lipid and energy value (Sullivan
with the first leaves. Male and female flowers 1993). Sassafras makes an attractive ornamental
usually occur on different plants (Sargent 1923). small tree and is widely, although not heavily
Fruits mature in August or September. Seed used. The fall foliage colors of yellow, orange, and
production begins when plants are about 10 years red are particularly beautiful. Sassafras heartwood
old (Griggs 1990). There are about 12,800 is pale brown to orange brown; sapwood is
seeds/kg (Bonner and Maisenhelder 1974) or yellowish-white. The wood is ring-porous, coarse-
according to Griggs (1990), 8,800 to 13,200 but straight-grained, brittle, soft, and has a spicy
seeds/kg. About 35 percent of the seeds are sound. aromatic odor. It is resistant to decay making it
Birds are the principal dispersers of seeds, with useful for fence posts and other items exposed to
gravity, water, and mammals playing minor roles. moisture. It is also useful for lumber, millwork,
Germination, which occurs in the spring, is furniture, and small boats. Technical details of the
hypogeal. Seeds may remain viable in the soil seed wood have been published (Center for Wood
bank for up to 6 years (Griggs 1990). Sassafras Anatomy Research 2003). The use of sassafras in
sprouts prolifically from stumps and suckers from herbal medicine has a long history beginning with
lateral roots. Native Americans who used it to treat colds, high
blood pressure, heart troubles, swelling, as a tonic,
Growth and Management.—Growth of sassafras to treat for worms, and to control fever (Moerman
is moderately fast. Sprouts can reach 1.2 m in the 1986). Sassafras tea is still widely drunk both for
first year and 4.5 m in 4 years (Floridata 2003). pleasure and as a tonic. Extracts and infusions of
Rare individual trees reaching the forest canopy the plant are used to treat a wide range of physical
may live as long as 300 years (TreeGuide 2003). complaints. Dry, powdered leaves, called file, are
Fruits are ripe and can be harvested when they turn used to thicken gumbo and other Cajun dishes
from green to dark blue and can be picked by hand (Katzer 1999). Oil of sassafras, extracted from the
or by flailing them from the trees or shrubs onto a root bark, was the original flavoring for root beer
tarp. The seeds can be cleaned of fruit tissue by and was used to flavor candy, chewing gum, and
rubbing against hardware cloth and washing away medicines. Because it contains safrole, a
the pulp. One kg of fruits will yield about 310 g of carcinogen, it is no longer permitted in food, but
seeds. Seeds should be placed in sealed containers may be used to add scent to cosmetics and soap
and stored at 2 to 5 °C. Before planting, seeds (TreeGuide 2003).
should be moistened and held at the above
temperatures for 120 days. Drilling in nursery beds References
in 20- to 30-cm-wide rows and covering with 6 to
12 mm of soil is recommended (Bonner and Bonner, F.T. and L.C. Maisenhelder. 1974.
Maisenhelder 1974). Sassafras can also be Sassafrass albidum (Nutt.) Nees sassafras. In:
reproduced from root cuttings but not from stem C.S. Schopmeyer, tech. coord. Seeds of woody
cuttings (Griggs 1990). Wildlings are difficult to plants in the United States. Agriculture
transplant. Sassafras shrubs and trees can be killed Handbook 450. U.S. Department of Agriculture,
by injecting 2,4-D, picloram, or glyphosate Forest Service, Washington, DC. p. 761-762.
(Sullivan 1993).
Center for Wood Anatomy Research. 2003.
Sassafras albidum. Technical Transsfer Fact
682
Sheet. U.S. Department of Agriculture, Forest Oberle, R. 2003. Sassafras albidum (Nutt.) Nees.
Service, Forest Products Laboratory, Madison, http://www.biologie.uni-ulm.de/systax/
Wisconsin. http://www2fpl.fs.fed.us/ dendrologie/sasalblv.htm. 1 p.
TechSheets/HardwoodNA/htmlDocs/sassafrasal
bi.html. 3 p. Plants For a Future. 2003. Plant portrait-Sassafras
albidum. http://www.scs.leeds.ac.uk/pfaf/
Floridata. 2003. Sassafras albidum. http:// sassafr.html. 3 p.
www.floridata.com/ref/s/sass_alb.cfm. 4 p.
Sanchez de Lorenzo-C., J.M. 2003. Árboles
Griggs, M.M. 1990. Sassafras albidum (Nutt.) ornamentals: Sassafras albidum (Nutt.) Nees.
Nees, sassafras. In: R.M. Burns, and B.H. http://www.arbolesornamentales.com/Sassafrasa
Honkala. Silvics of North America. Vol. 2., lbidum.htm. 1 p.
Hardwoods. Ag. Handb. 654. U.S. Department
of Agriculture, Forest Service, Washington, DC. Sargent, C.S. 1923. Manual of the trees of North
p. 773-777. America (exclusive of Mexico). Houghton
Mifflin, Boston, MA. 910 p.
Katzer, G. 1999. Sassafras [Sassafras albidum
(Nutt.) Nees]. http://www-ang.kfunigraz. Sullivan, J. 1993. Sassafras albidum. In: U.S.
ac.at/~katzer/engl/Sass_alb.html. 4 p. Department of Agriculture, Forest Service,
Moerman, D.E. 1986. Medicinal plants of Native Laboratory, Fire Effects Information System.
America. Technical Reports 19. University of http://www.fs.fed.us/database/feis/plants/tree/
Michigan Museum of Anthropology, Ann sasalb/all.html. 17 p.
Arbor, MI. 534 p.
TreeGuide. 2003. Sassafras. http://www.
Natural Resources Conservation Service. 2003. treeguide.com/Species.asp?SpeciesID=953. 3 p.
Plants profile: Sassafras albidum (Nutt.) Nees.
http://plants.usda.gov/cgi_bin/plant_profile.cgi? Vines, R.A. 1982. Trees of North Texas.
symbol=SAAL5. 6 p. University of Texas Press, Austin, TX. 466 p.
683
Schaefferia frutescens Jacq. Florida-boxwood
CELASTRACEAE
Synonyms: Schaefferia completa Sw.

Schaefferia frutescens Jacq. var. buxifolia DC.
Schaefferia buxifolia (DC.) Nutt.
John K. Francis
ends. They measure 4 to 7 cm in length and 1.2 to
2.5 cm in width. The flowers are small, pale green
or greenish-yellow or greenish-white, and borne in
small clusters at the leaf axiles. The fruits that
arise from them are spherical to ovoid, 4 to 8 mm
long, and red or orange-red. Each fruit contains
two light-brown seeds (Howard 1989, Little and
others 1974, Liogier 1994, Nelson 1996).
Range.—Florida-boxwood is native to southern

Florida, the Bahamas, the West Indies, Mexico
through Central America, Colombia, Venezuela,
and Ecuador (Liogier 1994). It is not known to
have naturalized outside its native range.
Ecology.—Florida-boxwood grows in soils

ranging from sandy to clayey and from mildly
alkaline to moderately acid derived from both
sedimentary and igneous rocks. In Puerto Rico,
the species inhabits areas receiving from 750 to
about 1700 mm of mean annual precipitation and
from near sea level to 600 m in elevation. In the
upper range of rainfall, it grows only on
excessively drained sites. Florida-boxwood is
General Description.—Florida-boxwood is a moderately shade tolerant. It is usually found in
shrub or occasionally a small tree. Florida- the understory of low-density forest where it
boxwood is also known as yellowwood, jiga, grows well and reproduces. After the forest
cafeíllo, cabra cimarrona, amansa, guapo, jasmín, overstory is removed, Florida-boxwood grows
limoncillo, fruta de paloma, petit bois blanc, bois well in full sun. In Puerto Rico, the species grows
capable, petit garcon, and merisier (Little and in remnant dry and moist forests and middle and
others 1974). It has multiple stems, branching near late secondary forests. In Florida, it grows in
the base, or a single stem, and usually reaches tidewater areas and hammocks. The species is
heights of 4 to 5 m. However, the Florida record drought tolerant and somewhat resistant to soil
tree measures 8 m in height and 18 cm diameter at salinity (Gilman 1999). Although the species is
breast height (Champion Tree Project 2001). The common in Puerto Rico and in much of its range,
bark is light gray and nearly smooth, becoming it is considered endangered in natural
increasingly furrowed in older shrubs and trees. environments in Florida (Institute of Systematic
The inner bark is light yellow. The roots are bright Botany 2001).
orange. There is a weak taproot supporting robust
and stiff lateral roots and fine 3rd and 4th order Reproduction.—Florida-boxwood flowers in
roots. Twigs of Florida-boxwood are slender, March through May in Florida (Nelson 1996).
slightly angled, hairless and pale yellow-green Fruits collected in Puerto Rico averaged 0.142 +
when young, turning light gray with age. The 0.002 g/fruit. Seeds separated from them averaged
branch habit is upright to partially drooping. The 0.011 + 0.000 g/seed or 98,000 seeds/kg. Twenty-
leaves are yellow-green, alternate, simple, one percent of these seeds germinated between 62
leathery, and elliptic to oval shaped with pointed
684
and 93 days of sowing in moist peat. Germination Cooperative Extension Service. 3 p.
is epigeal. The fruits are apparently eaten and the
seeds dispersed by birds. Howard, R.A. 1989. Flora of the Lesser Antilles,
Growth and Management.—Florida-boxwood is Arboretum, Harvard University, Jamaica Plain,
reported to have a high growth rate, withstands MA. 604 p.
heavy pruning, and can be formed into hedges and
shaped into small trees (Gilman 1999). Institute of Systematic Botany. 2001. Atlas of
Florida Plants: Schaefferia frutescens.
Benefits.—Florida-boxwood makes superior University of South Florida. http://
quality hedges, screens, and accent plantings. It is plantatlas.usf.edu/main.asp?plantID==2227. 1 p.
also used for reclamation plantings (Gilman
1999). The wood is light brown to yellow, hard Liogier, H.A. 1990. Plantas medicinales de Puerto
and moderately heavy. It has been used in the past Rico y del Caribe. Iberoamericana de Ediciones,
for wood engraving (Little and others 1974). It is Inc., San Juan, PR. 566 p.
also used, when available, for carving, for boxes
(Gilman 1999), and, to a limited extent, for fuel. Liogier, H.A. 1994. Descriptive flora of Puerto
The leaves have been used in the past as Rico and adjacent Islands. Vol. 3. Editorial de la
emergency soap for the body and cloths (Liogier Universidad de Puerto Rico. Río Piedras, PR.
1990). Almost nothing is known about its forage 461 p.
value except that Key deer (Odocolleus virginanus
clavium) will not eat it (Schaus and others 2001). Little, E.L., Jr., R.O. Woodbury, and F.H.
Decoctions are used in herbal medicine for colds, Wadsworth. 1974. Trees of Puerto Rico and the
flu, and chronic cough, and crushed leaves are Virgin Islands. Vol. 2. Agriculture Handbook
used as a soothing bath for skin irritations (Liogier 449. U.S. Department of Agriculture,
1990). Washington, DC. 1,024 p.
References Nelson, G. 1996. The shrubs and woody vines of

The Champion Tree Project. 2001. National 391 p.
champion trees: Florida. http://www.
championtrees.org/database/championsFL.htm. Schaus, C., S. Wade, and J. Dunan. 2001. Key
8 p. deer and plants they won’t eat.
http://monroe.ifas.ufl.edu/key_deer_plants.htm.
Gilman, E.F. 1999. Schaefferia frutencens. Fact 3 p.
Sheet FPS-540. University of Florida,
685
Schinus terebinthifolius Raddi Brazilian pepper tree
ANACARDIACEAE
Synonyms: Sarcotheca bahiensis Turcz.

Schinus antiarthriticus Mart. ex Marchand
Schinus mellisii Engl.
Schinus mucronulatus Mart.
Rhus terebinthifolia Schlecht. & Cham.
John K. Francis
small white flowers. Male and female flowers are

borne on different plants (dioecious). The fruits
are bright-red, fleshy drupes 4 to 6.5 mm in
diameter with an aromatic brown pulp and an
elliptic light-brown stone (Liogier 1988, Long and
Lakela 1976, National Parks Service 2002).
Range.—Brazilian pepper tree is a native of

Brazil, Argentina, and Paraguay (Liogier 1988). It
was introduced into Florida in 1898 and has since
been distributed widely. The species has escaped
cultivation and occupies huge areas in natural
stands (Nelson 1996). It has also naturalized in
Arizona, California, Hawaii, Puerto Rico, the U.S.
Virgin Islands, and many tropical and subtropical
countries (Little and others 1974, National Parks
Service 2002).
Ecology.—Brazilian pepper tree is an aggressive

pioneer species that quickly colonizes disturbed
areas. The species has an intermediate tolerance of
shade and can survive and grow slowly under
forest canopies until disturbance releases it.
General Description.—Brazilian pepper tree, also Although seedlings are quickly killed by
known as Christmas berry, Florida holly, pimienta inundation, large plants can withstand up to 6
de Brazil, copal, chichita, and aroeira pimenteira, months of flooding. It is very drought resistant.
is a multiple-stemmed evergreen shrub or small Brazilian pepper tree also survives fire well and
tree, usually 2 to 6 m in height and 3 to 12 cm in can withstand high winds without significant
stem diameter. A few individuals of the species damage. It is apparently at home in tropical,
develop large central stems and a dense growth of Mediterranean, and desert climates (National
low limbs and basal sprouts. The bark is smooth Parks Service 2002). In Puerto Rico, the species is
and gray. Wounds in the stem and twigs exude a most common in low-elevation, moist limestone
resinous sap that turns black on exposure to air. areas and nearby coastal plains. In Florida, it
The alternate, compound leaves have a rachis 3 to grows in mangrove associations, hammocks,
14 cm long, often winged, with five to nine pinelands, old fields, and disturbed areas (Long
leaflets. The leaflets are 1.5 to 7.5 cm long, and Lakela 1976).
lanceolate to elliptic, pointed at each end, with
entire to serrate edges. The midribs, rachis, and Reproduction.—Brazilian pepper tree flowers
petiole are often reddish, especially when young. most prolifically in the spring in Florida (Nelson
Crushed foliage smells like turpentine. The 1996). It flowers and fruits intermittently
inflorescences (panicles) are mostly born in the throughout the year in Puerto Rico (Little and
leaf axils near the twig ends and contain many others 1974). In Florida, the flowers are pollinated
by a native syrphid fly (Fire Sciences Laboratory
686
2002). Seeds from Brazil were reported to average plant species in suppressing several important
80,600 seeds/kg (Instituto de Pesquisas e Estudos pathogenic bacteria (Martínez and others 1996,
Florestais 2002). Germination of intact fruits Siddiqui and others 1995). Despite these benefits,
(without the exocarp digested or manually Brazilian pepper tree is often undesirable outside
removed) is minimal, and seeds do not remain its native range. It is so well adapted and
viable in the soil bank more than 9 months aggressive in Florida and Hawaii that it suppresses
(Panetta and Mckee 1997). Birds, particularly and replaces native vegetation, including
robins, mockingbirds, and cedar waxwings, are the endangered species. For this reason it is banned
chief dispersers of seeds in Florida (Fire Sciences from cultivation there (Fire Sciences Laboratory
Laboratory 2002). Brazilian pepper tree is 2002, Nelson 1996). Brazilian pepper tree is
propagated from both seeds and cuttings (Little reported to have an allelopathic effect on
and others 1974). It readily sprouts when cut. competing plants (National Parks Service 2002).
The plant causes skin irritation similar to poison
Growth and Management.—Brazilian pepper ivy (Toxicodendron radicans Linn.) and
tree is fast growing and in certain circumstances is respiratory difficulties in some people when it is in
capable of growing to large diameters. The U.S. bloom (Nelson 1996). It is a host of the black twig
Champion tree in Florida measures 134 cm in borer of coffee, Xylosandrus compactus (Eichoff)
diameter at breast height and 10.7 m in height (College of Tropical Agriculture and Human
(American Forests 2002). Brazilian pepper tree Resources 2002). The fruits are eaten by mammals
has shown large growth responses to inoculation and birds, but excessive feeding has been blamed
with mycorrhizal fungi and superphosphate for massive bird kills in Florida (National Parks
(Carneiro and others 1996). Hand pulling, Service 2002), and the unripe fruits can be fatal to
bulldozing, prescribed fire, and spraying with horses (Fire Sciences Laboratory 2002).
herbicides have all been used with varying degrees
of success to control Brazilian pepper tree. So far, References
biological control methods have failed (National
Parks Service 2002). American Forests. 2002. National register of big
trees. American Forests, Washington, DC. http://
Benefits and Detriments.—Widely introduced as www.americanforests.org/resources/bigtrees/
an ornamental, Brazilian pepper tree was popular regester.php?details=2297. 1 p.
for its red berries and bright green foliage. It is
still used for Christmas decorations. Because it Carneiro, M.A.C., J.O. Siqueira, A.C. Davide, L.J.
and the species growing in its understory do not Gomes, N. Curi, and F.R. do Vale. 1996.
burn readily (Fire Sciences Laboratory 2002), Mycorrhizal fungi and superphosphate and the
Brazilian pepper tree has been recommended for growth of tropical woody species. Scientia
planting as fire resistant barriers (Castronovo Forestalis 50: 21-36.
1997). Goats browse on the foliage with no ill
effects (National Parks Service 2002). The species Castronovo, T. 1997. Fire resistive landscaping
is a honey plant (Little and others 1974) and the can save your house and your life.
wood has been used for fuel, lumber, stakes, posts, http://www.themastergardenershow.com/
and railway sleepers (Fire Sciences Laboratory fire_resistive_landscaping_.htm. 5 p.
2002). The dried fruits are marketed in Brazil as a
substitute for black pepper (Piper nigrum) College of Tropical Agriculture and Human
(personal communication with J.A. Parrotta, Resources. 2002. Crop knowledge master:
USDA Forest service, Washington, DC). A Xylosandrus compactus (Eichoff). University of
decoction of bark is used in baths to relieve Hawaii. http://extento.hawaii.edu/kbase/Type/
rhumatisum and back pain (Liogier 1990). The xylosand.htm. 4 p.
essential oils from leaves and flowers are made up
of a mixture of a large number of chemicals, the
Fire Sciences Laboratory. 2002. Fire effects
most important of which are alpha-pinene (24
information database: Schinus terebinthifolius.
percent), limonene (12 percent), and p-cymene (14
Rocky Mountain Research Station, Missoula,
percent) (Singh and others 1998). Extracts of
MT. http://www.fs.fed.us/database/feis/plants/
Brazilian pepper tree were shown to be the most
tree/ schter/all.html. 9 p.
effective of a number aromatic and medicinal
687
Instituto de Pesquisas e Estudos Florestais. 2002. National Parks Service. 2002. Exotic weeds I.
Tabla de preços de sementes de espécies nativas http://www.nature.nps.gov/wv/ipm/exweeds1.ht
e exóticas. http://www.ipef.br/especies/ m. 9 p.
natexoticas.asp. 2 p.
Liogier H.A. 1988. Descriptive flora of Puerto Florida. Pineapple Press, Inc., Sarasota, FL.
Rico and adjacent islands, Spermatophyta. Vol. 391 p.
Río Piedras, PR. 481 p. Panetta, F.D. and J. McKee. 1997. Recruitment of
the invasive ornamental, Schinus
Liogier, H.A. 1990. Plantas medicinales de Puerto terebinthifolius, is dependent upon frugivores.
Rico y del Caribe. Iberoamericana de Ediciones, Australian Journal of Ecology 22(4): 432-438.
Siddiqui, R.R., Uzma-Zafar, S.S. Chaudhry, and
Little, E.L., Jr., R.O. Woodbury, and F.H. Hamid-Ahamad. 1995. Antimicrobial activity of
Wadsworth. 1974. Trees of Puerto Rico and the essential oils from Schinus terebinthifolius,
Virgin Islands. Vol. 2. Agriculture Handbook Cypress sempervirens, Citrus limon, Ferula
449. U.S. Department of Agriculture, assafoetida. Part 1. Pakistan Journal of
Washington, DC. 1,024 p. Scientific and Industrial Research. 38(9-10):
358-361.
Tropical Florida. University of Miami Press, Singh, A.K., J. Singh, K.C. Gupta, and J.J.
Coral Gables, FL. 962 p. Brophy. 1998. Essential oil of leaves and
inflorescence of Schinus terebinthifolius: an
Martínez, M.J., J. Betancourt., N. A. González., exotic plant of India. Journal of Essential Oil
and A. Jauregui. 1996. Screening of some Cuban Research 10(6): 697-699.
medicinal plants for antimicrobial activity.
Journal of Ethnopharmacology 52(3): 171-174.
688
Securidaca virgata Sw. bejuco de sopla
POLYGALACEAE
Synonyms: Elsota virgata (Sw.) Kuntze
John K. Francis
variety of soils over both sedimentary and igneous

rocks at lower and middle elevations. Annual
rainfall in its native areas ranges from about 900
to 2500 mm/year. It sustains itself on other
vegetation by twining around stems and branches
(Vélez and van Overbeek 1950). Although it
depends on trees and shrubs for support, it does
not smother them, tending to hang down from side
branches. Bejuco de sopla is intolerant of shade
and does not grow under closed, shady forest
canopies. It grows in brushy pastures, old fields,
fencerows, roadsides, canyons, gallery creek
bottoms, secondary forest, and low-density
remnant forests. The species does not form pure
stands or dense thickets, although is may grow in
mixture with other vines and shrubs in thickets.
General Description.—Bejuco de sopla, also
known as jaboncillo and maravedí, is a woody Reproduction.—Bejuco de sopla flowers and
vine-like shrub that ascends trees and other fruits sporadically throughout the year (Acevedo-
vegetation and can reach 9 m in length and 3 cm Rodrígiez 1985, Vélez and van Overbeek 1950). A
in basal stem diameter. The plants have a tap and collection of fruits from Puerto Rico weighed an
lateral root system of tough and flexible roots with average (air-dried) of 0.0728 + 0.0011 g/fruit. The
a moderate amount of fine roots. The stems give acorn-like seeds separated from the wing weighed
rise to few branches until they reach full or nearly an average of 0.0455 + 0.0012 g/seed or 22,000
full light after which there are abundant secondary seeds/kg. Sown in moist potting mix, 22 percent of
and tertiary branches. The greenish, cylindrical these seeds germinated in 32 days by which time
woody stems have pronounced annual rings. The some of the seedlings began to die from
fine branches are green and produce relatively few “damping-off” (fungal species unknown). The
leaves, which they lose during dry seasons. survivors were 10 cm tall 2 months after sowing
Foliage is found only on the current year’s growth. and ready to prick out. Fruits of bejuco de sopla
The alternate leaves are oval, have a 1 mm petiole, are mature and ready to harvest when they are dry
and are 10 to 20 mm long by 8 to 15 mm broad. on the tips or have a yellow color. They should
Small, 8 mm-long, pink to violate, fragrant then be air-dried before storage or planting. If the
flowers are borne near the ends of the branches in seeds are separated, it must be done by hand, a
paniculate racemes. The fruit, a reticulate-veined laborious process that probably is not necessary.
samara 2 to 4 cm long and about 1 cm broad, turns Bejuco de sopla vines layer (root) whenever they
from green to yellow to light tan as it ripens, and come in contact with the ground. Although
contains a single seed (Acevedo-Rodrígiez 1985, untested, this probably indicates that the species
author’s observations, Grisebach 1963, Liogier can be propagated by air layering and from
1988). cuttings. The seeds are dispersed as they spiral
sideways in the wind or still air. Seedlings are
Range.—Bejuco de sopla is native to Puerto Rico, usually well scattered.
its offshore island, Vieques, as well as Jamaica,
and Cuba (Acevedo-Rodrígiez 1985, Grisebach Growth and Management.—Bejuco de sopla has
1963, Liogier 1988). a moderate growth rate. Height growth of
established plants is 1 m or more per year and
Ecology.—Bejuco de sopla grows on a wide diameter growth is about 3 mm/year. Individual
689
stems live 5 to 10 years or more; plants may last P. Lohr, and D. Atha. 2000. Medicinal plants
longer by resprouting. Bejuco de sopla is not used by Latino healers for woman’s health
abundant, aggressive, or weedy. Should it be conditions in New York City. Economic Botany
necessary to eliminate individual plants, grubbing 54(3): 344-357.
them out will probably be effective.
Grisebach, A.H.R. 1963. Flora of the British West
Benefits.—Bejuco de sopla contributes to Indian Islands. J. Cramer-Weinheim, New York.
biodiversity, helps protect the soil, and furnishes 789 p.
wildlife cover. It is a honey plant (Marcano-F.
1973). The roots, principally, are used to treat flu, Liogier, H.A. 1988. Descriptive flora of Puerto
laryngitis, edema, urinary infections, and as an Rico and adjacent islands, Spermatophyta. Vol.
expectorant and a diuretic (Liogier 1990). The 2. Editorial de la Universidad de Puerto Rico,
plant is prescribed by traditional Latin healers in Río Piedras, PR 481 p.
New York City to treat uterine fibroids (Balick
and others 2000). Liogier, H.A. 1990. Plantas medicinales de Puerto
References Inc., San Juan, PR. 566 p.
Acevedo-Rodríguez, P. 1985. Los bejucos de Marcano-F., E. de J. 1973. La flora apícola de la

Puerto Rico. Vol. 1. General Technical Report República Dominicana. http://marcano.
SO-58. U.S. Department of Agriculture, Forest freeservers.com/nature/estudios/apicola/floraap1
Service, Southern Forest Experiment Station, .html. 7 p.
Vélez, I. and J. van Overbeek. 1950. Plantas
Balick, M.J., F. Kronenberg, A.L. Ososki, M. indeseables en los cultivos tropicales. Editorial
Reiff, A. Fugh-Berman, B. O’Connor, M. Roble, Universitaria, Río Piedras, PR. 497 p.
690
Senecio flaccidus Less. threadleaf groundsel
ASTERACEAE
Synonyms: Senecio douglasii DC.
James E. Nellessen
synonyms; S. douglasii DC var. longilobus

(Benth.) L. Benson, S. filifolius Nutt., S. flaccidus
Nutt. var. jamesii Torr. & Gray, and S. longilobus
Benth. This variety tends to be more uniformly
white hairy with the column of the flower head
narrower (8 to 10 mm wide at summit). Variety 2:
S. flaccidus Less. var. douglasii (DC) B. Turner &
T. Barkley, with two synonyms; S. douglasii DC
var. douglasii, and S. warnockii Shinners. This
variety is not white hairy throughout and the heads
are wider (10 to 15 mm). Variety 3: S. flaccidus
Less. var. monoensis (Greene) B. Turner & T.
Barkley, with two synonyms; S. douglasii DC var.
monoensis (Greene) Jeps., and S. monoensis
Greene. This variety is generally hairless
General Description.—Threadleaf groundsel is throughout, more herbaceous, and has a more
also known as threadleaf butterweed, Douglas comb-like leaf, the linear lobes curling upward.
groundsel or ragwort, felty groundsel, sand wash
groundsel, comb butterweed, old man, yerba cana, Range.—Threadleaf groundsel occurs from
squawweed, creek senecio, and cenicillo (Elmore western Texas, west through New Mexico,
1976, Epple 1995, Stubbendieck and others 1997). Arizona, to California, north to Wyoming and
This species commonly reaches heights of 80 to Nebraska, Utah, and Nevada, and south into
130 cm and under ideal conditions may reach 2 m. northern Mexico. It is a wide ranging species
It is probably best classified as a subshrub. It occurring in a variety of soil types and elevations
forms woody stems to a meter or so but often dies from about 760 to 2,440 m.
back to less than 50 cm from the ground-line. It
tends to become somewhat straggly as branches Ecology.—This species grows in short grass
may lean over in different directions, some prairies of the southwestern Great Plains and a
becoming almost decumbent. The leaves are wide variety of scrubland and grassland habitats in
alternate, pinnately compound (sometimes not the Southwest. It has been considered a diagnostic
divided), narrow and thread-like, up to 12 cm in or climax species for desert grassland (Dick-
length, and usually covered with whitish hairs Peddie 1993). It tends to become more abundant in
giving the plant an obvious grayish color (Carter disturbed habitats and overgrazed rangelands and
1997, Ivey 1995, Kearney and others 1951, Martin often grows in washes. It is a taprooted plant that
and Hutchins 1980-81). It may have some grows relatively quickly and tends to be a short-
tendencies to be evergreen in the southern portion lived perennial shrub to subshrub. Oxygen flux, as
of its range. it relates to photosynthesis, from leaves and twigs
was constant with increasing drought stress, up
Taxonomy.—There has been a profusion of until leaf drop occurred (Ehleringer and Cooper
taxonomic names and varietal subclassifications 1992). Twigs may serve to maintain some
attached to this species, making for a complicated photosynthesis through drought periods when new
taxonomy. The oldest specific name applied to this foliage cannot easily be produced, such as during
species complex is now known to be Senecio dry springs and early summers. Threadleaf
flaccidus Lessing (Turner and Barkley 1990). groundsel was one of several shrub and subshrub
Three varieties are recognized with the following species, studied in Mojave and Sonoran desert
synonymies (Allred 2002, Kartesz 1994). Variety areas that generated substantial net carbon gains
1: S. flaccidus Less. var. flaccidus, with four
691
from twig photosynthesis (Comstock and others singing during some ceremonial songs (Elmore
1988). 1976). The Navajo would also use the flower
heads to clean cactus fruits of their spines.
Reproduction.—Threadleaf groundsel generally
blooms in the late summer and fall but will bloom References
earlier in the summer if sufficient moisture is
available. It produces numerous yellow flower Allred, K.W. 2002. A Working Index of New
heads with eight to 17 ray florets, the rays 10 to 15 Mexico Vascular Plant Names. New Mexico
mm long (Carter 1997, Martin and Hutchins 1980- State University. http://web.nmsu.edu/~kallred/
81). Seeds (fruits) are wind-borne with a white herbweb/ [not paged].
fluffy pappus.
Carter, J.L. 1997. Trees and Shrubs of New
Growth and Management.—Threadleaf Mexico. Johnson Books, Boulder, CO. 534 p.
groundsel is a fast growing, short lived perennial.
Plants may live 3 to 6 years or more. It increases in Comstock, J.P., T.A. Cooper, and J.R. Ehleringer.
rangelands as a result of overgrazing pressure. Its 1988. Seasonal patterns of canopy development
abundance may be controlled through proper and carbon gain in nineteen warm desert shrub
grazing management and careful use of herbicides. species. Oecologia, Berlin 75(3): 327-335.
Spraying with 2,4-D has been used to control this
species. Cooper, R.A., R.J. Bowers, C.A. Beckham, and
R.J. Huxtable. 1996. Preparative separation of
Benefits and Disadvantages.—Because this pyrrolizidine alkaloids by high-speed counter
species will colonize open disturbed areas, it helps current chromatography. Journal of
achieve a quick ground cover in a natural Chromatography 732(1): 43-50.
succession process. The perennial nature of the
plant further helps stabilize soil for longer-lived Dick-Peddie, W.A. 1993. New Mexico
perennials to eventually become established. Vegetation: Past, Present and Future. University
Direct human uses are minimal, but it was once of New Mexico Press, Albuquerque, NM. 244 p.
used medicinally by Native Americans. It is a poor
forage plant for cattle and horses and is known to Ehleringer, J.R. and T.A. Cooper. 1992. On the
be toxic (Gay and Dwyer 1998, Stubbendieck and role of orientation in reducing photoinhibitory
others 1997, Warnock 1974) and causes liver damage in photosynthetic-twig desert shrubs.
damage when consumed in large quantities Plant Cell and Environment 15(3): 301-306.
(Kirkpatrick 1992). All parts of the plant are
poisonous, younger leaves more so, and poisoning Elmore, F.H. 1976. Shrubs and Trees of the
is more of a problem on drier range conditions. Southwest Uplands. Southwest Parks and
Symptoms in cattle may include a dry scaly nose, a Monuments Association, Tucson, AZ. 214 p.
rough coat, and continuous walking with no
direction (Phillips Petroleum Company 1963). Epple, A.O. 1995. A Field Guide to the Plants of
Because it has low palatability, it tends to be Arizona. Falcon Publishing Inc., Helena, MT.
avoided except during long droughts or in heavily 347 p.
overgrazed areas where it may increase in
abundance (USDA 1937). Toxicity of threadleaf Gay, C. W., Jr. and D. D. Dwyer. 1998 Reprint.
groundsel may be attributed to the pyrrolizidine New Mexico Range Plants. Cooperative
alkaloids (PA). About 50 percent of all identified Extension Service Circular 374. Revisions by:
PA are toxic, and some are carcinogenic in rodents C. Allison, S. Hatch, and J. Schickedanz. New
(Stegelmeier and others 1999). Threadleaf Mexico State University, Las Cruces, NM. 84 p.
groundsel contains the pyrrolizidine alkaloids:
senecionine, seneciphylline, florosenine, Ivey, R.D. 1995. Flowering Plants of New Mexico,
otonecine-based florosenine, and retrorsine 3rd Edition. Published by the author, NM.
(Cooper and others 1996). As with many toxic 504 p.
plants, concentration and preparation are critical
factors. For example, the Navajo would boil entire Kartesz, J.T. 1994. A Synonymized Checklist of
plants and drink the mixture to assist the voice and the Vascular Flora of the United States, Canada,
692
and Greenland, Vol. 1, 2nd Edition. Biota of Stegelmeier, B.L., J.A. Edgar, S.M. Colegate, D.R.
North America Program of the North Carolina Gardner, T.K. Schoch, R.A. Coulombe, and R.J.
Botanical Garden. Timber Press, Portland, OR. Molyneux. 1999. Pyrrolizidine alkaloid plants,
622 p. metabolism and toxicity. Journal of Natural
Toxins 8(1): 95-116.
Kearney, T.H., R. Peebles, and Collaborators.
1951 with 1960 supplement. Arizona Flora. Stubbendieck, J., S.L. Hatch, and C.H. Butterfield.
University of California Press, Berkeley, CA. 1997. North American Range Plants, 5th Edition.
1,085 p. University of Nebraska Press, Lincoln, NE.
501 p.
Kirkpatrick, Z. M. 1992. Wildflowers of the
Western Plains. University of Texas Press. Turner, B.L. and T.M. Barkley. 1990. Taxonomic
Austin, TX. 240 p. overview of the Senecio flaccidus complex in
North America, including Senecio douglasii.
Martin, W.C. and C.E. Hutchins. 1980-1981 Phytologia 69(1): 51-55.
(reprinted 2001). A Flora of New Mexico. Vol.
2. Bishen Singh Mahendra Pal Singh (India) and U.S. Department of Agriculture, Forest Service.
Koeltz Scientific Books (Germany). p. 1,277- 1937 (1988 Dover edition). Range Plant
2,591. Handbook. Dover Publications Inc., New York.
816 p.
Phillips Petroleum Company. 1963. Pasture and
Range Plants. Phillips Petroleum Co., Warnock, B.H. 1974. Wildflowers of the
Bartlesville, OK. 176 p. Guadalupe Mountains and the Sand Dune
Alpine, TX. 176 p.
693
Senecio spartioides Torr. & Gray many-headed groundsel
ASTERACEAE
Synonym: Senecio multicapitatus Greenm. ex Rydb.
James E. Nellessen
Range.—This is a wide-ranging species occurring

from western Texas, west through New Mexico,
northern Mexico, into Arizona, Utah, and
California, north to Wyoming, western Nebraska,
and southwestern South Dakota. Variety
spartioides tends to occur further north and west
within this range, while variety multicapitatus
tends to be more southern in its distribution
(Kearney and others 1951, Great Plains Flora
Association 1986). Within its range it occurs at
elevations from about 1,070 m to 2,745 m.
Ecology.—Many-headed groundsel occurs on

plains, open slopes, valleys, arroyos, and semi-
General Description.—Many-headed groundsel is stabilized dunes in piñon-juniper woodlands,
also known as broom groundsel, broom ponderosa pine forests, and desert areas. It tends to
butterweed, or grass-leaved ragwort (Elmore 1976, be an early colonizer of disturbed soil situations, is
Epple 1995). This plant produces many stems from a prolific seed producer because of the numerous
a large woody crown, in a tight to spreading flower heads, and is generally a short-lived (2 to 4
cluster, giving it a broom-like appearance. Plants years) perennial.
range in height from 20 cm to slightly more than 1
m. In large plants, some stems may lean over, Reproduction.—This species produces numerous
becoming almost decumbent. The leaves are bright yellow-flowered heads, the ray flowers numbering
green, both lobed (pinnately) or nonlobed, from four to eight per head (over 18 flowers with
alternate, linear, 4 to 10 cm long, and to 5 mm the disk flowers included). The heads are 7 to 12
wide, generally hairless or slightly pubescent mm high, 3 to 7 mm wide, subcylindric to
(Carter 1997, Great Plains Flora Association 1986, cylindric with about eight to 12 phyllaries, and the
Ivey 1995, Martin and Hutchins 1980-81, Weber rays are from 7 to 12 mm long. It generally blooms
and Wittman 2001a and 2001b). This plant is best in the late summer and fall but will bloom early in
classified as a subshrub because even though the the summer given sufficient moisture. Hence, the
stems do become woody, especially the bases, it full range of potential blooming runs from May
often dies back during the winter to less than 20 into early November, depending on local
cm above the ground-line. environmental conditions. A bright white fluffy
pappus enables easy dispersal of seeds by the
Taxonomy.—Two varieties are recognized: the wind.
typical variety, S. spartioides var. spartioides, and
S. spartioides var. multicapitatus (Greenm. ex Growth and Management.—Seeds will
Rydb.) S.L. Welch (Allred 2002). The leaves of germinate under moist conditions in the fall, the
variety spartioides are generally simple, or seedlings over-wintering. Growth continues slowly
infrequently with small lobes near the base, and in the spring but is rapid once summer rains begin.
the plant is hairless. The variety multicapitatus Many-headed groundsel is a short-lived perennial.
tends to have leaves irregularly pinnately divided, Most plants appear to survive only 2 to 4 years,
more hairs, and smaller, more numerous flower some perhaps slightly longer. This species,
heads. The latter variety is also synonymized with although toxic to livestock, is not palatable and is
the full species designation: S. multicapitatus generally not highly abundant or dominant in most
Greenm. ex Rydb. (Kartesz 1994). habitats to be a significant management problem.
694
Benefits.—Many-headed groundsel will colonize Ivey, R.D. 1995. Flowering Plants of New Mexico,
open disturbed areas and provide quick ground 3rd Edition. Published by the author, NM.
cover in a natural succession process. Although 504 p.
relatively short-lived the perennial nature of the
plant further helps stabilize soil for longer-lived Kartesz, J.T. 1994. A Synonymized Checklist of
perennials to eventually become established. the Vascular Flora of the United States, Canada,
Many-headed groundsel is toxic to cattle, but it is and Greenland, Vol. 1, 2nd Ed. Biota of North
infrequently eaten (USDA 1937, Kearney and America Program of the North Carolina
others 1951, Warnock 1974). This species should Botanical Garden. Timber Press, Portland, OR.
be utilized more in urban and native plant 622 p.
landscaping. When in bloom it bears, as the name
implies, numerous yellow flower heads, adding Kearney, T.H., R. Peebles, and Collaborators.
considerable color to a desert backyard. It is a 1951 with 1960 supplement. Arizona Flora.
prolific seed producer and will easily regenerate University of California Press, Berkeley, CA.
itself. This may pose a problem for some 1,085 p.
gardeners, managing the abundance of seedlings,
although many of the seedlings will not survive to Martin, W.C. and C.E. Hutchins. 1980-1981
maturity within a natural landscape anyway. (reprinted 2001). A Flora of New Mexico.
Bishen Singh Mahendra Pal Singh, India and
References Koeltz Scientific Books, Germany. Vol. 2, p.
1,277-2,591.
Allred, K.W. 2002. A Working Index of New
Mexico Vascular Plant Names. Available on a U.S. Department of Agriculture Forest Service.
New Mexico State University web page, 1937 (1988 Dover edition). Range Plant
http://web.nmsu.edu/~kallred/herbweb/ [not Handbook. Dover Publications Inc., New York.
paged]. 816 p.
Carter, J.L. 1997. Trees and Shrubs of New Warnock, B. H. 1974. Wildflowers of the
Mexico. Johnson Books, Boulder, CO. 534 p. Guadalupe Mountains and the Sand Dune
Elmore, F.H. 1976. Shrubs and Trees of the Alpine, TX. 176 p.
Southwest Uplands. Southwest Parks and
Monuments Association, Tucson, AZ. 214 p. Weber, W.A. and R.C. Wittman. 2001a. Colorado
Flora: Eastern Slope. University Press of
Epple, A.O. 1995. A Field Guide to the Plants of Colorado, Boulder CO. 521 p.
Arizona. Falcon Publishing Inc., Helena, MT.
347 p. Weber, W.A. and R.C. Wittman. 2001b. Colorado
Flora: Western Slope. University Press of
Great Plains Flora Association. 1986. Flora of the Colorado, Boulder CO. 488 p.
Great Plains. University Press of Kansas, KS.
1,392 p.
695
Senna atomaria (L.) Irwin & Barneby flor de San José
FABACEAE
Synonyms: Cassia atomaria L.

Cassia emarginata L.
Cassia arborescens Miller
Cassia triflora Vahl
Isandrina emarginata (L.) Britt. & Rose (and others: International
Legume Database and Information Service 2002)
John K. Francis
legumes 12 to 35 cm long and 8 to 12 mm broad,

and contain numerous brown seeds 4 to 5 mm
long (Howard 1988, Liogier 1988, Little and
others 1974, Secretaría de Medio Ambiente,
Recursos Naturales y Pesca 2002, Stevens and
others 2001).
Range.—Flor de San José is native to the

Bahamas, the West Indies, Mexico, Central
America, Colombia, Venezuela, and Guyana
(Howard 1988, Little and others 1974). It has been
planted in Hawaii and probably other areas outside
the natural range as an ornamental (Neal 1965).
General Description.—Flor de San José is also Ecology.—Flor de San José is common to rare in
known as yellow candlewood, senna-tree, palo de coastal and upland dry forests at elevations from
chivo, palo de burro, cañafístola cimarrona, near sea level to 1,100 m that receive annual
carbonera, alcaparro, chile perro, palo de zorrillo, rainfalls of about 600 to 1,000 mm. Soils, often
arguchoco, vainillo, chivato, carángano, platanillo, rocky and excessively drained, developed from
bois cabrite, casse marron, and petite casse. It is a both igneous and sedimentary parent materials
flowering, drought-deciduous shrub or small tree 2 with a wide range of textures and with pH’s that
to 12 m in height that occasionally exceeds 20 cm are mildly acid to mildly alkaline. Flor de San José
in stem diameter. The plant may or may not have is moderately intolerant of shade; it grows in the
multiple stems but almost always supports stout open, as a codominant in low forest, and in the
branches relatively low to the ground. The understory of open forests. The species may be
branches are brittle. It is supported by a tap and found in remnant and late secondary forests.
extensive lateral root system that has slender fine
roots. The roots are woody and flexible with a Reproduction.—Flor de San José flowers and
dark gray bark over a pale yellow interior. The fruits throughout the year in Nicaragua (Stevens
crown may be described as spreading and branchy and others 2001). Although flowers and fruits
but not dense. The bark is light gray. Leaves, have been observed at every season of the year in
twigs, and other vegetative parts have an Puerto Rico, individual trees normally flower and
unpleasant odor and are covered with soft hairs. fruit once per year. Because of extended
The alternate, pinnate leaves have petioles 2 to 5 flowering, flowers and fruits may be present on
cm long and two to five pairs of ovate to obovate the same tree. Seeds collected by the author from
entire leaflets, 2 to 11 cm long. The flower Puerto Rico weighed an average of 0.0238 +
clusters are corymbose racemes that bear many 0.0004 g/seed. Sown on moist filter paper, 96
flowers and arise from the bases of the upper percent germinated between 3 and 15 days after
leaves. The flowers have five unequal sepals and sowing. Germination is epigeal. A test of percent
five unequal yellow petals 8 to 15 mm long. The germination of sowings throughout the year in
fruits are flattened, linear, red-brown to black, Mexico resulted in emergence varying from 1.2 to
696
44 percent over 45 day periods depending on Arboretum, Harvard University, Jamaica Plain,
ambient temperature and precipitation (Vera and MA. 673 p.
Sánchez 1995). Reproduction may be common in
areas overgrazed by cattle. Seeds germinate after International Legume Database and Information
pods rot on the ground under mother trees Service. 2002. Senna atomaria (L.) H. Irwin &
(Stevens and others 2001) and may be dispersed to Barneby. http://www.ildis.org/LegumeWeb/
a limited degree by ruminants. 6.00/taxa/15514.shtml. 2 p.
Growth and Management.—Flor de San José Liogier, H.A. 1988. Descriptive flora of Puerto
has a moderate growth rate. Although planted as Rico and adjacent islands, Spermatophyta. Vol.
an ornamental, the author is not aware of any 2. Editorial de la Universidad de Puerto Rico,
reports of wildland plantings. Management and Río Piedras, PR. 481 p.
protection of existing stands are recommended
(Secretaría de Medio Ambiente, Recursos Little, E.L., Jr., R.O. Woodbury, and F.H.
Naturales y Pesca 2002). Wadsworth. 1974. Trees of Puerto Rico and the
Benefits.—Flor de San José is planted widely but 449. U.S. Department of Agriculture,
not abundantly as an ornamental for its beautiful Washington, DC. 1,024 p.
floral display and moderate size. The sapwood is
yellow and hard, and the heartwood is brown and Neal, M.C. 1965. In gardens of Hawaii. Special
hard with a density of 0.57 to 0.85 g/cm3 (Timyan Publication 50. Bernice P. Bishop Press,
1996). It is used for fuel and rude construction Honolulu, HI. 924 p.
(Holdridge and Poveda 1975, Secretaría de Medio
Ambiente, Recursos Naturales y Pesca 2002). Northern Prairie Wildlife Research Center. 2002.
Cows eat the pods but do not browse the leaves. Butterflies of North America: Yellow angled-
Some of the common names imply that goats sulphur (Anteos maerula). U.S. Geological
browse the species. Meal made from seed-free Survey. http://www.npwrc.usgs.gov.resource.
pods of flor de San José was eagerly consumed by distr/lepid/bflyusa/usa/723.htm. 3 p.
lambs in an acceptance test (Palma and Román
2002b); the meal contained 6.5 percent crude Olmos, F. 1997. Tapirs as seed dispersers and
protein, 6.8 percent fat, 4.5 percent ash, and 32.1 predators. UNCN/SSC Tapir Specialist Group.
percent crude fiber (Palma and Román 2002a). It http://www.tapirback.com/tapirgal/iucn-ssc/tsg/
has been demonstrated that Baird’s tapir (Tapirus action97/ap97-05.htm. 14 p.
bairdii) kills 100 percent of the flor de San José
seeds when it consumes the pods, apparently Palma, J.M. and L. Román. 2002a. Frutos de
because they germinate within the gut (Olmos especies arbóreas leguminosas y no
1997). Flor de San José is the principal or a key leguminosas para la alimentación de rumiantes.
larval host for the yellow angled-sulphur butterfly, Centro Universitario de Investigación y
Anteos maerula (Fabricius), and the large moth, Desarrollo Agropecuario. http://lead.
Sphingicampa montana (Packard) (Northern virtualcentre.org/es/ele/conferencia2/vbconfe15
Prairie Wildlife Research Center 2002, Savela .htm. 15 p.
2002). A tea made from the leaves is used as a
purgative (Secretaría de Medio Ambiente, Palma, J.M. and L. Román. 2002b. Prueba de
Recursos Naturales y Pesca 2002). selectividad con ovinos de pelo de harinas de
frutos de especies arbóreas. Centro
References Universitario de Investigación y Desarrollo
Agropecuario. http://www.cipav.org.co/
Holdridge, L.R. and L.J. Poveda A. 1975. Arboles redagrofor/memoirias99/PalmaJM.htm. 4 p.
de Costa Rica. Centro Cientifico Tropical, San
José, Costa Rica. 546 p. Savela, M. 2002. Sphingicampa Walsh, 1864.
http://www.funet.fi/pub/sci/bio/life/insecta/lepi
Howard R.A. 1988. Flora of the Lesser Antilles, doptera/ditrysia/bombycoidea/saturniidae/
Leeward and Windward Islands. ceratocampinae/sphingicampa/. 3 p.
697
Secretaría de Medio Ambiente, Recursos Naturales 1. Missouri Botanic Garden Press, St. Louis,
y Pesca. 2002. Especies forestales no MO. 943 p.
maderables y maderables no tradicionales de
zonas áridas y semiáridas: Senna atomeria (L.) Timyan, J. 1996. Bwa yo: important trees of Haiti.
Irwin et Barneby. http://www.semarnat.gob.mx/ South-East Consortium for International
pfnm3/fichas/senna_atomaria.htm. 2 p. Development, Washington, DC. 418 p.
Stevens, W.D., C. Ulloa-U., A. Pool, and O.M. Vera-S., F. and M. Sánchez-R. 1995. Epoca de
Montiel, eds. 2001. Flora de Nicaragua. siembra de doce especies forestales de Baja
Monographs in Systematic Botany Vol. 85, No. California Sur. Revista de Ciencia Forestal en
México 20(78): 59-78.
698
Senna occidentalis (L.) Link coffee senna
FABACEAE
Synonyms: Cassia occidentalis L.

Ditremexa occidentalis (L.) Britt. & Rose
John K. Francis
are ovate to ovate-lanceolate, pointed at the tip

and rounded at the base. Inflorescences are few-
flowered axillary racemes with yellow-petaled
flowers about 2 cm across. The legumes (pods) are
brown, flat, slightly curved and 5 to 12 cm long.
They contain 40 or more brown to dark-olive,
ovoid seeds about 4 mm long. The species has 2n
= 26, 28 chromosomes (Henty and Pritchard 1975,
Liogier 1988, Long and Lakela 1976, Stevens and
others 2001).
Range.—Coffee senna grows throughout the

tropics and subtropics (Liogier 1988, Stevens and
others 2001) including the United States from
Texas to Iowa eastward, Hawaii, the Pacific Island
Territories, Puerto Rico, and the U.S. Virgin
Islands (Natural Resources Conservation Service
2002). It appears to be of South American or New
World origin (Haselwood and Motter 1966, Henty
and Pritchard 1975, Raintree 2002).
Ecology.—In Cuba, coffee senna is said to be

most common in fertile, cultivated areas (Sánchez
and Uranga 1993) and in Nicaragua it also grows
in disturbed sites (particularly fertile ones), river
overflow areas, meadows, and forests from near
sea level to 900 m in elevation (Stevens and others
General Description.—Coffee senna is also 2001). The species occurs on pinelands in Florida
known as stinking weed, miki palaoa, hedionda, and grows better on near-neutral soils than acid
bucho, pico de pájaro, furrusca, bois puante, (pH 4.7) soils (Long and Lakela 1976). Coffee
fedegoso, and many other names. The species senna grows in moist areas (mean annual rainfall
varies from a semi-woody annual herb in warm from about 1000 to 2200 mm) in Puerto Rico.
temperate areas to a woody annual shrub or
sometimes a short-lived perennial shrub in frost- Reproduction.—Coffee senna flowers and fruits
free areas (Haselwood and Motter 1966, Henty throughout the year or seasonally, depending on
and Pritchard 1975, Holm and others 1997). It rainfall and cold seasons. Seeds collected in
usually matures at from 0.5 to 2.0 m in height. In Puerto Rico averaged 0.0158 + 0.0002 g/seed or
Brazil it is reported to reach 5 to 8 m in height 63,000 seeds/kg. Some 95 percent of scarified
(Raintree 2002). Coffee senna produces a hard, seeds sown in potting mix germinated between 5
woody tap root with relatively few laterals. It and 36 days after sowing (author’s observation).
usually has a single purplish stem and sparse Scarification is necessary for good seed
branching. Young stems are four-angled, germination (Institute of Food and Agricultural
becoming rounded with age. The crushed foliage Sciences 2002). Mechanical scarification, acid
has an unpleasant odor. Compound alternate treatment, and immersing in boiling water all
leaves have four to six pairs of glabrous leaflets worked well, giving 82 to 100 percent germination
and a gland near the base of the petiole. Leaflets (Demel and Teketay 1996). The seeds are
699
dispersed by grazing animals (Sánchez and Senna species from Ethiopia. New Forests
Uranga 1993). 11(2): 155-171.
Growth and Management.—In seasonally cold Guzmán, D.J. 1975. Especies útiles de la flora
or dry climates, the life cycle of coffee senna is salvadoreña. Ministerio de Educación, Dirección
complete in 6 to 9 months, but in warm, de Publicaciones, San Salvador, El Salvador.
continually moist areas plants may last a full year 703 p.
or grow through the second year. Perhaps in Brazil
where extraordinary heights are reached, the Haselwood, E.L. and G.G. Motter, eds. 1966.
species may live a third or fourth year. Growth is Handbook of Hawaiian weeds. Experiment
moderately rapid. Plants add 0.5 to 1.5 m during Station, Hawaiian Sugar Planters’ Association,
the first season. Although coffee senna is planted Honolulu, HI. 479 p.
to yield medicinal materials (Barbadine 2002),
because of its short life and weedy potential, it is Henty, E.E. and G.H. Pritchard. 1975. Weeds of
not advisable to plant it in wildlands. The species New Guinea and their control. Botany Bulletin
can be controlled with broadleaf herbicides (Henty 7. Division of Botany, Department of Forests,
and Prichard 1975). Lae, Papua New Guinea. 189 p.
Benefits and Detriments.—As the name implies, Holm, L., J. Doll, E. Holm, J. Pancho, and J.
the seeds of coffee senna are roasted and used as a Herberger. 1997. World weeds. John Wiley &
coffee substitute (Guzmán 1975). The plant’s Sons, New York. 1,129 p.
tissues contain a host of phytoactive chemicals
that may support its numerous applications in folk Institute of Food and Agricultural Sciences. 2002.
medicine. Extracts or powdered leaves are used as Coffee senna, Cassia occidentalis L. Institute of
an analgesic, antibacterial, anti-hepatotoxic, Food and Agricultural Sciences, Cooperative
antifungal, anti-inflammatory, antiseptic, Extension Service, University of Florida,
antispasmodic, antiparasitic, antiviral, Gainesville, FL. http://edis.ifas.ufl.edu/
carminative, diaphoretic, emmenagogue, BOFY_FW008. 3 p.
febrifuge, insecticidal, immunostimulant, laxative,
purgative, sudorific, and vermifuge. Several of Liogier H.A. 1988. Descriptive flora of Puerto
these effects have been demonstrated in laboratory Rico and adjacent islands, Spermatophyta. Vol.
and clinical tests (Raintree 2002). While coffee 2. Editorial de la Universidad de Puerto Rico,
senna can be a weed of cultivated fields and Río Piedras, PR. 481 p.
plantations, it is principally a problem because it
accumulates in heavily-grazed pastures (Henty Long, R.W. and O. Lakela. 1976. A flora of
and Pritchard 1975). The foliage is poisonous and Tropical Florida. Banyon Books, Miami, FL.
generally avoided by livestock (Stevens and others 962 p.
2001). Ingestion of large amounts of seeds has
been implicated in deaths of cows, horses, and Natural Resources Conservation Service. 2002.
goats (Raintree 2002). Poisoning of pigs fed Plant profile: Senna P. Mill, senna. U.S.
coffee senna seeds resulted in muscle necrosis Department of Agriculture, Natural Resources
(Timm and Riet-Correa 1997). Conservation Service, Washington, DC.
http://plants.usda.gov/cgi_bin/plant_profile.chi?
References symbol=SENNA. 6 p.
Barbadine. 2002. Senna occidentalis (L.). Rain-tree. 2002. Fedegoso. http://rain-tree.com/

Barbadine, Reunion Island, France. http://www. fedegosa.htm. 7 p.
barbadine.com/pages/senna_occ_lien.htm. 2 p.
Sánchez, P. and H. Uranga. 1993. Plantas
Demel, T. and D. Teketay. 1996. The effect of indeseables de importancia economica en los
different pre-sowing seed treatments, cultivos tropicales. Editorial Científico-Técnica,
temperature, and light on the germination of five Havana, Cuba. 166 p.
700
Stevens, W.D., C. Ulloa-U., A. Pool, and O.H. Timm, C.D. and F. Riet-Correa. 1997. Plants toxic
Montiel, eds. 2001. Flora de Nicaragua. to pigs. Ciencia Rural 27(3): 521-528.
MO. p. 1-943.
701
Senna polyphylla (Jacq.) Irwin & Barneby desert cassia
FABACEAE
Synonyms: Cassia polyphylla Jacq.

Peiranisia polyphylla (Jacq.) Britt. & Rose
John K. Francis
Range.—Senna polyphylla var. polyphylla Irwin

& Barneby is native to Puerto Rico and the Virgin
Islands and has been recorded under cultivation in
Florida, Grenada, Guyana, Surinam, Brazil, and
probably elsewhere. The variety montis-christi
Irwin & Barneby is an endemic of Hispaniola, and
the variety neglecta Irwin & Barneby is found
only on the island of Anegada, British Virgin
Islands (Liogier 1988).
Ecology.—In Puerto Rico, desert cassia grows on

a wide variety of well-drained soils that have
developed over igneous, metamorphic (including
ultramafics), and sedimentary (including
limestone) rocks. It is tolerant of salt spray
(Botanics Wholesale 2001). The species occurs at
elevations from near sea level to about 300 m
above sea level (Little and others 1974). Annual
rainfall ranges from about 750 mm to about 1200
mm. Because desert cassia is intolerant of shade
and grows slowly, it is usually found in areas
where competition is minimal—dry, very rocky,
General Description.—Desert cassia, also known excessively drained, overgrazed, and sites subject
in Spanish as hediondilla and retama prieta, is a to occasional fires. Bare ground is probably
shrub or occasionally a small tree of dry and moist necessary for reproduction.
forests of the middle Caribbean, now cultivated as
an ornamental. It is usually 2 or 3 m in height, but Reproduction.—Desert cassia flowers and fruits
rarely reaches 5 m in height and 12 cm in trunk throughout the year (Little and others 1974). The
diameter. The bark of older stems is dark gray and seeds from a Puerto Rican collection averaged
furrowed with short scaly plates. The inner bark is 20,400 seeds/kg. Placed on moist filter paper
light brown. Multiple stems sprouting from the without any pre-treatment, they began germinating
root crown are common, even in small plants. The in 2 days and gave 59 percent germination
stems are mostly clean, long and wand-like. (Francis and Rodríguez 1993). The principal mode
Desert cassia develops a root system with a strong of dispersal today is by livestock. Seedlings in the
taproot. The twigs are slender, warty, and light wild are scattered and not abundant.
green, maturing to brown. The leaves are alternate
or clustered three to five at the nodes. These Growth and Management.—Growth is slow.
pinnately compound leaves have three to 15 pairs About 30 cm of annual height increase is normal
of leaflets that are 4 to 10 mm long. The five- in Puerto Rico. Diameter growth ranges between
petaled flowers are yellow and about 3.8 cm about 1 and 4 mm/yr. Although no planting
across, grouped in axillary racemes of mostly two experience is published, plantations could
flowers. The legume is linear, 8 to 15 cm long, probably be established with containerized
slightly contorted, flattened between the seeds, seedlings followed by several years of weeding
and dark brown at maturity. The seeds are round, until the seedlings grew above herb and grass
flattened, and dark brown (Howard 1988, Liogier competition.
702
Benefits.—Desert cassia is grown by commercial Department of Agriculture, Forest Service,
nurseries and widely planted as a flowering Southern Forest Experiment Station, New
ornamental in the New World Tropics. It is Orleans, LA. 5 p.
employed as an accent plant or trained into a small
background tree. The wood is light brown and Howard R.A. 1988. Flora of the Lesser Antilles,
hard and used occasionally for fence posts and Leeward and Windward Islands.
fuel. Cows apparently do not readily eat the Dicotyledoneae, Part 1. Vol. 4. Arnold
foliage of desert cassia—the species is more Arboretum, Harvard University. Jamaica Plain,
common in heavily grazed rangeland than MA. 673 p.
elsewhere.
References Rico and adjacent islands, Spermatophyta. Vol.
Botanics Wholesale. 2001. http://www.botanics. Río Piedras, PR. 481 p.
com/Products/botanics_details.asp?NameAssoc
+153.htm. 1 p. Little, E.L., Jr., R.O. Woodbury, and F.H.
Francis, J.K. and A. Rodríguez. 1993. Seeds of Virgin Islands. Vol. 2. Agriculture Handbook
Puerto Rican trees and shrubs: second 449. U.S. Department of Agriculture,
installment. Research Note SO-374. U.S. Washington, DC. 1,024 p.
703
Serenoa repens (Bartr.) Small saw palmetto
PALMAE
Synonyms: None
Deborah K. Kennard, G.W. Andrews, Kenneth Outcalt, and Mary Carrington
(Myrica cerifera L.), staggerbush [Lyonia

ferruginea (Walter) Nuttall], oaks (Quercus spp.
L.), wiregrass (Aristida beyrichiana Trin. &
Rupr.), several bluestem species (Andropogon spp.
L.), and lop-sided Indian grass [Sorgastrum
secundum (Ell.) Nash.] (Tanner and others 1996).
Saw palmetto occurs on a wide range of soil types
but most commonly is found on seasonally
flooded, sandy, acidic spodosols typical of
flatwoods ecosystems in the Lower Coastal Plain.
Typical soil types are Leon fine sand, Myakka fine
sand, and Immokalee fine sand. Saw palmetto also
grows on deep, sandy entisols, on calcareous
General Description.—Saw palmetto, also sandy soils near coasts, and on limestone in
known as palmetto, is a low-growing palm southern Florida. Annual rainfall varies from 114
endemic to the Southeastern United States. Saw cm in the northern part of its range to over 150 cm
palmetto stems typically lie prostrate at the soil along the southeastern coast of Florida. Rainfall
surface but can grow upright and reach heights of also becomes more seasonal going from north to
5 to 7 m (Tanner and others 1996). Multiple, south, with increasingly more rainfall occurring
persistent, palmate leaves up to 1 m wide emerge during summer. In southern Florida, the
from the stem’s terminal buds. Short recurved southernmost part of its range, 64 percent of
spines line the petioles (Small 1933). Flowers are average annual rainfall occurs from June to
perfect and borne on paniculate inflorescences that September (Hilmon 1968).
emerge from the leaf bases. The fruit is a one-
seeded ellipsoid or subglobose drupe with a fleshy Reproduction.—Saw palmettos primarily
mesocarp, 16 to 25 mm long and 12 to 19 mm reproduce vegetatively through suckers from the
wide. Fruit color turns from green, to yellow, to main stem (Fisher and Tomlinson 1973). Each
orange, and then to bluish-black when fully ripe sucker has the capacity to become a ramet,
(Hilmon 1968). forming extensive clumps of genetically identical
clones. Saw palmettos must be at least 0.6 m in
Range.—Saw palmetto is endemic to the Coastal height to flower (Carrington and others 2000) after
Plain of the Southeastern United States. The which probability of flowering increases with plant
northern limits of its range extend from height. Inflorescences emerge from buds at the
southeastern Louisiana through Tifton Georgia to bases of previous season’s leaves in February to
Charleston County, South Carolina (Hilmon 1968, April, and flowering occurs from April to June
McNab and Edwards 1980). (Hilmon 1968). Flowers are insect pollinated
(Tanner and others 1996). While 34 insect species
Ecology.—Saw palmetto occurs as a major are known to pollinate saw palmetto flowers, the
understory plant in seasonally wet pine flatwoods, European honeybee (Apis mellifera L.) is the
well-drained scrubby flatwoods, and on sandy primary pollinator (Carrington and others in
berms and dunes along rivers and the coast review). Emerging saw palmetto inflorescences are
(Tanner and others 1996). Within these habitats, it subject to attack by cabbage palm caterpillars
can grow in a variety of conditions from shade to (Litoprosopus futilis G. & R.), and green fruits are
full sun (Small 1926). Common associates include: subject to anthracnose [Colletotrichum
gallberry [Ilex glabra (L.) Gray], wax myrtle gloeosporioides (Penz.) Penz. & Sacc. in Penz.]
704
(Carrington and others 2000). Fruits ripen in increase coverage of saw palmetto (Carrington and
August to November. Saw palmettos flower others 2000).
heavily about every 2 to 4 years. During these
years, individual ramets commonly produce two to Fire.—Saw palmetto naturally occurs in plant
three inflorescences. Due to high flower density communities adapted to periodic burning. Saw
(several thousand flowers/inflorescence), typically palmetto itself is highly flammable due to the
0.4 to 0.5 kg of fruit are produced on each accumulation of dead fronds that remain on the
inflorescence. In some cases, individual plants for several years (Carrington and others
inflorescences can produce up to 12 kg of fruit. 2000) and volatile waxes that cover green leaves.
Average fruit yield for a site is approximately 200 Fires commonly consume all above-ground foliage
kg/ha; however, yields can vary from less than 100 of saw palmettos, and therefore, prescribed fire is
kg/ha to more than 1,500 kg/ha (Carrington and used to reduce leaf coverage for short periods.
others 1997). Fruits are bird and mammal However, leaf growth usually begins a few days
dispersed (Tanner and others 1996). Seed after fires, and saw palmettos can regain 80
germination ranges from 20 percent after 15 percent of crown coverage the first year after
months in field conditions to 55 percent after 6 burning (Hilmon 1968). Fire also benefits fruit
months under lab conditions (Hilmon 1968). Seeds production of saw palmetto, especially within the
can remain viable for up to 1 year, albeit with first year following burns (Carrington and others
reduced germination rates (Carrington and others 2000). However, flowering may be reduced for
2000). Seed germination may be enhanced by several years after burns (Abrahamson 1999) as
passing through animal digestive systems (Tanner palmettos recover from both burning and heavy
and others 1996). fruit production. Frequent burning (every 1 to 4
years) curtails flowering and fruiting by keeping
Growth and Management.—Growth of saw carbohydrate reserves low (Hilmon 1968).
palmetto is slow, ranging between 0.6 and 2.2
cm/year in stem elongation (Abrahamson 1995, Detriments and Benefits.—Saw palmetto was
Hilmon 1968). Based on these estimates, some viewed by early settlers in the Southeastern United
saw palmettos may be 500 to 700 years old States as an obstacle to establishing agricultural
(Abrahamson 1995, Tanner and others 1996). In fields, cattle pastures, and home sites. Ranchers
some cases, land managers may want to reduce the continue to regard it as a competitor of native
density and cover of saw palmetto in order to forage grasses, and foresters cite that it inhibits
improve cattle forage and/or timber production or pine regeneration. Despite these detriments, saw
to reduce fire hazard. Because saw palmetto is palmetto is viewed as a significant pharmaceutical
fire-tolerant, mechanical or chemical treatments resource in Florida and southern Georgia due to
are often needed for control. Roller-chopping the effectiveness of certain fruit compounds (such
reduces abundance of saw palmetto (Hilmon and as free fatty acids, phytosterols) in treating benign
others 1963, Lewis 1970) by crushing vegetative prostatic hyperplasia (Tasca 1985, Braeckman
parts and removing terminal buds. This treatment 1994, Wilt and others 1998), or swelling of the
is particularly effective during periods of low soil prostate gland. In 1995, saw palmetto fruits
moisture (Moore 1974) or when followed by collected for pharmaceuticals sold for over $6/kg.
prescribed burning. Among herbicide treatments, Total estimated value of fruits sold in 1996 was
repeated cycles of 2,4,5-T followed by burning approximately $5 million (Carrington and others
will reduce palmetto cover (Altobellis and Hough 2000). In addition to its medicinal value, saw
1968), but this chemical is no longer available. palmetto can be a significant source of honey
Metsulfuron, often sold as Escort, is the most production (Bennett and Hicklin 1998). Saw
effective herbicide currently available for control palmetto is also a good native plant for
of saw palmetto (personal communication J.L. enviroscaping due to its natural drought- and
Michael, Southern Research Station, Auburn, AL). insect-resistance, and low requirements for
In other cases, land managers may want to fertilization (Tanner and others 1996). Saw
enhance saw palmetto flowering and fruiting. The palmettos serve as nesting and denning habitat for
most cost-efficient practice to increase fruit over 100 animal species, including: the
production is prescribed burning (Carrington and endangered Florida grasshopper sparrow
others 2000). Optimal burning frequency is every (Ammodramus savannarum Gmein), Florida
5 to 8 years. Soil fertilization has been used to panther (Felis concolor Linnaeus), Florida
705
woodrat (Neotoma floridana Ord), wild turkey Fisher, J.B. and P.B. Tomlinson. 1973. Branch
(Meleagris gallopavo Linnaeus), and white-tailed and inflorescence production in saw palmetto
deer (Odocoileus virginianus Zimmermann) (Serenoa repens). Principes 17: 10-19.
(Tanner and others 1996). Fruits are eaten by black
bears (Ursus americanus Pallas), white-tailed deer Hilmon, J.B. 1968. Autecology of saw palmetto
(Odocoileus virginianus), raccoons (Procyon lotor [Serenoa repens (Bartr.) Small]. Ph.D.
Linnaeus), turkeys (Meleagris gallopavo), bob- dissertation. Duke University, Durham, N.C.
white quail (Colinus virginianus L.), gray foxes 191 p.
(Urocyon cinereoargenteus Schreber), opossum
(Dasypus novemcinctus Linnaeus), and gopher Hilmon, J.B., C.E. Lewis, and J.E. Bethune. 1963.
tortoises (Gopherus polyphemus Daudin) (Maehr Highlights of recent results of range research
and Layne 1996). in Southern Florida. Society of American
Foresters Proceedings 1962: 73-76.
References
Lewis C.E. 1970. Responses to chopping and rock
Abrahamson, W.G. 1995. Habitat distribution and phosphate on south Florida ranges. Journal of
competitive neighborhoods of two Florida Range Management 23: 276-282.
palmettos. Bulletin of the Torrey Botanical
Club 122: 1-14. Maehr, D.S. and J.N. Layne. 1996. Florida’s all-
purpose plant the saw palmetto. Palmetto
Abrahamson, W.G. 1999. Episodic reproduction in (Fall): 16 –10, 15, 21.
two fire-prone palms, Serenoa repens and
Sabal etonia (Palmae). Ecology 80:100-115. McNab, W.H. and M.B. Edwards. 1980. Climatic
factors related to the range of saw-palmetto
Altobellis, A.T. and W.A. Hough. 1968. [Serenoa repens (Bartr.) Small]. American
Controlling palmetto with fire and herbicides. Midland Naturalist 103: 205-208.
Georgia Forestry Research Paper 52. Georgia
Forest Research Council, Macon, Ga. Moore, W. H. 1974. Some effects of chopping
saw-palmetto pineland threeawn range in
Bennett, B.C. and J.R. Hicklin. 1998. Uses of saw south Florida. Journal of Range Management
palmetto (Serenoa repens, Arecaceae) in 27(2): 101-104.
Florida Economic Botany 52: 381-393.
Small, J.K. 1926. The saw-palmetto–Serenoa
Braeckman, J. 1994. The extract of Serenoa repens repens. Journal of the New York Botanical
in the treatment of benign prostatic Garden 27: 193-202.
hyperplasia: A multicenter open study.
Current Therapy Research 55: 776-785. Small, J.K. 1933. Manual of the southeastern
flora. John Kunkel Small, New York.
Carrington, M.E., T.D. Gottfried, and J.J. 1,554 p.
Mullahey. (In review) Pollination biology of
saw palmetto (Serenoa repens: Palmae) in Tanner, G.W., J.J. Mullahey, and D. Maehr. 1996.
southwestern Florida. Palms. Saw-palmetto: an ecologically and
economically important native palm. Circular
Carrington, M.E., J.J. Mullahey, G. Krewer, B. WEC-109, Florida Cooperative Extension
Boland, and J. Affolter. 2000. Saw palmetto Service, Institute of Food and Agricultural
(Serenoa repens): an emerging forest resource Sciences, University of Florida, Gainesville,
in the southeastern United States. Southern FL. 7 p.
Journal of Applied Forestry 24(3): 129-134.
Tasca, A. 1985. Treatment of obstructive
Carrington, M.E., J.J. Mullahey, and F. Roka. symptomatology in prostatic adenoma with an
1997. Saw palmetto: A fountain of youth. extract of Serenoa repens. Minerva Urologica
Proceedings American Forage Grassland e Nefrologica 37: 87-91.
Council 6: 233-237.
706
Wilt, T., A. Ishani, and C. Mulrow. 1998. Saw Journal of the American Medical Association
palmetto extracts for treatment of benign 280: 1,604-1,609.
prostatic hyperplasia: a systemtatic review.
707
Serjania polyphylla (L.) Radlk. bejuco de corrales
SAPINDACEAE
Synonyms: Paullinia polyphylla L.

Paullinia triternata Jacq.
Serjania lucida Schum.
Serjania triternata (Jacq.) Willd.
Serjania dubia Spreng.
John K. Francis
than five. The branches are slender and flexible

with few bifurcations. Tendrils, which support the
vine on trees and other structures, grow at the leaf
axils. The alternate compound leaves divide by
threes. Leaflets, which number three to 19 or
more, are dentate from the middle to the tip, 1.5 to
6.0 cm long and 1.0 to 3.5 cm broad.
Inflorescences are axillary panicles and contain
many white flowers. The brown or red-brown
fruits are samaras, 1.5 to 2.5 cm long, that grow in
groups of three. Each samara contains a single
seed (Acevedo-Rodrígues 1985, Liogier 1994).
Range.—Bejuco de corrales is native to Puerto

Rico and offshore islands, the Virgin Islands, and
Hispaniola (Liogier 1994). It is not known to have
been planted elsewhere.
Ecology.—Bejuco de corrales grows in remnant

and secondary forests, on roadsides, fence rows,
river banks, and brushy pastures. It is usually
common but rarely abundant. The species is
moderately intolerant of shade, requiring full
sunlight to flower and fruit; yet it can survive in
the understory of low basal-area forest. Bejuco de
corrales grows on soils with textures from sand to
clay that are derived from igneous and
sedimentary (including limestone) rocks. In Puerto
Rico, the species grows from near sea level to over
General Description.—Bejuco de corrales, also 600 m in elevation and in areas that receive from
known as bejuco de canastas, bejuco de costillas, 750 to over 2000 mm of annual rainfall. In the
basket wood, and blackwith, is a woody vine that areas with the highest rainfall, it is confined to
ascends as much as 20 m into the crowns of trees. excessively drained sites. Bejuco de corrales
Bejuco de corrales is a Spanish name meaning tolerates drought and at least mild salt spray.
corral vine. The older stems are brown, rough, Frosts are not known within the native range.
with three main striations and up to 5 cm in
diameter. The stem cross section has a Reproduction.—Bejuco de corrales flowers and
characteristic pattern reminiscent of a string of fruits from November through June (Acevedo-
beads around a central core. The vines have a Rodrígues 1985). An infrutescence may contain
taproot, extensive lateral, and abundant fine roots, 25 or more fruits, and a plant may produce several
and all have a dark brown color. Periodic infrutescences per year. A collection of bejuco de
sprouting from the root crown replaces dying corrales samaras averaged 0.0248 g/fruit or 40,000
stems and gives rise to multiple stems, usually less fruits/kg. When the clusters break up, the samaras
708
spin and fly sideways as they descend and may be a diuretic and purifier of the blood in herbal
travel a considerable distance before reaching the medicine (Liogier 1990).
ground.
References
Growth and Management.—Seedlings grow
about 0.5 m in their first year. Older plants and Acevedo-Rodríguez, P. 1985. Los bejucos de
sprouts grow 2 or 3 m each year. Individual stems Puerto Rico. Vol. 1. General Technical Report
can live for several years; by sprouting, plants SO-58. U.S. Department of Agriculture, Forest
may survive for at least several decades. No Service, Southern Forest Experiment Station,
planting experience for the species has been New Orleans, LA. 331 p.
reported. In the past, farmers controlled bejuco de
corrales by repeated cutting with a machete. These Liogier, H.A. 1990. Plantas medicinales de Puerto
vines could probably be killed faster by spraying Rico y del Caribe. Iberoamericana de Ediciones,
the sprouts that arise after cutting with broadleaf Inc. San Juan, PR. 566 p.
weed killer.
Benefits.—The strong and flexible stems of Rico and adjacent Islands. Vol. 3. Editorial de la
bejuco de corrales are suitable for basketry, and Universidad de Puerto Rico, Río Piedras, PR.
some of the common names indicate that it was 461 p.
used for this purpose. Bejuco de corrales is
considered to be a honey plant in the Dominican Marcano Fondeur, E. de J. 1973. La flora apícola
Republic (Marcano Fondeur 1973). Livestock eat de la República Dominicana. http://
young plants and the foliage of older ones that are marcano.freeservers.com/nature/estudios/apicola
within their reach. Bejuco de corrales is reputed to /dicotsp.htm. 11 p.
709
Sesbania sericea (Willd.) Link silky sesban
FABACEAE
Synonyms: Coronilla sericea Willd.
John K. Francis
leaves are 5 to 25 cm long and support 10 to 20

leaflet pairs. The leaflets are oblong, rounded at
both ends, mucronate at the apex, with a silky
pubescence below. The racemes have two to eight
yellow, greenish-yellow, or orange flowers. The
brown pods that develop after flowering are 10 to
20 cm long, 3 mm broad, and contain 20 to 30
seeds (Howard 1988, Liogier 1988, Nelson 1996).
The species has 2n = 24 chromosomes (Long and
Lakela 1976).
Range.—There is confusion about the native

range of silky sesban. Howard (1988) states that it
is a native of Sri Lanka. Others refer to tropical
Asia (Long and Lakela 1976) or Africa and the
Caribbean (Evans and Rotar 1987) as the native
ranges. It is clear that the species has been widely
introduced. The current New World range covers
southern Florida, the Bahamas, the West Indies,
Trinidad, Surinam, the Guianas, Venezuela, and
Central America (Liogier 1988).
Ecology.—Silky sesban is highly intolerant of

shade and must have disturbed soil in which to
germinate. Once established, it competes
aggressively with grass and herbs, but it must
dominate all its life or die. Stands of silky sesban
vigorously self-thin. The species develops best on
moist alluvial soils. It will grow on soils with low
fertility, particularly when poorly or very poorly
General Description.—Silky sesban, also known drained. Modest amounts of salts in the soil are
as papagayo in Spanish, is a short-lived woody tolerated. Apparently, the northern extent of the
shrub of frequently disturbed areas. Its mature range is limited by cool weather and frost.
height may vary from 1 to 6 m with basal
diameters reaching as much as 10 cm under the Reproduction.—A collection of 45 air-dry pods
most favorable conditions. A single stem is of silky sesban weighed an average of 0.378 +
usually developed unless the plant sprouts after 0.0008 g/pod. The seeds separated from them
damage. Robust low side branches develop on averaged 0.0053 + 0.0001 g each or 189,000
open-grown individuals. Secondary branches on seeds/kg. Sown without any pretreatment on
stems are usually unbranched. The species is commercial potting mix, these seeds germinated at
unarmed or may occasionally have weak prickles. 99 percent between 3 and 24 days after sowing.
Stems are green when young, turning gray. There Germination is epigeal. Although the original
may or may not be a well-developed taproot, mode of seed dispersal has not been documented,
depending on the aerated rooting depth. The white in the Caribbean it seems to be opportunistic,
lateral roots are relatively thick (3 to 5 mm) and disbursing by wind, water, grazing animals, and
support many nodules with reddish centers and farm machinery. Seedlings may be abundant in the
abundant fine roots. The pinnately compound presence of a seed source on moist bare soil.
710
However, relatively few of the seedlings progress experimentally intercropped with maize and
past the early seedling stage. resulted in no depression in maize yield. In
Guyana, the species grows in upland rice fields
Growth and Management.—Plants of silky but does not occur in flooded rice paddies (Evans
sesban grow rapidly. Seedlings are about 10 cm and Rotar 1987). The stemwood of silky sesban is
tall at the end of 1 month and reach 2 to 6 m in brash, has a relatively low specific gravity, and
height in about 6 months. Silky sesban was offers few prospects for commercial use.
reported to yield 26.8 Mg/ha (fresh weight) in a
84-day growing period (Evans and Rotar 1987). References
After flowering, height growth of silky sesban
ceases. Plants live from about 8 months to a little Evans, D.O. and P.P. Rotar. 1987. Sesbania in
over a year, depending on conditions. When their agriculture. Tropical Agriculture Series 8.
seeds mature, the plants die. The species may Westview Press, Boulder, CO. 192 p.
grow and die in an annual cycle timed with wet
and dry seasons, or it may simply grow in Howard, R.A. 1988. Flora of the Lesser Antilles,
response to favorable soil conditions as they Leeward and Windward Islands.
occur. If land managers find it necessary to Dicotyledoneae, Part 1. Vol. 4. Arnold
establish silky sesban, sowing into a moist, Arboretum, Harvard University. Jamaica Plain,
prepared seedbed should be sufficient. MA. 673 p.
Benefits.—The species of Sesbania, including Liogier, H.A. 1988. Descriptive flora of Puerto
silky sesban, are used as green manure in India. Rico and adjacent islands, Spermatophyta. Vol.
Silky sesban is readily consumed by cattle in 2. Editorial de la Universidad de Puerto Rico,
pastures. The closely related S. sesban (L.) Merr. Río Piedras, PR 481 p.
has been shown to have moderately good
nutritional values. Some of the species of Sesbania Long, R.W. and O. Lakela. 1976. A flora of
have toxic saponins and canavanine in their seeds. Tropical Florida. Banyon Books, Miami, FL.
So far, the seeds of silky sesban have not been 962 p.
reported to be toxic. Crushed seeds of this species
were fed to chicks at 1 percent of body weight Nelson, G. 1996. The shrubs and woody vines of
each day for 3 days without any signs of toxicity Florida. Pineapple Press, Inc. Sarasota, FL.
(Evans and Rotar 1987). Silky sesban can be 391 p.
weedy but usually causes few problems. It was
711
Sida fallax Walp 'ilima
MALVACEAE
Synonyms: Anoda ovata Meyen

Sida cordifolia L.
Sida dielli Gray,
Sida fallax var. kauaiensis Hochr.
Sida ledyardii St. John
Sarah A. Taylor and Randy S. Senock
environments.
Ecology.―'Ilima displays numerous ecological

types, occurring from sea level to altitudes of 600
m. Beach forms are low growing, dense, spreading
mats, while upland types may grow erect and
shrub-like (Krauss 1998). Lowland types are
typically densely pubescent, while upland types
are glabrous. 'Ilima occurs on a variety of
substrate, from limestone reefs, dry lava fields, and
along sandy and rocky shores. Often 'ilima
represents the only ground cover, aside from
nonnative species, on arid, rocky coastlines. 'Ilima
appears intolerant of wildfire as adult plants
typically do not resprout within burned areas. In
General Description.―Sida fallax, known as montane dry shrublands or leeward Hawaii, 'ilima
'ilima, is an erect to prostrate perennial herb or re-established from seed within 4 years of burning
subshrub that grows to 0.2 to 1.5 m tall. The shrub (Sherry and others 1999).
varies from smooth to densely covered in woolly
hairs, the hairs usually stellate. 'Ilima has simple Reproduction.―The flower parts are typically in
leaves, the blades from 1 to 12 cm long, fours or fives and are true flowers with stamen and
lanceolate-ovate to oblong or elliptic. The upper pistils. The ovules are covered by ovaries. Seeds
surfaces are typically bright green and smooth, or are brown or black, obovoid, with three angles, 1.7
sparsely to densely pubescent, the lower surface to 2.2 mm long, minutely pubescent around the
densely pubescent. Flowers arise from an axil, raphe. Somatic chromosome number of 2n = 28
solitary or two to seven per node born in the has been identified (Wagner and others 1990). It is
pedicel, at the end branches or in the leaves. probably wind pollinated, though the native yellow
Corolla is usually yellow to orange-yellow, faced bee species Hylaeus has been observed
sometimes dark maroon at the base. Fruits split visiting the flowers of ‘ilima (Anonymous 2002).
into two or more carpels at maturity. Extensive
variation in the morphology of this species exists Growth and Management.―Easily grown from
in leaf size and shape, stature, pubescence, and seeds, 'ilima makes an excellent xeriscape plant.
arrangement of its inflorescence (Wagner and Seedlings germinate and grow quickly, while
others 1990). cuttings do less well. It requires full sunlight and
light to moderate watering. Upland varieties can be
Range.―Sida is a genus of 125 to 150 species of susceptible to nematodes, while lowland types are
tropical and subtropical regions of the world, from resistant. Leaves may become yellow spotted and
the Pacific Islands to Hawaii. 'Ilima has a series of chlorotic, and will respond well to fertilizer with
similar forms common to all islands but also micronutrients (Krauss 1998). It does well in
exhibits morphological variation from island to sandy or rocky soils.
island. The shrub exists on all the main islands of
Hawaii, including the atoll Midway and the Benefits.―'Ilima is an important plant to Hawaii
pinnacle Nihoa from coastal to subalpine and its people. Four separate types of 'ilima were
712
recognized. These included 'ilima-ku-kala, ilima- Anonymous. 2002. ttp://www.anglefire.com/hi4/
lei, 'ilima-ku-kahakai and 'ilima-koli-kukui nhps/news122 k. html [not paged].
(Stephens 2000). 'Ilima-ku-kala is the wild tall Krauss, B. 1998. How to plant a native Hawaiian
form found in the mountains. 'Ilima-ku-kahakai is garden. http://hawaii.gov/health/
the creeping form found along the coastline. oeqc/garden/index.html
'Ilima-lei were cultivated by the Hawaiians for use
in lei making. 'Ilima-koli-kukui was another Krauss, B. 2000. Native plants used as medicine in
cultivar with reddish brown flowers (Stephens Hawaii. http://library.kcc.hawaii.edu/~
2000). 'Ilima is considered a special flower of the soma/krauss/ilima.html. [not paged]
island of Oahu. 'Ilima leis are among the most
treasured leis and are difficult to make, requiring Sherry, K., J.M. Castillo, and R.B Shaw. 1999.
over a thousand blossoms. At one time, only Effects of wildfire on vegetation and rare plants
royalty were allowed to wear an 'ilima lei in arid montane shrubland. Pohakaloa Training
(Anonymous 2001). 'Ilima was also used Area, Hawaii. 1999 Hawaii Conservation
medicinally. The juice squeezed from the flowers Conference, Honolulu, HI.
was used as a mild laxative for children, and the
root bark mixed with flowers was used to treat Stephens, M. 2000. The comparative ecophsiology
asthma (Krauss 2000). Buds were chewed to of mountain and coastal populations of Sida
quench thirst on hot, dry days. Stems were used to fallax Walp. (Malvaceae). M.S. thesis.
make baskets. 'Ilima is said to be one of the forms University of Hawaii, Hilo, HI.
that Laka, the goddess of hula, could take at will
(Quensell 2000). Quensell, N. 2000. Native Hawaiian plants.
http://www.kcc.hawaii.edu/campus/tour/plants/p
Detrimental Effects.―‘Ilima is known to be host ilima.htm [not paged]
to nonnative rust, Puccinia heterospora, which is
now becoming established on native plants. Wagner, W.L., D.R. Herbst, and S.H. Sohmer.
Infection may be heavy and conspicuous. 1990. Manual of the Flowering Plants of
Hawaii. University of Hawaii Press, Honolulu,
References HI. p. 897-898.
Anonymous. 2001. Hawaii's Island symbols.

http://www.geobop.com/World/NA/US/HI/Islan
d.htm [not paged].
713
Sida rhombifolia L. arrowleaf sida
MALVACEAE
Synonyms: Sida hondensis Kunth in Humb.

Sida ruderata Macfad.
John K. Francis
pale yellow to yellow-orange petals. The fruits are

round, flattened schizocarps, 4 to 5 mm in
diameter and contain 10 to 14 small dark-brown
seeds. Both diploid (2n = 14) and tetraploid (2n =
28) forms exist (Holm and others 1997, Howard
1989, Liogier 1994).
Range.—Arrowleaf sida grows today in over 70

countries throughout the tropical, subtropical, and
warm temperate regions (Holm and others 1997,
Howard 1989). Its original range is not known, but
the presence of multiple subspecies and varieties
seems to indicate that it was from the Old World.
The species was introduced into the United States
in the late 1800’s as a promising fiber crop (Holm
and others 1997).
Ecology.—Arrowleaf sida grows only in disturbed

areas and in situations where overhead
competition is controlled or naturally light. It is
common in cultivated fields, pastures, abandoned
farmland, roadsides, vacant lots, construction
sites, landslides, and river overflow areas. It does
General Description.—Arrowleaf sida is known not survive in tall grass swards, tall brushlands,
by many common names: broom weed, paddy’s and closed forests. Arrowleaf sida grows on both
lucerne, Cuba jute, common sida, escoba dulce, fertile and degraded soils of all textures and
escoba blanca, escoba dura, malva de cochino, derived from most parent materials. It may be
huang hau mu, rhomboid ilima, antibala, idem, found growing from near sea level to 2,000 m in
afata, guanxuma, and basbasot (Holm and others elevation (Holm and others 1997). In Puerto Rico,
1997). It is a short-lived perennial subshrub the species grows in areas receiving from 900 to
(woody stem and herbaceous branches) commonly 3000 mm of precipitation.
growing to 60 cm, but sometimes reaching 1.5 m
in height. (Holm and others 1997, Howard 1989, Reproduction.—Arrowleaf sida flowers and
Liogier 1994). Arrowleaf sida develops a taproot fruits continuously starting at 3 or 4 months of age
and many lateral and fine roots. The plant usually in Puerto Rico. In Central India, plants flower
has a single stem unless disturbed, but may branch from September to December and fruit from
near the ground. The stems are woody, flexible, October to January (Parrotta 2001). A single plant
and tough, and may reach 1 cm or more in basal can produce as many as 11,600 seeds. Individual
diameter. The twigs are slender, green, and seeds have been reported to weigh between 0.0012
semiwoody. The alternate leaves are variable in and 0.0015 g (Holm and others 1997). Seeds
both shape and size. They have a 3- to 8-mm collected in Puerto Rico averaged 0.0016 g/seed or
petiole with broadly ovate to lanciolate (often 625,000 seeds/kg. Sown on moist filter paper
rhomboidal) blades 2 to 6 cm long and serrate at without any pretreatment, 33 percent germinated
the margins, especially from the middle to the tip. within 13 months (author’s observation). Fresh
The axillary flowers are solitary on slender seed is dormant, and dry storage, freezing, various
pedicels 2 to 3 cm long. The corolla contains five light regimes, and several chemical treatments
714
failed to induce germination. However, acid an alkaloid content in the root of 0.1 percent and
scarification, heating, and cold storage succeeded the presence of choline, pseudoephedrine, beta-
in breaking dormancy. Some 80 percent phenethylamine, vascin, hipaphorine and related
germination was obtained at the best soil depth for indole alkaloids (Shaman Australis
germination, which was 0.5 to 2 cm (Holm and Ethnobotanicals 2002). Perhaps because of these
others 1997). Arrowleaf sida plants sprout chemicals, arrowleaf sida is unpalatable to cattle
vigorously if cut. The seeds are dispersed by (Kuniata and Rapp 2001). Arrowleaf sida has
water, farm machinery, ruminant animals, and in significant medicinal applications for which it is
impure agricultural seed. Ants disperse the seeds cultivated throughout India. The pounded leaves
in Africa (Holm and other 1997). are used to relieve swelling, the fruits are used to
relieve headache, the mucilage is used as an
Growth and Management.—Arrowleaf sida emollient, and the root is used to treat rheumatism
plants may reach 0.5 m or more in their first year. (Parrotta 2001). Australian Aborigines use the
Growth is most rapid in warm conditions (days of herb to treat diarrhea. Leaves are smoked in
30 °C and nights of 25 °C). Growth nearly ceases Mexico and a tea is prepared in India for the
below 20 °C. Plants survive frosts and winters as stimulation it provides (Shaman Australis
far north as Tennessee (Holm and others 1997). Ethnobotanicals 2002).
Although the species is able to perpetuate itself as
an annual in difficult climates and under cultivated References
agriculture, the aerial portions of individual plants
in favorable climates may live 3 years, and Food and Agriculture Organization. 2002. Cultural
possibly more by resprouting. Control, but not practices. http://www.fao.org/ag/AGP/AGPC/
elimination of arrowleaf sida in plantations and doc/Publicat/FAOGUL2/B204.htm. 2 p.
pastures may be achieved by hand weeding and
herbicide spraying. Hand pulling and mowing are Guzmán, D.J. 1975. Especies útiles de la flora
only partially effective because arrowleaf sida is Salvadoreña. Ministerio de Educación,
difficult to pull and quickly sprouts after cutting. Dirección de Publicaciones. San Salvador, El
The weed problem from arrowleaf sida becomes Salvador. 703 p.
more severe in reduced tillage agriculture (Holm
and others 1997). Mowing or chain slashing is Holm, L., J. Doll, E. Holm, J. Pancho, and J.
recommended in pastures to suppress arrowleaf Herberger. 1997. World weeds. John Wiley and
sida and other unpalatable Sida species and allow Sons, Inc. New York. 1,129 p.
pasture forage plants to grow (Food and
Agriculture Organization 2002). Effective Howard, R.A. 1989. Flora of the Lesser Antilles,
biological control has been obtained in Australia Leeward and Windward Islands. Vol. 5. Arnold
and Papua New Guinea by introducing the leaf- Arboretum, Harvard University, Jamaica Plain,
eating beetle Calligrapha pantherina, which feeds MA. 604 p.
exclusively on three species of weedy Sida’s
(Kuniata and Rapp 2001). Kuniata, L. and G. Rapp. 2001. Biocontrol of Sida
rhombifolia in Papua New Guinea. http://w
Benefits.—Arrowleaf sida stems are used as ww.nt.gov.au/dpif/pubcat/agnotes/ 542.htm. 4 p.
rough cordage, sacking, and for making brooms.
The stems have a high quality fiber and were once Liogier, H.A. 1994. Descriptive flora of Puerto
exported from India and elsewhere as “hemp” Rico and adjacent Islands. Vol. 3. Editorial de la
(Guzmán 1975, Holm and others 1997). Chemical Universidad de Puerto Rico, Río Piedras, PR.
analysis revealed that the leaves contain 461 p.
respectable amounts of nutrients: 74,000 to
347,000 ppm protein, 94,000 to 475,000 ppm Parrotta, J.A. 2001. Healing plants of Peninsular
carbohydrates, 33,000 to 167,000 ppm fiber, India. CABI Publishing, Wallingford, UK and
14,000 to 71,000 ppm fat, and 16,000 to 81,000 New York. 917 p.
ppm ash. However, it was reported that the root
contained 450 ppm alkaloids and the presence of Shaman Australis Ethnobotanicals. 2002. Sida
ephedrine and saponin (Southwest School of rhombifolia, common sida. http://www.shaman-
Botanical Medicine 2002). Another source reports australis.com/Website/Sidarhombifolia.htm. 1 p.
715
Southwest School of Botanical Medicine. 2002.
Constituents. http://www.rt66.com/hrbmoore/
Constituents/Sida_rhombifolia.txt. 1 p.
716
Smilax domingensis Willd. bejuco de membrillo
SMILACACEAE
Synonyms: Smilax caudata Lundell

Smilax microscola (B.L. Rob.) Killip & C.V. Morton
John K. Francis
Garden 2003). At least one major authority

(Natural Resources Conservation Service 2003)
has bejuco de membrillo as conspecific under S.
smallii Morong that ranges across the Southern
United States. Time will tell whether this union is
reliable or whether further taxonomic adjustments
are necessary. It is not known to have been planted
or naturalized elsewhere.
Ecology.—Bejuco de membrillo has been

collected from near sea level to over 2,000 m
elevation in Costa Rica (Missouri Botanical
Garden 2002). It grows on most soils over nearly
all types of parent material in areas that receive
from about 1400 to over 3200 mm of
precipitation. Bejuco de membrillo is moderately
General Description.—Bejuco de membrillo, intolerant of shade and cannot sustain itself
also called bejuco de sarzaparilla and zarzaparilla, indefinitely under a closed forest canopy. It is
is an evergreen woody vine or scrambling shrub most often found in natural openings of secondary
that reaches 5 to 20 m of extension and 5 cm of and remnant forests. It may also grow in brushy
basal diameter and supports itself by means of pastures, fencerows, and roadsides.
tendrils. The roots produce rhizomes with
occasional tuber-like structures. The plant has a Reproduction.—In Puerto Rico, bejuco de
round, smooth stem, gray in older parts and membrillo blooms during July and August and
greenish-yellow in younger portions. The stem fruits in January and February (Acevedo-
wood does not have annual rings. Some specimens Rodríguez 1985). It is reported to flower and fruit
have small spines or prickles in their lower stems. throughout the year in Nicaragua (Stevens and
Bejuco de membrillo has few branches until it others 2001). A collection of fresh fruits in Puerto
reaches increased sunlight and forms a crown. The Rico weighed an average of 0.486 + 0.018 g/fruit.
alternate leaves are oval to lanceolate, glabrous, Seeds cleaned from these fruits averaged 0.105 +
and leathery. Leaves have short petioles and entire 0.002 g/seed or 9,500 seeds/kg. Planted on
blades 8 to 12 cm long. The inflorescences are commercial potting mix, 50 percent of the seeds
axillary cymes of minute, greenish, fragrant germinated between 65 and 94 days after sowing
flowers. The fruits are 7 mm in diameter, globose, (author’s observation). The seeds are dispersed by
juicy, maturing to dark purple or black, and vine extension and presumably by birds and
contain one seed each (Acevedo-Rodríguez 1985, animals. The plant suckers from rhizomes and
Stevens and others 2001). The fruits taste slightly sprouts heavily when cut.
sour and slightly sweet, somewhat like apples. The
seeds are shiny tan or light brown and flattened on Growth and Management.—The growth of
one side (author’s observation). established bejuco de membrillo sprouts is rapid,
more than 3 m/year. Individual stems live several
Range.—Bejuco de membrillo is native to the years, and plants, by suckering, live almost
Greater Antilles, and Mexico to Panama indefinitely. New plants can probably be
(Acevedo-Rodríguez 1985, Stevens and others established by planting the thickened rhizomes in
2001). Many herbarium specimens also exist from tree-fall gaps and other forest openings. Creating
Colombia, Ecuador, and Peru (Missouri Botanical the essential gaps could also promote the natural
717
establishment of the species. The seeds must be Rico y del Caribe. Iberoamericana de Ediciones,
collected by hand and can be cleaned by Inc., San Juan, PR 566 p.
maceration and wet sieving.
Missouri Botanical Garden. 2002. Flora de Costa
Benefits.—Bjuco de membrillo adds to Rica: Smilax domingensis Willd. http://mobot.
biodiversity, helps to protect the soil, and furnishes org/manual.plantas/051162/S051173.html. 10 p.
food and cover for wildlife. Extracts of the plant
are used as a tonic, a sudorific, and a purifier, and Missouri Botanical Garden. 3002. W3 specimen
to treat skin conditions, rheumatism, and venereal data base--28 Aug. 2003. http://mobot.
disease (Liogier 1990) mobot.org/cgi_bin/search_vast. [not paged].

References Plant profile: Smilax smallii Morong.
http://plants.usda.gov/ [not paged].
Puerto Rico. Vol. 1. General Technical Report Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
SO-58. U.S. Department of Agriculture, Forest Montiel, eds. 2001. Flora de Nicaragua.
Service, Southern Forest Experiment Station, Monographs in Systematic Botany Vol. 85, No.
New Orleans, LA. 331 p. 3. Missouri Botanical Garden, St. Louis, MO.
p. 1,911-2,666.
718
Solanum drymophilum O.E. Schulz erubia
SOLANACEAE
Synonyms: none
John K. Francis
stands are known. Erubia may actually have a

much wider ecological amplitude. The existing
natural stand is situated at 840 m on clayey soil
derived from volcanic rock. The area receives
about 1800 mm of mean annual rainfall. The
plantation is located on clay soil over porous
limestone in an area that receives about 2200 mm
of mean annual precipitation. The plants in the
plantation are healthy but do not appear to be
reproducing on the site. The species appears to be
intolerant of shade and requires disturbance to
become established. Before colonization, it
probably relied on landslides, tree tips, and other
natural disturbances to become established. The
known wild stand is confined to a brushy pasture.
General Description.—Erubia is a rare shrub that
reaches 3 to 6 m and 7.5 cm or more in basal Reproduction.—Erubia appears to flower and
diameter. The plant may have a single stem or fruit throughout the year (U.S. Fish and Wildlife
multiple branches from the base or near the base. service 2002). Flower and fruit production can be
It is supported primarily by semiflexible lateral abundant. The flowers are insect pollinated.
roots. The foliage is concentrated near the ends of Twenty-eight fruits collected from a plantation in
twigs. Alternate lanceolate to oblong leaves, 8 to Río Abajo, Puerto Rico weighed (fresh) an
20 cm long and 2 to 4 cm wide, are long-pointed average of 0.0504 + 0.0050 g. The variability in
at the tip and obtuse or subcordate at the base with weight is high (coefficient of variation = 52
petioles up to 1 cm long. There are sharp, stiff, percent). Air-dried seeds separated from them
yellow spines up to 1 cm long on the mid-vein of averaged 0.00092 g/seed or 1.1 million seeds/kg.
the leaves and sometimes on the twigs and stems. Sown on moist potting mix without pretreatment,
Five-lobbed white flowers about 2 cm across with 89 percent of the seeds germinated between 12
yellow anthers are grouped in lateral or and 50 days after sowing. Birds apparently
subterminal racemes. The fruits are spherical, disperse the seeds. Plants that are disturbed
shiny black berries about 6 mm in diameter that resprout.
contain many tiny seeds (Liogier 1995, Little and
others 1974, U.S. Fish and Wildlife Service 2002). Growth and Management.—Erubia has a
moderate growth rate and probably lives at least
Range.—Erubia is endemic to Puerto Rico. In 10 years. Nursery production appears to be easy,
1974, it was known only from the upper Cordillera but field planting must be accompanied by at least
forest of the eastern and central mountains (Little 2 years of weed control.
and others 1974). Since then, the range has
become constricted and fragmented until it is Benefits.—Erubia helps reforest disturbed sites
currently known from one natural site and one such as abandoned pastures and helps protect the
plantation (Miner Solá 1999) and has been soil. It contributes cover for wildlife, and
declared an endangered species (U.S. Fish and presumably, the fruits are eaten by birds. As most
Wildlife Service 2002). Deforestation and other members of the Solanaceae family, its
misguided weed control have contributed to the tissues probably contain physiologically active
species’ scarcity. chemicals that might be useful in herbal medicine
and control of pathogens.
Ecology.—The environment of only the extant
719
References Agriculture, Forest Service, Agriculture
Handbook. 449. Washington, DC. 1,024 p.
Rico and adjacent islands. Vol. 4. Editorial de la Miner Solá, E. 1999. Arboles y plantas en peligro
Universidad de Puerto Rico, Río Piedras, PR. de extinción en Puerto Rico. Puerto Rico
617 p. Ecológico Vol. 3. First Book Publishing of
Puerto Rico, San Juan, PR. 91 p.
Wadsworth. 1974. Trees of Puerto Rico and the U.S Fish and Wildlife Service. 2002. Species
Virgin Islands. Vol. 2. U.S. Department of accounts: Erubia (Solanum drymorphilum).
http://endangered.fws.gov/i/q/saq50.html. 2 p.
720
Solanum dulcamara L. bittersweet nightshade
SOLANACEAE
Synonyms: none
John K. Francis
the petioles 1 to 4 cm long. Inflorescences are

drooping lateral cymes of usually more than 10
flowers. The corolla is deeply five-lobbed (star
shaped), 12 to 16 mm in diameter, blue to violate
(occasionally white), with exerted yellow anthers
about 5 mm long. The ripe fruits (berries) are
bright red, ovoid to globose, 8 to 12 mm long, and
contain numerous flattened, pale-yellow seeds.
Chromosome number of the species is 2n = 24
(Abrams 1951, Stance 1997).
Range.—Bittersweet nightshade is native to

temperate Eurasia and northern Africa (Crossley
1974, Steyermark 1963). It is generally considered
to be naturalized well into Canada and the United
States including all the conterminous states except
Arizona, New Mexico, Texas, Oklahoma,
Arkansas, Louisiana, Mississippi, Alabama, and
South Carolina (Natural Resources Conservation
Service 2003). Its observation by early botanists
and its presence in native habit led one author to
conclude that it is native to North America (Dean
1940). The species has been widely planted for
ornamental and medicinal purposes.
Ecology.—Bittersweet nightshade often grows in

General Description.—Bittersweet nightshade is wet sites or sites close to water, but also grows in
also known as bitter nightshade, climbing moist upland sites: river banks, weedy ditches, sea
nightshade, woody nightshade, scarlet berry, shores, at the edges of lakes, bogs, and fens,
fellenwort, poison bittersweet, and dulcamara. The roadsides, fencerows, orchards, and open forests.
name bittersweet comes from the report that stems A wide variety of soils are colonized. The species
first taste bitter, then sweet when chewed. It is a tolerates dormant-season temperatures well below
vine or slender scrambling shrub up to 7 m in freezing but for unknown reasons has not invaded
extension and 2 or 3 cm in basal diameter. The tropical climates. Continuous soil moisture is
base of the plant is woody, and the upper required. Seedlings are tolerant enough to grow
herbaceous branches die back to the woody under a moderate forest canopy (Samodien and
portion each year. The slender branches are green others 2003). However, mortality of seedlings
becoming gray-brown and finely furrowed in the tends to be high (89 to 94 percent) (Kollmann and
lower stem. The wood, which has annual rings, is Grubb 1999). Plants in full sun or light shade
creamy-white, brittle, and of light to medium flower and fruit much more heavily than those in
density. The main root grows horizontally just moderate to medium shade. Bittersweet nightshade
below the surface and suckers frequently. Its bark climbs into small trees, shrubs, weeds, and fences
is tan, corky, and knotty. The crushed bark and or remains prostrate as opportunities avail
foliage have an unpleasant smell. The alternate, themselves. Plants of the species grow singly or in
dark-green leaves are ovate to lanceolate, entire, small patches.
simple or with small, opposite lobes or leaflets
near the base. The blades are 5 to 12 cm long and
721
Reproduction.—Bittersweet nightshade flowers glycoside dulcamarin (responsible for the bitter-
from May to September (Horticulture Purdue sweet taste). Decoctions, usually of the twigs, were
1998). Fruit and seed production can be abundant. once used to treat a variety of ailments. These have
Fresh fruits collected by the author in Utah (n = been largely discontinued because of severe side
50) averaged 0.441 + 0.013 g/fruit. They contained effects. The herb is used today to treat scaly skin
an average of 30.9 + 1.6 seeds/fruit, and the air- afflictions and deficient capillary circulation in the
dried seeds averaged 0.00149 + 0.00002 g/seed or skin (Herbdata New Zealand 2003).
670,000 seeds/kg. Crossley (1974) reports 938,000
seeds/kg. The fruits can be collected by hand, References
cleaned by macerating and wet screening, and
stored at 6 percent moisture for periods of less Abrams, L. 1951. Illustrated flora of the Pacific
than 1 year. When seeds are germinated under States. Vol. 3. Stanford University Press,
light, stratification is not necessary and total Stanford, CA. 866 p.
germination percentages of 61 to 98 percent may
be reached. Stems and branches layer (root) Crossley, J.A. 1974. Solanum dulcamara L. In:
whenever they come in contact with the ground. C.S. Schopmeyer, tech. coord. Seeds of woody
Plants coppice when they are cut or damaged. plants of the United States. Agriculture
Nursery propagation is usually by seeds, but the Handbook 450. U.S. Department of Agriculture,
species can be propagated asexually by root or Forest Service, Washington, DC. p. 777-778.
stem cuttings (Crossley 1974). Seeds of wild
plants are dispersed by birds (Samodien and others Dean, C.C. 1940. Flora of Indiana. Department of
2003). Conservation, Division of Forestry,
Indianapolis, IN. 1,236 p.
Growth and Management.—Bittersweet
nightshade branches grow and die back 1 to 3 m or Herbdata New Zealand. 2003. Solanum
more each year. Larger stems examined by the dulcamara, Linn. http://www.herbdatanz.com/
author had two or three growth rings. Plants Bittersweet.html. 4 p.
probably live much longer by resprouting,
suckering, and layering. Bittersweet nightshade Horticulture Purdue. 1998. Bitter nightshade.
usually is not abundant or aggressive enough to Purdue University, West Lafayette, IN. http://
need control. Occasional plants or patches in www.hort.purdue.edu/newcrop/herbhunters/
gardens, orchards, or landscaped areas can be nightshade.html. 1 p.
eliminated by grubbing or spot spraying with
broadleaf or broad-spectrum herbicides. Kollmann, J. and P.J. Grubb. 1999. Recruitment of
fleshy-fruited species under different shrub
Benefits and Detriments.—Bittersweet species: control by under-canopy environment.
nightshade contributes to the aesthetics of Ecological Research. http://dogwood.ag.
wildlands where it grows and furnishes food and utk.edu/literature/1999/99_kollmann.htm. 1 p.
cover for wildlife. It is a pretty plant, cultivated as
an ornamental for its colorful flowers and fruits Lambo Seeds. 2003. Genus: Solanum. http://www.
and dark green foliage. Several species of moths aros.net/~lambo/dulcamara/dulcamara01.htm.
use it for larval food (Savela 2003). The fruits are 8 p.
eaten by birds, including pheasants (Phasianus
colchicus). Although all parts are poisonous to Natural Resources Conservation Service. 2003.
cattle, horses, and sheep, which normally will not Plants profile: Solanum dulcamara L., climbing
eat it, the stems are eaten by muskrats (Ondatra nightshade. http://plants.usda.gov.cgi_bin/
zibethica). The berries have been used to poison plant_profile.cgi?symbol=SODU. 6 p.
rabbits and dogs (Steyermark 1963). Although the
fruits have an attractive appearance, the flavor is Samodien, J., M. Wood, K. Epps, and M. Thandy.
so disagreeable that it is doubtful that anyone 2003. An introduction to common exotic species
would mistakenly eat enough to be poisoned. Also, to British Columbia’s ecosystems. University
poisonous principals are low in ripe fruits (Lambo College of Fraser Valley, Abbotsford, BC,
Seeds 2003). Bittersweet nightshade contains the Canada. http://www.ucfv.bc.ca/biology/Biol210/
alkaloid solanine (the poisonous principal) and the 1999/Exotic/Exotic_plant.htm. 9 p.
722
Savela, M. 2003. Solanum L. http://www. Stance, C. 1997. New flora of the British Isles. 2nd.
funet.fi/pub/sci/bio/life/plants/magnoliophyta/ Ed. Cambridge University Press, Cambridge,
magnoliophytina/magnoliopsida/solanaceae/ UK. 1,130 p.
solanum/. 5 p.
Steyermark, J.A. 1963. Flora of Missouri. The
Iowa State University Press, Ames, IA. 1,725 p.
723
Solanum persicifolium Dunal berenjena de playa
SOLANACEAE
Synonyms: Solanum ignaeum parvifolium Vahl

Solanum persicifolium angustifolium Dunal in DC.
Solanum persicifolium belloni O.E. Schulz in Urban
Solanum persicifolium parvifolium (Vahl) O.E. Schulz
John K. Francis
are lateral and contain up to 15 small, blue, violet,
or rarely white flowers. The fruits are globose, red
berries about 5 mm in diameter with the calyx still
attached. Each contains several flattened, yellow
seeds (author’s observation, Liogier 1995).
Range.—Berenjena de playa is native to

Hispaniola, Puerto Rico, and the Virgin Islands
(Liogier 1995). It is not known to have been
Ecology.—Berenjena de playa is most common

near the sea shore but does occur several km
inland. It grows on the whole range of soil textures
on well-drained soils. In Puerto Rico, the species
colonizes areas that receive from 750 mm to about
1700 mm of mean annual precipitation from near
sea level to over 400 m of elevation. Berenjena de
playa is moderately intolerant of shade and grows
within low basal-area forest as well as in
openings. The plants are apparently avoided by
cattle and appear to benefit by overgrazing. Some
degree of disturbance appears to be necessary for
establishment. Vázquez and Kolterman (1998)
report it as “occasional” in pasture, shrubland, and
woodland. It may also be found on roadsides,
General Description.—Berenjena de playa, trails, fencerows, bluffs, and beach strands.
which means beach eggplant, is a semideciduous
shrub usually about 2 m in height with a basal Reproduction.—Berenjena de playa flowers and
diameter of 1.5 cm. The species is also known as fruits more or less continuously in moist habitat
rubia. It may have a single or multiple stems and but seasonally in dry habitat. A group of fruits
is supported by a stout taproot and many lateral collected in Puerto Rico averaged 0.0708 + 0.0035
roots. The most striking feature of the species is g/fruit. They contained from three to 20 seeds
the sharp yellow to orange prickles on the stem, each, which averaged (air-dried) 0.00137 + 0.0003
branches and midveins of the leaves. The prickles g/seed or 730,000 seeds/kg. The seeds are
may reach as much as 13 mm long. However, the apparently dispersed by birds that eat the fruits.
density of prickles is variable and some plants Plants sprout when they are broken or cut.
have none. The branches and twigs are slender and
form a thin crown. The alternate leaves, which Growth and Management.—Sprouts of
have 4- to 7-mm petioles, are thin, oblong to berenjena de playa grows about 0.5 to 1.0 m in the
linear-oblong, 5 to 13.5 cm long and 0.6 to 3 cm first year and slower thereafter. Seedling growth is
wide with pointed ends and edges rolled under undoubtedly slower. Stems apparently live about 3
(revolute). The leaves take on a wilted appearance to 10 years, but the rootstalks resprout and may be
during dry periods. The raceme-like inflorescences older than the top. No management experience has
724
been published. Small areas of disturbance in References
suitable habitat would probably encourage natural
establishment. The species is rarely common Liogier, H.A. 1995. Descriptive flora of Puerto
enough to control as a weed in pastures or Rico and adjacent islands. Vol. 4. Editorial de la
plantations. Universidad de Puerto Rico, Río Piedras, PR.
617 p.
Benefits.—Berenjena de playa contributes to the
biodiversity of habitat where it grows, helps to Vázquez, O.J. and D.A. Kolterman. 1998. Floristic
protect the soil, and furnishes food and cover for composition and vegetation types of the Punta
wildlife, particularly birds. Guaniquilla Natural Reserve—Cabo Rojo,
Puerto Rico. Caribbean Journal of Science 34(3-
4): 265-279.
725
Solanum torvum Sw. turkey berry
SOLANACEAE
Synonyms: Solanum ficifolium Ortega

Solanum daturifolium Dunal
Solanum torvum var. daturifolium (Dunal) Schulz
Solanum maccai L.C. Rich. ex Spreng.
John K. Francis
and contain numerous flat, round, brown seeds

1974).
Range.—Turkey berry apparently is native from

Florida and southern Alabama through the West
Indies and from Mexico through Central America
and South America through Brazil (Little and
others 1974). Because of its rapid spread as a
weed in disturbed lands, it is difficult to tell which
populations are native and which are introduced.
Turkey berry has been introduced and naturalized
throughout tropical Africa, Asia, Australia, and
General Description.—Turkey berry is an erect the Pacific Islands including Hawaii, Guam, and
spiny shrub that is also known by prickley American Samoa (Pacific Island Ecosystems at
solanum, shoo-shoo bush, devil’s fig, wild egg Risk 2001).
plant, boo, terongan, berenjena cimarrona,
berenjena de gallina, berenjena silvestre, tabacón, Ecology.—In Puerto Rico, turkey berry grows in
pendejera, tomatillo, bâtard balengène, zamorette, upland sites that receive from about 1000 to 4000
friega-platos, and many other names (Howard mm of annual precipitation. It also grows in
1989, Little and others 1974, Pacific Island riparian zones in drier areas. Turkey berry grows
Ecosystems at Risk 2001). It is usually 2 or 3 m in on all types of moist, fertile soil at elevations from
height and 2 cm in basal diameter, but may reach near sea level to almost 1,000 m in Puerto Rico
5 m in height and 8 cm in basal diameter. The (Little and others 1974) and 2,000 m in Papua
shrub usually has a single stem at ground level, New Guinea (Pacific Island Ecosystems at Risk
but it may branch on the lower stem. The stem 2001). Given an equal start after disturbance,
bark is gray and nearly smooth with raised turkey berry quickly overtops most herbs, grasses,
lenticels. The inner bark has a green layer over an and other shrubs. It grows best in full sunlight and
ivory color (Little and others 1974). The plants does well in light shade or shade for part of the
examined by the author, growing on firm soil, had day, but cannot survive under a closed forest
weak taproots and well developed laterals. The canopy. Turkey berry single plants, groups, and
roots are white. Foliage is confined to the growing thickets are most frequently seen on roadsides,
twigs. The twigs are gray-green and covered with vacant lots, brushy pastures, recently abandoned
star-shaped hairs. The spines are short and slightly farmland, landslides, and river banks.
curved and vary from thick throughout the plant,
including the leaf midrib, to entirely absent. The Reproduction.—Flowering and fruiting is
leaves are opposite or one per node, broadly ovate continuous after the shrubs reach about 1 to 1.5 m
with the boarder entire or deeply lobed. The in height. Ripe fruits collected in Puerto Rico
petioles are 1 to 6 cm long and the blades are 7 to averaged 1.308 + 0.052 g. Air dry seeds from
23 by 5 to 18 cm and covered with short hairs. The these fruits weighed an average of 0.00935 g or
flowers are white, tubular with 5 pointed lobes, 1,070,000 seeds/kg. These seeds were sown on
and grouped in corymbiform cymes. They are commercial potting mix and 60 percent
shed soon after opening. The fruits are berries that germinated between 13 and 106 days following
are yellow when fully ripe. They are thin-fleshed sowing. The seedlings are common in recently
726
disturbed ground. Frugivorous birds eat the fruits of the interspecific eggplant hybrid Solanum
and spread the seeds (Pacific Island Ecosystems at melongena x Solanum torvum.
Risk 2001). Turkey berry can be propagated http://www.actahort.org/books/467/467_17.htm.
vegetatively by placing branch cuttings, with or 1 p.
without leaves, in a mist chamber for one month
(Badola and others 1993). CPR Environmental Education Centre. 2001.
Medicinal plants. http://cpreec.org/edu/
Growth and Management.—Turkey berry grows medi-pln.htm. 4 p.
about 0.75 to 1.5 m in height per year. The species
is not long-lived; most plants live about 2 years. Horzog, F. and D. Gautier-Béguin. 2001.
Physical control of the shrub may be done by Uncultivated plants for human nutrition in Côte
grubbing out the plants; lopping will not kill them. d’Ivorie. http://www.fao.org/docrep/W3735e/
They can be killed by translocated herbicides w3735e10.htm. 12 p.
applied to the leaves or the cut stumps (Pacific
Island Ecosystems at Risk 2001). Howard, R.A. 1989. Flora of the Lesser Antilles,
Benefits.—The wood is soft and light and of little Dicotyledoneae. Part 3. Vol. 6. Arnold
use except for emergency fuel. The fruits are Arboretum, Harvard University, Jamaica Plain,
edible and used in Thai cuisine (Royal MA. 658 p.
Horticultural Society 2001) and incorporated into
soups and sauces in the Ivory Coast (Herzog and Liogier, H.A. 1995. Descriptive flora of Puerto
Gautier-Béguin 2001). Turkey berry contains a Rico and adjacent islands. Vol. 4. Editorial de la
number of potentially pharmacologically active Universidad de Puerto Rico, Río Piedras, PR.
chemicals including the sapogenin steroid, 617 p.
chlorogenin (Badola and others 1993). Aqueous
extracts of turkey berry are lethal to mice or Liogier, H.A. 1990. Plantas mediciales de Puerto
depress the erythrocytes, leukocytes and platelets Rico y del Caribe. Iberoamericana de Ediciones,
in their blood (Tapia and others 1996). A related Inc., San Juan, PR. 566 p.
chemical, cholecalciferol, is the active ingredient
in a number of commercial rodentacides Little, E.L., Jr., R.O. Woodbury, and F.H.
(American Board of Veterinary Toxicology 2001). Wadsworth. 1974. Trees of Puerto Rico and the
Extracts of the plant are reported to be useful in Virgin Islands. Vol. 2. Agriculture Handbook
the treatment of hyperactivity (Null 2001), colds 449. U.S. Department of Agriculture,
and cough (CPR Environmental Education Centre Washington, DC. 1,024 p.
2001), pimples, skin diseases, and leprosy (Liogier
1990). Turkey berry is being crossed with Null, G. 2001. The biochemical activity of plants.
eggplant in an attempt to incorporate genes for http://www.garynull.com/Documents/phytoche
resistance to Verticillium wilt into the vegetable micals/Phytochemicals5.htm. 15 p.
(Bletsos and others 2001).
Pacific Island Ecosystems at Risk. 2001. Invasive
References plant species: Solanum torvum Sw., Solanaceae.
http://www.hear.org/pier/sotor.htm. 2 p.
American Board of Veterinary Toxicology. 2001.
An overview of cholecalciferol toxicosis. Royal Horticultural Society. 2001. Fruits and
http://www.abvt.org/tow.htm. 6 p. vegetables: felty leaves.
http://www.rhs.org.uk/about/mn_pubs_journals_
Badola, K.C., P. Mohinder; H.C.S. Bhanderi, and garden_0898_aubergine_b.asp. 1 p.
M. Pal. 1993. Vegetative propagation of
ranbaigan (Solanum torvum Sw.) by rooting
branch cuttings. Indian Forester 119(12): 1,027-
1,028.
Bletsos, F.A., D.G. Roupakias, and C.C.

Thanassoulopoulos. 2001. Morphological traits
727
Tapia A., R., A. Astudillo V., R. Uribe H., 1996.
Resultados preliminaries del efetco de Solanum
torvum y Plantago major sobre la proliferación
de células hematopoyéticas in vivo e in vitro.
Resumen de Ponencias de Primer Congreso
Nacional de Plantas Medicinales de México. 24-
30 June 1996. Tlaxcala, Tlax., Mexico. 102-
104.
728
Sorbus scopulina Greene Greene’s mountain-ash
ROSACEAE
Synonyms: Sorbus sambucifolia non Roem.

Sorbus cascadensis G.N. Jones
John K. Francis
Conservation Service 2003, Treeguide 2003).

There are two varieties. The typical variety
occupies the entire inland range and var.
cascadensis (G.N. Jones) C.L. Hitchc. occupies
coastal habitat from British Columbia into
California (Natural Resources Conservation
Service 2003). In addition, an intergeneric hybrid
named Amelasorbus jackii Rehder between
Amelanchier alnifolia Nutt. and Greene’s
mountain-ash has been observed in Oregon and
Idaho (Love 2003). Greene’s mountain-ash has
been widely planted, but no reports of it
naturalizing outside its range are known.
Ecology.—Greene’s mountain-ash colonizes well-

drained soils on rocky hillsides, canyons, wooded
slopes, forest clearings, avalanche chutes, and
General Description.—Greene’s mountain-ash, along streams. It grows as dispersed individuals or
also known as western mountain-ash, is a in nearly pure (usually small) clumps. The sites are
deciduous shrub or small tree ranging in height moist; the species is not drought resistant.
from 1 to 6 m and up to 10 cm in stem diameter. It Elevations range from near sea level in the
usually has multiple stems with smooth yellowish northern portions of its range to 3,000 m in the
to grayish-red bark and slender light brown twigs Southern Rocky Mountains. Greene’s mountain-
that are white-hairy when young. Winter buds are ash is moderately intolerant of shade; it sometimes
glutinous and glossy. The alternate leaves are 10 to grows in the understory of open stands but usually
20 cm long, odd-pinnately compound with seven grows in openings. Low-vigor plants are also
to 15 lanceolate leaflets that are nearly sessile, common under closed, but not dense, conifer
pointed, and serrate on the margins. They are thin, stands (Alaska Department of Fish and Game
shiny-green above and paler beneath. 2001, Ibiblio 2003, Luna and Wick 2001, Umatilla
Inflorescences are much-branched corymbs, 6 to National Forest 2003, Utah State University
12 cm broad, that contain many 10-mm broad, Horticulture 2003).
white to cream, five-petaled flowers. Fruits, which
grow in clusters, are shiny, orange to red, berry- Reproduction.—Flowering of Greene’s
like, 5- to 10-mm-long, globose pomes with an mountain-ash occurs between May and July
attached calyx at the apex. Each contains up to depending on location (Stein 2003). The flowers
eight flattened, brown or red-brown seeds 3 to 4 are pollinated by insects (Ibiblio 2003). The fruits
mm long (Davis 1952, Viereck and Little 1972, ripen in July through September and remain on the
Welsh 1974). shrubs into the winter (Alberta Agriculture, Food
and Rural Development 2003). Fruits can be
Range.—Greene’s mountain-ash is native to broad collected in quantity by hand after the fruits turn
areas and disjuncts in the southern half of Alaska, orange or red and the seeds are brown. The seeds
disjuncts and broad areas of the Yukon, Northwest are extracted by maceration, washing, and
Territories, British Columbia, Alberta, screening. The weight of seeds has been reported
Saskatchewan, Montana, Idaho, Washington, and at 100 seeds/g (Luna and Wick 2001) and 180
scattered areas in Oregon, California, Wyoming, seeds/g (Browse-Shrub and Forb Committee of the
North Dakota, South Dakota, Utah, Colorado, Association of Official Seed Analysis 1985). In a
Nevada, and New Mexico (Natural Resources
729
collection made by the author in Utah, fresh fruits protect the soil, adds to the aesthetics of wildland
weighed an average of 0.333 + 0.009 g/fruit, and sites, especially with its yellow to orange-red fall
air-dried seeds weighed an average of 0.00386 + foliage and red-orange berries, and furnishes cover
0.0001 g/seed (259 seeds/g). There were 24 to 117 for wildlife. The foliage and twigs are browsed by
fruits/cluster and an average of 2.4 seeds/fruit. deer (Odocoileus spp.), elk (Cervus canadensis),
Seed will remain viable in sealed, refrigerated moose (Alces americana), and to a lesser extent,
containers at low relative humidity for 5 years. cattle. The fruits are eaten by black bears
Recommended pretreatment is a 3:1 (Euarctos americanus), martin (Martes
water/hydrogen peroxide soak for 10 minutes americana), fisher (Martes pennanti), rodents,
followed by a 25-hour water soak and a cold American robin (Turdus migratorius), hermit
stratification in peat at 3 °C for 90 to 120 days. (Catharus guttatus), gray-cheeked (Catharus
Germination varies from 90 to 100 percent (Luna minimus) and Swainson’s (Catharus ustulatus)
and Wick 2001). A test of viability can be made by thrushes, pine grosbeaks (Pinicola enucleator),
incubating imbibed, excised embryos at 20 °C for Bohemian waxwings (Bombycilla garrulus),
6 days. Viable embryos either retain their freshly northern flickers (Colaptes auratus), Steller’s jays
excised appearance or become deep green; (Cyanocitta stelleri), and blue grouse
nonviable embryos deteriorate or turn pale yellow (Dendragapus obscurus) (Alaska Department of
green (Umatilla National Forest 2003). In the wild, Fish and Game 2001, Gullion 1964, Stein, 2003,
seeds are dispersed mainly by birds. Greene’s Wier 2003). The fruits are edible to humans, fresh,
mountain-ash may be asexually reproduced using cooked, and dried, but it is necessary to wait until
cuttings. Late-summer semihardwood stem the bitterness disappears after multiple frosts
cuttings 15 to 25 cm long and 0.3 to 1.3 cm in (Moser 2003). A wine is also made from the
diameter are recommended. After hormone berries (Keller 2003). In herbal medicine,
treatment, cuttings placed in a bottom-heated infusions of bark have been used to reduce fevers
mistbed for 6 weeks yielded 47 percent rooted and as a tonic, and infusions of branches have been
cuttings (Luna and Wick 2001). given to children with bed-wetting problems
(Ibiblio 2003). Greene’s mountain-ash is planted
Growth and Management.—This species grows to a limited extent as an ornamental, especially in
slowly as seedlings and only at an intermediate naturalistic landscape settings. The wood is soft
rate later. Plants are able to flower and fruit after 3 and has a specific gravity of 0.59 g/cm3 (Ibiblio
to 5 years (Luna and Wick 2001). Greene’s 2003). It is probably useful for firewood, but other
mountain-ash is usually grown for revegetation uses are not known.
projects from seed in nurseries as container or
bare-root stock. Potted seedlings can be expected References
to reach 15 cm tall with a 2.0 cm caliper by the
end of the first year (Luna and Wick 2001). They Alaska Department of Fish and Game. 2001.
are outplanted as either 1- or 2-year-old stock. Native Alaskan and exotic plants used by
Greene’s mountain-ash planted in seven sites in wildlife. http://www.state.ak.us/adfg/wildlife/
Alberta averaged 1.2 to 2.3 m in height after 4 to 8 geninfo/birds.htm. 9 p.
years of growth (Alberta Agriculture, Food and
Rural Development 2003). Direct seeding is also Alberta Agriculture, Food and Rural Development.
possible. Drilling seed 2 mm deep in fertile, well- 2003. Evaluating woody plants for hardiness and
drained soil is recommended. Many seeds will not landscaping quality in Alberta: Sorbus sp.
germinate until the second or third year (Umatilla (mountain ash). http://www.agric.gov.ab.ca/
National Forest 2003). Although the species is crops/trees/rwptp/ sorbus.html. 3 p.
adapted to many soil conditions, it is sensitive to
high pH in the nursery and the field and is Browse-Shrub and Forb Committee of the
damaged by the pear slug [Caliroa cerasi (L.)] Association of Official Seed Analysis. 1985.
(Alberta Food and Rural Development 2003). Handbook on seeds of browse-shrubs and forbs.
Browsing keeps the plants compact in many Technical Publication R8-TP8. U.S. Department
locations (Clark 1976). The author knows of no of Agriculture, Forest Service, Southern Region,
published data on management of natural stands. Atlanta, GA. 246 p.
Benefits.—Greene’s mountain-ash is an important Clark, L.J. 1976. Wild flowers of the Pacific
component of the Western shrub community and Northwest. Gray’s Publishing, Limited. Sidney,
furnishes a number of benefits. The species helps BC, Canada. 604 p.
730
Davis, R.J. 1952. Flora of Idaho. Brigham Young ash. http://plants.usda.gov/cgi_bin/plant_profile.
University Press, Provo, UT. 836 p. cgi?symbol=SOSC2. 4 p.
Gullion, G.W. 1964. Wildlife uses of Nevada Stein, W.I. 2003. Sorbus L, mountain-ash. In: F.T.
flora. Contr. Toward a Flora of Nevada 49. U.S. Bonner, and R.G. Nisley, eds. Woody plant seed
National Arboretum, Washington, DC. 170 p. manual. U.S. Department of Agriculture, Forest
Service, Washington, DC. http://wpsm.net/
Ibiblio. 2003. Plants for a future: database search Sorbus.pdf. 12 p.
results: Sorbus scopulina. University of North
Carolina, Chapel Hill, NC. http://www.ibiblio. Treeguide. 2003. Greene mountain-ash, Sorbus
org/pfaf.cgi-bin/arr_html?Sorbus+scopulina& scopulina, Greene, Rosaceae.
CAN =LATIND. 6 p. http://www.treeguide.com/Species.asp?Region=
NorthAmerican&SpeciesID=1021. 2 p.
Keller, J. 2003. The winemaking home page:
mountain ash. http://winemaking.jackkeller.net/ Umatilla National Forest. 2003. Umatilla National
mtnash.asp. 2 p. Forest native plant species: Sorbus scopulina.
Love, R. 2003. Amelasorbus: An intergeneric Umatilla National Forest, Pendleton, OR. http://
hybred and a new taxon for the Oregon www.fs.fed.us/r6/uma/native/ts100.htm. 8 p.
checklist. http://www.oregonflora.org/ofn/v4n2/
amelasorbus.html. 3 p. Utah State University Horticulture. 2003. Sorbus
scopulina. http://www.usu.edu/natives/shrubs/
Luna, T. and D. Wick. 2001. Propagation protocol scopulina.htm. 1 p.
for production of container Sorbus scopulina
Greene var. scopulina Greene plants (172 ml Viereck, L.A. and E.L. Little, Jr. 1972. Alaska
containers), Glacier National Park, West trees and shrubs. Agriculture Handbook 410.
Glacier, Montana. http://www. U.S. Department of Agriculture, Forest Service,
nativeplantnetwork.org. 8 p. Washington, DC. 265 p.
Moser, R. 2003. Edible Sierra Nevada plants: Welsh, S.L. 1974. Anderson’s flora of Alaska.
Rosaceae-rose family, mountain ash. Brigham Young University Press, Provo, UT.
Backcountry rangers, Fresno, CA. http://www. 724 p.
backcountryrangers.com/edibles/SORBUS.html.
2 p. Wier, S.K. 2003. Greene’s mountain-ash of the
southern Rocky Mountains. http://home.
Natural Resources Conservation Service. 2003. earthlink.net/~swier/MtnAsh.html. 1 p.
Sorbus scopulina Greene, Greene’s mountain
731
Spermacoce verticillata L. botón blanco
RUBIACEAE
Synonyms: Borreria verticillata (L.) Meyer

Borreria podocephala DC.
Borreria stricta DC.
John K. Francis
amount of fine roots. Branching is bifurcate or

ternate. The leaves are opposite but appearing with
two or a cluster of smaller leaves in whorls at the
nodes. The leaves are sessile or nearly so, linear or
linear-lanceolate, 2 to 6 cm long, and pointed at
both ends. The tiny white flowers grow in heads or
glomerules in terminal or lateral positions. The
terminals continue to grow through the center of
the inflorescence so that the fruits develop at
nodes in mid-stem. The capsules are oblong or
subglobose with two carpels, each with one seed.
The seeds are ellipsoidal, brown, and about 1 mm
long (Correll and Johnston 1970, Howard 1989,
Liogier 1997).
Range.—Botón blanco appears to be a native of

the New World and possibly Africa, but the
original range is uncertain. It grows as a native or
naturalized species from Florida through the West
Indies, and Texas through Central and South
America to Argentina, and throughout the moist
portions of Tropical Africa and Madagascar
(Burkill 2000, Howard 1989, Instituto Botánico
Darwin 2002, Liogier 1997, Natural Resources
Conservation Service 2002). It has also been
reported from India (Kudremukh Wildlife
Foundation 2002).
General Description.—Botón blanco, which Ecology.—Botón blanco grows on moist soils,

means white button in Spanish (Puerto Rico), is both acid and alkaline, of all textures derived from
also known as shrubby false buttonwood, nearly all types of rocks. The species grows in
vassourinha, cardio de frade, poaia, borrerie areas that receive from about 750 to 3000 mm of
verticillée, éribun, and many other names (Burkill annual rainfall from near sea level to 600 m or
2000, Instituto de Biociências 2002, Natural more in elevation in Puerto Rico. It grows on sand
Resources Conservation Service 2002). It is a fine- and caliche in prairies and openings in Texas
stemmed scrambling shrub that may reach a few (Jones 1975). In Puerto Rico, it grows on
meters of lateral extension and 1.2 m in height as a roadsides, construction sites, old fields, and
free-standing plant. The square stems are pastures. Botón blanco competes with cultivated
herbaceous to semiwoody in their first year, crops and plantations in Brazil and Africa (Holm
becoming woody and more rounded in the and others 1997). It is one of the major invaders of
following year. The brown stems reach a abandoned pastures and slash-and-burn fields
maximum diameter of about 8 mm, have a solid (Ministério de Ciencia e Tecnología 2002). The
pith, and lack visible annual rings. Botón blanco species requires disturbance to establish itself and
produces a weak taproot, many important laterals must have full or good partial sunlight to survive.
that are pale yellow and flexible, and a moderate It competes well with disbursed grass and weeds,
732
but is overcome by dense, tall grass, brush, and treatment of schistosomiasis, and as an abortive.
trees. Because of grazing, mowing, and An essential oil extracted from the leaves has been
cultivation, most plants do not progress beyond shown to inhibit Escherichia coli and
the herbaceous stage. If allowed to grow, they will Staphylococcus aureus (Burkill 2000).
form dense clumps and mats.
References
Reproduction.—In Brazil botón blanco blooms
from February through August (Instituto de Animal Science at Cornell University. 2002. Mind
Biociêcias 2002). In Texas, it flowers from March bending beta-carbolines. http://www.ansci.
through May (Correll and Johnston 1970). cornell.edu/plants/toxicagents/betacarbolines/
Flowering is almost continuous in moist portions bcarb.html. 5 p.
of Puerto Rico. Plants begin blooming in the
nursery at about 9 months. The flowers are Burkill, H.M. 2000. The useful plants of West
pollinated by several species of bees (Instituto de Tropical Africa. Royal Botanic Gardens, Kew,
Biociências 2002). Seeds collected in Puerto Rico UK. 686 p.
averaged 0.00016 g each or 6,250,000 seeds/kg.
Sown on peat without pretreatment, these seeds Correll, D.S. and M.C. Johnston. 1970. Manual of
germinated at 49 percent beginning in 13 days and the vascular plants of Texas. Texas Research
ending at 74 days. The seeds are disbursed by Foundation, Renner, TX. 1,881 p.
grazing animals and farm equipment. Established
plants root readily at the nodes when covered by Environnement et Développement du Tiers-
soil or rotting plant material. monde. 2002. Reconnaisssez, Protégez et
utilisez: Borreria vericillata L. Rubiaceae.
Growth and Management.—Botón blanco http://www.enda.sn/plantesmed/borreria.html.
seedlings grow slowly at first but begin rapid 2 p.
growth after about 6 months. Twenty-three 9-
month-old nursery plants averaged 64 cm in Holm, L., J. Doll, E. Holm, J. Pancho, and J.
height with a maximum of 109 cm. Botón blanco Herberger. 1997. World weeds. John Wiley &
shrubs appears to live at least 4 years and probably Sons, Inc. New York. 1,129 p.
much longer in Puerto Rico. Botón blanco is
controlled in crops and pasture by cultivation, Howard, R. A. 1989. Flora of the Lesser Antilles,
mowing, and spraying with broadleaf herbicides. Leeward and Windward Islands. Vol. 6. Arnold
The importance value of botón blanco in a Arboretum, Harvard University, Jamaica Plain,
Colombian pasture was reduced by 39 percent by MA. 658 p.
simply fertilizing with potassium and sulfur to
increase the vigor of the pasture grasses (Tejos Instituto de Biociências. 2002. Borreria
1981). verticillata. Instituto de Biociêcias,
Universidade de São Paulo, São Paulo, Brazil.
Benefits.—Botón blanco is a forage plant, but not http://ib.usp.br/beeplant/bove.htm. 1 p.
one highly favored by livestock. African material
was found to contain 0.2 percent alkaloid Instituto Botánico Darwin. 2002. Catálogo de las
including borreine and borreverine (Burkill 2000). plantas vasculares de la Argentina. http://
At least part of the alkaloids are beta-carbolines www.darwin.edu.ar/Catalogo/rubiaceae.pdf.
and would represent a poisoning hazard if they 22 p.
were present in higher concentrations (Animal
Science at Cornell University 2002). Botón blanco Jones, F.B. 1975. Flora of the Texas Costal Bend.
has a number of uses in herbal medicine, most Mission Press, Corpus Christi, TX. 262 p.
frequently for skin conditions. In Africa, leaf
extracts are used to treat leprous conditions, Kudremukh Wildlife Foundation. 2002. Checklist
furuncles, ulcers, and gonorrheal sores (Burkill of non-woody plants reported from Kudremukh
2000, Environnement et Développement du Tiers- National Park. http://kudremukh.org/
monde 2002). A lotion is prepared to relieve skin kudremukh/nonwoody.html. 4 p.
itches (Liogier 1990). Other preparations are used
internally to treat diarrhea, as a diuretic in the
733
Liogier, H.A. 1997. Descriptive flora of Puerto http://www.mct.gov.br/prog/ppg7/projetos/
Rico and adjacent islands. Vol. 5. Editorial de la proj991.pdf. 16 p.
436 p. Natural Resources Conservation Service. 2002.
Plants profile: Spermacoce verticillata L.,
Liogier, H.A. 1990. Plantas medicinales de Puerto shrubby false buttonweed. http://plants.usda.
Rico y del Caribe. Iberoamericana de Ediciones, gov/cgi_bin/plant_profile.cgi?symbol=SPVE2.
Inc., San Juan, PR. 566 p. 2 p.
Ministério de Ciencia e Tecnología. 2002. Tejos, R. 1981. Efecto del potasio y azufre sobre el
Recuperação de areas de pastagems pastizal nativo de una sabana. 1. Producción
abandonadas e degradadas através de sistemas contenido de material seca y composicion
agroflorestais na Amazônia Ocidental. botánica. Agronomía Tropical 29(6): 503-516.
734
Strumpfia maritima Jacq. pride-of-Big-Pine
RUBIACEAE
Synonyms: none
John K. Francis
headlands, coastal cliffs, beach strands, and coastal

flats from near sea level to a few meters in
elevation. It frequently grows in cracks in rocks or
rocky rubble. The soils are salty, and the plant
tolerates sea water overwash and heavy salt spray.
Mean annual precipitation ranges from about 600
to 900 mm. Pride-of-Big-Pine grows singly,
scattered, or in small thickets. When growing in
severe habitats, it sometimes forms natural
bonsais.
Reproduction.—Pride-of-Big-Pine blooms and

fruits throughout the year (Nelson 1996). The
flowers are probably pollinated by insects. The
fresh fruits weigh about 0.02 g each. A collection
of air-dried seeds (n = 99) from Puerto Rico
General Description.—Pride-of-Big-Pine, also averaged 0.0052 + 0.0001 g/seed or 192,000
known as snowbank, rosemary, strumpfia, lirio, seeds/kg. The seeds were placed to germinate on
and rosarin-bord-de-mer, is an evergreen shrub the surface of moist peat, and 6 percent germinated
usually 1 m or less in height but sometimes between 4 and 6 months after sowing. Germination
reaching 2 m. Most plants have a profusion of gray is epigeous. Plants coppice when broken or cut
stems and branches. The twigs are slender (3 to 4 (author’s observation). The seeds are probably
mm in diameter), and prominently ringed from dispersed by birds.
persistent stipule bases. The whorled dark-green
foliage is crowded at the ends of branches. Leaves Growth and Management.—Pride-of-Big-Pine
are linear and leathery, 1 to 3 cm long with grows slowly. Planting experience for this species
petioles about 1 mm long, with an entire and has not been published. Because it is rare or
revolute margin. Inflorescences are axillary, few- uncommon and never aggressive, control is not
flowered racemes. The individual flowers are necessary. Probably the best management is to
white to pink, campanulate with a tube about 1 totally protect existing stands and suitable habitat
mm long and corolla 6 mm across with five lobes. from disturbance.
The fruit, a white, fleshy globose drupe, is 3 to 6
mm in diameter with a persistent calyx. It contains Benefits.—Pride-of-Big-Pine helps protect the soil
one or two seeds (Howard 1989, Liogier 1997, in coastal sites and provides cover for wildlife.
Nelson 1996). The Exuma Island iguana (Cyclura cychlura
figginsi), an endangered species from the
Range.—Pride-of-Big-Pine is native to the Florida Bahamas, feeds on fruits, flowers, buds, and leaves
Keys, the Bahamas, the Greater and Lesser of Pride-of-Big-Pine (Knapp 2002). Probably
Antilles, Grand Cayman, several islands north of many other species eat the fruits. It is a pretty plant
Venezuela, and Yucatan, Mexico (Howard 1989, and adds to the aesthetics of coastlines. If
Liogier 1997, Nelson 1996). The species is listed propagation and cultural techniques can be worked
as endangered in Florida (Florida Fish and out, it would make a valuable ornamental.
Wildlife Conservation Commission 1997). It is not Infusions of the leaves are reported to be a
known to have naturalized outside its native range. stimulant useful in treating poisonous bites, fever,
and stupor and weakness caused by fever (Liogier
Ecology.—Pride-of-Big-Pine is confined to 1990).
coastal areas. Types of habitat include rocky
735
References page. http://www.iucn-isg.org/actionplan/ch2/
exumaisland.php. 5 p.
Florida Fish and Wildlife Conservation
Commission. 1997. Florida’s endangered Liogier, H.A. 1990. Plantas medicinales de Puerto
species, threatened species, and species of Rico y del Caribe. Iberoamericana de Ediciones,
special concern. http://floridaconservation.org/ Inc., San Juan, PR. 566 p.
pubs/endanger.html. 18 p.
Howard, R.A. 1989. Flora of the Lesser Antilles, Rico and adjacent islands. Vol. 5. Editorial de la
Leeward and Windward Islands. Universidad de Puerto Rico, Río Piedras, PR.
Dicotyledoneae. Part 3. Vol. 6. Arnold 436 p.
MA. 658 p. Nelson, G. 1996. The shrubs and woody vines of
Knapp, C. 2002. Exuma Island iguana, Cyclura 391 p.
cychlura figginsi. Iguana Specialist Group home
736
Styrax americanus Lam. American snowbell
STYRACACEAE
Synonyms: Styrax pulverulentus Michx.
Kristina Connor
Range.—American snowbell is found in the

Southeastern United States, from Virginia south to
Florida and west to Louisiana (Bailey and Bailey
1976) and eastern Texas. It is also recorded in
McCurtain County in Oklahoma (Johnson and
Hoagland 1999). It will grow as far north as
southern Missouri, southern Illinois, Indiana, and
southern Ohio but is rare, endangered or possibly
extirpated in Ohio and Illinois (Ohio Department
of Natural Resources 2002, Iverson and others
1999).
Ecology.—American snowbell grows primarily in

rich, poorly-drained acidic soils along swamp
margins, stream beds and lake edges, oxbow lakes,
and in floodplains. (Johnson and Hoagland 1999).
It grows well in the shade but will tolerate full sun.
It is associated with trees commonly found in wet
areas, such as baldcypress and overcup oak. Plants
are particularly tender when young (Krüssmann
1986).
Illustration source: USDA—Soil Conservation Service, Reproduction.—American snowbell is named for
Southern Technical Center
its showy, fragrant, white flowers. Flowers are
complete and their placement hypogynous
General Description.—American snowbell, also
(Iverson and others 1999). They occur in 2.5 to 12
known as mock orange or storax, is a deciduous
cm long racemes, one to four flowers per raceme.
shrub or small tree with a widely branched crown.
Raceme stalks are pubescent. The flowers have
It reaches 3 to 5 m in height, and the stems can
five petals, and the white style extends beyond the
reach 7.5 cm in diameter (Johnson and Hoagland
10 stamens. The calyx is shallowly five-lobed and
1999, Plant Identification Resource 2001). While
has tiny, triangular teeth. The 6 to 8 mm
the bark on the stems is smooth and dark grey to
subglobose fruits are one-seeded drupes that
brown, branches range in color from green to grey
mature from July to October (Krüssmann 1986,
to red-brown. Young stems are pubescent,
Johnson and Hoagland 1999).
becoming glabrous with age (Johnson and
Hoagland 1999). The alternate, simple leaves are
Growth and Management.—This species is often
2.6 to 9 cm long, ovate to elliptic, entire to finely
planted and grows well in cultivation. Since it
serrate, glabrous above, and pubescent beneath
requires moist soils and prefers shade, it can be
(Bailey and Bailey 1976, Krüssmann 1986). The
seriously threatened by drainage and removal of
leaf apex is acute, venation is pinnate. The showy,
the forest canopy. Its major pollinators are bees
fragrant white flowers for which the plant is noted
(Iverson and others 1999), and the species relies
form in racemes from April to June (Iverson and
on seed dispersal for regeneration. It is, however,
others 1999, Krüssmann 1986). American
easily propagated from softwood cuttings (Johnson
snowbell is distinguishable from bigleaf snowbell
and Hoagland 1999).
(S. grandifolius) by its smaller leaves and shorter
flowering racemes with fewer flowers per raceme
(Brown and Kirkman 1990).
737
Benefits.—American snowbell is an attractive Johnson, F.L. and B.W. Hoagland. 1999. Styrax
landscape plant. Its leaves are also a preferred food americanus. Catalog of the woody plants of
of the caterpillar of Callosamia promethea, the Oklahoma, Oklahoma Biological Survey Home
promethea moth (Oehlke [no date]). Page, http://www.biosurvey.ou.edu. 1 p.
References Krüssmann, G. 1986. Manual of cultivated broad-

leaved trees and shrubs. Volume III, Pro-Z.
Bailey, L.H. and E.Z. Bailey. 1976. Hortus Third: Timber Press, Beaverton, OR. 510 p.
A Concise Dictionary of Plants Cultivated in the
United States and Canada. McMillan Publishing Oehlke, B. [no date]. The Promethea Moth
Co., Inc., New York. 1,312 p. Callosamia promethes (Drury, 1773).
http://www3.
Brown, C.L. and L.K. Kirkman. 1990. Trees of islandtelecom.com/~oehlkew/scpromet.htm. 5 p.
Georgia and adjacent states. Timber Press,
Portland, OR. 292 p. Ohio Department of Natural Resources. 2002.
Styrax americanus. Ohio Division of Natural
Iverson, L.R., D. Ketzner and J. Karnes. 1999. Areas and Preserves. Rare Native Ohio Plant
Styrax americanus. Illinois Plant Information 2000-2001 Status List. 2 p.
Network. Database at
http://www.fs.fed.us/ne/delaware/lpin. html. Plant Identification Resource. 2001. Styrax
Illinois Natural History Survey and USDA Forest americanus. Auburn University College of
Service. 3 p. Agriculture, Landscape/Horticulture webpage,
http://www.ag.auburn.edu/landscape/386.html.
3 p.
738
Suaeda suffrutescens S. Wats. desert seepweed
CHENOPODIACEAE
Synonyms: Dondia suffrutescens (Wats.) Heller

Suaeda nigrescens I.M. Johnst.
James E. Nellessen
(2) S. nigrescens I.M. Johnst., and (3) S.

nigrescens var. glabra I.M. Johnst. (Allred 2002,
Great Plains Flora Association 1986, Kartesz
1994). The latter variety, S. suffrutescens var.
detonsa (S. nigrescens var. glabra) is distinguished
by having smooth, or relatively smooth (hairless)
stems and leaves. This variety is more common in
Mexico, but extends into southern Texas.
Range.—Desert seepweed occurs from western

Oklahoma and Texas across central and southern
New Mexico into Arizona and California, and
south into Mexico. It occurs at elevations from
about 750 m to 1,830 m.
Ecology.—Desert seepweed is a perennial shrub

of saline or alkaline soils in plains and valleys.
These habitats include the margins of ephemeral
lake beds, salt flats, and flood plains of rivers and
streams. Some moisture needs to be relatively near
the surface for part of the growing season. This
species is classified as a halophyte, or salt tolerant
plant. Desert seepweed may dominate alkali sink
habitats in association with other salt tolerant
species, including other species of Suaeda
(seepweeds), Salicornia spp. (glasswort or
General Description.—Desert seepweed is also saltwort), Distichlis spicata (L.) Green var. stricta
known as inkweed, iodinebush, and quelite salado. (Torr.) Beetle (inland saltgrass), Allenrolfea
Desert seepweed has stout stems and branches, occidentalis (Wats.) Kuntze (burro weed or
woodiness greatest toward the base, side branches quinine bush), and Heliotropium curassavicum L.
generally short, although the plant is highly (salt heliotrope), to name a few (Hendrickson
branched. It could be classified as a subshrub. The 1974, Dick-Peddie 1993).
plant is erect and ascending, or spreading. Desert
seepweed ranges in height from about 50 to 120 Halophyte Ecology and Physiology.—Although
cm, often forming mounds as broad as 150 cm. little if anything is reported about desert seepweed,
Young stems and leaves are soft and felty with much is known about the common salt tolerant
numerous hairs. Leaves are simple, alternate, characteristics of the genus Suaeda. Chapman
numerous, fleshy, linear to linear-lanceolate, 5 to (1942) used the term halophyte in describing
20 mm long, narrower at the base, semirounded to species that could grow in an environment with
somewhat flat (Correll and Johnston 1970, Great greater than 0.5 percent sodium chloride (NaCl).
Plains Flora Association 1986, Ivey 1995, Kearney Halophytes can be divided into two groups: (1)
and others 1951, Martin and Hutchins 1980). Two those requiring salt for survival and best growth,
varieties have been recognized: S. suffrutescens and (2) those tolerating or resisting salty
var. suffrutescens, and S. suffrutescens var. environments (Waisel 1972). The second group,
detonsa I.M. Johnston. The latter variety (detonsa) the salt tolerators, have adapted to saline habitats
has three synonyms: (1) S. duripes I.M. Johnst., and survive there because salt-sensitive plants
739
(glycophytes) cannot survive there. Halophytes members of this genus required prior cold
accumulate salts obtained from the soil solution stratification of the seeds with optimal daytime
within their tissues during the transpiration germination temperatures of 25 to 27 °C
process. Many members of the genus Suaeda (summarized in Baskin and Baskin 2001). NaCl
readily translocate salts to upper portions of the concentrations ranging from 0.17 to 0.90 molar
plant. This second group of halophytes can be have depressed maximum germination of 75 to
subdivided into three more categories of tolerance. 100 percent down to 10 percent in four different
Salt enduing species simply tolerate the presence species of Suaeda (summarized in Baskin and
of salt in their cells. Salt excluding species may Baskin 2001).
excrete salt from shoots, including those with
specialized hairs on the plant, or transport them Growth and Management.—Desert seepweed
back to the roots. Salt evading species have can grow relatively quickly. It has been observed
mechanisms to prevent transport to the leaves or to resprout readily in managed and mowed areas
prevent absorption from the soil solution (Waisel adjacent to levees along the lower Rio Grande
1972). Some members of the genus Suaeda fall floodplain in New Mexico. This species is
into the first group requiring salt for survival, restricted to saline and riparian habitats and is not
while others are salt enduring halophytes. Salt highly browsed. Consequently, it does not pose as
(NaCl) and other ions (including sulfate and a major management concern on most rangelands.
potassium) may be sequestered into cell vacuoles It does not typically form dense stands.
where they do not interfere with normal cell
biochemistry. These plants will generally have Benefits.—Desert seepweed leaves and young
higher water potential gradients between the soil shoots have been added to vegetable greens or
and the leaves, which may add to succulence. cacti fruits by Native Americans during cooking to
Waisel (1972) proposed a new order to the add flavor. It has also been added between layers
classification of halophytic plant communities: of cacti fruit and sotol butts (stems of Dasylirion
Suaedetalia--persistent communities in dry desert sp.) during the 24 to 48 hour cooking process in
saline habitats, dominated by annual and perennial mescal pits (Warnock 1974). Pima Indians would
members of the family Chenopodiaceae (as well as eat the plant for greens and the roasted seeds,
other families). Zahran (1982) likewise includes a called pinole. Coahuila Indians extracted a black
Suaeda community classification. Some shrubby dye from the plant used for artwork (Dodge 1985).
members of the genus (S. fruticosa Forsk. and S. This species is not browsed much by livestock but
monoica Forsk.) are reported to accumulate large may be utilized when other vegetation is in short
amounts of salt annually when in dense stands, as supply (Warnock 1974, Dodge 1985).
much as 2 to 4 tons/ha (Waisel 1972). In this
function they may help to desalinate some saline References
habitats.
Allred, K.W. 2002. A Working Index of New
Reproduction.—Flowers are small and greenish, Mexico Vascular Plant Names. http://web.
and occur in spiked clusters of three to nine nmsu.edu/~kallred/herbweb. [not paged].
flowers in the leaf axils. The corolla is absent. The
five-lobed calyx has all parts equal, may be hood Baskin, C.C. and J.M. Baskin. 2001. Seeds:
shaped, and lacks appendages, that is, it is not Ecology, Biogeography, and Evolution of
winged. Seeds are small, black, and 0.7 to 1 mm Dormancy, and Germination. Academic Press,
wide. Desert seepweed blooms from March to Sand Diego, CA. 666 p.
September, depending on local environmental
conditions (Correll and Johnston 1970, Great Chapman, V.J. 1942. The new perspective in the
Plains Flora Association 1986, Kearney and others halophytes. Quarterly Review of Biology 17:
1951, Martin and Hutchins 1980). Many of the salt 291-311.
tolerant members of the Chenopodiaceae have
physiological seed dormancy (Baskin and Baskin Correll, D.S. and M.C. Johnston. 1970. Manual of
2001). Although no seed germination reports the Vascular Plants of Texas. Texas Research
regarding this particular species were located, Foundation, Renner, TX. 1,881 p.
numerous studies have been performed on other
members of the Suaeda genus. Three of four
740
Dick-Peddie, W.A. 1993. New Mexico and Greenland, Vol. 1, 2nd Ed. Biota of North
Vegetation: Past, Present and Future. America Program of the North Carolina
University of New Mexico Press, Albuquerque, Botanical Garden. Timber Press, Portland, OR.
NM. 244 p. 622 p.
Dodge, N.N. 1985. Flowers of the Southwest Kearney, T.H., R. Peebles, and Collaborators.
Deserts. Southwest Parks and Monuments 1951 with 1960 supplement. Arizona Flora.
Association, Tucson, AZ. 136 p. University of California Press, Berkeley, CA.
1,085 p.
Great Plains Flora Association. 1986. Flora of the
Great Plains. University Press of Kansas, KS. Martin, W.C. and C.E. Hutchins. 1980 (reprinted
1,392 p. 2001). A Flora of New Mexico, Vol. 1. Bishen
Singh Mahendra Pal Singh (India) and Koeltz
Hendrickson, J. 1974. Saline habitats and Scientific Books (Germany). 1,276 p.
halophytic vegetation in the Chihuahuan Desert
region. In: R.H. Wauer and D.H. Riskind, eds. Waisel, Y. 1972. Biology of Halophytes.
Transactions of the Symposium of the Academic Press, NY. 395 p.
Biological Resources of the Chihuahuan Desert
Region, United States and Mexico. U.S. Warnock, B.H. 1974. Wildflowers of the
Department of Interior, National Park Service Guadalupe Mountains and the Sand Dune
Transactions and Proceedings Series, No. 3, Country, Texas. Sul Ross State University,
U.S. Goverment Printing Office, Washington Alpine, TX. 176 p.
DC. p. 289-314.
Zahran, M.A. 1982. Ecology of halophytic
Ivey, R.D. 1995. Flowering Plants of New vegetation of Egypt. In: D.N. Sen and K.S.
Mexico, 3rd Edition. Published by the author, Rajpurohit, eds. Contributions to the Ecology of
NM. 504 p. Halophytes.. Dr. W. Junk Publishers, The
Hague, Netherlands. p. 3-20.
Kartesz, J.T. 1994. A Synonymized Checklist of
the Vascular Flora of the United States, Canada,
741
Suriana maritima L. bay-cedar
SURIANACEAE (formerly in SIMAROUBACEAE)
Synonyms: none
John K. Francis
Indies, the eastern side of Mexico through Brazil,

East Africa, tropical Asia, Australia, and many of
the islands of the Indian and Pacific Oceans (not
Hawaii) (Howard 1988, Stevens and others 2001).
The species has been planted widely as an
ornamental. It is listed as endangered in the wild in
Florida (Smithsonian Institution 2002).
Ecology.—Bay-cedar is strictly costal. It grows on

beaches, dunes, sandy thickets (Nelson 1996), and
rocky headlands. It tolerates moderately salty soils,
storm-surge overwash, heavy salt spray, blowing
sand, high surface heat, drought, and strong winds
(Wildflower Nervana 2002). If planted and given
protection from competition, it will grow well on
inland sites (Nelson 1996) on a wide range of
well-drained soils. The species requires full sun to
grow and flower. Bay-cedar is rare to uncommon
in much of its habitat, but occasionally forms
thickets.
Reproduction.—Bay-cedar flowers and fruits

intermittently throughout the year (Little and
General Description.—Bay-cedar, is also known others 1974). The fruits are buoyant and remain
as tassel-plant, guitarán, temporana, jovero, viable for long periods in sea water enabling
cuabilla, pantsil, cucharo, palo corra, oseille bord colonization of tropical beaches throughout the
de mer, romarin noir, and crises marine. It is an world (Nelson 1996).
evergreen shrub usually 1 to 2 m in height but
sometimes reaching 6 m. Multistemmed and Growth and Management.—Growth rate of bay-
much-branched, it has a prostrate to ovoid crown. cedar is described as moderate. The species is
Stems have a dark brown, rough and flakey bark. currently being propagated by seeds to produce
Heartwood is dark red or reddish brown; the nursery plants. The recommended spacing for
sapwood being somewhat lighter. The wood is ornamentals is 90 to 150 cm. Irrigation is
hard, heavy, fine-textured, strong, and durable. recommended during establishment but afterwards
The gray-green, succulent, downy foliage has a is not needed (Gilman 1999).
salty taste and a cedar-like fragrance when
crushed. Leaves are crowded at the ends of the Benefits.—Bay-cedar helps stabilize beaches and
branches, sessile, narrowly obovate, and 1 to 4 cm costal dunes, and furnishes food and cover for
long. Inflorescences, solitary or few-flowered wildlife. It is widely planted as an ornamental for
cymes, are almost hidden among the leaves. The hedges, screens, borders, accents, seminatural
small, five-merous, bisexual flowers are yellow covers, and potted plants. The fruits attract birds
and develop into clusters of five hard, 3- to 4-mm, that feed on them (Gilman 1999). The foliage
dry drupes surrounded by five gray sepals. furnishes food for blue and hairstreak butterfly
(Gilman 1999, Howard 1988, Liogier 1988, Little larva (Schaefer and others 2002). Key deer
and others 1974). (Odocolleus virginanus clavium) and probably
other ruminants will not eat the foliage of bay-
Range.—Bay-cedar is native to the shores of cedar (Schaus and others 2002). Herbalists employ
southern Florida, Bermuda, the Bahamas, the West
742
extracts of the leaves and bark to treat rheumatism Nelson, G. 1996. The shrubs and woody vines of
and skin ulcers and to stop bleeding (Liogier Florida. Pineapple Press, Inc., Sarasota, FL.
1990). 391 p.
References Schaefer, J., C.N. Huegel, and F.J. Mazzotti. 2002.

Butterfly gardening in Florida. Cooperative
Gilman, E.F. 1999. Suriana maritima. Cooperative Extension Service, University of Florida,
Extension Service, University of Florida, Gainesville, FL. http://edis.ifas.ufl.edu/
Gainesville, FL. 3 p. BODY_UW057. 24 p.
Howard R.A. 1988. Flora of the Lesser Antilles, Schaus, C, S. Wade, and J. Dunan. 2002. Key deer
Leeward and Windward Islands. and plants they won’t eat. Monroe County
Dicotyledoneae, Part 1. Vol. 4. Arnold Extension Service, Key West, FL.
Arboretum, Harvard University, Jamaica Plain, http://monroe.ifas.ufl.edu/key_deer_plants.htm.
MA. 673 p. 4 p.
Liogier H.A. 1988. Descriptive flora of Puerto Smithsonian Institution. 2002. Special status
Rico and adjacent islands, Spermatophyta. Vol. species in the Indian River Lagoon. Smithsonian
2. Editorial de la Universidad de Puerto Rico, Marine Station at Fort Pierce, Smithsonian
Río Piedras, PR. 481 p. Institution, Washington, DC. http://www.
sms.si.edu/IRLSpec/ListedSpec.htm. 3 p.
Liogier, H.A. 1990. Plantas mediciales de Puerto
Rico y del Caribe. Iberoamericana de Ediciones, Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
Inc., San Juan, PR. 566 p. Montiel, eds. 2001. Flora of Nicaragua.
Little, E.L., Jr., R.O. Woodbury, and F.H. 3. Missouri Botanical Garden Press, St. Louis,
Wadsworth. 1974. Trees of Puerto Rico and the MO. p. 1,911-2,666.
449. U.S. Department of Agriculture, Wildflower Nervana. 2002. Suriana maritima—
Washingtion, DC. 1,024 p. guitarán or temporana (bay-cedar). http://
www.wfnirvana.com/pr/surimari/html. 3 p.
743
Symphoricarpos albus (L.) Blake common snowberry
CAPRIFOLIACEAE
Synonyms: Symphoricarpos racemosus Michx.

Symphoricarpos rivularis Suksdorf
Viburnum album L.
John K. Francis
Range.—Common snowberry is native from Nova

Scotia to southern Alaska, throughout the northern
half of the United States, and in California, Utah,
Colorado, New Mexico, Tennessee, and North
Carolina (McWilliams 2000, Natural Resources
Conservation Service 2003). It does not occur in
Nevada and Kansas, and is reported to be only
naturalized in Utah (Welsh and others 1987). The
species was introduced to Great Britain in 1817
and has naturalized widely in the United Kingdom,
Ireland, and northern Europe (Binggeli 1998,
Gilbert 1995). There are two varieties, albus and
laevigatus (Fern.) Blake. Variety laevigatus, a
coarser and taller (up to 2 m) plant, and has a
Western distribution. Variety albus occurs
intermittently across the Continent (Natural
Resources Conservation Service 2003). Both are
cultivated and naturalized in many places, making
determination of distribution somewhat
General Description.—Common snowberry, also complicated.
known as snowberry, waxberry, and white
coralberry, is a deciduous shrub. The slender, Ecology.—Common snowberry grows best in full
branchy, multiple stems are usually low (0.5 to 1 sunlight or light shade on well-drained, moist,
m) and arching or straight, but sometimes reaching fertile soils. It does survive and grow slowly in
as much as 2 m in height. The stems rarely exceed medium shade. The species grows in many types
2.5 cm in thickness. The bark is yellow-gray in of open forests, forest edges, shrublands, riparian
young stems, tan to grayish-brown in older ones, vegetation, and fencerows. It does not grow in
becoming scaly or splitting lengthwise in larger swampy sites, deserts, and mountain tops above
stems. Stem wood is soft and greenish-white. The the tree line. Common snowberry grows in soils
opposite-branching twigs are slender and flexible. with mildly acidic pH (6.0) and mildly alkaline
Thin, papery leaves are opposite, simple, short- conditions (pH’s of 7.8) and does very well on
stalked, and round to oblong-elliptic with margins soils derived from limestone; less aggressively on
entire, lobbed, or wavy-toothed. Petioles are 3 to 4 soils derived from granite. It tolerates poor fertility
mm long. The perfect flowers are solitary to and has been used to revegetate disturbed sites.
several in terminal or lateral spikes. The five- Common snowberry in interior Western United
lobbed corolla is pink and white, tubular or bell- States grows at elevations from 800 to 2,800 m. It
shaped, and about 6 mm long. Fruits are white, grows at much lower elevations in other areas of
spongy, berry-like drupes 6 to 12 mm in diameter. North America. Because it sprouts readily from
Each contains two white, flattened, elliptical rhizomes, common snowberry is resistant to fire
nutlets with one seed per nutlet. Chromosome and browsing. It may increase dramatically in
number is 2n = 54 or 72 (Gilbert 1995, Grimm density after logging opens the forest canopy
1966, McWilliams 2000, Soper and Heimburger (Gilbert 1995, McWilliams 2000, Natural
1982, Stephens 1973, Welsh and others 1987). Resources Conservation Service 2003).
744
Reproduction.—Common snowberry blooms are on the shrubs. Containing 4 to 13 percent
between early May and late July depending on protein, depending on tissue and season, it
location and matures fruits in August to October provides important early-season browse for cattle
(Grimm 1966). The flowers are pollinated by a and domestic sheep. It is also browsed by most
range of bees, wasps, and syrphids (Gilbert 1995). wild ungulates. It is generally unpalatable to
Fruit and seed production can be abundant, horses and moose. The fruits are consumed by
especially when plants grow in full sun on fertile both black and grizzly bears and are eaten by a
sites. Normally a good seed crop is produced every number of birds and small mammals (McWilliams
year. Fruits remain on stems until midwinter or 2000). Some sources refer to the fruits as being
early spring, if not eaten. Numbers of seeds per toxic to humans (Gilbert 1995, Moerman 1986).
gram vary from 119 to 250 for var. albus and from The toxic agent appears to be the isoquinoline
86 to 144 for var. laevigatus. Germination may be alkaloid chelidonine and causes mild symptoms of
as high as 87 percent if exacting conditions of vomiting, dizziness, and sedation in children
pretreatment are met. Germination is epigeal (Canadian Biodiversity Information Facility 2003).
(Walker 2003). Seeds are dispersed by birds and The fruits were reported to have been eaten but not
mammals. Seedlings are relatively uncommon. favored by Native Americans (McWilliams 2000).
However, vegetative reproduction is common by Various tissues were used by Native Americans as
sprouting of underground stems that run a diuretic, to treat gonorrhea, and for sore eyes
horizontally at a depth of 2 to 5 cm for up to 60 cm (Moerman 1986). Common snowberry is planted
before turning up to form new tops. Stems layer as an ornamental and in conservation plantings for
when they come in contact with the ground, and wildlife amenity, soil stabilization, and strip mine
severed stems lying on the ground will also take reclamation in temperate areas around the world
root (Gilbert 1995). (Gilbert 1995).
Growth and Management.—Planted common References

snowberry reached 30 cm in height during the first
year (Center for Urban Horticulture 2002). Shoot Binggeli, P. 1998. Plants of the Pacific Northwest
extension of plants in England is reported at 0.6 m in Western Europe. Botanical Electronic News
per year. Individual stems have been aged up to 34 ISSN 1188-603X. http://www.ou.edu/cas/
years old in England (Gilbert 1995). By suckering, botany-micro/ben/ben195.html. p. 1-4
plants may survive indefinitely. Fruits may be
collected by hand picking or flailing the plants Canadian Biodiversity Information Facility. 2003.
with a tarp spread underneath. The fruit is then Notes on poisoning: Symphoricarpos albus.
macerated and water is used to float off the pulp http://sas.agr.gc.ca/pls/pp/ppack.info?p_psn=122
and empty seeds. The residue is then dried and &p_type=all&p_sci=sci&p_x=px. 5 p.
cleaned. The air-dried seeds can be stored at 5 °C
for up to 5 years. Pretreatments are necessary for Center for Urban Horticulture. 2002. Does mulch
acceptable germination. Warm stratification for 3 improve plant survival and growth in restoration
to 4 months at 22 to 30 °C followed by cold sites? Fact sheet 38. University of Washington,
stratification at 5 °C for 4 to 6 months is College of Forest Resources. http://www.cfr.
recommended. Sowing in the late fall can be used washington.edu/Research/fact_sheets/38-mulch.
instead of the cold treatment. Sown seeds should pdf. 2 p.
be covered with about 6 mm of soil and 2 cm of
mulch. Seedlings for planting can be grown Gilbert, O.L. 1995. Symphoricarpos albus (L.)
successfully from cuttings, wildlings, and pieces of S.F. Blake (S. rivularis Suksd., S. racemosus
stem with roots collected in the spring. Transplant Michaux). Journal of Ecology 83(1): 159-166.
establishment success of bareroot and
containerized nursery stock is as high as 90 Grimm, W.C. 1966. Recognizing native shrubs.
percent when done properly (Walker 2003). The Starkpole Company, Harrisburg, PA. 319 p.
Benefits.—Common snowberry helps protect the McWilliams, J. 2000. Symphoricarpos albus. In:
soil, provides food and cover for wildlife, browse U.S. Department of Agriculture, Forest Service,
for livestock, and adds beauty to the landscape, Rocky Mountain Research Station, Fire Sciences
especially in fall and winter when the white fruits Laboratory. Fire Effects Information System.
745
http://www.fs.fed.us/database/feis/plants/shrub/ Stephens, H.A. 1973. Woody plants of the North
symalb/all.html. 32 p. Central Plains. The University Press of Kansas,
Lawrence, KS. 530 p.
America. Technical Reports 19. University of Walker, S.C. 2003. Symphoricarpos Duham.,
Michigan Museum of Anthropology, Ann snowberry. U.S. Department of Agriculture,
Arbor, MI. 534 p. Forest Service, Washington, DC. http://wpsm.
net/Symphoricarpos.pdf. 10 p.
Plants profile: Symphoricarpos albus (L.) Blake Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
http://plants.usda.gov/cgi_bin/plant_profile. Higgins, eds. 1987. A Utah flora. Great Basin
cgi?symbol =SYAL. [not paged]. Naturalist Memoirs 9. Brigham Young
Soper, J.H. and M.L. Heimburger. 1982. Shrubs of
Ontario. Royal Ontario Museum, Toronto,
Ontario, Canada. 495 p.
746
Tabebuia haemantha (Bertol. ex Spreng.) DC. roble cimarrón
BIGNONIACEAE
Synonyms: Bignonia haemantha Bert. ex Spreng.

Tecoma haemantha (Bert.) Griseb.
Spathodea portoricensis Bello
John K. Francis
to 11 cm long and contain many membranous two-

winged seeds 1.9 cm long (author’s observations,
Range.—Roble cimarrón is endemic to the island

of Puerto Rico. Specifically, the species is found
in the dry forests in the southern and eastern parts
of the island and the moist slopes above them,
including serpentine areas in western Puerto Rico,
and on the limestone hills in the northern coast
and inland (Little and Wadsworth 1964, Vélez and
van Overbeek 1950).
Ecology.—Roble cimarrón grows on loamy and

clayey soils with pH’s from about 5.5 to 7.5. It
often occurs in sites that have been eroded and
partially compacted by grazing animals. The soils
where it grows today are usually steep and rocky.
The species formerly inhabited all types of
topography from near sea level to about 400 m in
elevation. These sites receive from about 700 to
General Description.—Roble cimarrón, also 2100 mm of precipitation. In areas that receive the
known as roble colorado and roble bobo, is a higher rainfall of the range, roble cimarrón tends
shrub or a small tree up to 8 m in height and 15 to grow on excessively drained sites. The species
cm in basal diameter. Commonly, plants are 3 to 5 is frequently common in overgrazed pastures
m in height and 4 to 8 cm in diameter. Young indicating that it is unpalatable to cattle. Roble
roble cimarrón usually have a single stem, until cimarrón is moderately intolerant of shade. It
they have been damaged mechanically or by fire. grows well in the open and in low basal area forest
Older plants often develop multiple stems by stands. Adult plants are usually found in
spontaneously sprouting just above the ground codominant and intermediate crown positions in
level. The bark is gray, smooth except for a slight dry forests. It is top-killed by fires but sprouts
fissuring. The inner bark is nearly white and vigorously afterwards.
slightly bitter. There are relatively few branches.
The twigs are light gray and slightly flattened Reproduction.—Flowering occurs almost
below the nodes. The species develops a tap and throughout the year according to Little and
lateral root system. The roots are brown and Wadsworth (1964). However, most shrubs in
flexible. Roble cimarrón is evergreen with particular areas appear to flower synchronously
palmately compound leaves. There are three or (author’s observation). Hummingbirds, other bird
five stiff, leathery leaflets on a stout 2.5- to 5.0-cm species, and insects pollinate the flowers. Capsules
petiole. The 3- to 15-cm long leaf blades are form and mature about 6 weeks after flowering.
elliptic or ovate with entire edges and rounded to They are green, turning tan at maturity, after
pointed at the tip. The inflorescences are panicles which they quickly split open. The seeds adhere to
of several to many flowers on short branches. The a flat central placenta but detach and become
flowers have a crimson, tubular corolla 3 to 5 cm airborne in the slightest breeze (Vélez and van
long with five irregular lobes. The capsules are 6 Overbeek 1950). A sample of 17 capsules
747
collected in Salinas, Puerto Rico contained an environmental restoration plantings within its
average of 59.9 ± 2.01 seeds/capsule. The air-dry native range. The light brown, hard wood is
seeds averaged 0.0148 + 0.0003 g/seed or 68,000 suitable for firewood and charcoal but is little used
seeds/kg. Sown on peat moss, these seeds because of the plant’s small size and relative
germinated at 90 percent between 7 and 31 days scarcity.
after sowing. The species grows and survives well
in the nursery, similar to other Tabebuia seedlings. References
In the wild, seedlings are well scattered and
uncommon. Liogier, H.A. 1995. Descriptive flora of Puerto
Growth and Management.—Roble cimarrón Universidad de Puerto Rico, Río Piedras, PR.
grows about 0.5 m/yr in early years after it is well 617 p.
established. It does not appear to be a long-lived
species. Unfortunately, no plantations are known Little, E.L., Jr. and F.H. Wadsworth. 1964.
and no management experience has been reported. Common trees of Puerto Rico and the Virgin
Islands. Agriculture Handbook 249. U.S.
Benefits.—Roble cimarrón would probably be Department of Agriculture, Forest Service.
suitable as an ornamental for background Washington, DC. 548 p.
plantings in moist and dry frost-free areas. It
would have the advantage of being drought-hardy Vélez, I. and J. van Overbeek. 1950. Plantas
and tolerant of infertile and partially compacted indeseables en los cultivos tropicales. Editorial
soils and should be a good species to plant in Universitaria, Río Piedras, PR. 497 p.
748
Tamarix chinensis Lour. tamarisk
TAMARICACEAE
Synonyms: Tamarix pendandra Pallas

Tamarix ramosissima Ledebour
Tamarix gallica L. var. micrantha Ledebour
Tamarix pentandra Pallas ssp. tigrensis (Bge.) Hand.-Mazz.
more obscure characters.” For most species, the

tiny floral organs and the use of a microscope, or
at least hand lens, are essential for identification
(Allred 2002). Tamarix pentandra var, pentandra
Pallus has been mistakenly applied to this species
within North America. Actually, T. pentandra
Pallus is synonymous with T. hispida Willd. that is
distributed throughout central Asia (Baum 1978).
Other confusion has resulted from this species
being referred to as a sub-species of T. gallica L.,
which is a unique taxon originally from Europe,
and now also naturalized within the United States
(Allred 2002, Baum 1978). Although Baum’s
treatment (1978) described T. chinensis Loureiro
and T. ramosissima Ledebour as unique taxa
placing them in two different sections, recent
chloroplast and nuclear DNA studies suggest the
two entities should be merged (Allred 2002).
These two names are synonymized in the Utah
Flora (Welsh and others 1993). Tamarix chinensis
has a chromosome number of 2n = 24 (Welsh and
others 1993).
General Description.—Tamarix chinensis
Loureiro is commonly known as tamarisk or Range.—Tamarisk is a common, wide-ranging
saltcedar. Occasionally it grows with a defined species that has been collected across Asia from
trunk as a tree but generally it grows as a multiple- western Turkey through Mongolia, Afghanistan,
stemmed shrub up to 6 m tall. The bark of larger and China to Korea and Japan. It was introduced
trunks, or stems, is reddish-brown to dark brown into the Western United States and has become
or blackish. The leaves, that are only 1.5 to 3.5 naturalized.
mm long and appear scale-like, are deciduous. The
tiny flowers are five-merous, having five sepals, Ecology.—Tamarisk grows in generally moist
five petals, five stamens, and a five carpellate sites at seeps, streams, and along river banks and
pistil, and are arranged on a flowering stalk that is roadsides (Dick-Peddie 1993). It tolerates saline
2 to 7 cm long. The petals are 1 to 2.3 mm long, soils and also contributes to salinization of the
and persistent in fruit. They are frequently a shade soils in which it grows. It forms equally dense
of pink but may be white or red. The sepals are 0.5 stands in both the Western United States and in
mm to 1.3 mm long. The scale-like leaves and central China where it is an apparent native
shape of the shrub are reminiscent of a juniper or (author’s personal observation). It has an extensive
cedar and this accounts for the common name root system and uses such large volumes of water
“saltcedar.” This shrub has been the subject of that streams and ponds become dry. Studies in the
considerable taxonomic confusion. McClintock Southwestern United States show that tamarisk
(1951) stated “there is probably not another genus stands have far less biodiversity than stands of
of plants as well known as the tamarisks in which native riparian vegetation. Miles of riverbank can
the species is so poorly understood or separated on be almost monotypic with dense tamarisk stands
749
that have 50 percent fewer small mammal species insects have been investigated--from grasshoppers
as well as fewer reptile and amphibian species than to leaf hoppers. Two species specific to tamarisk, a
native riparian stands (Olson 1999). Although bees mealybug (Trabutina mannipara) from Israel and
visit the flowers (Epple 1995) the diversity of a leafbeetle (Diorhaba elongata) from China, have
insects is also low, and nestling birds rely on received advance testing (DeLoach and others
insects brought to the nests by their parents. 1996, DeLoach, 1994). Physical cutting and
Although some birds can nest in the shrubs the removal of tamarisk will only be successful if the
trunks are not large enough to provide habitat for roots are removed or killed. By itself, cutting
animals such as wood-peckers, owls, chickadees, tamarisk by hand (that is by chain saw, weed eater,
and squirrels. The tiny, hairy seeds provide poor or axe) is not often an effective treatment. Cutting
nutrition and are largely indigestible. tamarisk tends to just cause more shoots to grow.
However, there was a report of successful control
Reproduction.—Tamarisk generally flowers in where large tree stumps (20 to 25 cm diameter)
April through August. The pink, fragrant flowers had been cut back to the soil surface and then
are insect pollinated. The seeds, which are small, covered by a black, liner material (Cinnamon
mature in summer and fall and are dispersed by 1990). Purely mechanical removal is also
birds and by water. However, the seed may have sometimes successful if there is enough standing
limited longevity under humid conditions. Seeds vegetation to substantially shade the resprouting
retain high viability if stored in sealed containers tamarisk. For example, in Nevada (Busch and
with desiccant at temperatures between 3 to 27 oC Smith, 1995) when tamarisk had been completely
but lose viability if stored at “moderate humidity” cleared above ground in a stand of willows, there
(Wilgus and Hamilton 1962, Baskin and Baskin was essentially no grow back during the 4 years
2001). Germination temperature appears to be follow up. Tamarisk is sensitive to numerous
flexible. Freshly matured seeds germinated from herbicides (Kunzman and Bennett 1990, Sisneros
87 to 98 percent when kept at constant 1991). At the present time, the most successful
temperatures between 19 to 43 oC. Percent approach to control tamarisk is by cut-stump
germination did not appear related to temperature; herbicide or cut-stump/frill herbicide treatments
96 percent germinated at 19 oC and 91 percent at (Neill 1990, Hughes 1996). This approach is quite
43 oC (Wilgus and Hamilton 1962, Baskin and costly but it is very controlled, does not affect non-
Baskin 2001). Vegetative reproduction by suckers target organisms, and causes the least
and rooting branches is copious. environmental impact. The tamarisk shrubs should
be cut to within 2.5 cm of the ground surface
Growth and Management.—Tamarisk is an (Cinnamon 1990) and a systemic herbicide applied
aggressive, invasive species. Its invasive behavior to the stump within a few minutes after cutting.
is illustrated by its colonization of the Pecos River The time between cutting and herbicide treatment
Valley in New Mexico (Allred 2002). In 1912, a should be as short as possible. Leaving the cut
few seedlings were observed at Lake McMillan. stump for 2 to 5 days before herbicide treatment
By 1915 the shrub covered 600 acres and extended reduces success rate from greater than
up and down the riverbanks. By 1926 it covered approximately 88 percent tamarisk death to less
12,300 acres and extended to 57,000 acres by 1960 than 10 percent (Hays and Mitchell 1990). Fire is
(Allred 2002). It out-competes many native often ineffective. The high water and salt content
species and is difficult to eradicate once of tamarisk makes it difficult to burn. Too light
established (USDA Forest Service 1988). The (low temperature) a fire will permit, or even
extremely deleterious impact of this species on the encourage, tamarisk to re-sprout and become even
riparian and wetland areas of the Southwest has denser. If the fire burn is so hot it kills the
led to intense study on how to control and reverse tamarisk, it is likely that the fire will leave the soil
its spread. There are four general methods-- bare and in such a poor condition that it cannot
biological control, physical removal by hand or by support grasses and desirable vegetation. In the
machine, herbicides, and fire--that can be used to horticultural trade, tamarisk is propagated from
control tamarisk, and each has shortcomings. The hardwood cuttings in the spring or softwood
most appropriate method, or combination of cuttings in the summer (Tykač 1990). Plants
methods, depends upon the particular site. grown for a long time in containers often do not
Biological controls for tamarisk are not yet establish after transplanting (Tykač 1990).
commercially available, although a variety of
750
Benefits.—Tamarisk were once planted for Group for the Introduction of Biological
erosion control and as wind breaks (Baum 1978, Control Agents of Weeds, USDA-APHIS. 21
Allred 2002). It is not significantly grazed or March.
browsed. The tannin substances in the vegetation
are likely to make it unpalatable. Tamarisk has DeLoach, C.J., D. Gerling, L. Fornasari, R.
commercial value in the landscape and Sobhain, S. Myartseva, I.D. Mityaev, Q.G. Lu,
horticultural trade. It is a popular ornamental shrub J.L. Tracy, R. Wang, J.F. Wang, A. Kirk, J.W.
although its use in the United States is now Pemberton, V. Chikatunov, R.V. Jashenko, J.E.
generally discouraged. Many cultivars exist that Johnson, H. Zheng, S. L. Jiang, M.T. Liu, P.A.
have been chosen for flower color and growth Liu, and J. Cisneros. 1996. Biological control
form (Bailey and others 1976, Tykač 1990). program against salt cedar (Tamarix spp.) in the
Tamarix species, although not specifically T. U.S.A. In: V. C. Moran and J.H. Hoffman, eds.
chinensis, have been used for fuel and building Proc. of the IX International symposium on the
materials by Native American tribes in the control of weeds, Cape Town, South Africa. p.
Western United States (Moerman 1998). 253-260.
References. Dick-Peddie, W.A. 1993. New Mexico vegetation

– past, present, and future. University of New
Allred, K. 2002. Identification and taxonomy of Mexico Press, Albuquerque, NM. 244 p.
Tamarix (Tamaricaceae) in New Mexico.
Desert plants 18(2): 26-31 Epple, A.O. 1995. A field guide to the plants of
Bailey, L.H., E.Z. Bailey, and the staff of the MT. 347 p.
Liberty Hyde Bailey Hortorium. 1976. Hortus
Third. Macmillian Publishing Company, New Hays, F. and J. Mitchell. 1990. Tamarisk
York. 1,290 p. eradication in Zion national Park. In: Tamarisk
control in Southwestern United States. Special
Baskin, C.C. and J.M. Baskin. 2001. Seeds, Report 9. Cooperative National Park Resources
ecology, biogeography, and evolution of Studies Unit. p. 36-38.
dormancy and germination. Academic Press,
New York. 666 p. Hughes, D. 1996. Restoring native riparian
vegetation. In: Desired future conditions for
Baum, B.R. 1978. The genus Tamarix. The Israel South western riparian ecosystems: Bringing
Academy of sciences and humanities, interests and concerns together. General
Jerusalem, Israel. 209 p. Technical Report RM-GTR-272. U.S.
Department of Agriculture, Forest Service, Fort
Busch, D.E. and S.D. Smith. 1995. Mechanisms Collins, CO.
associated with decline of woody species in
riparian ecosystems of the Southwestern U.S.A. Kunzman, M.R., and P.S. Bennett. 1990. Arsenal
Ecological Monographs. 65(3) 347-370. as a control agent for Saltcedar (Tamarix). In:
Tamarisk control in Southwestern United
Cinnamon, S.K. 1990. Wupatki National States. Special Report 9. Cooperative National
Monument tamarisk and camelthorn eradication Park Resources Studies Unit. p. 82-90.
program 1983-1988. In: Tamarisk control in
Southwestern United States. Special Report 9. McClintock, E. 1951. Studies in California
Cooperative National Park Resources Studies ornamental plants. 3. The tamarisks. California
Unit. p. 20-24. Horticulture Society Journal 12: 76-83
DeLoach, C.J. 1994. Petition to release into the Moerman, D.E. 1998. Native American ethno-
field the leaf beetle Diorhaba elongata from botany. Timber Press, Portland, OR. 927 p.
China for biological control of saltcedar,
Tamarix ramosissima, a weed of riparian areas Neill, W.M. 1990. Control of Tamarisk by cut-
of the Western United States and Northern stump herbicide treatment. In: Tamarisk
Mexico. Submitted to the Technical Advisory control in Southwestern United States. Special
751
Report 9. Cooperative National Park Resources Tykač, J. 1990. The illustrated guide to ornamental
Studies Unit. p. 91-98. shrubs. Treasure Press, London. 224 p.
Olson, BE. 1999. Impacts of noxious weeds on United States Department of Agriculture, Forest
ecologic and economic systems. In: R.L Sheley Service. 1988. Range Plant Handbook. Dover
and J.K. Petroff, eds. Biology and Management Publications, Inc. New York. 838 p.
of noxious rangeland weed. Oregon State
University Press, Corvallis, OR. 528 p. Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
Higgins. 1993. A Utah flora. Brigham Young
Sisneros, D. 1991. Herbicide analysis: Lower University, Provo, UT. 986 p.
Colorado River saltcedar vegetation
management study. Publication R-91-06. Wilgus, F. and K.C. Hamilton. 1962. Germination
Bureau of Reclamation, Denver, CO. 165 p. of salt cedar seed. Weeds 10: 332-333.
752
Tamonea boxiana (Moldenke) Howard crow broom
VERBENACEAE
Synonyms: Ghinia boxiana Moldenke

Tomonea spicata sensu Urban
Ghinia spinosa sensu Britt. & Wils.
John K. Francis
horns and contain a four-celled central cavity with

1-mm oblong seeds (Howard 1989, Liogier 1995).
Range.—Crow broom is native to Puerto Rico,

the Virgin Islands, Barbuda, and Antigua (Howard
1989, Liogier 1995). It is not known to have been
Ecology.—Crow broom grows in dry habitats that

receive from about 700 to 900 mm of annual
precipitation. It is normally found within a few km
of the coast at elevations from near sea level to
about 450 m. Soils are variable in texture, pH, and
parent material, but are always well drained. Crow
broom is intolerant of shade and heavy
competition and apparently requires disturbance to
become established. It is usually open-grown but
also occurs in savannas. Crow broom is not eaten
or is browsed little by cattle and appears to benefit
from control of competition in overgrazed
pastures. Vasquez and Kolterman (1998) list it as
“occasional” in pasture and shrublands in
Southeastern Puerto Rico. It is also found on
roadsides, disturbed areas, and on rocky ridges
and hill sides.
Reproduction.—Crow broom blooms during the

dry season (March and April). It is insect
pollinated. The fruits mature during the wet season
and are released at the end of the wet season
(December through February). The seed-like fruits
are dispersed by sticking to animals or clothing,
General Description.—Crow broom, also known and after being released, germinate (presumably
as coast broom and cardero, is a branchy shrub 40 during the wet season) and grow. A collection of
cm to 100 cm in height. The stems are slender and air-dried fruits from Puerto Rico weighed an
woody with many bifurcating branches. Crow average of 0.0108 + 0.0003 g/seed or 93,000
broom plants are supported by robust taproots fruits/kg.
with laterals through their lengths. The short-
petioled lower leaves are oblong, 0.8 to 1 cm long Growth and Management.—Crow broom plants
and may have three to five lobes; the upper leaves grow 20 to 30 cm/year and live about 4 or 5 years.
are linear and up to 2.5 cm long. Leaves and twigs No management experience has been reported.
are yellow-green. The plant is deciduous or There appears to be no reason to encourage the
semideciduous during the dry season. Tiny purple species and possibly motivation in rangeland to
or whitish flowers are grouped in terminal control it. However, it is seldom common enough
racemes. The fruits are dry and shiny with four to warrant control measures.
753
Benefits and Detriments.—Crow broom Liogier, H.A. 1995. Descriptive flora of Puerto
contributes to biodiversity in the habitat where it Rico and adjacent islands. Vol. 4. Editorial de
grows, helps protect the soil, and furnishes cover la Universidad de Puerto Rico, Río Piedras, PR.
for wildlife. It may become a minor weed in 617 p.
rangelands and the clinging seeds can be a
nuisance. Vasquez, O.J. and D.A. Kolterman. 1998. Floristic
composition and vegetation types of the Punta
References Guaniquilla Natural Reserve—Cabo Rojo,
Howard, R.A. 1989. Flora of the Lesser Antilles. 4): 265-279.
Vol. 6. Arnold Arboretum, Harvard University,
754
Taxus brevifolia Nutt. Pacific yew
TAXACEAE
Synonyms: Taxus baccata L. ssp. brevifolia (Nutt.) Pilger

Taxus baccata var. canadensis Benth.
Taxus boursieri Carrière
Taxus lindleyana A. Murray
John K. Francis
different trees (dioecious). The fruit (seed) is two-

to four-angled, ovoid, 5 to 6.5 mm long, enclosed
in a cup-like, fleshy red aril about 10 mm in
diameter. Seeds have a large oily endosperm and a
small embryo (Abrams 1940, Bolsinger and
Jaramillo 1990, Earle 2002, Rudolf 1974, Viereck
and Little 1972).
Range.—Pacific yew is native to the coastal

forests from southern Alaska through northern
California and a moist interior corridor from
southern British Columbia and Alberta to
northwestern Montana, north-central Idaho, and a
little of eastern Washington and Oregon (Bolsinger
and Jaramillo 1990). Although the species has
been planted to a limited extent in Europe, it is not
known to have naturalized outside its native range.
Ecology.—Pacific yew is very tolerant according

to the five-step tolerance rating. Seedlings grow,
develop, and eventually fruit under heavy conifer
forest canopies. The species is most often found in
deep ravines, along streams, and under coniferous
General Description.—Pacific yew, also known stands (Sargent 1923). It grows as an understory
as western yew and mountain mahogany, is an dominant in most of the forest types where it
evergreen large shrub or small tree. In good sites occurs. Under closed stands, the limbs are often as
in coastal areas and a few inland areas, old long as the trees or shrubs are tall (Bolsinger and
specimens sometimes reach 60 cm in diameter at Jaramillo 1990, Termenstein 1990). Leaves persist
breast height and 15 m in height. In high elevation on the twigs 4 to 5 years (Abrams 1940). Although
and poor sites, Pacific yew usually grows to only a initially shocked, it is able to adapt to full sunlight
few meters in height and a few centimeters in after overstory removal by logging, and it
diameter, sometimes forming low mats. The bark sometimes grows in the open in avalanche chutes,
of stems is thin (6 mm) and has a multihued and on ridges and rocky slopes at high elevations
purplish scale pattern. Bark on young twigs is rose (Bolsinger and Jaramillo 1990, Termenstein 1990).
colored and on older ones, reddish-brown. The Plants growing in the interior of the continent are
sapwood is yellow and the heartwood red to more often shrubby than those near the coast
reddish brown. Roots are deep and wide- (Sargent 1923). It typically grows as an understory
spreading. The green leaves are linear, 8 to 35 mm tree 3 to 5 m tall near the coast and as a shrub
long and 1 to 3 mm broad, acute at the tip, with a about 1 m tall east of the Cascade Mountains. It
median ridge on the upper surface and two yellow- grows from near sea level to about 2,200 m in
green bands on the lower surface. They are elevation (Earle 2002).
whorled but bent to give a two-ranked appearance.
Inconspicuous male and female strobili are borne Reproduction.—Pacific yew flowers mostly in
on the underside of branches and occur on June, matures fruits August to October, and
755
releases seeds in October. The pollen is dispersed mauls, splitting wedges, gunstocks, boat decking,
by the wind. There are 32,000 to 36,000 seeds/kg fence posts, musical instruments, carved figurines,
(Washington source). About 50 to 99 percent will novelty items, furniture, oriental ceremonial
eventually germinate under proper conditions. The “Toko” poles, turnery, cabinetry, and firewood
seeds are dispersed primarily by birds. Natural (Bolsinger and Jaramillo 1990, Termenstein 1990).
germination usually takes place in the second Infusions, decoctions, and poultices of leaves,
spring after dispersal (Rudolf 1974). Development twigs, and bark were used by Native Americans to
of seedlings is slow. Pacific yew sprouts from treat lung problems, stomachache, wounds, and
stumps and rootstalks after being cut or top-killed. pain (Moerman 1986). Pactitaxel (often called by
It commonly layers when branches or stems are the trade name Taxol) was discovered during a
pressed to the ground by snow or falling debris massive anticancer activity screening program by
(Bolsinger and Jaramillo 1990). the National Cancer Institute during the 1960’s
(21cecPharm 2003). Paclitaxel is a white
Growth and Management.—Growth of Pacific crystalline powder extractable from any of the
yew is slow. Trees in Idaho took 25 years to reach Taxus species. It is a cytotoxic anticancer drug
a stem diameter (15 cm above the ground) of 2.5 used to treat ovarian, breast, and lung cancers, and
cm and 100 years to reach 15 cm. Height growth is Kaposi’s sarcoma (SFT Enterprise 2003).
correspondingly slow (Bolsinger and Jaramillo Harvesting for peclitaxel extraction threatened to
1990). Even in moist, rich soils, few plants live decimate the species’ populations until it was
long enough to reach tree sizes. Fruits should be discovered that other Taxus species were better
picked by hand as soon as they are ripe. The fruit sources and that the drug could be produced by
pulp is removed by maceration and floating off the semi-synthesis and by cell cultures (21cecPharm
pulp, or by soaking in water at room temperature 2003). Pacific yew is used to a limited extent as an
for 4 or 5 days, rubbing the fruits on screens, and ornamental shade tree, foundation plant, hedge,
washing away the pulp. The seeds should be air- and topiary plant (Bolsinger and Jaramillo 1990).
dried for 1 to 2 weeks and should be sown or
stored as soon as drying is complete. Air-dried References
seeds can be stored for 5 to 6 years in sealed
containers at 1 to 2 °C. Seeds sown in nursery beds 21cecPharm. 2003. Paclitaxel story. http://www.
should be covered by about 1 cm of mineral soil 21cecpharm.com/px/story.htm. 4 p.
and mulched. Germination primarily occurs in the
second spring afterwards. To obtain germination a Abrams, L. 1940. Illustrated flora of the Pacific
shorter time after sowing, seeds should be States. Stanford University Press, Stanford, CA.
stratified for 90 to 210 days at 16 °C followed by 538 p.
60 to 120 days at 3 to 6 °C (Rudolf 1974).
Bolsinger, C.L. and A.E. Jaramillo. 1990. Taxus
Benefits.—Pacific yew contributes to the brevifolia Nutt., Pacific yew. In: R.M. Burns,
aesthetics of the forest, helps protect the soil, and and B.H. Honkala, eds. Silvics of North
provides food and cover for wildlife. It is a America. Agriculture Handbook 654. Forest
preferred moose browse. Although eaten in all Service, U.S. Department of Agriculture,
seasons, during the winter, moose will eat all the Washington, DC. p. 573-579.
available leaves and twigs, and even strip the bark.
Deer, elk, and rabbits also browse the species. Earle, C.J. 2002. Taxus brevifolia Nuttall 1849.
Livestock make limited use of it during the winter http://www.botanik.uni-bonn.de/conifers/ta/ta/
and when other food is lacking. The fruit is sweet brevifolia.htm. 3 p.
and eaten by many species of birds (Tirmenstein
1990). Plants growing along streams help protect Moerman, D.E. 1986. Medicinal plants of Native
against streambank erosion and shade the water America. Technical Reports 19. University of
and help maintain cool stream temperatures Michigan Museum of Anthropology, Ann
(Termenstein 1990). Pacific yew wood is fine- Arbor, MI. 534 p.
grained, heavy, hard, elastic, and strong. It has
been used and still is used to a limited extent (due Rudolf, P.O. 1974. Taxus L., yew. In: C.S.
to scarcity) for archery bows, harpoon shafts, Schopmeyer, tech. coord. Seeds of woody plants
canoe paddles, household utensils, tool handles, in the United States. Agriculture Handbook 450.
756
U.S. Department of Agriculture, Forest Service, Tirmenstein, D.A. 1990. Taxus brevifolia. In: U.S.
Washington, DC. p 799-802. Department of Agriculture, Forest Service,
Sargent, C.S. 1923. Manual of the trees of North Laboratory, Missoula, MT. Fire Effects
America (exclusive of Mexico). Houghton Information System. http://www.fs.fed.us/
Mifflin, Boston, MA. 910 p. database/feis/plants/tree/ taxbre/all.html. 19 p.
SFT Enterprise. 2003. Natural pactitaxel, full Viereck, L.A. and E.L. Little, Jr. 1972. Alaska
information. http://www.21cep.com/sft/ trees and shrubs. Agriculture Handbook 410.
pxsft.htm. 9 p. U.S. Department of Agriculture, Forest Service,
757
Taxus canadensis Marsh. Canada yew
TAXACEAE
Synonyms: Taxus minor (Michaux) Britt.

Taxus procumbens Loddiges
Taxus baccata L. var. canadensis Gray
Taxus baccata L. var. minor Michx.
Taxus baccata L. var. procumbens Loud
Gerry Moore and John K. Francis
winter, the leaves can be slightly reddish. Being a

gymnosperm, Canada yew lacks true flowers or
fruits. It does produce pollen and seeds, on
separate plants (dioecious) or on the same plant
(monoecious) in axillary cones. The pollen is
produced by the male cones, which appear as
stalked heads and mature in the first season. Each
male cone is a compound of four to 16 reduced,
stalked, specialized leaves called sporophylls. The
stalk is attached to the center of the sporophyll (i.e.
they are peltate). Each sprophyll produces two to
nine pollen sacs (sporangia) where the pollen is
produced. The pollen is spherical and lacks wings.
The female cones are reduced, subtended by a
series of small, inconspicuous bracts. Each cone
produces one ovule that develops into a single
seed, maturing in one season. The seeds are
brown, flattened, slightly broader than long, 4 to 5
mm long. Each seed is surrounded by a fleshy,
scarlet, cup-shaped aril that is open at the top,
exposing the seed. Inside the aril is a clear,
mucilaginous liquid called albumen (Fernald 1950,
Drawing source: Britton and Brown 1913 Pilger 1903, Soper and Heimburger 1982).
General Description.—The common name most Range.—Canada yew is native to Eastern Canada
often used for Taxus canadensis is Canada yew. It (Manitoba, New Brunswick, Newfoundland, Nova
is also known as American yew and ground Scotia, Ontario, Prince Edward Island, Quebec)
hemlock; a common French name used in Quebec and the Northeastern United States (Connecticut,
for this species is buis de sapin. This evergreen Illinois, Indiana, Iowa, Kentucky, Maine,
species is usually a low-growing, straggling non- Maryland, Massachusetts, Michigan, New
aromatic shrub, with ascending branches up to 2 m Hampshire, New Jersey, New York, North
tall. Branches often extend two-thirds of their Carolina, Ohio, Pennsylvania, Rhode Island,
length laterally before curving upwards. The bark Tennessee, Vermont, Virginia, West Virginia,
is thin, scaly and red to reddish-brown. Twigs are Wisconsin) (Hils 1993, Natural Resources
green and alternately arranged, and winter buds are Conservation Service 2003). It becomes rare above
composed of imbricate (overlapping), lanceolate, 50 °N (Soper and Heimburger 1982).
keeled scales. The leaves are needle-like, on short
stalks, and persist for several years. The leaf Systematic Botany.—The genus name Taxus is
blades are spirally arranged, flattened, often in two from the Greek word, taxos, for yew.
ranks, linear, abruptly narrowed into a fine point, Approximately eight species of Taxus are
1.2 to 2.2 cm long, with pale green bands on the recognized, and all of them are similar
under surface. The midrib is slightly elevated on morphologically, even though they are well-
the top surface. The leaves lack resin ducts. In separated geographically (Hils 1993). Canada yew
758
is the only native yew throughout its range. are suggested to promote more rapid germination
However, it can be confused with two other yew of seeds (Dirr and others 1987). Seeds may be
species--English yew (T. baccata L.) and Japanese sown in nursery beds in the fall and covered by 10
yew (T. cuspidata Sieb. & Zucc.)--and a hybrid to 12 mm of mineral soil topped by mulch.
between them, (T. media Rehd.) that occasionally However, fall-sown seeds usually germinate the
escapes from cultivation. Canada yew can be second spring (Rudolf 1974). Seedlings should be
distinguished from these by its low-growing, transplanted while still small. However, they are
straggling nature. Immature saplings are slow growing and they must be maintained for 2
unidentifiable to species. Canada yew has 2n = 24 years in nursery beds or pots at a minimum.
chromosomes (Hils 1993). Seedlings should be outplanted in late spring or
early summer after the last frost has occurred.
Ecology.—Canada yew, a slow-growing species, Established plants will tolerate pruning (Brand
is tolerant of shade, which gives it a competitive 2003).
advantage over other species with which it must
compete. It grows best in at least partial shade Benefits.—Canada yew is an important
(Sullivan 1993). Canada yew is found in rich soil component of the forest understory in Northeastern
in a variety of habitats including bogs, swamps, forests, helping to protect the soil and imparting
and rocky banks at elevations up to 1,500 m. It beauty to forest landscapes. The species is used as
prefers soils that are well-drained and that have an ornamental for ground cover and mass
pH’s between 5.0 and 7.5. The climate is humid plantings, and for parental stock in yew breeding
and cool. Exposure to winds from the ocean is not (Sullivan 1993). The toxic compound that is
tolerated. The species grows in many forest present in the plant is the alkaloid taxine
associations and is taken as an indicator of old- (Kingsbury 1964). The seeds and the dried foliage
growth forest. It does not occur in seral have been fatal to livestock, but the fresh foliage is
communities. Canada yew is apparently easily browsed by deer (Fernald 1950, Hils 1993). Moose
killed by fire. Wildfires and other influences that also browse the species and have seriously reduced
open the forest canopy reduce the competitive Canada yew abundance on Isle Royale, Michigan,
ability of the species (Sullivan 1993). since their introduction. The fleshy aril is eaten by
numerous species of birds (Sullivan 1993). The
Reproduction.—Canada yew flowers in April and pulp of the arils is sweet and edible to humans
May and is wind pollinated. Individual plants are (Fernald 1950) but has a slimy texture (Soper and
not self-fertile (Plants For a Future 2003). It Heimburger 1982). A number of Native American
produces seed almost every year (Sullivan 1993). groups made decoctions of leaves and twigs to
The seeds mature in late summer and autumn. treat rheumatism. Infusions and decoctions were
There are 33,000 to 62,400 seeds/kg (Rudolf also used to treat numbness of fingers and legs,
1974). The seeds are dispersed by birds (Sullivan colds, gonorrhea, and as a diuretic (Moerman
1993). Canada yew spreads vegetatively by 1986). New foliage and green stems are
layering (Rook 1998). The connections between commercially harvested for the extraction of
plants of a clonal group usually rot away (Sullivan paclitaxel (Public Forest Council 2003). Paclitaxel
1993). Small plants tend to be males and browsing is a white crystalline powder extractable from any
and other stresses on a population tends to increase of the Taxus species. It is a cytotoxic anticancer
the proportion of male plants in the population drug used to treat ovarian, breast, and lung
(Sullivan 1993). cancers, and Kaposi’s sarcoma (SFT Enterprise
2003).
Growth and Management.—Canada yew is slow
growing (Public Forest Council 2003). It is the References
hardiest of all yew species; however, it is the least
attractive yew species used as an ornamental due Brand, M.H. 2003. Plant UConn database of trees,
to its low-growing, straggling habit (Rehder, shrubs and vines: Taxus canadensis.
1940). The species may be propagated with seeds http://www.hort.uconn.edu/plants/t/taxcan/
and by cuttings of half-hardened or hardened taxcan3/html. 2 p.
terminal shoots that root with good success (Plants
For a Future 2003). Warm stratification for 4 Britton, N.L. and A. Brown. 1913. Illustrated flora
months followed by 4 months of cold stratification of the northern states, Canada, and the British
759
possessions. Vol. 2, 2nd Ed. Scribner, New York. Public Forest Council. 2003. Ground hemlock
735 p. (Taxus canadensis). Public Forest Council Fact
Sheet. Charlottetown, Prince Edward Island,
Dirr, M.A., M.W. Heuser, Jr., and B.L. Dirr. 1987. Canada http://www.gov.pe.ca/af/agweb/
The reference manual of woody plant library/factsheets/ground_hemlock.pdf. 2 p.
propagation. Varsity Press, Athens, GA. 239 p.
Rehder, A. 1940. Manual of cultivated trees and
Fernald, M.L. 1950. Gray’s manual of botany. shrubs. Macmillan Publishing Co., New York.
American Book Co., New York. 1,632 p. 996 p.
Hils, H.J. 1993. Taxaceae. In: Flora of North Rook, J.S. 1998. Taxus canadensis, Canada yew.
America Editorial Committee, eds. Flora of http://www.rook.org/earl/bwca/nature/shrubs/
North America. Vol. 2. Oxford University Press, taxuscan.html. 3 p.
New York. p. 423-427.
Rudolf, P.O. 1974. Taxus L., yew. In: C.S.
Kingsbury, J.M. 1964. Poisonous Plants of the Schopmeyer, tech. coord. Seeds of woody plants
United States and Canada. Prentice-Hall, Inc., in the United States. Agriculture Handbook 450.
Englewood Cliffs, NJ. 626 p. U.S. Department of Agriculture, Forest Service,
Washington, DC. p. 799-802.
America. Technical Reports 19. University of SFT Enterprise. 2003. Natural pactitaxel, full
Michigan Museum of Anthropology, Ann information. http://www.21cep.com/sft/
Arbor, MI. 534 p. pxsft.htm. 9 p.
Natural Resources Conservation Service. 2003. Soper, J.H. and M.L. Heimburger. 1982. Shrubs of
Plants profile: Taxus canadensis Marsh. http:// Ontario. Royal Ontario Museum, Toronto,
plants.usda.gov/cgi_bin/plant_search.cgi?mode= Ontario, Canada. 495 p.
Scientific+Name&keywordquery=Ta...
[not paged]. Sullivan J. 1993. Taxus canadensis. In: U.S.
Pilger, R.K.F. 1903. Taxaceae. In: H.G.A. Engler, Rocky Mountain Research Station, Fire Sciences
ed. Das Pflanzenreich. V. 18(IV,5). Laboratory, Fire Effects Information System.
p. 1-124. http://www.fs.fed.us/database/feis/plants/shrub/
taxcan/all.html. 10 p.
Plants For a Future. 2003. Taxus canadensis.
http://www.ibiblio.org/pfaf/chi-bin/arr_html?
Taxus+canadensis. 7 p.
760
Tecoma stans (L.) Juss. ex Kunth ginger-thomas
BIGNONIACEAE
Synonyms: Bignonia stans L.

Gelseminum stans (L.) Kuntze
Stenolobium stans (L.) Seem.
John K. Francis
northern Argentina and from Florida and the

Bahamas to Trinidad in the Caribbean. It has
naturalized in much of tropical and subtropical
Africa, Asia, the Pacific islands, and Australia.
Ecology.—Ginger-thomas does not tolerate heavy

frost. However, it will grow in most well-drained
soils, including calcareous fill, infertile sands,
acidic Ultisols, and volcanic regolith in areas
receiving from 700 to 1800 mm of rainfall. The
species is described as a water spender that is able
to convert to a water saver (Tipton 1994). It
requires minimal competition and nearly full
sunlight to survive (shade intolerant). Roadsides
and disturbed areas are the most common habitats.
Given a start in secondary forest, it is usually
present for only 10 to 20 years following
disturbance. Many species of insects feed on
ginger-thomas, and it is attacked by the genera of
parasitic plants, Cuscuta, and a number of disease
organisms; but, nothing seems to pose a serious
threat to the species.
Reproduction.—Ginger-thomas produces flowers

that have functional male and female parts. The
tubular flowers are fragrant, 4 to 5 cm in length,
Illustration source: USDA-Forest Service collection, and pollinated by bees and other insects and
Hunt Institute hummingbirds. The 10- to 25-cm pods (capsules)
develop in about 1 month and liberate large
General Description.—Ginger-thomas, also numbers of papery-winged seeds. The species may
known as roble amarillo, saíco amarillo, bois flower and fruit nearly throughout the year (Little
caraîbe, trumpet flower, yellow-elder, and many and Wadsworth 1964) in climates without strong
other common names (Little and Wadsworth seasonal change, or flower heavily in autumn in
1964), is a medium-sized shrub with many seasonal climates such as southern Florida.
branches and basal stems. The leaves are opposite, Developing pollen becomes sterile when
pinnately compound with 5 to 13 saw-toothed temperatures rise above 34 °C. This leads to seed
leaflets. The twigs are green, turning brown, and failures during summer months in many areas
the older bark is light gray and very furrowed. A (Kumar and Singh 1988). Natural reproduction
profusion of bright yellow flowers has made the may vary from dense to scattered. In nurseries,
species much loved throughout the tropics. Ginger- ginger-thomas is usually propagated from seed,
thomas is the official flower of the U.S. Virgin although greenwood cuttings can also be rooted
Islands. (Baily 1941). A collection of seeds from Puerto
Rico averaged 208,000 seeds/kg. Germinating
Range.—The native range extends from southern began in 3 days and finished with 97 percent
Texas, New Mexico, and Arizona to Bolivia and
761
germinated (Francis and Rodríguez 1993). No Bailey, L.H. 1941. The standard cyclopedia of
pregermination treatments are necessary. horticulture. Vol. 3. The MacMillan Company,
New York. p. 2,423-3,639.
Growth and Management.—Early growth is
relatively rapid. About 1 m of height growth can be Francis, J.K. and A. Rodríguez. 1993. Seeds of
obtained during the first year. Depending on the Puerto Rican trees and shrubs: second
environment, maximum height of plants may vary installment. Research Note SO-374. U.S.
from 1.5 m to 7 m, and diameters of 8 cm are Department of Agriculture, Forest Service,
sometimes achieved. Establishment activities may Southern Forest Experiment Station, New
include removal of competition and disturbance of Orleans, LA. 5 p.
the soil to encourage natural regeneration, planting
(more reliable for smaller numbers of plants), and Kranz, W.M. and T. Passini. 1997. Amarelinho:
control of competing vegetation. When control of biologia e control. Informe da Pesquisa
ginger-thomas is needed, cutting, grubbing out No.121.Instituto Agronomico do Parana.
stumps, or poisoning is recommended. In Brazil, Londrina, PR, Brazil. 19 p.
tebuthiuron was found to effectively kill plants, but
picloram and 2, 4-D were not (Passini and Kranz Kumar, R. and G. Singh. 1988. Investigations into
1997). the cause of sterility. Tecoma stans L. Letters
Botaniques (France) 135(2): 131-135.
Benefits.—Ginger-thomas is planted as an
ornamental throughout the topics and subtropics. It Little, E.L., Jr. and F.H. Wadsworth. 1964.
is especially prized as a flowering hedge plant. The Common trees of Puerto Rico and the
shrub is planted and managed to enhance the VirginIslands. Agriculture Handbook 249. U.S.
beauty of green belts and natural forests used for Department of Agriculture, Forest Service.
recreation. Used in combination with trees, ginger- Washington, DC. 548 p.
thomas contributes to effective windbreaks and
sound breaks. Growing in thick patches that shade Liogier, H.A. 1990. Plantas medicinales de Puerto
out grass, the species has become a serious weed in Rico y del Caribe. Iberoamericana de Ediciones,
Brazilian pastures (Kranz and Passini 1997). Inc., San Juan, PR. 566 p.
Browsing cattle and goats in Mexico consume up
to 20 percent of the leaves and 100 percent of the Lozoya-Meckes, M. and V. Mellado-Campos.
available flowers (Susano Hernandez 1981). The 1985. Is the Tecoma stans infusion an anti-
foliage in one study in India contained 17 percent diabetic remedy? Journal of Ethnopharmacology
crude protein, 6 percent ash, 18 percent fat, 25 14(1): 1-9.
percent fiber, and 14 percent total polyphenols
(Nag and others 1994). Ginger-thomas leaves, Neg Datta, A., A.K. Lehiri, S. Matai, and L. Si.
bark, and roots contain many biologically active 1994. Proximate composition and polyphenol
chemicals, and extracts from those tissues have content of some tree leaves. Indian Forester
been used in traditional folk medicine to treat many 120(12): 1,122-1,125.
diseases and conditions (Liogier 1990). Perhaps the
most promising compounds are monoterpine Passini, T. and W.M. Kranz. 1997. Eficacia de
alkaloids, which have been shown to effectively herbicidas no controle de amarelinho (Tecoma
reduce the symptoms of diabetes mellitus in rats, stans) em pastagem. Planta Daninha 15(2): 190-
dogs, and mice (Aguilar and others 1993, Lozoya- 197.
Meckes and Mellado-Campos 1985, Perez and
others 1984) Perez G., R.M., A. Ocegueda Z., J.L. Muñoz L.,
J.G. Avila A., and W.W. Morrow. 1984. A study
References of the hypoglycemic effect of some Mexican
plants. Journal of Ethnopharmacology 12(3):
Aguilar, L.C., S. Macias, A. Chagoya, A. 253-262.
Cardenas, P. Díaz, and J M. Cantu. 1993.
Antidiabetic activity of Tecoma stans in rats. Susano Hernandez, R. 1981. Especies arboreas
Fitoterapia 64(4): 304-305. forestales susceptibles de aprovecharse como
forraje. Ciencia Forestal 6(29): 31-39.
762
Tipton, J.L. 1994. Relative drought resistance
among selected southwestern landscape plants.
Journal of Agriculture 20(3): 150-155.
763
Tetrapterys inaequalis Cav. bejuco de paralejo
MALPIGHIACEAE
Synonyms: Tetrapteris citrifolia (Sw.) Pers.

Banisteria bracteata DC.
Triopteris citrifolia Sw.
Tetrapteris paniculata Bello
John K. Francis
(Howard 1988). This differs from Liogier (1988)

and Acevedo-Rodríguez (1985) who use the
synonym T. citrifolia and add Jamaica, Hispaniola,
and Tobago to the range. Howard states: “The
flowering specimen from Tobago that Niedenzu
called T. citrifolia is almost certainly T. discolor
(G. Meyer) DC. ...Niedenzu considered [T.
inaequalis] a synonym of Tetrapterys citrifolia
(Sw.) Pers. They are certainly closely related, both
belonging to the difficult group called Section
Lophogynixa by Niedenzu. However, T. citrifolia
from Jamaica, the type locality, has a well-
developed dorsal wing on the samara and young
stems that are tightly sericeous with straight
appressed hairs. These differences lead me to
General Description.—Bejuco de paralejo, also consider T. citrifolia endemic to Jamaica and apply
known as brjuco de sopla and aile à ravet, is a the later name T. inaequalis to the plant of Puerto
woody vine or climbing shrub that may extend 15 Rico and the Lesser Antilles. ...The species of the
m laterally and reach 10 m into the crowns of Lesser Antilles seems not to occur on
trees, with stems as thick as 8 cm. The root system Hispaniola...” In their latest checklist, Liogier and
is superficial, especially roots arising from layered Martorell (2000) now concur with Howard.
stems. The stems are cylindrical, smooth, and have
rings at the nodes. The wood is moderately hard Ecology.—Bejuco de paralejo colonizes a wide
and has growth rings. The stems tend to be long range of soils derived from both sedimentary and
and little branched except near the extremities. igneous rocks. It requires mean annual rainfall of
The simple, opposite ovate or elliptical leaves from about 1200 to about 2500 mm. The species
have petioles 1 to 1.5 cm long, and blades 5 to 17 grows at elevations between 110 and 500 m in the
cm long and 5 to 8 cm broad, rounded to cordate Lesser Antilles (Howard 1988) and from a few
at the base and pointed at the tip. Inflorescences meters above sea level to 600 m or more in Puerto
are panicles of four-flowered cymes flanked by Rico. Bejuco de paralejo does not tolerate shade
leaves of reduced size that often shrivel as the well. It will survive and grow slowly in light shade
fruits mature. The five-merous, yellow-petaled but requires full or nearly full sunlight to flower
flowers are about 2 cm in diameter. The fruits and fruit. The species may be found on roadsides,
consist of four samaras united at the base. Each along rivers, in brushy pastures, early secondary
has an upper lateral wing 17 to 28 mm by 6 to 10 forests, and clearings in secondary and remnant
mm and a lower lateral wing about half as large forests.
and a medial ridge protruding about 1 mm
(Acevedo-Rodríguez 1985, Howard 1988, Liogier Reproduction.—Bejuco de paralejo usually
1988). flowers in Puerto Rico in October and November
and fruits in July to October (Acevedo-Rodríguez
Range.—The range of bejuco de paralejo includes 1985). Howard (1988) notes collections in the
Puerto Rico and the offshore island of Vieques, St. Lesser Antilles with flowers and fruits in every
Thomas, and St. Croix in the U.S. Virgin Islands, month except May and June. Hundreds of seeds
Antigua, Guadaloupe, Martinique, and St. Vincent can be produced by a large plant. Air-dried
764
samaras from Puerto Rico collected by the author References
in April averaged 0.0953 + 0.0014 g/fruit. Planted
on commercial potting mix without any Acevedo-Rodríguez, P. 1985. Los bejucos de
pretreatment, 79 percent germinated between 34 Puerto Rico. Vol. 1. General Technical Report
and 82 days after sowing. Samaras spiral sideways SO-58. U.S. Department of Agriculture, Forest
a few meters from fruiting adult plants. While Service, Southern Forest Experiment Station,
young plants are relatively common in many New Orleans, LA. 331 p.
areas; fruiting plants are relatively rare. Layering
(rooting in contact with the ground) is common Howard R.A. 1988. Flora of the Lesser Antilles,
and appears to be an important means of Leeward and Windward Islands.
perpetuating established plants. Dicotyledoneae, Part 1. Vol. 4. Arnold
Growth and Management.—Bejuco de paralejo MA. 673 p.
vines extend rapidly from sprouts. A large vine
(7.7 cm in diameter) cut by the author had 27 Liogier, H.A. 1988. Descriptive flora of Puerto
growth rings. Although large plants are capable of Rico and adjacent islands, Spermatophyta. Vol.
smothering small trees, the species causes few 2. Editorial de la Universidad de Puerto Rico,
problems. No planting or management experience Río Piedras, PR. 481 p.
has been published.
Liogier, H.A, and L.F. Martorell. 2000. Flora of
Benefits.—Bejuco de paralejo contributes to the Puerto Rico and adjacent islands, a systematic
biodiversity of forests, helps protect the soil, and synopsis. 2nd ed. Editorial de la Universidad de
furnishes cover for wildlife. Puerto Rico, Río Piedras, PR. 382 p.
765
Thrinax morrisii H. Wendl. brittle thatch palm
ARECACEAE
Synonyms: Sampsonia microcarpa (Sarg.) O.F. Cook

Thrinax keyensis Sarg.
Thrinax microcarpa Sarg.
Thrinax ponceana O.F. Cook
Thrinax praeceps O.F. Cook
John K. Francis
long, split at the base. There are deeply incised leaf

segments 33 to 75 cm long and 2.3 to 4.8 cm
broad. The inflorescences (spadixes or panicles)
are 55 to 100 cm long, arching or straight and
usually extend beyond the leaves. Small, white
flowers are borne in large numbers. The fruits are
white (turning yellow at maturity) drupes, 3.5 to 8
mm in diameter, containing one hard, spherical
seed 2.7 to 4.2 mm in diameter. Chromosome
number is 2n = 36 (Gilman and Watson 1994,
Howard 1979, Little and others 1974, Long and
Lakela 1976, Nelson 1996)
Range.—Brittle thatch palm is native to southern

Florida, the Bahamas, Cuba, Hispaniola, Puerto
Rico, the Virgin Islands, Anguilla, and Barbuda
(Howard 1979, Little and others 1974). It is
considered endangered in Florida (Institute of
Systematic Botany 2002). The species is planted as
an ornamental within and outside its natural range
(Hoyos-F. and Braun 2001, Little and others 1974)
but has not been reported to have naturalized.
Ecology.—Brittle thatch palm grows in sites

where it suffers a minimum of competition. It
grows along the edges of hammocks and in
General Description.—Brittle thatch palm is also pinelands in the Florida Keys (Nelson 1996). In
known as Key thatch palm, small-fruited thatch Puerto Rico, it grows primarily on cliffs, and
palm, broom palm, buffalo-tip, palma de escoba, ridges of dry and moist limestone and ultramafic
yaray, pandereta, palma de petate, palma de rocks at elevations from near sea level to 300 m
cogollo, guano de sierra, miraguano, and palmita. (Little and others 1974). The soils are sands, or are
It is an evergreen, single-stemmed shrub or small loams or clays shallow over fractured rocks. Brittle
tree 1 to 6 m in height. Although most plants in thatch palm is moderately intolerant of shade. It
natural stands do not live long enough to develop a occurs singly, in small groups or in nearly pure
discernable stem, when formed, it is brown or thickets in the open, in small openings, or under
gray, 7 to 13 cm in diameter at breast height, with open stands, usually of low trees. The species can
many vertical cracks and prominent leaf scars in withstand temperatures as low as -4 °C (Desert-
the clean portion, covered with old leaf bases near tropicals 2002), is tolerant of drought and salt
the top, and thick with rootlets at the base. Brittle spray, and resists breakage in storms (Gilman and
thatch palms maintain about 20 fan-shaped, pale Watson 1994).
blue-green or yellow-green leaves, spirally
arranged on the trunk. The petioles are 27 to 84 cm Reproduction.—Brittle thatch palm in natural
stands flowers seasonally or irregularly in response
766
to favorable conditions in Puerto Rico (Little and squirrel (Sciurus aureogester (Cowley 2002) and
others 1974) and flowers in the spring in Florida the eastern box turtle (Terrapene carolina) (Liu
(Gilman and Watson 1994). The flowers are insect and others 2002), and in the Bahamas by the
pollinated. A collection of seeds in Puerto Rico Bahamas amazon parrot (Amazona leucocephala
averaged 0.0396 + 0.0007 g/seed or 25,000 bahamensis) (Attril 2002).
seeds/kg. Sown without pretreatment on moist
blotter paper, 85 percent germinated in 17 months References
(author’s observation). The seeds are reported to
begin germination 50 to 70 days after sowing Attril, R. 2002. Bahamas wildlife pages: The status
(Hoyos-F. and Braun 2001). The seeds can stand and conservation of the Bahamas amazon.
storage at -40 °C and 66 °C for at least 1 week http://www.bahamaswildlife.fsnet.co.uk/
without loss of viability (Broschat and Meerew parrotconservation.htm. 8 p.
2000). Because seeds have such a long dormant
period, they can be collected by hand from plants Broschat, T.K. and A.W. Meerew. 2000.
or from the ground at almost any time of the year. Ornamental palm horticulture. University Press
The seeds are dispersed by animals. Surviving of Florida, Gainesville, FL. 255 p.
seedlings are not usually common, but the plants
are long-lived and populations can build to high Cowley, M. 2002. Florida native plants: palm
levels. family. http://www.nsis.org/garden/family/palm.
html. 3 p.
Growth and Management.—Brittle thatch palm
is slow growing. A specimen in the Fairchild Desert-tropicals. 2002. Broom palm. http://desert-
Tropical Garden, Florida was 42 years old and tropicals.com/Palm/Thrinax_morrisii.html. 3 p.
1.16 m in height (Zona and Maidman 2000). Plants
mature and begin flowering and fruiting with only Gilman, E.F. and D.G. Watson. 1994. Thrinax
a rosette of basal leaves, long before they develop morrisii, Key thatch palm. Fact Sheet ST-628.
a discernable stem. Plants with long stems are U.S. Department of Agriculture and Southern
uncommon and are probably more than 50 years Group of State Foresters. Winder, GA. 3 p.
old. Wildland plantings are not documented but
presumably could be done with container stock. Howard, R.A. 1979. Flora of the Lesser Antilles,
Either they must be established in poor sites where Leeward and Windward Islands. Vol. 3. Arnold
few other species will grow, or weed protection Arboretum, Harvard University, Jamaica Plain,
must be given almost in perpetuity. Management MA. 586 p.
of existing stands should consist mainly of
protection from fire, development, or heavy Hoyos-F., J. and A. Braun. 2001. Palmas en
grazing. Venezuela. Monograph 47. Sociedad de
Ciencias Naturales La Salle, Caracas,
Venezuela. 424 p.
Benefits.—Brittle thatch palm helps protect the
soil, contributes to the aesthetics of natural stands, Institute of Systematic Botany. 2002. Thrinax
and furnishes food and cover for wildlife. The morrisii. Institute of Systematic Botany,
species is grown and sold as an ornamental. It is University of South Florida, Tampa, FL.
used as a container plant and for buffer strips in http://www.plantatlas.usf.edu/main.asp?plantID
parking lots and medians on highways as well as =555. 2 p.
background and specimen plantings (Gilman and
Watson 1994). Leaves are still used to make Little, E.L., Jr., R.O. Woodbury, and F.H.
brooms, thatch, and mats (called petate) (Osvaldo Wadsworth. 1974. Trees of Puerto Rico and the
2002), although now for ornamentation rather than Virgin Islands. Vol. 2. Agriculture Handbook
necessity. The fruit pulp contains 4,083 µg/g of 449. U.S. Department of Agriculture,
oxalate—capable of causing a burning sensation Washington, DC. 1024 p.
on the skin of sensitive people—that may reduce
its palatability to some species of wildlife Liu, H., S.G. Platt, and C.K. Borg. 2002. Seed
(Broschat and Meerew 2000). However, it is eaten dispersal by the eastern box turtle (Terrapene
in Florida by the introduced Mexican red-bellied carolina) in subtropical pine rockland of the
767
Lower Florida Keys. Botany 2002 Conference, Florida. Pineapple Press, Inc. Sarasota, FL.
August 2-4, Madison, WS. Abstract. 391 p.
http://www.botany2002.org/section3/abstracts/3
6.shtml. 1 p. Osvaldo, J. 2002. El petate. http://joseosvaldo.
freeyellow.com/Petate/html. 12 p.
Tropical Florida. Banyan Books, Miami, FL. Zona, S. and K. Maidman. 2000. Growth rates of
962 p. palms in Fairchild Tropical Garden. Palms
45(3): 151-154.
768
Toxicodendron radicans (L.) Kuntze eastern poison ivy
ANACARDIACEAE
Synonyms: Rhus radicans L.

Rhus littoralis Mearns
Rhus blodgettii Kearney
Philostemon radicans Raf.
Toxicodendron blodgettii (Kearney) Greene
Rhus scandens Salisb. (and others: Institute of Systematic Botany 2003)
John K. Francis
1992, Secretaría de Madio Ambiente y Recursos

Naturales 2003).
Range.—Eastern poison ivy is native to Prince

Edwards Island and from Nova Scotia to southern
Ontario of Canada, all of the Eastern and
Midwestern States of the United States (except for
North Dakota), Arizona, Mexico, Guatemala, the
Bahamas, Bermuda, China, Taiwan, and Japan
(Brooks 2003, Griffiths 1994, Natural Resources
Conservation Service 2003, Pavek 1992). The
species has been planted as an ornamental and
medicinal plant and presumably naturalized in
Great Britain, Europe, and Australia (Schwartz
1999).
There are at least seven subspecies of
eastern poison ivy. Subspecies divericatum occurs
in Arizona, ssp. eximium ranges from Texas to
Guatemala, ssp. negundo grows in the Midwestern
and Northeastern sections of North America,
General Description.—Eastern poison ivy, also pubens is native to the Southwestern States, ssp.
known as climbing poison ivy, common poison radicans ranges across the Southern and Eastern
ivy, poison ivy, poison vine, and bemberecua, is a portions of U.S., ssp. verrucosm occurs in Texas,
deciduous vine or shrub capable of ascending into Oklahoma, and Missouri, and ssp. barkleyi
tree crowns 20 m or more and along the ground for inhabits the Far East portion of the range (Griffiths
as much as 45 m. The stems sometimes reach 15 1994, Natural Resources Conservation Service
cm in diameter. The older stems are brown and 2003).
covered with brown adventitious roots by which
they cling to tree trunks and other surfaces and Ecology.—Eastern poison ivy grows in soils of all
layer (root) whenever they come in contact with textures from nearly all parent materials. Although
the ground. The small stems are lighter colored it grows best in fertile, near-neutral, moist but well
and covered with dark brown dots. The alternate drained soils, it tolerates a wide range of fertility,
leaves have long (to 15 cm) petioles and three pH, and other conditions. Mean annual
ovate to lanceolate leaflets 4 to 20 cm long with precipitation ranges from 391 mm to 1,450 mm in
entire, coarsely toothed, or lobed margins. Small the United States and Canada. It grows from sea
white or greenish white flowers grow in axillary level on the Eastern and Gulf Coasts to 2,158 m in
panicles. Clusters of white or greenish-white elevation in New Mexico. In Florida swamps, it
drupes 3 to 6 mm in diameter develop later and survives inundation by fluctuations in water level
remain on the vine well after the leaves fall. The and tolerates mildly saline water. An average
leaves are dark green during the summer, turning growing season of 150 days with a mean annual
yellow to red in the fall. The species has 2n = 30 temperature of 5.3 °C occurs at the northern extent
chromosomes (Long and Lakela 1976, Pavek
769
of its range. There is an average growing season of Poison Effects.—The pale yellow oil called
240 days in southern Florida, and a mean annual urushiol, present in all plant parts, is the source of
temperature of 19 °C in Arizona (Pavek 1992). all the allergenic pain and suffering associated
The mean annual temperatures are probably with this species. Even in minute amounts, if it
higher, precipitation greater, and growing seasons stays in contact with human skin for more than 5
may approach 365 days in Central America. minutes, it will be absorbed and metabolized. The
Eastern poison ivy is moderately tolerant of shade. metabolites bind with proteins, forming
It is often the most important understory plant, compounds that the immune system recognizes as
climbs nearly every tree in many Southern river “foreign” and attacks. The immune response
floodplain forests, and is a component of most results in rash, blisters, and an intense itch
forest and vegetation types in its ample range. It (Borialforest.org 2003) that lasts for 1 to 3 weeks.
also grows in full sunlight in forest openings, Sensitivity can vary over the life of a person. A
fencerows, bluffs, railroad tracks, and vacant lots. few people have life-threatening reactions to the
Eastern poison ivy grows in old-growth forests in allergen (Schwartz 1999). The chemical is the
some portions of its range and is an early invader same and the effect (called rhus dermatitis) is the
during succession in many environments. It is same for all Toxicodendrons. Ten to 15 percent of
tolerant of fire, browsing, insects, and disease, and the U.S. population is completely immune to rhus
competes aggressively with weeds and brush. dermatitis, 25 to 35 percent react only to high
doses, and 50 percent have a consistent reaction to
Reproduction.—Eastern poison ivy blooms from relatively small doses of the allergen (Family
May through June in Virginia (Von Essen 2000) Practice Notebook.com 2003). The best defense is
and from March through May in Mexico to avoid contact with the plant, especially with the
(Secretaría de Madio Ambiente y Recursos sap or smoke from burning Toxicodendron
Naturales 2003). The start of flowering material. The next line of defense is to quickly
corresponds with leaves being about half formed wash the exposed skin with soap and copious
in the spring (Pavek 1992). The flowers are amounts of water, or if not available, with mud
pollinated by insects, including honey bees (Apis and water or an alkaline material such as baking
mellifera L.) (Pollen Lab 2003). Good crops of soda or wood ashes and water. After a rash
fruits and seeds are produced annually. Seeds are develops, the sufferer is advised to apply
dispersed by birds and mammals and sometimes hydrocortisone cream or any of several creams
by water. Seeds that have been regurgitated from sold over-the-counter for the treatment of rhus
the crop or passed through the digestive tract of dermatitis. A folk remedy mentioned in the
birds demonstrate excellent germination (Pavek literature (Felter 1922) and found by the author to
1992). Eastern poison ivy spreads extensively by almost instantly and completely (but temporarily)
means of scandent stems that root along their stop the itch and irritation of rhus dermatitis is to
whole lengths and below-ground stems (rhizomes) rub the mashed juicy stems of jewel weed
that send up shoots frequently. (Impatiens spp.) on the affected area.
Growth and Management.—Eastern poison ivy Benefits.—Eastern poison ivy, for its dark green
grows rapidly after establishment and assumes foliage during the summer and bright fall colors,
several forms. It scrambles along the ground and adds considerably to the beauty of forests and
sometimes forms thick mats. When given the shrublands where it grows. It has been used to a
opportunity, it ascends trees as a woody vine. limited extent as an ornamental and for
Plants take 3 years to reach the flowering stage conservation plantings. Eastern poison ivy is an
from seed (Pavek 1992). It often forms diffuse important browse species. It is one of the seven
stands of short upright stems. It can be partially most important food species for white-tailed deer
controlled by grazing, but there is a resurgence of in Indiana (Pavek 1992). Goats are said to like
Eastern poison ivy after secession of grazing eastern poison ivy and have been used to control it
because of the reduction in competition. Spot (Schwartz 1999). The fruits are an important fall,
spraying or wiping with the herbicide glyphosate winter, and early spring food for upland game
during the active growing season can effectively birds and many species of song birds (Pavek
control it (Pavek 1992). Grubbing and mowing 1992). Rabbits and rodents also eat the leaves and
offers some degree of control but puts the worker fruits. In herbal medicine, extracts of the leaves
at risk of contracting dermatitis. have been and still are used to treat herpetic
770
eruptions, palsy, and rheumatism. Small internal Natural Resources Conservation Service. 2003.
doses will act as a sedative but must be used with Plants profile: Toxicodendron radicans (L.)
care (Felter 1992, Grieve 2003). Kuntze. http://plants.usda.gov/cgi_profile.
cgi?symbol=TORA2. 6 p.
References Pavek, D.S. 1992. Toxicodendron radicans. In:

Borialforest.org. 2003. Toxicodendron radicans, Rocky Mountain Research Station, Fire Sciences
poison ivy. http://www.lakeheadu.ca/~borfor/ Laboratory, Fire Effects Information System.
world/herbs_shrubs/poison_ivy.htm. 2 p. http://www.fs.fed.us/database/feis/plants/vine/
toxrad/all.html. 29 p.
Brooks, B. 2003. The Toxicodendrons: poison ivy,
poison oak and poison sumac. http://nac. Pollen Lab. 2003. Pollen descriptions:
tamu.edu/x075bb/caddo/frameidx.html. 7 p. Toxicodendraon radicans. http://pollen.usda.
gov/Pollen_descriptions/Toxicodendron_radican
Family Practice Notebook.com. 2003. Rhus s.htm. 1 p.
dermatitis. http://www.fpnotebook.com/
Der10.htm. 4 p. Schwartz, D. 1999. Poison ivy (Toxicodendron
radicans (Rhus radicans). http://www.cloudnet.
Felter, H.W. 1922. The eclectic material medica, com/~djeans/FlwPlant/PoisonIvy.htm. 4 p.
pharmacology and therapeutics: Rhus
toxicodendron. http://www.ibiblio.org/herbmed/ Secretaría de Madio Ambiente y Recursos
eclectic/felter/toxicodendron-radi.html. 6 p. Naturales. 2003. Especies con usos no
maderables en bosques tropicales y
Grieve, M. 2003. A modern herbal: ivy, poison. subtropicales: Toxicodendron radicans (L.)
http://www.botanical.com/botanical/mgmh/i/ Kuntze http://www.semarnat.gob.mx/pfnm2/
ivypoi17.html. 3 p. fichas/toxicodendron_radicans.htm. 2 p.
Griffiths, M. 1994. Index of garden plants. Timber Von Essen, T. 2000. Poison Ivy, Toxicodendron
Press, Portland, OR. 1,234 p. radicans. Wildwood Park, Radford, VA.
http://www.radford.va.us/Community/wildwood
Long, R.W. and O. Lakela. 1976. A flora of /today/Plant_articles/Poison_ivy.htm. 2 p.
Tropical Florida. Banyan Books, Miami, FL.
962 p.
771
Toxicodendron rydbergii (Small ex Rydb.) Greene western poison ivy
ANACARDIACEAE
Synonyms: Rhus radicans L. var. rydbergii (Small ex Rydb.) Rehd.

Rhus radicans L. var. vulgaris (Small ex Rydb.) Rehd.
Rhus toxicodendron L. var. vulgaris Michx.
Toxicodendron desertorum Lundell
John K. Francis
is similar in appearance to eastern poison ivy [T.

radicans (L.) Kuntze] except that it is a shrub and
rarely and weakly climbs. It is known to hybridize
with eastern poison ivy (Rook 1998).
Range.—Western poison ivy is native across

southern Canada from Nova Scotia to British
Colombia and throughout the United States except
California and New Jersey and from Missouri,
Kentucky, and South Carolina southward
(McMurray 1988, Natural Resources Conservation
Service 2003). Although it has been planted
outside its natural range (botanic gardens, etc.),
there are not reports of it escaping.
Ecology.—Western poison ivy grows on a wide

range of sites--on river bottoms, streambanks,
disturbed upland sites such as roadsides,
fencerows, sand dunes, logged areas, orchards,
talus slopes in subalpine areas, and around seeps
and on shaded canyon sides in cold desert areas. It
grows from 600 to 2,500 m elevation in the
interior West and probably near sea level in coastal
General Description.—Western poison ivy, also areas. It tolerates short-term flooding. The species
called poison oak, poison ivy, and Ryberg’s poison grows in both full sun and under forest canopies. It
ivy, is a deciduous, normally upright shrub, colonizes all soil textural types derived from most
occasionally a scrambling vine, usually 20 to 100 parent materials with pH’s from mildly acid to
cm in height with stems 4 to 10 mm in diameter. mildly alkaline. Western poison ivy often grows
The woody stems, which arise from underground on skeletal soils, rocky slopes, and cliff sides. The
stems that extend up to 2 m from the parent plant, species invades quickly after disturbance. Bare
are sparingly branched and have smoothish, brown mineral soil is the best surface for seed
to gray bark, with dark brown lenticels, and wooly, germination and seedling establishment (Moye
brown, winter buds. The alternate, bright-green 1999). It often forms clonal thickets. Western
leaves are trifoliate with long petioles. Leaflets are poison ivy tolerates fire well by sprouting from
ovate to rhombic, 3 to 15 cm long, the lateral underground stems but does not increase in density
leaflets usually somewhat oblique and shorter than as a result (McMurry 1988, Welsh and others
the central leaflet. Leaflet margins are lobbed, 1987).
rounded-dentate, or entire. Small yellowish
flowers are grouped in dense axillary panicles. The Reproduction.—Flowering occurs in late spring
fruits, which remain attached throughout the through early summer. Plants on the Great Plains
winter, are ivory-colored, globose drupes 4 to 6 sometimes flower a second time in August or
mm in diameter and contain one seed each. There September (McMurry 1988). Good fruit and seed
are 2n = 30 chromosomes (Lasica 2002, St. John crops occurs almost every year. The seeds are
1963, Welsh and others 1987). Western poison ivy dispersed by birds, mammals, and gravity.
772
Seedlings are necessary to establish new colonies almost instantly and completely (but temporarily)
but vegetative (by sprouts from underground stop the itch and irritation of rhus dermatitis is to
stems) reproduction is more important for local rub the mashed juicy stems of jewel weed
spread. (Impatiens spp.) on the affected area.
Growth and Management.—Western poison ivy Benefits.—Western poison ivy helps protect the
grows at a moderate rate. Clones are relatively soil and adds beauty in the summer with its dark
stable over time. Although it is not as invasive as green leaves and in autumn with bright colors to
eastern poison ivy, it is often desirable to eliminate forest and shrublands. Although it causes browsing
it from areas of heavy human use. Spot spraying or animals no ill effects, western poison ivy is low in
wiping with the herbicide glyphosate during the protein and energy and consequently is only
active growing season can effectively control occasionally grazed by domestic animals and wild
eastern poison ivy (Pavek 1992) and probably will ungulates. However, the fruits are eaten by quail,
be effective for western poison ivy. Grubbing and wild turkeys, and some songbirds. It does provide
mowing offers some degree of control but puts the cover for small mammals and birds (McMurray
worker at risk of contracting dermatitis so that 1988). In herbal medicine, extracts of the leaves of
protective clothing should be worn during the this and other Toxicodendron species have been
operation. Branches and roots should be piled to and still are used to treat herpetic eruptions, palsy,
rot in an out-of-the-way place or bagged and sent and rheumatism. Small internal doses will act as a
to the landfill, never burned. sedative but must be used with care (Grieve 2003).
Poison Effects.—The pale yellow oil called References

urushiol, present in all plant parts, is the source of
the allergenic pain and suffering associated with Borialforest.org. 2003. Toxicodendron radicans,
this species. If even a minute amount (nanograms) poison ivy. http://www.lakeheadu.ca/~borfor/
the oil stays in contact with human skin for more world/herbs_shrubs/poison_ivy.htm. 2 p.
than 5 minutes, it will be absorbed and
metabolized. The metabolites bind with proteins, Family Practice Notebook.com. 2003. Rhus
forming compounds that the immune system dermatitis. http://www.fpnotebook.com/
recognizes as “foreign” and attacks. The immune Der10.htm. 4 p.
response results in rash, blisters, and an intense
itch (Borialforest.org 2003) that lasts for 1 to 3 Felter, H.W. 1922. The eclectic material medica,
weeks. Sensitivity can vary over the life of a pharmacology and therapeutics: Rhus
person. A few people have life-threatening toxicodendron. http://www.ibiblio.org/herbmed/
reactions to the allergen (Schwartz 1999). The eclectic/felter/toxicodendron-radi.html. 6 p.
chemical is the same and the effect (called rhus
dermatitis) is the same for all Toxicodendrons. Ten Grieve, M. 2003. A modern herbal: ivy, poison.
to 15 percent of the U.S. population is completely http://www.botanical.com/botanical/mgmh/i/
immune to rhus dermatitis, 25 to 35 percent react ivypoi17.html. 3 p.
only to high doses, and 50 percent have a
consistent reaction to relatively small doses of the Lasica, P. 2002. A flora of Glacier National Park,
allergen (Family Practice Notebook.com. 2003). Montana. Oregon State University Press,
The best defense is to avoid contact with the plant, Corvallis, OR. 512 p.
especially with the sap or smoke from burning
Toxicodendron material. The next line of defense McMurray, N.E. 1988. Toxicodendron rydbergii.
is to quickly wash the exposed skin with soap and In: U.S. Department of Agriculture, Forest
copious amounts of water, or if not available, with Service, Rocky Mountain Research Station, Fire
mud and water or an alkaline material such as Sciences Laboratory, Fire Effects Information
baking soda or wood ashes and water. After a rash System. http://www.fs.fed.us/database/feis/
develops, the sufferer is advised to apply plants/shrub/ toxryd/all.html. 15 p.
hydrocortisone cream or any of several creams
sold over-the-counter for the treatment of rhus Moye, H. 1999. The Toxicodendrons: poison ivy,
dermatitis. A folk remedy mentioned in the poison oak, and poison sumac.
literature (Felter 1922) and found by the author to
773
http://chiron.valdosta.edu/rgoddard/biol4900/mo Rook, E.J.S. 1998. Toxicodendron rydbergii,
ye/moye.html. 7 p. poison ivy. http://www.rook.org/earl/
bwca/nature/shrubs/toxicodendron.html. 4 p.
Plants profile: Toxicodendron rydbergii (Small Schwartz, D. 1999. Poison ivy (Toxicodendron
ex Rydb. Greene. http://plants.usda.gov/ radicans (Rhus radicans). http://www.cloudnet.
cgi_profile.cgi?symbol=TORA2. 5 p. com/~djeans/FlwPlant/PoisonIvy.htm. 4 p.
Pavek, D.S. 1992. Toxicodendron radicans. In: St. John, H. 1963. Flora of Southeastern
U.S. Department of Agriculture, Forest Service, Washington and adjacent Idaho. Outdoor
Rocky Mountain Research Station, Fire Sciences Pictures, Escondido, CA. 583 p.
Laboratory, Fire Effects Information System.
http://www.fs.fed.us/database/feis/plants/vine/ Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
toxrad/all.html. 29 p. Higgins, eds. 1987. A Utah flora. Great Basin
774
Trema lamarckianum (J.A. Schultes) Blume West Indian trema
ULMACEAE
Synonyms: Celtis lamarckiana Roem. & Schult.

Sponia lamarckiana (Roem. & Schult.) Decne.
Celtis lima Lam.
Trema lima authors, not Blume
John K. Francis
flowers are clustered at the leaf axils. The fruits

are rounded, fleshy, 1- to 3-mm pink drupes. The
fruits have little flavor and each contain one
brown seed (Howard 1988, Liogier 1985, Little
and Wadsworth 1964).
Range.—West Indian trema is native to Florida,

Bermuda, the Bahamas, Grand Cayman, and the
Greater and Lesser Antilles south to St. Vincent
(Liogier 1985, Little and Wadsworth 1964).
Ecology.—West Indian trema is a pioneer species.

It colonizes disturbed sites on a wide variety of soil
types over both sedimentary and igneous rocks. It is
especially common in ultramafic (serpentine) areas.
Areas in Puerto Rico with West Indian trema
populations receive annual rainfall ranging from
about 1200 to about 3000 mm at elevations ranging
from a few meters above sea level to about 900 m.
The species is intolerant of shade and usually grows
in areas with a sparse to moderate cover of herbs,
shrubs and trees. It does not grow under a closed
forest canopy. Common sites are road cuts and fill,
abandoned roads, unstable slopes, landslides,
General Description.—West Indian trema, also mechanically disturbed sites, and hammocks
known as Lamarck trema, pain-in-back, cabrilla, (author’s observation, Long and Lakela 1976).
memizo de majagua, capulí cimarrón, and orme
petites feuilles, is an evergreen shrub or small tree Reproduction.— After West Indian trema reaches
to 8 m in height and 20 cm in diameter at breast about 1 m, it flowers and fruits continuously
height. The plant is supported by a system of tan- throughout its life. During periods of favorable
colored, stiff, but flexible, tap, lateral, and fine moisture, plants produce fruits and seeds in large
roots. There is usually a single main stem. It is numbers. A collection of fruits from Puerto Rico
covered by smoothish, light brown to gray bark weighed an average of 0.0077 + 0.0001 g/fruit.
with many tiny warty lenticels over a light brown Air-dried seeds cleaned from them averaged
to pinkish, fibrous inner bark. The species has a 0.0024 + 0.0000 g/seed or 416,000 seeds/kg.
monopodial branching pattern, in which the main Placed in commercial potting mix, 38 percent of
stem produces a continuous progression of fine the seeds germinated between 60 and 120 days
lateral branches (Massey and Murphy 1996) some after sowing. Birds disperse the seeds. Seedlings
of which may thicken and become major branches are common on disturbed ground near seed-
in older plants. The alternate leaves, which are bearing plants. Young plants sprout when cut.
attached to twigs by petioles 8 to 10 mm long,
have ovate-lantiolate, green to gray-green blades Growth and Management.—Early seedling
that are rough on both surfaces and have fine- growth is slow. Later, growth reaches about 1
toothed edges. The 2-mm diameter, greenish m/year, slowing as the plants get old. West Indian
775
trema lives about 10 to 20 years. The species has Arboretum, Harvard University, Jamaica Plain,
not been reported as weedy. Reproduction MA. 673 p.
probably can be promoted by scarifying the soil
near seed-bearing plants before the start of Liogier H.A. 1985. Descriptive flora of Puerto
seasonal rains. Rico and adjacent islands, Spermatophyta. Vol.
Benefits.—West Indian trema is an important Río Piedras, PR. 352 p.
colonizer of disturbed areas and helps protect the
soil from erosion. It has not been used for, but Little, E.L., Jr. and F.H. Wadsworth. 1964.
might be useful for, site stabilization plantings. Common trees of Puerto Rico and the Virgin
West Indian trema is listed as a nitrogen-fixing Islands. Agriculture Handbook 249. U.S.
species (Winrock International 2002). The wood is Department of Agriculture, Forest Service,
soft and seldom used. The fruits are one of the Washington, DC. 548 p.
most important foods of the endangered Puerto
Rican plain pigeon, Columba inornata wetmorei Long, R.W. and O. Lakela. 1976. A flora of
(Division of Endangered Species 2002). Tropical Florida. University of Miami Press,
Coral Gables, FL. 962 p.
References
Massey, J.R. and J.C. Murphy. 1996. Vascular
Division of Endangered Species. 2002. Puerto plant systematics. Section B. General characters
Rican plain pigeon, Columba inornata and character states: 9. Patterns. http://www.
wetmorei. U.S. Fish and Wildlife Service. ibiblio.org/botnet/glossary/b_ix.html. 2 p.
http://endangered. Fws.gov/i/b/sab40.html. 3 p.
Winrock International. 2002. Nitrogen fixing trees
Howard R.A. 1988. Flora of the Lesser Antilles, and shrubs. http://www.winrock.org/forestry/
Leeward and Windward Islands. factpub/nftlist.htm. 29 p.
776
Trichilia hirta L. broomstick
MELIACEAE
Synonyms: Trichilia spondioides Jacq.

Trichilia wawrana antillana C. DC.
John K. Francis
seeds are covered by a thin, scarlet to orange aril

(Howard 1988, Liogier 1988, Little and
Wadsworth 1964, Stevens and others 2001).
Range.—Broomstick is native to the West Indies

and from Mexico through Central America to
Brazil and Bolivia in South America (Killeen and
others 1993, Little and Wadsworth 1964). It has
been planted in southern Florida (Little and
Wadsworth 1964).
Ecology.—Broomstick is relatively common and

widely scattered through most of its range. It does
not form stands but tends to be scattered in open
forests, roadsides, brushy pastures, and disturbed
areas. Although the species is found in primary
remnants, it is more common in secondary forests,
usually with low basal areas. Broomstick may be
found in Puerto Rican forests that receive from
750 to about 1700 mm of mean annual
600 m. It grows in Nicaragua in deciduous to
General Description.—Broomstick is also known moist forests from 0 to 1,000 m elevation (Stevens
as broomwood, cabo de hacha, garbancillo, guaita, and others 2001). Broomstick is reported in
jojobán, jabillo, molinillo, palo de anastasio, mountain rainforests up to 3,000 m in elevation in
retamo, canalete, conejo colorado, mata piojo, Bolivia (Killeen and others 1993). Although the
carrapeta, and mombin bâtard. It is an evergreen to species grows on a wide range of soil types, it is
semideciduous shrub or small tree ranging in more often seen on rocky and infertile sites where
mature height from 3 to 15 m in different parts of competition is less severe. Broomstick can grow in
its range. The shrubs usually grow with a single partial shade but requires good overhead sunlight
stem in closed stands but often have multiple to flower and fruit.
stems when open-grown or when they have been
damaged. The bark is brown or gray, rough and Reproduction.—Broomstick flowers and fruits
lightly furrowed or scaly. Sapwood is yellowish throughout most of the year (Little and Wadsworth
white, and the heartwood is reddish brown with 1964). The species is reported to flower from
darker streaks. Twigs are green, becoming brown March through May and fruit from January
with age. The plants are supported by a tap and through April in Nicaragua (Stevens and others
extensive lateral root system of flexible, somewhat 2001) and flowers from May through July in
brittle roots with thin, reddish-brown bark. The Panama. The flowers are insect pollinated (Center
alternate leaves are pinnately compound, 20 to 35 for Tropical Forest Science 2002). Plants receiving
cm in length with nine to 21 ovate to lanceolate ample sunlight can produce several hundred seeds
leaflets. The inflorescences are unisexual axillary per year. A collection of fresh fruits from Puerto
racemes. However, these plants are dioecious. The Rico weighed an average of 0.129 + 0.0019 g/fruit.
flowers are small and greenish white. The two- to Surface-dry seeds separated from them weighed an
four-chambered capsules split open to expose one average of 0.0891 + 0.0020 g/seed or 11,000
or two seeds per chamber. The 6- to 8-mm ovular seeds/kg. Sown without pre-treatment on moist
777
blotter paper, 100 percent of them germinated Institution, Washington, DC. http://ctfs.si.edu/
between 17 to 232 days after sowing. Germination webatlas/spanish/tri2hi.html. 2 p.
is hypogeal, and they appear to be recalcitrant
(lose viability if dried beyond a certain point) Howard, R.A. 1988. Flora of the Lesser Antilles,
(author’s observation). In Panama, the seeds are Leeward and Windward Islands.
animal-dispersed (Center for Tropical Forest Dicotyledoneae, Part 1. Vol. 4. Arnold
Science 2002). In Puerto Rico, seedlings are Arboretum, Harvard University, Jamaica Plain,
relatively common but only an occasional MA. 673 p.
individual lasts more than a few weeks or months.
Killeen, T.J., E. García-E., and S.G. Beck, eds.
Growth and Management.—Broomstick in 1993. Guía de árboles de Bolivia. Herbario
Puerto Rico appears to have a moderate growth Nacional de Bolivia and the Missouri Botanical
rate and to live about 10 to 30 years. Management Garden, La Paz, Bolivia and St. Louis, MO.
information is not available. 958 p.
Benefits.—Broomstick adds to secondary forest Liogier, H.A. 1988. Descriptive flora of Puerto
biodiversity, helps protect the soil, and furnishes Rico and adjacent islands, Spermatophyta. Vol.
food and cover for wildlife. The species is used to 2. Editorial de la Universidad de Puerto Rico,
a limited extent for landscaping and is considered Río Piedras, PR. 481 p.
a honey plant. The wood is of medium density
(specific gravity 0.5), moderately hard and Little, E.L., Jr. and F.H. Wadsworth. 1964.
somewhat brash, and is resistant to attack by dry- Common trees of Puerto Rico and the Virgin
wood termites (Criptotermes brevis (Walker)). Islands. Agriculture Handbook 249. U.S.
Another source lists the wood density as 0.48 to Department of Agriculture, Forest Service,
0.96 at 12 to 15 percent moisture content (World Washington, DC. 548 p.
Agroforestry Centre 2002). It has been used for
oars, broom handles, and rustic furniture; today the Núñez-Meléndez, E. 1982. Plantas medicinales de
wood is mainly used for fuel, stakes, and fence Puerto Rico. Editorial de la Universidad de
posts (Little and Wadsworth 1964). An infusion of Puerto Rico, Río Piedras, PR. 498 p.
leaves, flowers, and roots has been used as a
treatment for skin ulcers and as an emetic that has Stevens, W.D., C. Ulloa-U., A. Pool, and O.M.
the unfortunate side effect of causing abortions Montiel, eds. 2001. Flora de Nicaragua.
and bleeding. Young branches were strewn on Monographs in Systematic Botany Vol. 85, No.
floors to control fleas and lice (Núñez Meléndes 2. Missouri Botanical Garden Press, St. Louis,
1982). MO. p. 945-1,910.
References World Agroforestry Centre. 2002. Trichilia hirta.

http://www.worldagroforestrycentre.org/sea/
Center for Tropical Forest Science. 2002. Arboles AgroModels/Dbases/WD/asps/DisplayDetail.asp
del area de Canal de Panama. Smithsonian ?SpecID=3487. 1 p.
778
Trichostigma octandrum (L.) H. Walt. hoop vine
PHYTOLACCACEAE
Synonyms: Rivinia octandra L.

Rivinia humilis var. scandens L.
Rivinia scandens (L.) Miller
Trichostigma rivinoides A. Rich. in Sagra
Villamilla octandra (L.) Hook. f. in Benth. & Hook.
John K. Francis
unpleasant aftertaste. The seeds are 4 to 5 mm in
diameter, black, and shiny (Acevedo-Rodríguez
1985, Howard 1988, Liogier 1985).
Range.—Hoop vine is native to southern Florida,

throughout the West Indies, and from Mexico to
northern Argentina (Anonymous 2002, Howard
1988, Liogier 1985). Although relatively common
in most other areas, the species is listed as
critically endangered in Florida (Gann and others
2001).
Ecology.—Although hoop vine has no specialized

climbing structures, it often ascends into the lower
and mid-crowns of trees. The shrub also covers
windfalls, fences, rocks, and forms mounds of
tangled stems to 2 m high in open areas. Hoop
vine grows on soils of all textures in a wide range
of pH’s derived from both igneous, metamorphic
(including ultramafics) and sedimentary (including
General Description.—Hoop vine, which is also limestone) rocks. In Puerto Rico, the species
known as basket wiss, bejuco de paloma, bejuco occurs in areas receiving from 900 mm to 2400
de nasa, bejuco de palma, murette, liane à mm of precipitation. In Costa Rica, herbarium
barriques, and liane à terre, is a scrambling shrub specimens have been collected from near sea level
that may extend 10 m laterally or 6 m up into the to 1,100 m in elevation (Anonymous 2002). Hoop
crowns of trees and develops stems with diameters vine is moderately intolerant of shade. It invades
of 5 to 15 cm. These plants produce weak early successional forest and continues at least
taproots, extensive lateral roots and many fine into the late secondary forest stage, and also
roots. The stems are gray, cylindrical, smooth, and occurs in remnant forests, especially in tree-fall
relatively stiff. Older plants often have several gaps and on the edges of natural openings. In
branches near the base and then few branches until farms and urban areas, the species grows on
near the growing tips. Young branches of hoop roadsides, fencerows, woodlots, brushy pastures,
vine are cylindrical, slender, and covered with brushy vacant lots, and stream bottom galleries.
elongated lenticels. The leaves are alternate,
elliptic to lanceolate, 5 to 15 cm long and 1.5 to 6 Reproduction.—In Puerto Rico, hoop vine plants
cm broad. The petioles and young leaves are flower from April to August and again in October
reddish. The inflorescence is a many-flowered (Acevedo-Rodríguez 1985). In Costa Rica,
raceme about 15 cm long, often terminal on a herbarium specimens were collected with fruit and
leafless stem. The flowers are about 10 mm flowers from November through June (Instituto
across, greenish-white, becoming red as the fruits Nacional de Biodiversidad 2002). Fruits collected
develop. Hoop vine fruits are elliptic, about 5 to 6 in Puerto Rico weighed an average of 0.178 +
mm in diameter, and purple to black when fully 0.007 g/fruit. Air-dried seeds separated from them
ripe. They have little flavor but leave a slightly averaged 0.0324 + 0.0004 g/seed or 31,000
779
seeds/kg. Sown in commercial potting mix, 32 Anonymous. 2002. Contribución a la flora polínica
percent of these seeds germinated between 10 and de Noreste Argentina. http://www.unne.edu.ar/
71 days after sowing. Birds apparently disperse cyt/2001/6-Biologicas/B-048.pdf. [not paged].
the seeds. Natural seedlings are not common.
Stems layer (root) whenever they are covered with Educational Concerns for Hunger. 2002. Tropical
soil or organic material. vegetables. http://www.echonet.org/eln&herbs/
eln_catalog/veggies.htm. 4 p.
Growth and Management.—Growth of hoop
vine is rapid from sprouts, as much as 3 m/year. Gann, G.D., K.A. Bradley, and S.W.
Established plants appear to live for several Woodmansee. 2001. Status of South Florida
decades, about as long as the trees they use for flora. Institute of Regional Conservation.
support. No management experience has been http://www.Regionalconservation.org/CH2.htm.
published. 10 p.
Benefits.—The young leaves of hoop vine are Howard R.A. 1988. Flora of the Lesser Antilles,
cooked and eaten as a vegetable after the cooking Leeward and Windward Islands.
water is discarded to remove the bitter flavor Dicotyledoneae, Part 1. Vol. 4. Arnold
(Educational Concerns for Hunger 2002). Both the Arboretum, Harvard University, Jamaica Plain,
split stems and the bast fibers of hoop vine are MA. 673 p.
used for making baskets. The stems are also used
for making bent furniture and craft items (Ruiz Instituto Nacional de Biodiversidad. 2002. Lista de
2002). Juice from the fruits leaves a purple stain especimenes de Trichostigma octandrum. http://
on whatever it contacts and may have potential as www.inbio.ac.cr.bims/k03/p13/c045/o0245/f015
a natural stain for cloth and crafts. The species 56/g008574/s026880.htm. 2 p.
contributes to biodiversity and soil stability, and
provides food and cover for wildlife. In herbal Liogier H.A. 1985. Descriptive flora of Puerto
medicine, the vegetative parts are used as a Rico and adjacent islands, Spermatophyta. Vol.
treatment for heart palpitations, and a powder of 1. Editorial de la Universidad de Puerto Rico,
the root bark is applied topically to treat colds and Río Piedras, PR. 352 p.
water retention (Liogier 1990).
References Rico y del Caribe. Iberoamericana de Ediciones,
Puerto Rico. Vol. 1. General Technical Report Ruiz, D.L. 2002. Cuba. Food and Agriculture
SO-58. U.S. Department of Agriculture, Forest Organization of the United Nations. http://
Service, Southern Forest Experiment Station, www.fao.org/docrep/t2354s/t2354s0r.htm. 4 p.
780
Triumfetta semitriloba Jacq. burweed
TILIACEAE
Synonyms: Triumfetta hirta Vahl

Triumfetta ovata DC.
Triumfetta tiliifolia Vahl
Heliocarpus hirta (Vahl) R.O. Williams & Sandw.
John K. Francis
which are located at the leaf axils, have yellow

petals about 5 mm long. The fruits, round burs 6 to
8 mm in diameter with numerous hooked spines,
are borne in groups of two or three. Each fruit has
three compartments and contains three seeds, if
complete (Howard 1989, Liogier 1994, Long and
Lakela 1976).
Range.—Burweed is native to southern Florida,

Bermuda, the West Indies, Mexico, Central
America, and South America (Howard 1989,
Liogier 1994, Long and Lakela 1976). It was
thought that burweed had been recently introduced
to Rapa Nui (Easter Island), but fossil pollen
indicates that it has been present for at least
35,000 years (National Geographic Society 2001).
The species has naturalized in Hawaii, Guam,
Western Samoa and a number of other Pacific
Islands, and in fact is reported to be a pantropical
weed (Pacific Island Ecosystems at Risk 2001).
Ecology.—Burweed grows in pinelands and

hammocks in Florida (Long and Lakela 1976) and
in roadsides, neglected pastures, and wastelands in
Puerto Rico. It grows in sand to clay soils with pH
values from about 5.5 to 8.0. In Puerto Rico,
burweed grows from near sea level to about 700 m
General Description.—Burweed is also known as in elevation. This diverse habitat receives annual
Sacramento bur, black bush, cadillo de perro, precipitation from about 900 to 2200 mm. In
cousin-petit, petit mahot-cousin, mahot-cousin Hawaii, it grows from sea level to 1,067 m in
rouge, tête à nègre, and nedyah (Howard 1989, elevation in areas with annual rainfalls of 760 to
Liogier 1994, Pacific Island Ecosystems at Risk 1500 mm (Haselwood and Motter 1966). The
2001). It is a single-stemmed, short-lived shrub species needs disturbance for establishment. If
that reaches 1 to 2 m in height and 3 cm in stem given an equal start, it competes well with other
diameter. The stems have gray, smooth bark and a herbs, grasses, and shrubs in full sun or partial
woody center. Burweed plants are supported by a shade. Burweed may be found growing in small
weak taproot and many lateral roots. The roots are patches and as single, dispersed plants.
tan colored, flexible, and have a fleshy outer layer
over a woody center. The leaves have petioles Reproduction.—Burweed blooms and fruits
from 0.8 to 9.5 cm long, growing progressively continuously (Long and Lakela 1976), beginning
shorter toward the plant apex. The leaves, 3 to 8 at about 6 months of age. Seed production is
cm long, are broadly ovate to rhombic-ovate, and guaranteed by self-pollination, but cross-
sometimes three-lobed, nearly hairless above, but pollination increases seed set and probably quality
densely stellate-pubescent below. The flowers, (Collevatti and others 1997a). The species has a
781
chromosome count of 2n = 32 (Long and Lakela Collevatti, R.G., L.A.O. Campos, and J.H.
1976). In Brazil, solitary and social bees visit and Schoereder. 1997b. Foraging behaviour of bee
pollinate burweed (Collevatti and others 1997b). It pollinators on the tropical weed Triumfetta
is an abundant seed producer. Fruits collected in semitriloba: departure rules from flower patches.
Puerto Rico averaged 0.0040 + 0.0016 g/fruit. Insectes Sociaux 44(4): 345-352.
Seeds separated from those fruits averaged 0.0039
+ 0.0001 g/seed or 256,000 seeds/kg. Placed on Haselwood, E.L. and G.G. Motter, eds. 1966.
filter paper, 41 percent germinated between 10 and Handbook of Hawaiian weeds. Hawaiian Sugar
119 days from sowing. Germination is epigeal. Planters Association. Honolulu, HI. 479 p.
Seeds are dispersed when they cling to passing
animals. Howard, R.A. 1989. Flora of the Lesser Antilles,
Growth and Management.—Burweed lives Arboretum, Harvard University, Jamaica Plain,
between 1 and 3 years. In seasonally dry habitat, MA. 604 p.
this species behaves as an annual. The plants
grow about 1 m the first year, with growth slowing Liogier, H.A. 1994. Descriptive flora of Puerto
in subsequent years. Because of its weedy nature, Rico and adjacent Islands. Vol. 3. Editorial de la
planting burweed is unlikely, and ill-advised Universidad de Puerto Rico, Río Piedras, PR.
outside its native range. Indeed, control is often 461 p.
needed in croplands and pastures. Treatments
follow normal practice for coarse broadleaf weeds: Long, R.W. and O. Lakela. 1976. A flora of
mechanical cultivation or grubbing and spraying tropical Florida. Banyan Books, Maimi, FL.
with herbicides such as 2,4-D. 962 p.
Benefits.—The stem wood has moderate strength Moanalua Gardens Foundation. 1999. The mystery
and hardness, but because of the plant’s small size of Rapa Nui. Moanalua Gardens Foundation,
the wood is of little value. Fibers from the stem Honolulu, HI. http://gamma.mhpcc.edu/voyage/
bark were once used by the inhabitants of Rapa moanalua/6_rapa.pdf. 20 p.
Nui to make fishing lines (Moanalua Gardens
Foundation 1999). Burweed is reported to have National Geographic Society. 2001. Rapa Nui and
little or no forage value for domestic livestock Sala-y-Gomez subtropical broadleaf forests.
(Haselwood and Motter 1966). National Geographic Society, Washington, DC.
http://nationalgeographic.com/wildworld/
References profiles/terrestrial/oc/oc0111.html. 3 p.
Collevatti, R.G., M.E.C. Amaral, and F.S. Lopes. Pacific Island Ecosystems at Risk. 2001. Invasive
1997a. Role of pollinators in seed set and a test plant species: Triumfetta semitriloba Jacaquin,
of pollen limitation hypothesis in the tropical Tiliaceae. http://www.hear.org/pier/trsem.htm.
weed Triumfetta semitriloba (Tiliaceae). Revista 2 p.
de Biologia Tropical 45(4): 1,401-1,407.
782
Turbina corymbosa (L.) Raf. Christmas vine
CONVOLVULACEAE
Synonyms: Rivea corymbosa (L.) Hall.f.

Convolvulus corymbosus L.
Convolvulus sidaefolius HBK.
Ipomoea sidaefolia (HBK.) Choisy
Ipomaea corymbosa (L.) Roth.
John K. Francis
Australia (Pacific Island Ecosystems at Risk

2002), and some parts of the Old World Tropics
(Howard 1989).
Ecology.—Christmas vine grows as single plants

or matted patches of vines in secondary forest
openings, old fields, neglected pastures, road
sides, stream banks, and vacant lots. In Florida, it
may be found in hammocks and brushy areas
(Long and Lakela 1971). It demands full or nearly
full sunlight and will not grow under a closed
forest canopy. Christmas vine will grow on most
well-drained soils. It is reported to bloom more
profusely on “red” and limey soils (Woman’s
Club of Havana 1952). The species has been
observed by the author growing in areas of Puerto
Rico that receive from 1000 to 2000 mm of annual
General Description.—Christmas vine, also precipitation.
known as Christmas wreath, aguinaldo blanco,
corona de novia, ololiuqui, and badoh, is a woody Reproduction.—A collection of fruits from
vine that may extend 5 m or more laterally and Puerto Rico weighed an average of 0.0389 +
into the crowns of trees and shrubs. Older stems 0.0004 g/fruit. Seeds separated from those fruits
reach about 2.5 cm in diameter. The gray, three- weighed an average of 0.0265 + 0.0002 g/seed or
sided stems have many lenticels and may be 38,000 seeds/kg. Sown in commercial potting mix,
grooved. The young stems are cylindrical and 95 percent of the seeds germinated between 33
tough. The foliage is concentrated on current and 48 days after sowing (author’s observation).
year’s growth. The leaves, with slender petioles, Well-established plants root whenever vines touch
have cordate blades 5 to 8 cm long with an the soil. Natural seedlings tend to be widely
elongated point. The inflorescences are corymbose scattered. Besides spreading by wind, water, and
cymes that arise from leaf axils. The 2.5- to 3-cm lateral vine extension, seeds of the species are now
corolla is trumpet shaped, white with a red or widely sold, exchanged, and grown for the
purple throat and green or greenish gray radiating narcotic the seeds contain.
stripes. The ellipsoidal capsule has three long and
three short sepals that cause it to spin and glide Growth and Management.—Seedlings of
laterally when released. Each capsule contains one Christmas vine grow slowly at first and develop a
brown, pubescent seed (Acevedo-Rodríguez and strong tap and lateral root system. Leaders of well-
Woodbury 1985, Howard 1989, Liogier 1995). established plants may extend 2 m or more per
year. The plants can be pruned back to the woody
Range.—Christmas vine is native to the West stems or thinned in the spring after blooming is
Indies, Mexico through Central America, and the complete (Woman’s Club of Havana 1952). No
tropical portion of South America (Liogier 1995). specific recommendations are given for
It has naturalized in Florida (Correll and Johnston controlling the species when it grows as a weed in
1970), Hawaii, a few other Pacific Islands, agricultural settings. Cutting the vines near the
783
ground, and spraying the resulting sprouts with New Orleans, LA. 331 p.
broad-leaf weed killers, should be effective.
Correll, D.S. and M.C. Johnston. 1970. Manuel of
Benefits.—Christmas vine is sometimes grown as the vascular plants of Texas. Texas Research
an ornamental for the clusters of white, scented Foundation. Renner, TX. 1,881 p.
flowers it produces during the early winter. The
nectar gathered from its flowers makes one of the Howard, R.A. 1989. Flora of the Lesser Antilles,
finest honeys (Woman’s Club of Havana 1952). Leeward and Windward Islands.
Extracts from the seeds are used as an analgesic in Dicotyledoneae. Part 3. Vol. 6. Arnold
herbal medicine (Schultes and Hoffmann 1992). Arboretum, Harvard University, Jamaica Plain,
MA. 658 p.
Narcotic Properties.—The seeds of Christmas
vine were valued as a sacred hallucinogen by Liogier, H.A. 1995. Descriptive flora of Puerto
Chinantec, Mazatec, Mixtec, Zapotec, and other Rico and adjacent islands. Vol. 4. Editorial de la
groups in Southern Mexico in Pre-Columbian Universidad de Puerto Rico, Río Piedras, PR.
times and are still cultivated and used today as 617 p.
aids in divination and witchcraft. It was
administered by grinding about 13 seeds, adding Long, R.W. and O. Lakela. 1971. A flora of
water, filtering, and drinking the filtrate in a quiet, Tropical Florida. University of Miami Press,
secluded place. Hallucinations follow that last Coral Gables, FL. 962 p.
about 3 hours, sometimes with aftereffects. The
active ingredients are the ergoline alkaloids, Pacific Island Ecosystems at Risk. 2002. Invasive
lysergic acid amide, and lysergic acid plant species: Ardisia elliptica Thunberg,
hydroxyethylamide that are closely related to LSD Myrsinaceae. http://www.hear.org/pier/
(Schultes and Hoffmann 1992). arell.htm. 2 p.
References Shultes, R.E. and A. Hoffmann. 1992. Plants of the

gods. Healing Arts Press, Rochester, VT. 192 p.
Puerto Rico. Vol. 1. General Technical Report Woman’s Club of Havana. 1952. Flowering plants
SO-58. U.S. Department of Agriculture, Forest from Cuban Gardens. Criterion Books, New
Service, Southern Forest Experiment Station, York. 365 p.
784
Urena lobata L. Caesar weed
MALVACEAE
Synonyms: Urena americana L. f.

Urena grandiflora DC.
Urena trilobata Vell.
Urena lobata L.
Urena diversifolia Schumach.
(and others: Institute of Systematic Botany 2003).
John K. Francis
ivory-colored, tough and flexible roots. Grayish-

green, discolorous, alternate leaves are stellate-
pubescent both above and below, ovate, angulate
or shallowly lobbed, 1 to 12 by 1 to 12 cm, and
have serrate margins. Axillary flowers are usually
solitary and have five pink petals. The flowers are
about 1 cm broad. The fruits are 8- to 10-mm
globose capsules that break into five fine-barbed
(glochidate) mericarps. (Howard 1989, Liogier
1994, Stevens and others 2001)
Range.—The original range of Caesar weed is

probably Asiatic. Today it grows throughout moist
tropic and subtropical regions including American
Samoa, Florida, Guam, Hawaii, Louisiana, Puerto
Rico, and the U.S. Virgin Islands (Natural
Resources Conservation Service 2003, Pacific
Island Ecosystems at Risk 2003).
Ecology.—Caesar weed readily invades disturbed

areas, especially poorly managed pastures,
scarified and eroded areas, and perennial crop
plantations but is much less of a problem in annual
crops. The species does not compete well in tall
grass swards and brushlands and does not grow
under forest canopies. Ceasar weed grows on a
wide variety of soils of varying fertility derived
General Description.—Caesar weed is also from most parent materials. It tolerates salt spray
known as hibiscus bur, aramina, pink Chinese and a moderate amount of salt in the soil but does
burr, bur mallow, grand cousin, cadillo, carrapicho not grow in saturated soils. The species ranges to
do mata, malva, mahot cousin, cousin petit, cousin elevations of 1,500 m (Pacific Island Ecosystems
rouge, jut africain, cooze mahot, dadangsi, and at Risk 2003). In Puerto Rico, it occurs in areas
mautofu. It is a subshrub 0.6 to 3 m in height and that receive from about 1400 to 3000 mm of mean
up to 7 cm in basal diameter. The species usually annual precipitation, forming thickets in favorable
has a single stem emerging from the ground but sites.
normally produces several stems or major
branches low on the stem and many branches Reproduction.—Caesar weed flowers and fruits
throughout. The nearly smooth bark is tough and throughout the year (Stevens and others 2001). A
fibrous, brown on the outside and green within. Its collection of seeds from Puerto Rico averaged
pale yellow wood is of medium density. Plants are 0.0239 + 0.0003 or 41,800 seeds/kg. Placed on
supported by a tap and lateral root system of tan or moist blotter paper without pretreatment, just 3
785
percent germinated in 9 months. Germination is References
epigeal (author’s observation). Sulfuric acid
scarification was tested as a means of breaking Bautista, L.M.A. 2000. Inquiry into the anti-
dormancy of seed from Sierra Leone. The best inflammatory activity of the syrup from the
treatment (18 molar solution for 90 minutes) gave glycosides of the leaves of kulutkulutan (Urena
96 percent germination as compared with distilled lobata, Linn., family Malvaceae). Centro
water that gave 4 percent germination (Harris Escolar Universitario, Mandiola, Philippines.
1986). Seeds are dispersed by clinging to fur and http:// www.ceu.edu.ph/research1.htm. 3 p.
clothing. Seedlings are common in disturbed sites
near seed sources. Câmara de Comércio e Indústria Portugal-Angola.
2003. Caracterização económico social de
Growth and Management.—Caesar weed grows Angola. http://www.cciportugal-angola.pt/
rapidly and can reach 0.5 to 2 m by the end of the caract.htm. [not paged].
first year. In Puerto Rico, it can live 2 years,
usually dying back to midheight after the first Fagundes, M.H. 2003. Sementes de juta e malva:
growing season. It is not known whether the algumas observações. Companhia Nacional de
shrubs sprout from the roots multiple times. A Abastecimento, Brazil. http://www.conab.gov.
fiber crop yielding 1,800 kg/ha is ready to harvest br/politica_agricola/Conjunturasemanal/Especiai
after 6 to 7 months and seed crops of 300 to 500 s/SEMENTES%20JUTA%MALVA.doc. 10 p.
kg/ha are ready after 7 or 8 months (Fagundes
2003). Fiber crops are established with seed. FAO. 2003. Food composition table for use in
Because of the aggressive nature of the species, Africa. Food and Agriculture Organization of
wildland plantings are not recommended. The the United Nations, Rome. http://www.fao.
author knows of no published specific control org/docrep/003/x6877e/X687705.htm#ch5. [not
measures. paged].
Benefits and Detriments.—Caesar weed Forest Research Institute of Malaysia. 2003. Plants
colonizes disturbed areas and helps to protect the information: Urena lobata Griff., pulut-pulut,
soil while furnishing cover for wildlife. It has Malvaceae. Forest Research Institute of
attractive flowers and contributes to aesthetics of Malaysia, Kuala Lumpur, Malaysia. http://www.
areas it has colonized. Tens of thousands of tons of frim.gov.my/tu/Urena.htm. 1 p.
a jute-like fiber from Caesar weed called aramina
fiber and Congo jute are produced in Brazil Freedman, R. 1998. Famine foods: Malvaceae.
(Fagundes 2003) and Africa (Câmara de Comércio Perdue University, Perdue, IN. http://www.hort.
e Indústria Portugal-Angola 2003). Various perdue.edu/newcrop/faminefoods/ff_families/M
extracts of leaves and roots are used in herbal ALVACEAE.html. 3 p.
medicine to treat such diverse ailments as colic,
malaria, gonorrhea, fever, wounds, toothache, and Harris, P.J.C. 1986. Dormancy of Urena lobata L.
rheumatism (Forest Research Institute of Malaysia seeds. I. Development of sulphuric acid
2003). A semipurified glycoside obtained from scarification techniques. Ghana Journal of
Caesar weed leaves was 86 percent as effective an Agriculatural Science 14-19: 79-84.
anti-inflammatory as aspirin in rats (Bautista
2000). The leaves and flowers are eaten as famine Howard, R.A. 1989. Flora of the Lesser Antilles,
food in Africa (Freedman 1998). Raw leaves are Leeward and Windward Islands. Vol. 5. Arnold
reported to contain 81.8 percent moisture, 54 cal, Arboretum, Harvard University, Jamaica Plain,
3.2 g of protein, 0.1 g fat, 12.8 g carbohydrates, MA. 604 p.
1.8 g fiber, and 2.1 g ash, 558 mg calcium, and 67
mg of phosphorous per 100 g (FAO 2003). Institute of Systematic Botany. 2003. Atlas of
However, the plant is little browsed by cattle and Florida vascular plants. University of South
can become a severe weed in pastures and Florida, Tampa, FL. http://plantatlas.usf.
plantations. Burs that collect on clothing and in edu/synonyms.asp?plantID=1364&genus=Urena
animal fur are a nuisance. &species=lobata. [not paged].
786
Liogier, H.A. 1994. Descriptive flora of Puerto Pacific Island Ecosystems at Risk. 2003. Urena
Rico and adjacent Islands. Vol. 3. Editorial de la lobata L., Malvaceae. http://www.hear.org/
Universidad de Puerto Rico, Río Piedras, PR. Pier/urlob.htm. 3 p.
461 p.
Natural Resources Conservation Service. 2003. Montiel, eds. 2001. Flora de Nicaragua.
Plant profile: Urena lobata L., Caesarweed. Monographs in Systematic Botany Vol. 85, No.
http://plants.usda.gov/cgi_bin/plant_profile.cgi? 2. Missouri Botanical Garden, St. Louis, MO.
symbol=URLO. 5 p. p. 945-1,910.
787
Urera baccifera (L.) Gaud. ortiga brava
URTICACEAE
Synonyms: none
John K. Francis
poorly drained soils of all textures are colonized.

Being moderately shade intolerant, disturbance is
generally required for the species to get started,
and it disappears after the forest understory
becomes dark. A high importance value of ortiga
brava in a Brazilian forest and an unbalanced
diameter distribution was used to infer past
disturbance (Bertoni and others 1988).
Reproduction.—Minute male and female flowers

are borne in pink or purple, branched clusters on
the stem on different plants (dioecious). The fruits
are white or pinkish, spongy and watery, with one
greenish-black seed per fruit (Little and others
General Description.—Ortiga brava means “bad 1974). A Puerto Rican collection of seeds
nettle” in Spanish. The species is known by a host averaged 4.36 + 1.17 g each (Francis and
of other common names, among which are: Rodríguez 1993) or 500,000 seeds/kg. They began
stinging nettle, ortiga, pringamoza, mala mujer, germinating in 26 days with a final germination of
chichicaste, nigua, guaritoto, ishanga, manman 49 percent (Francis and Rodríguez 1993). Ortiga
guêpes, and urtiga bronca (Little and others 1974). brava built up to high populations in a fire-
Ortiga brava is a weak-stemmed woody shrub that disturbed forest in the Atlantic forest in southern
sometimes becomes a small tree. Individual plants Brazil through a high recruitment rate and low
may have one or several stems, branching near the mortality (Nascimento and others 1999). The seeds
ground. Probably the most recognizable feature of are disbursed by birds.
the plant is the presence of many sharp, stinging
spines on the leaves, branches, and stem. The Growth and Management.—Individual stems of
alternate leaves are 12.5 to 25 cm in length and 7.5 ortiga brava may grow for 5 years or more and
to 12.5 cm in width, coarse-toothed with thin reach 5 m in height. Heights of 1.5 to 2 m are more
blades and stout petioles. The stem bark is smooth, typical. Ortiga brava stems cut during site
light gray with green-over-white inner bark, and management activities quickly sprout and regain
thick greenish pith. There are raised leaf scars and their former height. Whether sprouting from the
lenticels on the stems and branches. rootstalk occurs after senescence and death of
individual stems is not known.
Range.—Ortiga brava ranges throughout the
Greater and Lesser Antilles, Trinidad, and Tobago Benefits and Detriments.—In Central America,
and on the mainland from Mexico to Brazil, ortiga brava has been planted in hedges and fences
Argentina, Bolivia, and Peru (Little and others to make an almost impenetrable barrier (Little and
1974). The species is not known to have others 1974). Although not generally sought after
naturalized outside its native range. for food under normal conditions, ortiga brava is
one of the first plants to fruit after hurricanes, and,
Ecology.—Ortiga brava grows in the higher as such, is an important factor in the survival of
rainfall portion of the moist forest (>1600 mm frugiverous birds (personal communication with
precipitation), especially along drainages, and Joseph Wunderle, IITF, Río Piedras, PR). The
throughout the wet and rain forests (2000 to 4000 stems of ortiga brava were used by Aztec and
mm of precipitation). Alluvial, coluvial, and Otomi Indians in Mexico for making paper (Hagen
residual soils from acidic and calcareous rocks 1943). The plant is considered a weed in shaded
make suitable habitat. Well-drained and somewhat coffee plantations. (Francis and Rodríguiz 1993).
788
All parts of the plant are covered with the spine- Bertoni, J.E. de A., F.R. Martins, J.L. de Moraes,
like hairs that cause considerable pain when they and G.J. Shepherd. 1988. Composicão floristica
penetrate the skin. Normal clothing does not e estrutura fitossociologica do parque de
protect against the stinging effect. However, the Vacununga, Santa Rita do Passa Quatro, Sao
pain is usually short-lived, lasting from a few Paulo, gleba Praxides. Boletim Tecnico do
seconds to a few minutes. Although no permanent Instituto Florestal, Sao Paulo 42: 149-170.
injury results, some people may experience rash or
blisters; swelling, fever, and ulcers can occur in Francis, J.K. and A. Rodríguez. 1993. Seeds of
extreme cases (Allen 1943). The plant has long Puerto Rican trees and shrubs: second
been used in folk medicine. Recent laboratory tests installment. Research Note SO-374. U.S.
using rats have shown anti-inflamatory and Department of Agriculture, Forest Service,
analgesic activity of aqueous extracts of ortiga Southern Forest Experiment Station, New
brava (Badilla and others 1999). Amerindians in Orleans, LA. 5 p.
Costa Rica chastised themselves with branches of
ortiga brava in a toughening ritual (Badilla and von Hagen, V.W. 1943. Mexican paper-making
others 1999). plants. Journal of the New York Botanical
Garden 44: 1-10.
References
Allen, P.H. 1943. Poisonous and injurious plants Wadsworth. 1974. Trees of Puerto Rico and the
of Panama. Journal of Tropical Medicine Virgin Islands. Vol. 2. Agriculture Handbook
23(suppl.): 3-76. 449. U.S. Department of Agriculture,
Badilla, B., G. Mora, A.J. Lapa, and J.A. Silva
Emim. 1999. Anti-inflammatory activity of Nascimento, H.E.M., A. da S. Dias, A.A.J.
Urera baccifera (Urticaceae) in Sprague-Dawley Tabanez, and V.M. Viana. 1999. Tree
rats. http://www.ots.duke.edu/tropibiojnl/claris/ population and dynamics of a semidecidous
47-2/badilla.html. seasonal forest fragment in the Piracicaba
region, Sao Paulo State, Brazil. Revista
Brasileira de Biologia 59(2): 329-342.
789
Vernonia albicaulis Pers. Santa María
ASTERACEAE
Synonyms: Vernonia vhliana Less.

Vernonia thomae Benth.
Cacalia thomae (Benth.) Kuntze
Vernonia longifolia vahliana (Less.) Urban
Vernonia longifolia sintenisii Urban
Vernonia sintenisii (Urban) Gleason
Lepidaploa glabra (Willd.) H. Rob.
John K. Francis
1989). A second subspecies, longistylis Keeley, a

taller and more robust plant, has been described on
Guadeloupe (Howard 1989). Santa María is not
elsewhere.
Ecology.—Santa María is moderately intolerant to

intermediate in tolerance of shade. It grows at
forest edges, small openings, under low basal-area
stands, and on difficult terrain with few trees. This
may be in secondary or remnant forests. The
species is most common in limestone areas from
near sea level to 600 m elevation (Liogier 1997)
but also occurs in terrain underlain by igneous
rocks. Mean annual precipitation varies from
about 900 to 2400 mm. The soils are well-drained,
usually clayey or loamy, and have pH’s from
about 5.0 to 8.0.
General Description.—Santa María, also known Reproduction.—Seed production is abundant. A

as tabac à jacot, is a shrub to 2 m tall and stem sample of seeds from Puerto Rico averaged 3.9
basal diameter to 1 cm. The ribbed, woody, gland- million per kg. They were sown on the surface of
dotted, hairy stems are upright or arching. Several wet peat but failed to germinate. The seeds are
stems may arise from the root crown. The lateral dispersed by the wind. Seedlings are common to
root system has brown bark and a moderate widely scattered.
amount of fine roots. The alternate leaves are
evenly dispersed on the branches and have Growth and Management.—Santa María has a
petioles 1 to 11 mm long. The blades are oblong to moderate growth rate. Individual stems live about
elliptic, rounded at the apex and pointed at the 2 to 4 years, and plants may live much longer by
base, and have entire to weakly sinuate margins. resprouting from the roots. No planting or
Inflorescences are cymes of heads of 10 to 23 management experience has been published.
florets having lavender to white corollas 5 to 10 Because Santa María appears to benefit from
mm long. The fruits (achenes) are 3 mm long with disturbance to existing forest, thinnings and other
pale brown to white pappus with bristles 0.5 to 1 silvicultural activities may help maintain it.
mm long (Howard 1989, Liogier 1997).
Benefits.— Santa María contributes to the
Range.—Santa María is native to Puerto Rico, the diversity and beauty of the forest, helps protect the
Virgin Islands, Anguilla, St. Martin, St. Barts, soil, and furnishes cover for wildlife.
Antigua, Saba, St. Eustatius, St. Kitts, Monserrat,
Guadeloupe, La Désidade, and Dominica (Howard
790
Howard, R. A. 1989. Flora of the Lesser Antilles, Universidad de Puerto Rico, San Juan, PR.
Leeward and Windward Islands. Vol. 6. Arnold 436 p.
MA. 658 p.
791
Vernonia borinquensis Urban Puerto Rican ironweed
ASTERACEAE
Synonyms: Vernonia borinquensis var. resinosa Gleason

Vernonia borinquensis var. hirsuta Gleason
John K. Francis
stiff, have a short petiole, and blades in an ovate to

lanceolate form. They have a rounded base and a
pointed tip. Leaves remain on the twigs 1 year and
twigs remain green for an additional year.
Inflorescences are leafy cymes of sessile
campanulate heads, 5.5 to 8 mm long, of 13 to 22
florets. The corolla is white or pale lavender. At
maturity, the fruits (achenes) are 1 to 2 mm long,
ribbed, and have a brown pappus 5 to 6.5 mm long
(Acevedo-Rodríguez 1985, Liogier 1997).
Range.—Puerto Rico ironweed is endemic to

Puerto Rico and mainly found in the Cordillera
Central (Acevedo-Rodríguez 1985). The species is
elsewhere.
Ecology.—Puerto Rico ironweed grows on well-

drained soils with a wide range of textures derived
from most parent materials. Fertility is usually
moderate to good, and pH’s range from about 5 to
7. Exposed subsoils are not colonized. Mean
annual precipitation ranges from about 900 to
2400 mm and at elevations from a few to about
800 m above sea level. Puerto Rico ironweed has
an intermediate tolerance to shade. It grows in
openings but is often found under low to medium
basal-area forest canopies. It is resistant to grazing
and/or avoided by cattle enough to survive in
light- to moderately-grazed semiforested range
and brushy pastures. The species may be found in
both remnant and secondary forests.
General Description.—No local name is known
for this species in Puerto Rico. The Natural
Reproduction.—Puerto Rico ironweed flowers
Resources and Conservation Service (2002) has
from November to July and fruits from February
assigned it the name Puerto Rico ironweed. The
to September (Acevedo-Rodríguez 1985).
plant is a vine-like scrambling shrub reaching 5 to
Presumably, flowers are insect pollinated. In a
10 m of lateral extension and sometimes as much
sample of seeds collected near Calley, Puerto
as 3 m into the crowns of trees and shrubs.
Rico, about 3 percent of the seeds were filled.
Younger plants often have a single stem, but older
Sorted by examination under a microscope,
plants may form clumps and mats. The root system
apparently filled seeds weighed an average of
consists mainly of slender lateral roots and
0.00098 g/seed or about 1 million seeds/kg. After
abundant, fleshy, fine roots. The principal
being placed on moist filter paper, 76 percent
branches are cylindrical, about 0.5 cm thick, and
germinated within 24 days. The seeds are wind-
covered with gray bark. Fine branches are densely
dispersed. Although seed production can be heavy,
hairy, striated, and have a zig-zag form. Leaves,
established seedlings are relatively uncommon.
concentrated near the branch ends, are somewhat
792
Disturbance is probably necessary for seedling makes it somewhat more difficult to walk through.
establishment. The plants layer (root) whenever
stems come in contact with the soil. References
Growth and Management.—Puerto Rico Acevedo-Rodríguez, P. 1985. Los bejucos de

ironweed grows at a moderate rate. Established Puerto Rico. Vol. 1. General Technical Report
plants extend stems about 0.5 m/year. Although SO-58. U.S. Department of Agriculture, Forest
individual stems probably do not live more than 2 Service, Southern Forest Experiment Station,
to 4 years, plants may live much longer by New Orleans, LA. 331 p.
sprouting and layering. No management
experience has been published; thinning the forest Liogier, H.A. 1997. Descriptive flora of Puerto
overstory with accompanied soil disturbance is Rico and adjacent islands. Vol. 5. Editorial de la
recommended to encourage regeneration. Universidad de Puerto Rico, Río Piedras, PR.
436 p.
Benefits.—Puerto Rico ironweed helps protect the
soil and furnishes cover for wildlife. It adds a Natural Resources Conservation Service. 2002.
pretty aspect to trailsides and forest edges but Plants database. http://plants.usda.gov/cgi_
gives the understory a “closed” appearance and bin?topics.chi?earl=dl_all.html. [not paged].
793
Vernonia proctori Urbatsch Proctor’s vernonia
ASTERACEAE
Synonyms: none
John K. Francis
is endangered and has an estimated population of

950 individuals in an area of a few hectares (U.S.
Fish and Wildlife Service 2002).
Ecology.—Proctor’s vernonia grows in exposed

rock crevices and in shallow cherty clay loam soil
up to 30 cm deep that has developed over
weathered siliceous rock. Mean annual
precipitation is about 900 mm/year with a fall wet
season and spring dry season. Humidity is high
and breezes are almost continuous. Elevations
where the population grows ranges from 270 to
300 m above sea level (U.S. Fish and Wildlife
Service 2002). Proctor’s vernonia is intolerant of
shade. It grows in the open or in broken stands of
low trees, but not under closed-canopy forest.
Bare ground and 3 to 4 weeks of almost daily
precipitation is probably necessary for
establishment. The endemic area was once
severely grazed and cut over for fuelwood, but has
been much less disturbed in recent years.
Reproduction.—Proctor’s vernonia flowers and

General Description.—Proctor’s vernonia (a fruits during April and May (U.S. Fish and
name assigned by the author) is an upright gray- Wildlife Service 2002). The plant produces seed in
green shrub to 1.5 m in height and 1 cm in basal abundance. A small quantity of seeds collected by
stem diameter. Older plants may have as many as the author averaged 0.00038 g/seed or about 2.6
50 stems arising from the root crown. The plants million seeds/kg. Placed on moist filter paper, 13
have many lateral and abundant fine roots. percent germinated within 15 days. The seeds are
Branches are greenish gray, turning brown or gray dispersed by the wind. Seedlings are infrequent.
with age. The foliage is crowded on new twig
growth. Alternate leaves have short petioles and Growth and Management.—Proctor’s vernonia
blades that are broadly elliptic to suborbicular, 1.2 appears to grow slowly as a seedling and take
to 3.5 cm long, and rounded at the tip. They are many years to attain large plant size. Individual
dark green above and white beneath and have a stems arising from sprouts grow about 20 cm/year.
densely ciliate margin. The inflorescences are Although individual stems last only a few years,
terminal, sessile clusters of usually three heads the plant, renewing itself with new sprouts,
with 16 to 22 florets. The corollas are purple or appears to live for many years. Propagation
blue. At maturity, the heads contain 2- to 3-mm techniques have not been published. Complete
long achenes with a white pappus (Liogier 1997, protection of the Cerro Mariquita site from
Miner-Solá 1999, U.S. Fish and Wildlife Service development, grazing, and fire is critical.
2002).
Benefits.—Proctor’s vernonia is a pretty plant and
Range.—Proctor’s vernonia is endemic to Puerto certainly contributes to the aesthetics of the area
Rico and found only at the summit of Cerro where it grows. It protects the soil and furnishes
Mariquita in the range of hills known as Sierra cover for wildlife.
Bermeja in southwestern Puerto Rico. The species
794
References de extinción en Puerto Rico. Puerto Rico
Ecológico Vol. 3. First Book Publishing of
Liogier, H.A. 1997. Descriptive flora of Puerto Puerto Rico, San Juan, PR. 91 p.
Universidad de Puerto Rico, Río Piedras, PR. U.S. Fish and Wildlife Service. 2002. Species
436 p. accounts: Vernonia proctorii. http://endangered.
fws.gov/i/q/saqa9.html. 2 p.
Miner-Solá, E. 1999. Arboles y plantas en peligro
795
Vernonia sericea L.C. Rich. long-shoot
ASTERACEAE
Synonyms: Lepidaploa phyllostachya Cass.

Vernonia berteriana DC.
Conya portoricensis Bert. ex DC.
Vernonia arborescens lessingiana Griseb.
Cacalia sericea (L.C. Rich.) Kuntze
Cacalia arborescens lessingiana (Griseb.) Kuntze
John K. Francis
leaves are linear to elliptic, 2.5 to 10 cm long, with

an entire edge, and pointed at the tip and
somewhat rounded at the base. The inflorescences
are elongated scorpioid cymes with leafy bracts
and contain lavender to white, sessile flower heads
about 6 mm long with 11 to 18 florets. They
produce achenes 2 mm long tipped with a brown
pappus about 4 mm long (author’s observation,
Liogier 1997).
Range.—Long-shoot is native to Cuba,

Hispaniola, Puerto Rico, and the Virgin Islands
(Liogier 1997). It is not known to have been
Ecology.—Long-shoot grows on a variety of soil

types, mostly of medium to heavy texture with
pH’s from 5 to 7 that are derived from
metamorphic parent materials. These are well
drained to somewhat poorly drained soils in areas
that receive from about 850 to 2200 mm of mean
annual precipitation at elevations from near sea
level to about 700 m. Long-shoot is moderately
intolerant of shade and grows in openings and
under the canopy of low basal-area forest. It
competes well with forbs and grasses if the
vegetation is not too heavy. It is browsed by cattle,
but not preferentially, so that it can survive and
reproduce in lightly and moderately grazed
General Description.—Long-shoot, also known
pastures and range. Long-shoot grows on
as escobilla, huye-que-te-coge, tapa caminos, and
abandoned agricultural land, secondary forest,
yerba socialista, is an upright shrub usually 1 to
stream banks, open remnant forests, roadsides, and
1.5 m (occasionally 2 m) in height and 1 cm in
fence rows.
basal stem diameter. It usually has a single main
stem, although older and previously damaged
Reproduction.—Long-shoot blooms seasonally,
plants sometimes have multiple stems branching
probably at the latter part of the rainy season
from the root collar or from near the base. The
(November through January). The abundant seeds
wood of the stems is stiff and hard. The plant is
produced are dispersed by wind during the dry
supported by a tap and lateral root system of stiff,
season (March and April). A collection of seeds
orange to tan roots. Smaller roots are tough and
from Puerto Rico weighed an average of 0.000309
flexible. Branches and twigs are slender. The
796
g/seed or 3.2 million seeds/kg. Placed on moist to the aesthetic beauty of forests, and furnishes
filter paper, 69 percent germinated over the course cover for wildlife. The species is one of the nectar
of 2 months, beginning 7 days after sowing. sources for the butterfly, Eurema leuce antillarum
Hall (Torres Bauzá 1999).
Growth and Management.—Long-shoot grows
between 0.5 and 1 m in the first year from sprouts. References
The growth rate of seedlings is undoubtedly much
slower. Individual stems live about 3 years, but Liogier, H.A. 1997. Descriptive flora of Puerto
plants sprout and live through a second, perhaps Rico and adjacent islands. Vol. 5. Editorial de la
several, cycles of tops. No propagation or Universidad de Puerto Rico, Río Piedras, PR.
management experience has been published. 436 p.
Presumably, conditions for healthy populations
can be created by maintaining broken or low Torres Bauzá, J.A. 1999. Ciclo de vida de Eurema
basal-area forest stands. leuce antillarum Hall (Lepidoptera: Pieridae) en
Benefits.—Long-shoot helps protect the soil, adds 4): 195-200.
797
Waltheria indica L. sleepy morning
STERCULIACEAE
Synonyms: Waltheria americana L.

Waltheria elliptica Cav.
John K. Francis
stems and leaves are covered with a gray, velvety

pubescence. The alternate leaves are narrowly
ovate or oblong with a rounded to subcordate
base, irregularly serrate edges, and a rounded to
acute tip. The petioles are 0.5 to 3.3 cm long and
the blades are 2 to 12 cm long and 1 to 7 cm
broad. Axillary inflorescences are usually dense
glomerules that contain fragrant, yellow to orange
flowers. Each 2-mm capsule holds one tiny, black,
obovoid seed (Howard 1989, Liogier 1994,
Tropilab, Inc. 2001).
Range.—Sleepy morning now grows throughout

the tropics and warmer subtropics (Liogier 1994).
It apparently naturalized in Hawaii soon after the
arrival of nonnative colonists (Haselwood and
Motter 1966). Howard (1989) indicates that the
species is native to the New World where it occurs
from Florida and Texas to Brazil (Dick 2001,
Nelson 1996, Texas A & M University 2001).
Ecology.—Sleepy morning grows in disturbed dry

and well drained, moist habitat. In Puerto Rico,
the species occupies areas that receive from 750 to
about 1800 mm of annual precipitation from near
sea level to more than 400 m of elevation. The
species grows on sites up to 1,220 m in elevation
in Hawaii (University of Hawaii 2001). It
colonized a wide variety of soils in areas with
General Description.—Sleepy morning is also igneous, metamorphic (including ultramafics) and
known as velvet leaf, marsh-mallow, monkey sedimentary (including limestone) rocks. The
bush, boater bush, leather coat, buff coat, basora species may be found on old fields, construction
prieta, malvavisco, hierba de soldado, guimauve, sites, roadsides, burned forest and grasslands, and
mauve-gris, moto-branco, hioloa, fulutafu, kafaki, stream overflow areas in Puerto Rico. In Florida, it
and many other names (Haselwood and Motter grows in open pinelands, hammocks, and
1966, Howard 1989, Liogier 1994, Burkill 2000). disturbed sites (Long and Lakela 1976). It is
It is a short-lived shrub or subshrub sometimes intolerant of shade and will not survive under a
reaching 2 m in height and 2 cm in stem diameter. closed tree canopy and cannot compete with grass
Sleepy morning develops a weak taproot, robust in dense swards. It withstands drought, salt spray,
lateral roots, and abundant fine roots. The roots and mildly salty soils.
are brown and flexible. This shrub usually has a
single, strong stem emerging from the ground, but Reproduction.—Sleepy morning plants begin
frequently branches near the ground. Sleepy flowering at about 6 months of age and bloom
morning usually has an upright and somewhat more or less continuously for the rest of their
branchy form. However, in some environments, it lives. A collection of seeds made in Puerto Rico
may grow in a semiprostrate habit. The young averaged 0.0013 g/seed or 764,000 seeds/kg.
798
These seeds were sown without any pretreatment ethnobotanical dictionary. http://www.ars-
on filter paper and after 16 weeks yielded 13 grin.gov/duke/dictionary/tico/w.html. 4 p.
percent germination. Reproduction is by seeds.
The seeds are disbursed by water, agricultural Burkill, H.M. 2000. The useful plants of West
equipment (Sánchez and Uranga 1993), and Tropical Africa. Royal Botanic Gardens, Kew,
grazing animals. Seedlings are relatively common UK. 686 p.
in disturbed habitat.
Centro de Desarrollo de Agronegocios. 2000. Guia
Growth and Management.—Sleepy morning prictica de manejo de plagas y enfermedades del
plants in Puerto Rico live for 1 or 2 and chile. CDA Boletín Técnico de Producción 13.
occasionally 3 years. Death usually occurs during http://www.hondurasag.org/fintrac-cda/pubs/
the dry season. Perennial growth is more likely in Prod_13_Esp.pdf. 4 p.
continuously moist habitat. Stands of sleepy
morning in agricultural plantations can be Dick, C. 2001. Amazon plant list. Ecotour
controlled by cultivation and probably by Expeditions, Inc. http://www.naturetours.com/
broadleaf herbicides. Am_lispl.html. 8 p.
Benefits and Detriments.—In the Turks and Faftine, O., A. Alage, and J.P. Muir. 2001.
Caicos Islands, sleepy morning is used to make an Characterization of forage selected by cattle on
herb tea (Wood 2001). The plant produces a fiber communal range in Manhição, Mozambique.
that was formerly used for making cords, sacking, http://stephenville.tamu.edu/~jmuir/faftine.htm.
padding, and sandals (Guzmán 1975). Durawhite, 6 p.
an extract of sleepy morning, is used in a
commercial cosmetic for its ability to inhibit Guzmán, D.J. 1975. Especies útiles de la flora
melanin synthesis and whiten the skin (Janssen Salvadoreña. Ministerio de Educación,
Cosmeceutical Care 2001). The plant contains Dirección de Publiciones, San Salvador, El
steroid derivatives and alkaloids of the adouetine Salvador. 703 p.
group that perhaps make it physiologically active.
Various extracts are used as standard febrifugal, Haselwood, E.L. and G.G. Motter. 1966.
purgative, emollient, tonic, analgesic, and Handbook of Hawaiian weeds. Experiment
astringent herbal medicines in Africa (Burkill Station/Hawaiian Sugar Planter’s Association.
2000). In Hawaii, the root is chewed to relieve Honolulu, HI. 479 p.
sore throat (Neal 1965). Stems are used as a chew
stick, and extracts of the plant are used as an eye Howard, R.A. 1989. Flora of the Lesser Antilles,
bath and a remedy for hemoptysis in Panama Leeward and Windward Islands. Vol. 5. Arnold
(Agricultural Research Service 2001). Seeds are Arboretum, Harvard University, Jamaica Plain,
sold commercially, and the species is cultivated in MA. 604 p.
gardens as a medicinal plant. The plant is browsed
by all types of livestock, especially when young Janssen Cosmeceutical Care. 2001. Supreme
(Burkill 2000). Sleepy morning forage in a secrets. http://www.janssen-beauty.com/
Mozambique valley during the rainy season supreme.htm. 11 p.
contained 6.4 percent crude protein, 0.12 percent
phosphorus, and 0.51 percent calcium (Faftine and Lastres, L.. 2000. Guia practica de manejo de
others 2001). Sleepy morning is considered a plagas y enfermedades del chile. CDA Technical
weed in much of its range, but it is seldom Production Bulletin 13. Centro de Desarrollo de
aggressive enough to be a major problem Agronegocios. http://www/jpmdirasag/org/
(Sánchez and Uranga 1993). It is host for a fintrac-cda/pubs/Prod_13_Esp.pdf. 4 p.
number of insects harmful to agricultural crops
(Centro de Desarrollo de Agronegocios 2000, Liogier, H.A. 1994. Descriptive flora of Puerto
Lastres 2000, Sánchez and Uranga 1993). Rico and adjacent Islands. Vol. 3. Editorial de la
References 461 p.
Agricultural Research Service. 2001. Tico Long, R.W. and O. Lakela. 1976. A flora of
799
Tropical Florida. Banyon Books, Miami, FL. plant checklist: Sterculiaceae. Texas A & M
962 p. University, Department of Botany.
http://www.csdl.tamu.edu/FLORA/ftc/dft/
Neal, M.C. 1965. In gardens of Hawaii. Special ftc_str.htm. 1 p.
Press, Honolulu, HI. 924 p. Tropilab, Inc. 2001. Waltheria indica L.
http://www.tropilab.com/sleepy_waltheria.html.
Nelson, G. 1996. The shrubs and woody vines of 2 p.
391 p. University of Hawaii. 2001. Plants of Hawaii
Volcanoes National Park.
Sánchez, P. and H. Uranga. 1993. Plantas http://www.botany.edu/b308/bigisland/species/
indeseables de importancia economica el los waltheria/waltheria.htm. 3 p.
cultivos tropicales. Editorisal Científico-
Técnica. La Habana, Cuba. 166 p. Wood, K.M. 2001. Roadside wildflowers.
http://www.timespub.tc/Natural%History/
Texas A & M University. 2001. Texas vascular Archive/Summer2001/wildflowers.htm. 2 p.
800
Wedelia reticulata DC. manzanilla de monte
ASTERACEAE
Synonyms: Seruneum reticulatum (DC.) Kuntze

Stemmodontia reticulata (DC.) Cook & Collins
John K. Francis
long (author’s observations, Liogier 1997).
Range.—Manzanilla de monte is native to Puerto

Rico and the island of Hispaniola (Liogier 1997).
It is not known to have been planted or naturalized
elsewhere.
Ecology.—Manzanilla de monte grows on soils of

all textures, pH’s from mildly acid to mildly
alkaline, over parent materials of igneous,
sedimentary (including limestone), and
metamorphic (including ultramafic) rocks. Soils
are usually rocky or shallow over bedrock. The
species grows from near sea level to elevations of
600 m or more in Puerto Rico (Breckon 2001,
Francis and others 1998). Annual rainfall ranges
from about 900 to 2000 mm. Manzanilla de monte
tolerates salt spray and mild salinity in the soil. It
is drought tolerant. During drought, the leaves wilt
but usually do not fall off. Manzanilla de monte is
intolerant of shade. It is open-grown or grows in
small openings in low forest. The species is hardy
and competes well with grass and herbs of similar
height. Well-established stands become clumps
General Description.—Manzanilla de monte is and thickets that tend to shade out plants under
an evergreen shrub to 1.5 m in height and 1.5 cm them.
in basal diameter. Adult plants have few-to-many,
slender, brown, brittle, woody stems arising from Reproduction.—Manzanilla de monte blooms
the root crown. The stems have a 1-mm pith and and fruits continually. Seed production can be
discernible annual rings. The root system is abundant although unknown insects sometimes
dominated by stiff and brittle, slender, dark tan- destroy most of the seeds. Other than short-
over-green, wide spreading laterals. Also, distance transport by wind, gravity, and water,
adventitious roots grow from the bases of specific means of dispersal are unknown.
branches. Branching is often bifurcate or Seedlings are relatively uncommon. Suckers at the
trifurcate, sometimes forming a tangled structure edge of clumps are common, and plants
in older plants. However, foliage tends to occasionally layer.
concentrate at the periphery of the crown. Leaves
are opposite, narrowly oblong ovate to oblong Growth and Management.—Manzanilla de
lanceolate, short-pointed at the tip and rounded to monte is slow growing in natural environments.
subcordate at the base, 3 to 10.5 cm long, with a Sprouts grow 10 to 30 cm/year. Individual stems
0.5- to 1.0-cm petiole and fine serrate edges. The persist about 4 to 10 years. Through continually
leaves are green to yellow-green. Compound suckering and layering, plants can last almost
flowers are terminal, solitary, or occasionally in indefinitely.
two’s or three’s. The yellow involucral bracts
(“petals”) are 4 to 7 mm in length and numerous Benefits.—Manzanilla de monte contributes to the
florets. The fruits are four-sided achenes 3 mm aesthetics of natural areas where it grows, helps
801
protect the soil, and furnishes cover and possibly Francis, J.K., S. Alemañy, H.A. Liogier, and G.R.
food for wildlife. Although somewhat rough in Proctor. 1998. The flora of Cañón de San
appearance during dry periods, it has pretty Cristóbal, Puerto Rico. General Technical
flowers and should make a fine ornamental for Report IITF-4. U.S. Department of Agriculture,
natural landscaping, especially in coastal areas and Forest Service, International Institute of Tropical
rocky terrain. Forestry, Río Piedras, PR. 37 p.

Breckon, G.J. 2001. Preliminary check list for Universidad de Puerto Rico, Río Piedras, PR.
Maricao Forest. http://www.uprm. 436 p.
edu/biology/profs/breckon/herbarium/FLORAM
ARICAO.htm. [not paged].
802
Ziziphus obtusifolia (Hook. ex Torr. & Gray) Gray graythorn
RHAMNACEAE
Synonyms: Condalia obtusifolia (Hook.) Weberbauer

Condalia lycioides (Gray) Weberbauer
Zizyphus lycioides Gray
James E. Nellessen
var. lycioides; Condalia obtusifolia (Hook. ex

Torr. & Gray) Weberb.; Zizyphus lycioides Gray;
and Condaliopsis lycioides (Gray) Suess. (Allred
2002, Kartesz 1994).
Range.—This species occurs from just a few

counties in southwestern Oklahoma (McCoy
1981), portions of central and western Texas,
through southern New Mexico to northwestern
Arizona and southeastern California and south
well into Mexico. It occurs at a wide elevational
range from about 300 m to 1,525 m. Populations
in the more western portions of its range are
variety canescens (Kearney and others 1951).
General Description.—Graythorn is also known
as lote bush, southwestern condalia, prairie bush,
Ecology.—Graythorn will grow in a variety of
or thornbush. This shrub reaches heights of 2 to 3
situations from silty and sandy floodplains and
m but is often shorter. Stems are grayish green due
washes to gravelly slopes and plains in desert
to a whitish or grayish wax-like bloom, which
grasslands and scrublands. This species tends to be
helps to distinguish Ziziphus from Condalia. The
diffusely distributed within its local habitat, but
stems are robust, grooved, highly branched,
may also form thickets. This species is considered
although terminal branches are short, and bear
a characteristic component of Chihuahuan desert
stout spine-tips. The stem branching pattern gives
scrub and a diagnostic climax indicator species of
this shrub an irregular shape. Side branches tend to
desert grassland (Dick-Peddie 1993). In a northern
come out at right angles from the main axis. The
Texas study, although graythorn had a minimal
leaves are alternate, simple, generally less than 2.5
influence on overall grass cover, there appeared to
cm long, range from 0.4 to 0.7 cm in width, are
be differences in species distributions around
ovate, oblong, or elliptic with three main veins.
graythorn bushes (Foster and others 1984).
Leaf shape is variable which has lead to some
Buffalo grass [Buchloe dactyloides (Nutt.)
confusion in identification. Leaves are dropped
Engelm.] was more abundant near graythorn than
during dry periods (Carter 1997, Correll and
in shrub free zones, while Texas winter- or
Johnston 1979, Great Plains Flora Association
needlegrass (Stipa leucotricha Trin. & Rupr.)
1986, Kearney and others 1951, Martin and
cover and standing crop was greater away from
Hutchins 1980-81).
graythorn. A study conducted in a savanna
parkland habitat in southern Texas showed that
Taxonomy.—This shrub has a somewhat
graythorn is a deciduous shrub with an average
complicated proliferation of taxonomic names.
leaf lifespan measured at 66 days (Nelson and
Two varieties are recognized: Ziziphus obtusifolia
others 2002). Supplemental watering had no
var. canescens (Gray) M.C. Johnston; and the
detectable effect on leaf longevity. Root infection
typical variety Z. obtusifolia var. obtusifolia.
with symbiotic strains of Frankia and Rhizobium
Synonyms for variety canescens are: Condalia
did not occur on four species of Rhamnaceae from
lycioides (Gray) Weberb. var. canescens (Gray)
southern Texas, including graythorn, when
Trel.; Zizyphus lycioides Gray var. canescens
planted in pots in the laboratory, in a 1:1 native
Gray; and Condaliopsis lycioides var. canescens
soil (with symbionts) to vermiculite substrate
(Gray) Suess. Synonyms for variety obtusifolia
(Zitzer and others 1996).
are: Condalia lycioides (Gray) Weberb., including
803
Reproduction.—Flowers of graythorn are small, potassium, and iron were at levels to meet adult
inconspicuous, and occur in axillary cymes of two goat requirements during all seasons. Phosphorus
to seven flowers. The five-petalled flowers are was low in all seasons except the spring. Native
greenish yellow. Graythorn may bloom at various Americans have used the plant for medicinal
times during the growing season, from May purposes. Pima Indians have steeped the roots in
through September, depending on the availability water and applied them to sore eyes. Seri Indians
of moisture. The fruit is a dark blue to black have powdered the roots for application to skin
globose drupe, 6 to 10 mm long, covered with a and scalp sores. Solutions made from roots also
whitish bloom, and matures from July into early served as a soap substitute (Bowers and Wignall
October. Seeds are nondormant, but the seed coats 1993, Epple 1995, Kearney and others 1951).
are impermeable to water, hence are classified as
physically dormant (Baskin and Baskin 2001). References
Physically dormant seeds need some type of
scarification of the seed coat to initiate water Allred, K.W. 2002. A Working Index of New
penetration and germination. Optimal germination Mexico Vascular Plant Names. Available on a
occurs at 25 °C. Both light and prechilling New Mexico State University web page:
maximized the percent germination (Speer and http://web.nmsu.edu/~kallred/herbweb. [not
Wright 1981, Young and Young 1992). Graythorn paged].
seeds from the lower Rio Grande valley in
southern Texas did not need pretreatment for Baskin, C.C. and J. M. Baskin. 2001. Seeds:
germination (Vora 1989). Ecology, Biogeography, and Evolution of
Dormancy, and Germination. Academic Press,
Growth and Management.—Graythorn stem San Diego, CA. 666 p.
regeneration generally occurs just below the cut or
removed portion. When stems are removed to the Bowers, J.E. and B. Wignall. 1993. Shrubs and
ground line, regeneration will occur from the root Trees of the Southwest Deserts. Southwest
crown, and stems can be regenerated from roots, Parks and Monuments Association. Tucson,
but generally only when the root crown itself is AZ. 140 p.
removed (Flinn and others 1992). Graythorn may
demonstrate rapid growth when resprouting from Carter, J.L. 1997. Trees and Shrubs of New
cut stumps or during periods of good moisture but Mexico. Johnson Books, Boulder, CO. 534 p.
is generally a slow growing shrub. Although this
species may on occasion become locally dense and Correll, D.S. and M.C. Johnston. 1970. Manual
form thickets, it is generally a species that is of the Vascular Plants of Texas. Texas Research
diffusely distributed within its habitat, such that Foundation, Renner, TX. 1,881 p.
management should not be a problem.
Benefits.—The fruits are readily eaten by birds Vegetation: Past, Present and Future.
such as Gambel's quail and white-winged doves. University of New Mexico Press, Albuquerque,
Graythorn was among several shrubs in a northern NM. 244 p.
Mexico study testing the preferred browse
composition and digestibility of foliage. Graythorn Epple, A.O. 1995. A Field Guide to the Plants of
was grouped with species having a high crude Arizona. Falcon Publishing Inc., Helena, MT.
protein content, lower cell wall content, and 347 p.
smoother or waxy leaf surfaces with the absence of
trichomes. Range goats and white-tailed deer Flinn, R.C., C.J. Scifres, and S.R. Archer. 1992.
included graythorn as an important part of their Variation in basal sprouting in co-occurring
diets (Ramirez and others 1997). In another shrubs: implications for stand dynamics.
nutrient study in northern Mexico (Ramirez and Journal of Vegetation Science 3(1): 125-128.
others 2001) graythorn was among several shrubs
studied for seasonal mineral levels. General Foster, M.A., C.J. Scifres, and P.W. Jacoby, Jr.
mineral concentrations were higher in the spring 1984. Herbaceous vegetation - lotebush
and summer, but only calcium, magnesium, (Ziziphus obtusifolia var. obtusifolia)
804
interactions in north Texas USA. Journal of Ramirez, R.G., G.F.W. Haenlein, and M.A.
Range Management 37(4): 317-320. Nunez-Gonzalez. 2001. Seasonal variation of
macro and trace mineral contents in 14 browse
Great Plains Flora Association. 1986. Flora of the species that grow in northeastern Mexico. Small
Great Plains. University Press of Kansas, KS. Ruminant Research 39(2): 153-159.
1,392 p.
Ramirez, R. G., J.L. Pineiro-Hernandez, and R.K.
Kartesz, J.T. 1994. A Synonymized Checklist of Maiti. 1997. Nutritional profile and leaf surface
the Vascular Flora of the United States, Canada, structure of some native shrubs consumed by
and Greenland. Vol. 1, 2nd Edition. Biota of small ruminants in semiarid regions of
North America Program of the North Carolina northeastern Mexico. Journal of Applied
Botanical Garden. Timber Press, Portland, OR. Animal Research 11(2): 145-156.
622 p.
Speer, E.R. and H.A. Wright. 1981. Germination
Kearney, T.H., R. Peebles, and Collaborators. requirements of lotebush (Ziziphus obtusifolia
1951 with 1960 supplement. Arizona Flora. var. obtusifolia). Journal of Range Management
University of California Press, Berkeley, CA. 34: 365-368.
1,085 p.
Vora, R.S. 1989. Seed germination characteristics
Martin, W.C. and C.E. Hutchins. 1980-1981 of selected native plants of the lower Rio
(reprinted 2001). A Flora of New Mexico. Vol. Grande valley Texas, USA. Journal of Range
1. Bishen Singh Mahendra Pal Singh, India and Management 42(1): 36-40.
Koeltz Scientific Books, Germany. 1,276 p.
Young, J.A. and C.G. Young. 1992. Seeds of
McCoy, D. 1981. Roadside Trees and Shrubs of Woody Plants in North America. Dioscorides
Oklahoma. University of Oklahoma Press, Press, Portland, OR. 407 p.
Norman, OK. 116 p.
Zitzer, S.F., S.R. Archer, and T.W. Boutton. 1996.
Nelson, J.A., P.W. Barnes, and S. Archer. 2002. Spatial variability in the potential for symbiotic
Leaf demography and growth responses to N-2 fixation by woody plants in a subtropical
altered resource availability in woody plants of savanna ecosystem. Journal of Applied Ecology
contrasting leaf habit in a subtropical savanna. 33(5): 1125-1136.
Plant Ecology 160(2): 193-205.
805
Zuckia brandegei (Gray) Welsh & Stutz ex Welsh siltbush
CHENOPODIACEAE
Synonyms: Grayia brandegei Gray var. brandegei

Grayia brandegei Gray var. plummeri Stutz & Sanderson
Zuckia arizonica Standley
Nancy L. Shaw, Rosemary L. Pendleton, and Emerenciana G. Hurd
dorsiventrally compressed and mostly horizontal

with six-keeled bracts. Flowers of all three
varieties develop in small clusters in bract axils.
Staminate flowers consist of four or five stamens
and a four- or five-lobed perianth. Pistillate
flowers contain a single pistil enveloped by two
united bracts. Embryos are well developed.
Range.—Siltbush is a narrowly distributed

edaphic endemic, largely restricted to the Colorado
River drainage. G. b. var. brandegei occurs
primarily in south-central Utah and northeastern
Arizona. G. b. var. plummeri grows in isolated
populations in northeastern Utah, south-central
Wyoming, western Colorado, and northwestern
New Mexico. G. b. var. arizonica occurs in
scattered populations from northern Arizona to
northeastern Utah (Goodrich and Neese 1986,
Shaw and others 2001, Welsh and others 1987).
Ecology.—Siltbush grows in isolated monotypic

populations on shale outcrops in desert shrub and
lower juniper communities at elevations of 1,280
to 2,240 m (Goodrich and Neese 1986). Soils on
these outcrops are poorly developed, fine textured,
and characterized by moderate levels of soluble
General Description.—Siltbush or spineless
salts, and high levels of exchangeable sodium and
hopsage is a deciduous shrub or subshrub ranging
potassium. Micronutrient levels are generally high.
from 0.1 to 0.8 m in height (Goodrich and Neese
Siltbush tissues accumulate sodium and
1986). Stems of the current year are thornless and
magnesium, but potassium levels and the calcium
ascending to erect, branching from a persistent,
and magnesium ratio are low (Pendleton and
woody base. Leaves are alternate, gray-scurfy and
others 1996).
entire to lobed. Overwintering leaf buds are
prominent, axillary, and globose (Welsh and others
Reproduction.—All siltbush varieties are
1987). There are three varieties (Welsh 2000).
monoecious and heterodichogamous (Pendleton
Diploid (2x = 18) G. brandegei (Gray) Welsh &
and others 1988) with individual plants either
Stutz var. brandegei is small with narrow, linear
protogynous (producing pistillate, then staminate
leaves. Tetraploid (4x = 36) G. b. (Gray) Welsh &
flowers) or protandrous (producing staminate, then
Stutz var. plummeri (Stutz & Sanderson) Dorn
pistillate flowers). Flowering occurs in late spring
(Plummer’s siltbush) is larger with ovate to
or summer and fruits ripen in mid to late summer
lanceolate leaves. Utricles of both varieties are
or fall (Pendleton and others 1988). Protogynous
obcompressed and vertical with two-winged
plants generally produce more seed, but
samara-like bracts (see illustration). Utricles of the
protandrous plants may be equally productive in
third variety, G. b. (Gray) Welsh & Stutz var.
years when precipitation is high and seed
arizonica (Standley) Welsh (Arizona siltbush), are
806
predation is low (Pendleton and others 2000). Between population differences in dormancy
Wind and gravity disperse the fruits over time; and response to temperature. In: Proceedings,
some fruits generally remain on the plant until late Symposium on cheatgrass invasion, shrub die-
winter. There are about 625,000 seeds/kg off, and other aspects of shrub biology and
(Pendleton and others 2000). Germination management. U.S. Department of Agriculture,
experiments conducted with seeds of Z. b. var. Forest Service, General Technical Report INT-
brandegei and Z. b. var. plummeri showed that 276. Intermountain Research Station, Ogden,
seeds of warm winter populations germinated over UT. p. 187-292.
a wide range of constant temperatures (15 to 30
o
C) (Meyer and Pendleton 1990). Seeds of cold- Pendleton, R.L., D.C. Freeman, E.D. McArthur
winter populations were dormant at fall and winter and S.C. Sanderson. 2000. Gender specialization
temperatures and required exposure to overwinter in heterodichogamous Grayia brandegei.
chilling to promote germination. Germination American Journal of Botany. 87:508-516.
generally increased with duration of wet
prechilling at 1 oC for up to 8 weeks, dry Pendleton, R.L., E.D. McArthur, D.C. Freeman,
afterripening for up to 14 months, or bract removal and A.C. Blauer. 1988. Heterodichogamy in
(Meyer and Pendleton 1990). Germination is Grayia brandegei (Chenopodiaceae): Report
epigeal. from a new family. American Journal of Botany.
75: 267-274.
Growth and Management.—Few data are
available. Siltbush is a stress-tolerant species Pendleton, R.L., S.D. Nelson, and R.L. Rodriguez.
capable of establishing on steep, eroded sites 1996. Do soil factors determine the distribution
where the combination of salinity, aridity, and fine of spineless hopsage (Grayia brandegei)? In:
soils preclude establishment of other species Proceedings, Symposium on shrubland
(Pendleton and others 1996). Seedling growth rate ecosystem dynamics in a changing climate.
and palatability of mature plants are moderate General Technical Report INT-GTR-338. U.S.
(personal communication with Steve Monsen, Department of Agriculture, Forest Service,
USDA-FS-RMRS, Provo, UT, and Howard Stutz, Intermountain Research Station, Ogden, UT. p.
Brigham Young University, Provo, UT). Plants 205-209.
may attract rodents, other small animals, and deer.
Shaw, N.L., R.L. Pendleton, and E.G. Hurd. 2001.
Benefits.—Siltbush is a potential revegetation and Zuckia brandegei (Gray) Welsh & Stutz ex
soil stabilization species for disturbances resulting Welsh. In: F.T. Bonner and R.G. Nisley, eds.
from mining and other human activities on sites Woody Plant Seed Manual. U.S. Department of
where it is native. Few other species will grow on Agriculture, Forest Service, Washington, D.C.
these outcrops (Pendleton and others 1996). http://wpsm.net/index. html. 8 p.
References Welsh, S.L. 2000. Zuckia. Flora of North America.

http://hua.huh.harvard.edu/FNA/zuckia.ed2.html
Goodrich, S. and E. Neese. 1986. Uinta Basin 3 p.
flora. U.S. Department of Agriculture, Forest
Service, Intermountain Region, Ashley National Welsh, S.L., N.D. Atwood, S. Goodrich, and L.C.
Forest and USDI Bureau of Land Management, Higgins, eds. 1987. A Utah flora. Great Basin
Vernal District, Ogden, UT. 320 p. Naturalist Memoirs 9. Brigham Young
Meyer, S.E. and R.L. Pendleton. 1990. Seed
germination biology of spineless hopsage:
807
Glossary of Technical Terms Used in this Volume
Achene—an indehiscent one-seeded fruit type common in Asteraceae.

Alfisols—order of moderately leached soils, developed under forests, with relatively high native fertility,
and having well developed subsurface horizons with clay accumulation.
Allele—one of an array of genes possible at a certain position (locus) on a given chromosome.
Allelopathy—the negative influence of plants on each other generally understood to arise from released
metabolic products.
Alpine—sites in mountains above the timberline or vegetation characteristic of those areas.
Anthesis—the time at which a flower comes into full bloom.
Arborescent—having a treelike habit.
Aridisols—order of soils of arid regions that contain free CaCO3 and have at least some subsurface horizon
development.
Aril—an appendage or outer covering of some seeds, often fleshy or pulpy.
Armed—having spines or thorns.
Awn—bristle-like appendage of a flower and fruit usually seen in grasses.
Axillary—borne in the axils (or inside the angles of attachment) of leaves.
Beach strand—narrow strip of vegetation between a beach and inland formations.
Berry—fleshy fruit containing one or usually more seeds.
Biomass—the dry weight of the plants and animals (usually just plants in vegetation studies) in a unit area.
Bonsai—a tree or shrub grown in miniature form through cultural practices; the term is sometimes used for
naturally miniature trees or shrubs.
Bract—modified leaves associated with flowers.
Calyx—the whorl of green leaf- or bract-like sepals.
Campanulate—bell shaped.
Canescent—hoary, gray-colored pubescence.
Canopy—the more or less continuous cover of tree crowns in a forest.
Capsule—a fruit arising from a multicarpellate ovary that usually dries and splits open.
Cartaceous—papery texture (usually used to describe leaves).
Catkin—a drooping elongated inflorescence of unisexual flowers typical of willows.
Cespitose shrubs—very short shrubs with a cushion-like habit.
Chaining—dragging a heavy chain between two bulldozers to kill unwanted brush and small trees and to
scarify the soil.
Clambering—habit of growing over obstacles or into low vegetation without tendrils or other specialized
climbing structures.
Claviculate—club shaped.
Clearcut—silvicultural harvesting method in which all the trees are removed at one time and usually
unwanted material is felled as well.
Climax—terminal stage of ecological succession in which the vegetation association remains stable over a
relatively long period.
Codominant—a crown class in a tree or shrub canopy in which a plant shares the main canopy level with
many other individuals of similar height.
Colluvium (colluvial soil)—fractured rock and soil accumulated at the foot of a slope or the soil developed
from it.
Coppice—reproduce by sprouts after being cut or damaged.
Cordate—heart shaped.
Coriaceous—thickened and leathery.
Corolla—the whorl or aggregate of the petals of a flower.
Corymb—a flat-topped inflorescence in which the outer flowers open first.
Crenate—margin with rounded, broad teeth.
Cyme—a more or less flat-topped inflorescence in which the central flowers open first
d.b.h.—stem diameter at breast (1.37 m above the ground) height.
Decumbent—branching habit that is flattened, growing along the ground with ends turned up.
Dioecious—having staminate (male) and pistillate (female) flowers on different plants.
Diploid—having two sets of chromosomes in a somatic nuclei.
808
Disjunct population—a population isolated from the main part of the range.
Diurnal—occurring (blooming) during the day.
Dominant—a crown class in forest or shrub canopy in which the individual is taller than the general
canopy level; may also refer to species with a preponderance of numbers or biomass.
Drupe—fleshy, one-seeded fruit whose seed is encased in a stony endocarp.
Ecotone—a zone of transition between adjacent plant communities.
Ecotype—a population within a species that is genetically adapted to a certain habitat distinct from that
which is general for the species.
Edaphic—pertaining to the soil and its ecological relationships.
Endosperm—starchy or oily tissue in many seeds.
Entire—without teeth or indentations on the margin.
Epigeal—germination and emergence in which the cotyledons raise above the soil surface.
Escapes—introduced populations now established and competing in the wild and spreading beyond the
original point of introduction.
Excessively drained soil—water runs off, infiltrates, or evaporates rapidly so that the soil tends to be
droughty even with ample rainfall.
Floret—individual, usually minute, flower of a compound flower.
Fruticose—being woody or shrub-like.
Galleries—small patches of vegetation (usually trees or shrubs) in protected locations distinct from
surrounding vegetation.
Germinative capacity—the percentage of seeds that germinate during the normal period of germination.
Glabrous—not hairy; hairless or nearly so.
Glaucous—with a waxy bloom or whitish substance that rubs off.
Habit—the growth form or appearance of a plant.
Head—a compact, multiple inflorescence, often disk-shaped.
Hammock—a fertile area in Florida and Southeastern United States that is elevated above the surrounding
terrain.
Hypanthium—a floral tube formed by the fusion of the basal portions of the sepals, petals, and stamens
from which the rest of the floral parts arise.
Hypogeal—germination and emergence in which the cotyledons remain below the soil surface.
Inceptisols—order of soils that are young, frequently rocky, and have a poorly developed profile.
Intermediate—a crown class of tree or shrub canopies in which the individual plant crown is within and
does not quite rise to the level of the top of the canopy.
Lanate—soft, woolly pubescence.
Layering—rooting at a point on a stem where the stem comes in contact with the soil or with artificial
treatment called air layering.
Legume—one to multiple-seeded fruit (pod) common in the Fabaceae family.
Lenticels—lenticular or diamond-shaped dots, pits, or corky protrusions on stems.
Litter—organic debris on the surface of the soil or the uppermost layer of the forest floor organic horizon.
Maceration—the process of soaking and gently grinding fruits to remove the pulp from seeds.
Mesic—moist soil or environment; neither excessively wet nor dry.
Mine spoil—dirt and rock left after a mining operation.
Monoecious—having staminate (male) and pistillate (female) flowers on the same plant.
Naturalized—naturally reproduced progeny of horticultural plants or wildland plantings outside their
native range.
Nerve—vein, usually used in relation to leaves.
Nut—a one-seeded fruit enclosed in an involucre (diminuative=nutlet).
Old growth forest—characterized by old, usually large, and often decadent trees.
Open forest—a forest characterized by low basal area, a large amount of space between crowns, and a lot
of sunlight reaching the forest floor.
Open-grown—growing without significant competition from plants of similar size or taller.
Panicle—an inflorescence with indeterminate branching, the primary axis bearing branched secondary axes
and pedicellate flowers.
Pappus—hairs attached to small seeds that together function as a wing.
Perfect flower—a flower with both stamens (male parts) and carpels (female parts).
Pericarp—the wall of a ripened ovary (fruit).
809
pH—a measure of soil acidity or alkalinity; values above 7.0 are alkaline and values below are acid.
Phenotype—a plant as it exists; the product of the interaction of genetics and environment.
Pioneer—shrub or other plant that establishes on or has the ability to colonize newly disturbed habitat.
Pith—spongy central cylinder of parenchyma tissue in stems.
Ploidy—the degree of repetition of the basic chromosome number.
Poorly drained soil—water runs off or evaporates so slowly that the soil remains wet for a large part of the
year.
Provenance—the original geographic source of a plant’s genetic material.
Puberulent—covered with soft, very fine hairs.
Raceme—an inflorescence with indeterminate single axis and pedicellate flowers.
Rachis—the axis of a pinnately compound leaf.
Remnant (forest)—lightly to moderately disturbed pieces or stands of the original forest.
Reticulate—with a net-like structure.
Rhizome—modified underground stem.
Riparian vegetation—plant associations and stand types growing along streams.
Samara—a one-seeded, winged fruit.
Savannah—tropical or subtropical grassland with scattered trees; usually maintained by periodic fires.
Scabrous—covered with scattered, short, stiff hairs.
Scandent—vine-like habit.
Scarify—to perforate or make a seed coat permeable by mechanical or chemical means.
Scrub—vegetation consisting of shrubs and sometimes trees stunted because of poor site conditions.
Secondary forest—forest that has grown up after clearing or lesser disturbance.
Selfing—pollination of a plant with its own pollen.
Seral—transitional as a site or vegetation association moves toward a climax state.
Serrulate—finely serrate.
Sessile—connected directly to the stem or other structure without a stalk.
Shade intolerant—requiring full or nearly full sunlight to survive, grow and reproduce.
Shade tolerant—the ability to survive, grow, and reproduce in the shade of taller vegetation, as under a
tree canopy.
Skeletal soil—substrate in which stones form the structure of the soil with sand, silt, and clay filling the
voids between.
Slash—the cut stems and limbs of trees and brush, or to cut unwanted trees and brush leaving it in place.
Soil texture—classes defining the proportion of sand, silt, and clay making up a soil.
Spike—indeterminate inflorescence of sessile flowers arranged linearly along a central axis.
Stamen—male or pollen-producing part of the flower.
Stipule—a small leaf-like structure found at the base of some leaf petioles, usually present in pairs.
Stolon—a stem or shoot that bends to the ground and takes root after contact.
Stratify—breaking the physiological dormancy of certain seeds with a period of cold or warm incubation.
Succulent—class of plants with fleshy leaves and usually stems that store water in their tissues and usually
have special metabolic pathways (crassulan acid metabolism).
Sucker—root sprouts or to reproduce by root sprouts; sometimes used to mean an epichormic branch.
Suffruticose—perennial plant with stems that are woody near their base.
Suppress—inhibit the growth (and often eventually kill) by shading and competition for water and
nutrients.
Synonym—an equivalent but superseded species name.
Testa—the outer coat of a seed.
Ultisols—a family of highly weathered soils found in subtropical and tropical environments.
Ultramafic rocks—metamorphic rocks (including serpentine) characterized by low percentages of silica
and elevated percentages of iron and magnesium.
Variety—a subdivision of a species that has one or more inherited morphological characteristics.
Well-drained soil—water runs off, infiltrates, or evaporates readily but not rapidly.
Xeric garden—a garden planted with drought-resistant vegetation that is not irrigated.
810
References Consulted
Burns, R.M. and B.H. Honkala, eds. 1990. Silvics of North America. Vol. 2. Agriculture Handbook 654.
U.S. Department of Agriculture, Forest Service, Washington, DC. 877 p.
Editorial staff. 1984. Webster’s ninth new collegiate dictionary. Marriam-Webster Inc., Publishers,
Springfield, MA. 1,563 p.
Lawrence, G.H. 1955. An introduction to plant taxonomy. The Macmillan Company, New York. 179 p.
Soil Survey Staff. 1951. Soil survey manual. Agriculture Handbook 18. Agriculture Research
Administration, U.S. Department of Agriculture, Washington, DC. 503 p.
811
Common Name Index
Common name page Common name page
‘a’ali’i 293 amourette-grand-bois 487
‘a’ali’i ku makani 293 amourette-rivière 489
‘a’ali’i-ku ma kua 293 amouretto 489
abeyelo 235 angelica tree 40
abrojo Colorado 412 angel’s trumpet 36, 286
abuta 212 angel’s whisper 299
abutilon espinoso 540 anguila 327
acacia bush 21 añil 386
acacia courant 289 añil de pasto 386
acacia savane 289 añil falso 252
acacia zarza 19 anisette 567
aceituna Americana 117 anisillo 564, 567
aceituno 154 annatto 112
acequia willow 664 antelope bitterbrush 610
achiote 112 antelope brush 610
afata 714 antelope buckbrush 610
aguacero 127 antibala 714
aguedita 561 Antilles velvetshrub 570
ague tree 681 Apache-plume 337
aguinaldo blanco 783 aperta ruão 564
agüiste 489 applebush 361
‘aheahea 190 aramina 785
aile à ravet 764 arabo carbonero 323
aji 147 arazá-puitá 597
aji de gallina 144 árbol de navidad 352, 617
aji picante 147 arbre de soie 136
akalakala 658 arctic willow 660
akeake 293 arête-boeuf 529
akulikuli-kai 107 argythamnia 45
alcaparro 696 Arizona queen-of-the-night 181
alcotá 212 Armenian blackberry 652
alcotán 567 Aroeira pimenteira 686
alderleaf mountain-mahogany 178 aroma 14
alfilerillo 459 aromo 21
algodón de seda 137 arrowleaf sida 714
algorobo 580 arrowwood 379
alkali sagebrush 66 ash barberry 464
allamanda 36 Asian bittersweet 161
aloes vert 350 ati popa’a 43
alpine dryad 295 atiu 43
alpine juniper 401 auguey 475
amansa 684 ‘aweoweo 190
American angelica tree 40 azahar 508
American beautyberry 135 azota-caballo 299
American bittersweet 164 azúcares 395
American burning bush 333 baby tea 196
American ivy 532 badoh 783
American plum 586 Bahaman wild coffee 608
American snowball 737 balsam 473, 619
American yew 758 balsaminha 142
amor ardiente 393 bálsamo 372
amourette 19 basbasot 714
812
Common Name Index
bitter bush 196, 561, 619
bamboo piper 564 bitter cane 370
banglin 489 bitter cherry 589
baquiña 432 bitter nightshade 721
barbasco 395 bittersweet nightshade 721
basam 275 bizcocho 395
bashful plant 489 black ambret 487
basket willow 664 blackbead 575
basket wiss 779 blackbead tree 235
basket wood 708 blackberry 43, 652
basora 247 black bush 781
basora prieta 798 black hawthorn 254
bastard guava 329 black mimosa 489
bastard sirio 30 black sage 247, 676
bâtard balengène 726 black sagebrush 72
bayberry 502 black sumac 625
bay-cedar 742 black torch 319
bay grape 229 black widow 473
bayonette 125 black willow 668
beach creeper 321 blackwith 708
beach plum 591 blind-eye-tree 374
beautyberry 135 boater bush 798
bébèl 575 bog myrtle 511
bejuco de berac 192 bois-anisette 432
bejuco de canastas 708 bois bolibri 354
bejuco de corrales 708 bois cabrit 32, 524, 696
bejuco de costilla 538, 708 bois cabroit 32
bejuco de fuego 359 bois candelle 38, 319
bejuco de gato 456 bois canon 137
bejuco de guajanilla 538 bois capable 684
bejuco de indio 359 bois caraibe 761
bejuco de membrillo 717 bois-champignon 473
bejuco de mona 213 bois codine 115
bejuco de nasa 779 bois crabbe 575
bejuco de palma 779 bois de fer 32, 235
bejuco de paloma 538, 779 bois de l’encore 524
bejuco de paralejo 764 bois énivrant 219
bejuco de sarsaparilla 717 bois flambeau 38, 186, 319
bejuco de sopla 359, 689, 764 bois fleche 341
belly-ache bush 399, 619 bois flot 376
bemberecua 769 bois-foufou 354, 524
berenjena cimmarona 726 bois-guillaume 365
berenjena de gallina 726 bois jaune 149
berenjena de playa 724 bois mêche 140
berenjena silvestre 726 bois montagne 561
Bermuda mulberry 135 bois-noir 140
Bigelow sagebrush 50 bois petites feuilles 327
big sagebrush 85 bois poison 561
bija 112 bois-puant 140
bilimbi 505 bois pini 38
birchleaf cercocarpus 178 bois puante 699
birchleaf mountain-mahogany 178 bois tan 154
bird pepper 144, 147 bola de coral 393
biscuitwood 149 bomba 137
bonbon bleu 223
813
Common Name Index
bon bon mél 223 buttonball 170

bonbon rouge 259 buttonbush 170
boo 726 button mangrove 240
bord-de-mer 154 button sage 417
borrerie verticillée 732 button willow 170
botón blanco 732 buttonwood 240
botoncillo 240 bwa kabuit 32
bow-pigeon 225 cabai nache 617
box-brier 617 cabo de hacha 777
boxleaf stopper 327 cabra cimarrona 684
box thorn 453 cabrilla 775
Brazilian pepper tree 686 cabrito 363
Brazilian jasmine 397 cachimbo 219, 524, 604
bread-and-cheese 575 cachimbo peludo 647
brilla 107 cadillo 243, 785
brisselet 325 cadillo espinoso 540
britónica afelpada 473 cadillo de perro 781
britónica prieta 471 Caesar weed 785
brittlebush 302 café 232
brittle thatch palm 766 cafecillo 121, 151, 341
broom 275 café cimarrón 341
broombush 365 café bâtard 606
broom butterweed 694 café de la India 508
broom dalea 600 cafeillo 684
broom groundsel 694 café marron 606, 608
broom indigobush 600 café silvestre 151
broom palm 766 caïmite marron 201
broom snakeweed 367 caimitillo 201
broomstick 777 caimitillo cimarrón 201
broom tops 275 caimitillo de perro 201
broomweed 367, 473, 714 cají torcido 21
broomwood 473, 777 calico bush 407
brownweed 367 California brickellbush 123
bucho 699 California broom 445
buckbrush 158, 280 California buckwheat 314
budsage 82 California rosebay 621
buenas tardes 286 California sagebrush 52
buenda 192 calotrope 137
buffalo current 637 camasey 223
buffalo-tip 766 camasey almendro 469
buff coat 798 camasey blanco 477
buis de sapin 758 camasey colorado 219, 475
bun 508 camasey de costilla 475
bur mallow 785 camasey de felpa 480
burning bush 335 camasey de paloma 219
burning love 393 camasey peludo 221, 223
bursting heart 333 camasey racimoso 481
burro 140 cambrón 14
burro weed 388 cambronera 453
burweed 781 cammy cuphia 266
bush cinquefoil 280 camphire 107
bush monkey flower 495 caña brava 370
bush penstemon 545 Canada yew 758
busunuvo 372 cañafistola cimarrona 696
814
Common Name Index
canalete 777 chichita 686

canario 36 chico 425, 453
candleberry 502 chile perro 696
candlewood 38, 186, 319 chili pepper 147
cane 91, 555 China box 508
cangá 449 Chinese honeysuckle 442
canique grise 129 Chinese inkberry 184
canote-macaque 223 chocolate blanco 115
capaillo 32 chokecherry 594
caper 140 cholla 520
capim elefante 542 Christmas berry 686
capulí cimarrón 775 Christmas bush 196
caracolillo 149 Christmas cherry 329
carbonera 696 Christmas vine 783
cardero 753 Christmas wreath 783
cardinal’s guard 516 cigüillo 363
cardio de frade 732 cina-cina 529
cariaquillo 196 cinnamon wood 681
Caribbean spiritweed 32 ciruela 586
carrapata 777 citron 214
carrapateira 643 citronnier 186
carrapicho do mata 785 clavelón de playa 119
carricillo 518 clidemia 223
carruzo 219 climbing mimosa 487
cascabelillo axiliar 261 climbing nightshade 721
cascalata 235 climbing orange root 164
casie 14 climbing poison ivy 769
casse marron 696 clove current 637
Castle Valley clover saltbush 101 coast broom 753
Castle Valley saltbush 101 coast rhododendron 621
castor bean 643 cocaine falsa 323
castor-oil plant 643 cockspur 156
catclaw 16 coco-plum 199
catclaw acacia 16 coco-ravet 149
catclaw mimosa 482, 489 cocuisa 350
cat’s claw 456, 575 coffee 232
cattle tongue 577 coffee colubrine 235
cautiva 36 coffee senna 699
Cayenne pepper 147 coin vine 278
cedar 404 collier-diable 575
ceibey 535 colorín 140
celadonia 115 comb butterweed 691
cenicillo 691 common broom 275
cenizoso 477 common buttonbush 170
cerezo 149 common juniper 401
cerise de sable 591 common poison ivy 769
chaico blanco 286 common rabbitbrush 203
chamizo 93 common reed 555
chaparral broom 103 common sage 417
chaparro 268 common sassafras 681
chapeau g’leau 432 common sida 715
cheezefruit 505 common snowberry 744
chew-stick 359 common sumac 628
chichicaste 788 conchombre 286
815
Common Name Index
conejo colorado 777 cucubano liso 363

Congo pea 132 cudweed sagewort 69
contite 417 cuentas de oro 299
cooze mahot 785 cuernecillo 374
copal 686 cuero de sabana crucillo 363
coquelicot 540 cuneate saltbush 101
coral 259 cuphia 266
corazón de paloma 235 curarador 115
corcho blanco 149 curarina 212
cordobán 475 cure-for-all 577
cordobancillo 645 curl-leaf mountain mahogany 175
cordobancillo peludo 647 curumo 140
cordoncillo 564, 567 dabel-da-dama 36
corita 34 dadangsi 785
cornicopio 286 dama de día 184
corona de novia 783 dama de noche 127, 186
correosa 631 damson plum 201
cortaduro 518 David bush 286
corta-lengua 151 David’s root 192
coscorrón 154 day cestrum 184
cossie 21 day jasmine 184
costilla de vaca 93 deerbush 158, 178, 610
cotorrerillo 149 deer horn cactus 181
cotton-france 137 deerweed 445
cottonrat 374 desert cassia 702
cough bush 321 desert cedar 404
cousin-petit 781, 785 desert Christmas cactus 520
cousin rouge 785 desert mock orange 551
cowbush 374 desert seepwood 739
cowfoot leaf 432 desert spoon 283
cowheel bush 432 desert sumac 631
cowitch 374 desert sweet 188
cow’s foot 564 desmanthus 289
cow-tobacco 545 desmanto 289
coyote bush 103 devil’s claw 573
coyote willow 664 devil’s fig 726
crab’s eye 12 devil’s trumpet 286
---
crabwood 225 devil’s walking stick 40
crack-open 151 dhal 132
creambush oceanspray 379 ditay paye 142
creek senecio 691 doctor-bar 561
creeping barberry 464 doctor long 575
creeping mahonia 464 Doña Julia 372
creeping Oregon grape 464 dorme dorme 492
creosote bush 419 dormidera 492
criaquillo 414 dormilona 489
crises marine 742 dos caras 475
croc-chien 487, 617 Douglas groundsel 691
crow broom 753 Douglas hackberry 167
cruz de Malta 393 Douglas hawthorn 254
cuabilla 38, 742 Douglas maple 24
Cuba jute 714 Douglas rabbitbrush 209
cubra prieta 319 Douglas ragwort 691
cucharo 742 Douglas thorntree 254
816
Common Name Index
dulcamara 721 figus nut 399

duranta 299 filigrana 414
duro-duro 149 firebush 372
Dutch myrtle 508 firespike 516
dwarf American cherry 591 fire willow 668
dwarf juniper 401 fishing rod 617
dwarf koa 289 fivefinger 280
dwarf mountain-mahogany 173 five fingers 617
dwarf Oregon grape 464 five-leaved ivy 532
dwarf sumac 625 flameleaf sumac 625
early sagebrush 66 flame of the woods 393
eastern poison ivy 769 flat sagebrush 50
eastern swamp privet 343 fleur-corail 372
eastern wahoo 335 fleur de ben-aimé 19
elephant grass 542 fleuret-Noël 196
elfin herb 266 fleur-sorleil bord de mar 119
emajagua 376 flor de San José 696
endurece maiz 140 Florida bitterbush 561
enegro 401 Florida boxwood 684
English broom 275 Florida crossopetalum 259
English myrtle 511 Florida holly 686
equisósea 393 Florida privet 345
éribun 733 Florida swamp privet 345
ernodia de playa 321 flowering current 637
erubia 719 fortuga caspi 149
escambrón 573 fourwing saltbush 93
escoba blanca 714 fragrant current 637
escoba dulce 714 framboisier 658
escoba dura 714 French cotton 137
escoba negra 275 French mulberry 135
escobilla 796 fresa de montaña 658
escobón 327 friega-platos 726
espinillo 529 frijol de árbol 132
espino blanco 14 fringed sage 63
espinosa amarilla 105 fringed sagebush 63
estafiante 69 fringed sagewort 63
Everglade velvetseed 363 fruta de paloma 299, 684
false bittersweet 164 frutilla 417
false cocaine 323 fulutufa 798
false coffee 341 furrusca 699
false heather 266 galán arbóreo 30
false kava 564 galán de día 184
false mallow 467 galán de monte 186
fausse-cane à sucre 542 gallinero 30
fedegoso 699 gandul 132
fellenwort 721 gandure 132
felty groundsel 691 garbancillo 777
female karata 350 garden datura 286
fern bush 188 garricillo 604
fever nut 128 gato 374
fever twig 164 gato soga 374
few-flowered ninebark 558 gatura 482
fèy a kèy 432 genevrier commun 401
feyukke friudem 505 geranium-leaf eupatorium 194
817
Common Name Index
giant cabuya 350 ground hemlock 758

giant cane 91 groven eye 575
giant laurel 623 guacamayo 115
giant milkweed 136 guacimilla de canario 553
giant mimosa 489 guacoco 140
giant reed 555 guairaje 327
giant reedgrass 555 guaita 777
giant sensitive plant 489 guaje 437
gilia penstemon 545 guamis 419
ginger-thomas 761 guano de sierra 766
giroflé 449 guano pea 132
girofle-ma 449 guanxuma 714
glandular mesquite 580 guapo 684
globe-flower 170 guarema 561
goat-bush 156 guaritoto 788
goat horn 617 guassatonga 151
goat-weed 142 guatama indigo 386
gobernadora 419 guava 597
gobiabiera 597 guayaba 597
Gold Coast jasmine 397 guayaba silvestre 329
golden creeper 321 guayabilla 151
golden current 637 guayabillo prieto 363
golden dewdrop 299 guayabo 597
golden hardback 280 guayarote 154
golden trumpet 36 guayayo 564, 567
goose plum 586 guazumilla 121
gordolobo 115 guen léglise 12
goyavier 598 guerit-tout 577
goyavier bâtard 329 guerrero 257
goyavier de montague 329 guía mansa 149
granadilla 535 guimauve 798
granadillo bobo 477 Guinea pepper 147
grand cousin 785 guitará 742
grande chelidoine 115 gull feed 119
grand giroflé 449 gumbo file 681
grão de galo 299 gurgeon stopper 327
grass-leaved ragwort 694 gypsum scalebroom 434
grate-jambe 487 hardhack 379
great morinda 505 hava de San Antonio 129
gray ephedra 307 Havard oak 613
grayia 361 Havard shin oak 613
gray nicker bean 129 hearts-a-bursting 333
Gray’s saltbush 361 heather 266
graythorn 803 hedionda 699
gray willow 664 hediondilla 419, 702
green aloe 350 herbe-à-crâbes 107
green broon 275 herbe à elephants 542
green ephedra 310 Hercules’ club 40
greenheart 235 hibiscus bur 785
green rabbitbrush 209 hierba de peso 212
Green’s mountain-ash 729 hierba de soldado 798
griffe á chatte 456, 575 higuereta cimarrona 399
grosse-peau 154 higüerillo 643
ground cedar 401 higüerito 643
818
Common Name Index
higuillo 564, 569 jackass breadnut 219

higuillo de hoja menuda 564 jackswitch 243
higuillo de limo 567 jalapón 527
Himalayan blackberry 652 Jamaica loostrife 449
Hioloa 798 Jamaican trash 365
hoary sagebrush 57 Japanese honeysuckle 442
hog apple 505 jasmin à bouguet 397
hog plum 586 jasmin blanc 397
hoja de pasmo 475 jasmín de canario 397
hoja menuda 121, 327 jasmín de estrella 341
hold back 129 jasmín del monte 127
hollyleaved barberry 461 jasmín de trapo 397
hollow heart 30 jaune d’oeuf 149
honey bush 508 jasmine 397, 684
honey mesquite 580 jasmín oloroso 397
honteuse 492 jayajabico 319
hoop vine 779 Jerusalem-thorn 529
hop bush 293 jeucon 374
hopseed bush 293 jigá 325, 684
horn-of-plenty 286 jointfir 310
horse bean 529 jointwood 564, 567
horsebrush 361, 365 jojobán 777
huang hau mu 714 jovero 742
hueso 561 jumbie apple 149
huesito 323 jumbie bean 12
hueso 342 jumbie breatfruit 505
huevo de gato 374 jungle flame 393
huisache 14 jungle-geranium 393
humble plant 492 jut africain 785
hummingbird bush 370 kafaki 798
huye-que-te-coge 796 kak mé 223
icaco 199 kalmia 497
icaque ponne 199 kamole 449
idem 714 karatas 125
‘ilima 712 Key lime 214
Indian apple 286 Key thatch palm 766
Indian arrowwood 548 koa haole 437
Indian mulberry 505 ko’oko’olau 110
Indian Valley false mallow 467 Koster’s curse 223
indigo 386 Labrador tea 429
indio 323 Lakana 414
inkberry 617 Lamark trema 775
ink-bush 184, 345 lambedora 518
inkweed 739 lampuaye 293
iodine bush 739 lantana 414
ipil-ipil 437 la péchita 580
Irish tops 275 laurel 407
ishanga 788 leadtree 437
ivy 407 leandra 427
ivybush 407 leather coat 798
ixora 393 lechecillo 647
jabillo 777 lemonade sumac 628, 634
jaboncillo 689 leucaena 437
jacatirão 477 Lewis mock orange 548
819
Common Name Index
Lewis syringe 548 mahot fin 247

liana fragante 291 mahot jaune 540
liane á barriques 779 maidenberry 259
liane à lait 36 majagua 376
liane amère 212 majagüilla de costa 374
liane á terre 779 mahoe daux 376
liane cordé 212 mala mujer 788
liane des sorciers 192 malimbé 567
liane molle 213 mallet 243
liane-savon 359 mallow-leaved ninebark 558
liane-séche 321 mallow ninebark 558
licorice plant 12 malva 785 ---
lija 245 malva colorada 471
lila 299 malva de cochino 714
lilikoi 535 malva loca 467
lima 215 malva silvestre 467
limestone snakevine 384 malvavisco 243, 798
limón agria 214 mameyuelo 43
limón boba 214 mangle bobo 117
limoncaspi 149 mangle médaille 178
limón criollo 214 mangle negro 240
limoncillo 323, 684 mangle prieto 240
limoneria 508 manglillo 319
limonejo 259 mangue 240
linguam 140 manman guêpes 788
linmangkon 535 manto 259
lintentwa 518 many-headed groundsel 694
lintisco 178 many-seed 449
lipstick plant 112 manzanilla de monte 801
lirio 735 maracuyá 535
little leaf cercocarpus 173 maraimaray 278
littleleaf mock orange 551 maravedí 259, 689
littleleaf mountain-mahogany 173 marble tree 154
little-leaf sumac 631 marie-honte 492
little rabbitbrush 209 marieniera 30
llorasangre 115 marker grass 542
lluvia 299 marsh-mallow 798
long-leaved willow 664 mascapalo 359
long-shoot 796 mataburro 529
lote bush 803 mata de mariposa 354
Lousiana sagewort 69 mata gallina 30
Louisiana wormwood 69 mata pelandok 43
low juniper 401 mata piojo 777
low rabbitbrush 209 matchbrush 367
low sagebrush 45, 66 mate prieto 154
lyenn-savon 359 matico 564
macanabo 201 mato de playa 129
macata 437 mat saltbush 99
machacomo 149 matscale 99
madam naiz 529 Mauritius hemp 350
maguey 34, 350 Mauritius raspberry 658
maguey criollo 350 Mautofu 785
mahot-cousin 781, 785 mauve 471
mahot-cousin rouge 781 mauve d’Amérique 467
820
Common Name Index
mauve-gris 798 netleaf hackberry 167

maya 125, 475 Nevada ephedra 307
mayhont 492 Nevada jointfir 307
meadow fern 511 Nia 514
médicinier bâtard 399 night-blooming cereus 181
médicinier noir 399 nigua 223, 788
médicinier rouge 399 nigüto 30
medsinnyé benni 219 ninebark 280, 558
membrillo 259 no-eye pea 132
memizo de majagua 775 noni 505
mengkudu 505 nono 505
merisier 684 Nuttall willow 668
mesquite 580 oakleaf sumac 634
Mexican lime 214 ocotillo 347
miki palaoa 699 ojos de cangrejo 12
milk tree 619 old man 691
mille fleurs 414 oldman sage 60
miraguano 766 oldman sagebrush 60
mitten tree 681 oliba 117
mock orange 508, 548, 737 olivier bard 117
molinillo 777 olivo 140
mombin bâtard 777 olivo macho 140
mondururu preto 477 ololiuqui 783
monkey bush 798 orange jasmine 508
monkey’s hand 432 orange peel 217
monval 437 orange peel clematis 217
morinda 505 oregano 417
morito 508 Oregon grape 461, 464
morivivi 492 Oregon grape-holly 461
mormon tea 307, 310 Oregon holly-grape 461
moth penstemon 545 oreja de mula 475
moto-branco 798 oreja de rató 212
mountain avens 295 orelia 36
mountain balm 157 Oriental bittersweet 161
mountain cercocarpus 178 Oriental virginsbower 217
mountain grape 461 orme petites feuilles 775
mountain juniper 401 ortiga 788
mountain laurel 407 ortiga brava 788
mountain mahogany 755 oseille bord de mer 742
mountain maple 27 oxeye daisy 119
mountain ninebark 558 paaloe pretae 140
mountain sorrel 655 pachaca 140
mountain spray 379 Pacific rhododendron 621
mountain sumac 625 Pacific yew 755
mountain willow 668 paddy’s lucerne 714
mouse’s pineapple 505 pain-in-back 775
mullaca colorado 477 pain killer 505
murette 779 pale lyceum 453
napier grass 542 palétuvier 240
naranjo jasmine 508 pale wolfberry 453
naranjuelo 140 palma Christi 643
narrowleaf willow 664 palma de cogollo 766
nedyah 781 palma de escoba 766
negra loca 227 palma de petate 766
821
Common Name Index
palmetto 704 pico de paloma 259, 327

palmita 766 pigeonberry 299
palo amarillo 115 pigeon pea 132
palo blanco 149 pigeon tree 40
palo corra 742 pigra 489
palo de anastasio 777 pimiento de Brazil 686
palo de burro 696 pink Chinese bur 785
palo de cachimbo 602 pink plains penstemon 545
palo de chivo 696 pionia colorada 417
palo de cotorra 617 pitillo de monte 425
palo de hierro 323 placa chiquitu 259
palo de paloma 259 planta de sal 107
palo de paz 561 plume poppy 115
palo de peje 561 pimiento 144
palo de perico 247 pingouin 125
palo de pollo 278 pinguin 125
palo de rayo 529 pinipinche de sabana 154
palo de toro 115, 341 pipewood 149
palo de vaca 121 poaia 732
palo de zorrillo 696 pockhout 227
palo moro 608 poirier 323
palo pelado 354 pois à mabou 140
palo verde 529 pois cajan 132
palo zapo 140 poison bittersweet 721
pan cimarrón 115 poison cherry 259
pandereta 766 poison ivy 769, 772
panilla 115 poison oak 772
pantsil 742 poison tree 154
papagayo 710 poison vine 769
papelite 151 polyodon 411
parcha 535 pomme de singe 505
pareira 212 pomme épineuse 286
Parry rabbitbrush 206 pomme-macaque 505
parrot apple 319 pompon blank 289
parrot weed 115 pompons jaunes 21
passionfruit 535 ponasi 524
pata de pájaro 372 pondberry 440
paz y justicia 456 popian 278
peetsch-kitam 617 porcupine flower 105
pega palo 291 pork-fat-apple 199
pelúa 219 Pottawattami plum 586
penah heriah 43 prairie bush 803
peronías 12 prairie mesquite 580 ---
petit bois blanc 684 prairie penstemon 545
petit casse 696 prairie sagewort 63
petit coco 617 prairie-weed 280
petit garcon 684 prickle bush 617
petit mahot 247 prickly ash 40
phlox penstemon 545 prickly elder 40
picanier jaune 105 prickly mampoo 573
pickleweed 107 prickly sage 414
picklewood 363 prickly solanum 726
pick tree 40 pride of Big Pine 735
pico de pájaro 699 primrose willow 449
822
Common Name Index
pringamoza 788 riverbank willow 664

Proctor’s vernonia 794 riverbush 170
prostrate bundleflower 289 river hawthorn 254
prostrate juniper 401 river plum 586
Puerto Rican ironweed 792 roble agalla 561
Puerto Rico girdlepod 500 roble amarillo 761
pull-back-and-hold 573 roble bobo 747
punta de sarvia 475 roble de guayo 121
punteral 363 roble cimarrón 747
purga del fraile 399 roble colorado 747
purple datura 286 rocío 325
purple sage 600 rockspirea 379, 382
pygmy sagebrush 75 Rocky Mountain maple 24
pyramid bush 473 Rocky Mountain sumac 628
Queensland raspberry 658 rolón 575
Quelite salado 739 romarin noir 742
quiebrahacha 329 rope mangrove 376
quina del país 561 rosarin-bord-de-mer 735
quininebrush 610 roseau 555
raba stokki 140 roseau cane 555
rabbit thorn 453 rosebay rhododendron 623
rabo de ratón 354 roseleaf raspberry 658
raboe 617 rosemary 735
raichie 473 roseta 459
raisin de la mar 229 rothrock sagebrush 78
raisin marron 527 roucou 112
rajana 393 rough-leaf tree 278
rascagarganta 527 round-leaved bittersweet 161
raspalengua 121 rubber rabbitbrush 203
ratón 325 rufiana 184
rattlebox 263 Ryberg’s poison ivy 772
rat-wood 325 sabina 404
red gram 132 sabina morena 404
red ixora 393 sacagarganta 527
redosier dogwood 249 Sacramento bur 781
red palicourea 524 saffron-tree 201
red pepper 144, 147 sageweed 69
red physic nut 399 saico Amarillo 761
red plum 586 salmonberry 655
red sage 414 saloop 681
red sumac 628 saltbrush 361
redwood 323 saltcedar 749
reina-de-la-noche 181 saltwood 514
remaroogu 608 saltwort 107
resin-weed 367 salvadilla 142
retama 156, 529 salvia 577
retama de escobas 275 salzbush 374
retama falsa 412 sandbar willow 664
retama prieta 702 sand cherry 591
retamo 777 sand penstemon 545
rhacoma 259 sandsage 60
rhomboid ilima 714 sand sagebrush 60
rhubarbe caraïbe 505 sand wash groundsel 691
ricin commún 643 Santa María 196, 247, 790
823
Common Name Index
Santa María negra 564 silky sesban 710

santo selele bélétére 477 siltbush 806
sapo prieto 140 silver sagebrush 57
saragüero 247 silver sea oxeye 119
saraguero 247 siyou 30
sardine 477 skunkbush 634
sarnilla 151 skyflower 299
sassafras 619, 681 sleeping grass 492
satinleaf 201 sleepy morning 798
satinwood 508 slender willow 664
saúco 115 smallcain 425
saúco tintóreo 184 small crown flower 137
sauge 414 small-fruited thatch palm 766
saw palmetto 705 smelling stick 682
scabland sagebrush 77 smooth sumac 628
scarlet berry 721 snakebark 237
scarletbush 372 snowbank 735
scarlet sumac 628 snowberry 744
Schouler’s willow 668 snowbush 158
Scotch broom 275 snowbush ceanothus 158
screwtree 374 soap bush 223
sea amyris 38 Sodom apple 137
sea daisy 119 soldier’s-bush 365
seagrape 229 Sonadora 518
sea hibiscus 376 sonzonte 235
sea mulberry 240 soot-soot 567
sea ox-eye 119 souge rouge 577
sea sage 417 sourbush 135, 577
sea-shore ardisia 43 southern spicebush 440
seaside grape 229 southwestern condalia 803
seaside mahoe 376 sow-berry 135
seaside tansy 119 Spanish elder 569
Seminole balsamo 608 Spanish mulberry 135
Senna-tree 696 Spanish stopper 327
sensitive plant 492 spiked malvastrum 467
sereno 365 spiked pepper 564
shadscale 96 spineless hopsage 806
shallow ninebark 558 spiny abutilon 540
shambu 112 spiny hopsage 361
shining sumac 625 spiny sage 361
shinnery oak 613 spoonleaf 283
shin oak 614 spoon plant 283
shiny-leaf ceanothus 158 spoon tree 154
shiny-leaf wild coffee 608 spoonwood 407
shiny sumac 625 springsage 82
shoebutton ardisia 43 squaw current 641
shoo-shoo bush 726 squawweed 691
shot bush 40 staff-tree 164
showy crotalaria 263 sticky laurel 158
showy rattlebox 263 sticky monkeyflower 495
shrubby cinquefoil 280 stiff sagebrush 77
shrubby false buttonwood 732 stinging nettle 788
Siam weed 196 stinking weed 699
Siberian juniper 401 stool 283
824
Common Name Index
stopper bush 327 Texas night-blooming cereus 181

storax 737 thé du pays 142
strawberry bush 333 thicket creeper 532
strumpfia 735 thicket wild coffee 606
sulfato 529 thimbleberry 655
supple jack 538 thin-leafed erythroxylon 323
sureau gros 32 thornapple 286
swamp privet 343 thornbush 803
swamp redwood 323 thorny sensitive plant 489
sweet acacia 14 threadleaf butterweed 691
sweet-briar 21 threadleaf groundsel 691
sweet gale 511 threetip sagebrush 88
sweet fern 237 tibisi 425
sweet potatoe cactus 181 ti bonbon 259
sweetscent 577 ti café marron 608
sweetspire 391 ti coco 617
sweet yellowcrown 584 ti mawi 492
switchcane 91 timberline sagebrush 79
syringa 548 tintero 617
tabac à jacot 790 tintillo 617
tabac djab 30, 577 tobacco bush 158
tabaco del monte 30 togovao 293
tabacón 726 tomatillo 453, 726
tabaquillo 115 torchwood 395
taiche 140 torciopelo 219
tall mahonia 461 toothache bush 40
tallowbush 178 toothbrush tree 359
tallow shrub 502 torch 186
tamarisk 749 torchwood 38
tantan 437 tostado 149
tapacamino 524, 796 touch-me-not 492
tapaculo 374 tree oxeye 119
tapaculop 149 trios côtes 475
tapehuiste 485 trombeto 115
tarro de chivo 319 trompete du jugement 286
tarweed 266 true mountain-mahogany 178
tassel-plant 742 trumpet flower 761
tassel-white 391 túatúa 399
tautuba 399 tuchima 219
tea 38, 319 turkey berry 726
tea bush 473 turma de toro 219
tearblanket 16 turpentine-weed 367
te bois-lait 619 turtleweed 107
te café 386 twisted acacia 21
té del pais 142 tzazahuistli 520
te kafé mawon 608 Uganda grass 542
temporana 742 uña de gato 16, 456, 484, 575
tendra à caulilou-rivière 575 upland cotton 356
terciopelo 480 uri 129
terete 417 urtiga branca 788
terongan 726 urucú 112
teta de burro 201 urupagüita 156
tête à nègre 781 Utah juniper 404
Texas mimosa 16 uva de playa 229
825
Common Name Index
uva grass 370 wild cherry 149

uverillo 227 wild chili 144
uvero 229 wild clove 449
uvillo 227 wild-coffee 151, 235, 561, 608
vaillant garcon 561 wild cotton 356
vara prieta 140 wild egg plant 726
varnish-leaf ceanothus 158 wild grape 225
vassourinha 732 wild guava 329
vega blanca 127 wild honey-tree 149
velvet leaf 212, 798 wild indigo 386
velvetseed 363 wild jasmine 184
Virginia creeper 532 wild nutmeg 154
Virginia sweetspire 391 wild olive 117, 345
Virginia tea 391 wild pepper 144
Virginia willow 391 wild plum 586
viuda 291 wild pomegranate 321
wait-a-bit 129, 482 wild poponax 21
wait-a-while 16 wild raspberry 658
waraia 477 wild salve 374
waxberry 502, 744 wild sage 417
wax current 641 wild star-apple 201
waxwork 164 wild tamarind 437
western hackberry 167 wild tantan 289
western mountain-ash 729 wild tea 142
western mugwort 69 wild tobacco 30, 577
western poison ivy 772 winged burning bush 331
western sumac 628 winged spindletree 331
western syringa 548 winged sumac 625
western thimbleberry 655 witch’s broom 275
western thimble raspberry 655 woodbine 442, 532
western thorn apple 254 wood balsam 608
western yew 755 Wood’s rose 649
West Indian lime 214 woody nightshade 721
West Indian snow-berry 192 wooly corchorus 243
West Indian trema 775 yagrumito 115
Wheeler stool 283 yaray 765
whisky current 631 yaría de costa 325
white alling 117 yambush 365
white coalberry 744 yellow bell 36
white indigo berry 617 yellowbrush 209
white leadtree 437 yellow bush lupine 451
white mountain avens 295 yellow candlewood 696
white sage 69, 417, 671 yellow cane 555
white sagebrush 57 yellow cedar 524
white sassafras 681 yellow-elder 761
white sumac 628 yellow palicourea 524
white watting 327 yellow plum 586
white willow 140 yellow-root 164, 505
white wood 225 yellow rose 280
widdy 280 yellow sage 414
wild bamboo 425 yellow-top 367
wild bougainvillea 573 yellow weed 367
wild cane 370 yellowwood 684
wild cassava 399 yerba cana 691
826
Common Name Index
yerba de clavo 449 zarcilla 437

yerba de vibora 367 zarsa brava 19
yerba elephante 542 zarzaparilla 717
yerba pelada 354 zicate 199
yerba socialista 796 zombie 286
zamorette 726 zówèy mouten 577
zarca 487 z’yeux à chatte 129
827
Authors of Thamnic Descriptions
G.W. Andrews, USDA Forest Service, Forestry Sciences Laboratory, 520 Devall Drive, Auburn, AL
36830
Eric L. Berlow, Research Scientist, University of California San Diego, White Mt. Research Station, 3000
E. Line St., Bishop, CA 93514
Don C. Bragg, Research Forester, USDA Forest Service, Southern Research Station, P.O. Box 3516 UAM,
Monticello, AR 71656
Mary Carrington, Assistant Professor, Science Division, Governors State University, 1 University
Parkway, University Park, IL 60466
Barton D. Clinton, Research Ecologist, USDA Forest Service, Southern Research Station, Asheville, NC
28802
Kristina Connor, Plant Physiologist, USDA Forest Service, Center for Bottomland Hardwoods Research, Box
9681, Mississippi State, MS 39762
Ann M. DeBolt, Botanist, USD Forest Service, Rocky Mountain Research Station, Boise, ID 83702
Margaret S. Devall, Research Ecologist, USDA Forest Service, Center for Bottomland Hardwoods
Research, Stoneville, MS 38776
John K. Francis, formerly Research Forester, U.S. Department of Agriculture, Forest Service, International
Institute of Tropical Forestry, Jardín Botánico Sur, 1201 Calle Ceiba, San Juan PR 00926-1119, in
cooperation with the University of Puerto Rico, Río Piedras, PR 00936-4984, now retired and volunteering
with the USDA Forest Service, Shrub Sciences Laboratory, 735 N 500 E, Provo, UT 84606
Marshall R. Haferkamp, Rangeland Scientist, Fort Keogh Livestock and Range Research Laboratory,
USDA Agricultural Research Service, Miles City, MT 59301
Emerenciana G. Hurd, Botanist, Rocky Mountain Research Station, U.S. Department of Agriculture,
Forest Service, Boise, ID 83702
Deborah K. Kennard, Research Ecologist, USDA Forest Service, Forestry Sciences Laboratory, 320
Breen Street, Athens, GA 30602
Stanley G. Kitchen, Research Botanist, USDA Forest Service, Rocky Mountain Research Station, Shrub
Sciences Laboratory, 735 N. 500 E. Provo, UT 84606
Catherine E. Koehler, Reserve Manager, University of California Davis-McLaughlin Reserve, 26775

Morgan Valley Rd., Lower Lake, CA 95457-9411
Juanita A. R. Ladyman, Botanist, JnJ Associates, 6760 S. Kit Carson Cir. E., Centennial, CO 80122
Ariel E. Lugo, Director, International Institute of Tropical Forestry, U.S. Department of Agriculture,
Forest Service, Ceiba 1201, Jardín Botanico Sur, Río Piedras, PR 00926
Jinshuang Ma, Research Taxonomist, Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn,
NY 11225
E. Durant McArthur, Research Geneticist, USDA Forest Service, Rocky Mountain Research Station, Shrub
Sciences Laboratory, Provo, UT 84606
828
Paul A. McMillan, Coordinator, Biology Department, Capilano College, 2055 Purcell Way, North Vancouver,
British Columbia, Canada V7J 3H5
W. Henry McNab, Research Forester USDA Forest Service, Southern Research Station, Asheville, NC
28802
Ernesto Medina, Investigador Titular, Centro de Ecología, Instituto de Investigaciones Científicas,

Caracas, Venezuela and Adjunct Scientist, International Institute of Tropical Forestry, USDA Forest
Service, Ceiba 1201, Jardín Botanico Sur, Río Piedras, PR 00926
Stephen B. Monsen, retired and volunteering with the USDA Forest Service, Shrub Sciences Laboratory,
735 N 500 E, Provo, UT 84606
Arlee M. Montalvo, Plant Population Biologist, Department of Botany and Plant Sciences, University of
California, Riverside, CA 92521-0124
Gerry Moore, Research Taxonomist, Brooklyn Botanic Garden, 1000 Washington Avenue, Brooklyn, NY
11225
James E. Nellessen, Botanist, Biologist and Environmental Scientist, Taschek Environmental Consulting,
8901 Adams St., NE, Albuquerque, NM 87113
Kenneth Outcalt, Research Ecologist, USDA Forest Service, 320 Green Street, Athens, GA 30602
Paula M. Pijut, Plant Physiologist, USDA Forest Service, North Central Research Station, Hardwood Tree
Improvement and Regeneration Center, 195 Marsteller Street, West Lafayette, IN 47907
John A. Parrotta, Biological Scientist, Research & Development, USDA Forest Service, Washington, DC
20090-6090
Rosemary L. Pendleton, Research Ecologist, USDA Forest Service, Rocky Mountain Research Station,
Albuquerque, NM 87102
Sally A. Reynolds, Botanist, U.S. Fish and Wildlife Service, San Francisco Bay National Wildlife Refuge
Complex, P.O. Box 524, Newark, CA 94560
Christopher Ross, Natural Resource Specialist, Bureau of Land Management, Nevada State Office, P.O. Box
12000, Reno, NV 89520
Stewart C. Sanderson, Research Geneticists, USD Forest Service, Rocky Mountain Research Station,
Shrub Sciences Laboratory, 735 N. 500 E., Provo, UT 84606
Nathan M. Schiff, Research Entomologist, USDA Forest Service, Center for Bottomland Hardwoods
Research, Stoneville, MS 38776
Randy S. Senock, Assistant Professor of Tropical Forestry, College of Agriculture, Forestry and Natural
Resource Management, University of Hawaii at Hilo, 200 West Kawili St., Hilo, HI 96720
Nancy L. Shaw, Research Botanist, USDA Forest Service, Rocky Mountain Research Station, Boise, ID
83702
Amanda J. Stevens, student, College of Agriculture, Forestry and Natural Resource Management,
University of Hawaii at Hilo, 200 West Kawili St., Hilo, HI 96720
829
Jeffrey R. Taylor, formerly Biological Technician, USDA Forest Service, Rocky Mountain Research
Station, Shrub Sciences Laboratory, 735 N. 500 E., Provo, UT 84606, now student, University of Utah, Salt
Lake City, UT
Sarah A. Taylor Student, College of Agriculture, Forestry and Natural Resource Management, University
of Hawaii at Hilo, 200 West Kawili St., Hilo, HI 96720
Bruce L. Welch, Plant Physiologist, USDA Forest Service, Rocky Mountain Research Station, Shrub
Sciences Laboratory, 735 N. 500 E., Provo, Utah 84606
830
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Building, 1400 Independence Avenue, SW, Washington, DC 20250-9410 or call (202) 720-5964 (voice or
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Keywords: Asteraceae, Rosaceae, Fabaceae, Cactaceae, woody plants, hardwoods, shrubs,

Pesticide Precautionary Statement

This publication reports research involving pesticides. It does not

CAUTION: Pesticides can be injurious to humans, domestic animals,

John K. Francis, Editor

Lawrence, G.H.M. 1955. An introduction to plant taxonomy. The Macmillan Company,

Merriam-Webster Inc. 1984. Webster’s ninth new collegiate dictionary. Merriam-Webster

McMinn, H. 1951. An illustrated manual of California shrubs. University of California

Morley, T. 1969. Spring flora of Minnesota. The University of Minnesota Press,

Synonyms: Abrus abrus (L.) W.F. Wright

Ecology.—In Puerto Rico, crab’s eye grows in

Reproduction.—A sample of crab’s eye seed

Synonyms: Acacia cavenia Bert.

America, in South America as far south as Chile

Ecology.—Sweet acacia is a drought-hardy, fire-

Synonyms: Senegalia greggii (Gray) Britt. & Rose.

Range.—Catclaw acacia grows in New Mexico,

Ecology.—Catclaw acacia is a drought resistant,

Synonyms: Acacia westiana DC

petioles and rachises are armed with recurved

Range.—Zarza brava is native to Hispaniola,

Ecology.—Zarza brava grows in habitat that

Growth and Management.— Seeds are dispersed References

Synonyms: Mimosa tortuosa L.

Range.—Twisted acacia is native to Jamaica,

Ecology.—Twisted acacia is intolerant of shade

References Nelson, G. 1996. The shrubs and woody vines of

Synonyms: Acer tripartitum Nutt.

Range.—Rocky Mountain maple occurs from

Ecology.—Rocky Mountain maple is moderately

lesser extent. Staminate flowers drop soon after the

Range.—Mountain maple occurs from

Synonyms: Atropa arborescens L.

cluster, yellow or orange, globose, 5 to 11 mm in

Range.—Gallinero is native to the Greater

Ecology.—Gallinero is widespread and rare to

Benefits.—Gallinero is used occasionally as an Liogier, H.A. 1995. Descriptive flora of Puerto

Synonyms: Aegiphila glandulifera Moldenke

Howard 1989, Liogier 1995, Little and others

Ecology.—Caribbean spiritweed is rare in Puerto

Synonyms: Agave portoricensis Trel.

elliptical or globose and 2.5 to 3.8 cm long and

Range.—Corita is native to Puerto Rico and the

Ecology.—Corita grow naturally in well-drained to

Growth and Management.—Corita grows at a Missouri Botanical Garden. 2002. W3 tropicos.

Synonyms: Echites verticillata Sessé & Moç.

branches. The bright yellow flowers are 5 to 7.5 cm

Range.—Allamanda is apparently native to

Ecology.—Wild and naturalized allamanda grows

Synonyms: Amyris maritima Jacq.

are aromatic and contain one brown seed (Howard

Range.—Torchwood is native to Florida, the

Ecology.—Torchwood is moderately intolerant of

Synonyms: Aralia leroana K. Koch.

Range.—Devil’s walking stick is found in the

Synonyms: Ardisia ketoensis Hayata

and A. humilis Vahl under A. elliptica (Center for

Ecology.—Shoebutton ardisia principally grows

Synonyms: Tournesol candicans M. Gómez

Range.—Howard (1989) lists the range of

Ecology.—Argythamnia inhabits areas of Puerto

Synonyms: Artemisia tridentata var. arbuscula (Nutt.) McMinn

E. Durant McArthur and J.R. Taylor