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A G e n e r i c S y s t e m f o r t h e
S y n a n d r o u s M i m o s a c e a e o f t h e A m e r i c a s
P a r t I I I . C a l l i a n d r a
T H E L u E S T H E R T. K E R T Z LIS!
T H E N E W Y O R K B O T A N I C A L G A R D E N
S i l k T r e e , G u a n a c a s t e , M o n k e y ' s E a r r i n g
M e m o i r s of T h e N e w Y o r k Botanical G a r d e n
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RT.fv'-ERTZ L I B R A R Y
T H E M 3 N Y O R K B O T A N I C A L G A R D E N
S i l k T r e e ,
G u a n a c a s t e ,
M o n k e y ' s E a r r i n g
A G e n e r i c S y s t e m f o r t h e
S y n a n d r o u s M i m o s a c e a e o f t h e A m e r i c a s
P a r t I I I . C a l l i a n d r a
R u p e r t C . B a r n e b y
Issued: 3 0 June 1 9 9 8
M e m o i r s of T h e N e w York Botanical Garden
V o l u m e 7 4 , Part III
T H E N E W Y O R K B O T A N I C A L G A R D E N
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Library of Congress Cataloging in Publication Data
Barneby, Rupert C.
Silk tree, Guanacaste, Monkey's earring : a generic system for the
synandrous Mimosaceae of the Americas / Rupert C. Barneby
p. cm.—(Memoirs of the N e w York Botanical Garden, ISSN
0077-8931 ; v. 74)
Includes bibliographical references (v. 3, p. ) and index.
Contents: pt. 3. Calliandra
ISBN 0-89327-420-8 (alk. paper)
1. Mimosaceae—West Indies—Classification. 2. Mimosaceae—South
America—Classification. I. Title.
II. Series
QK1.N526 Vol. 74
[QK495.M545]
581 s—dc20
[583'.321] 95-39082
CIP
S i l k T r e e , G u a n a c a s t e , M o n k e y ' s E a r r i n g
MEMOIRS OF THE N E W YORK BOTANICAL GARDEN 74(3): 1-223
Silk T r e e , G u a n a c a s t e , M o n k e y ' s E a r r i n g : A G e n e r i c S y s t e m
for the S y n a n d r o u s M i m o s a c e a e of the A m e r i c a s .
P a r t III. Calliandra
Rupert C. Barneby
C o n t e n t s
Abstract 1
Introduction 1
Definition of Calliandra, Past and Present 2
Infrageneric Classification of Calliandra 3 Calliandra 3
Conspectus of Genus Calliandra Bentham 5
Regional Keys to Species of Calliandra . . 7 Species Descriptions 21
Acknowledgments 201
Literature Cited 202
Species Incertae vel Minus Cognitae 204
Numerical List of Taxa Listed as Exsiccatae 204 List of Numbered Exsiccatae 206
Index to Scientific Names 215
Abstract
Barneby, R. C. (The New York Botanical Garden, Bronx, NY 10458-5126, U.S.A.). Silk tree, guana-
caste, monkey's earring: A generic system for the synandrous Mimosaceae of the Americas. Part III. A
revision of Calliandra (Mimosaceae tribe Ingeae). Mem. New York Bot. Gard. 74(3): 1-223. 1998.—
This alphataxonomic revison of genus Calliandra Bentham continues systematic work on tribe Ingeae
of the New World recently published (Barneby & Grimes, 1996). No comprehensive account of the
genus has appeared since 1875, and its definition and content have become incoherent, an obstacle to
floristic work presently in preparation. As redefined, the genus is endemic to the Americas, extending
from New Mexico to Chile, the so-called calliandras of Africa and Asia being excluded, as is the
recently segregated American genus Zapoteca H. Hernandez. The genus comprises 132 species, some
composed of two to several varieties, that are assigned to five sections and 14 series, all newly de
scribed. Of 168 specific and varietal taxa recognized, 36 are new. Each taxon is fully described, its
nomenclature is analyzed in the context of current rules of nomenclature, and its distribution recorded
verbally or in a map. Thirty-four taxa are illustrated, and a list of exsiccatae examined is provided.
I N T R O D U C T I O N
The last comprehensive account of the mimosace-
ous genus Calliandra, indeed the only one, appeared
in Bentham's summary classification of suborder
Mimoseae, published in 1875. Since that year many
new species of Calliandra have been discovered and
described, but the taxonomy of the genus has been
studied only piecemeal, by country, or by continent.
Step-by-step definition of the genus has been modi
fied, either purposely or inadvertently, by additions
and subtractions scattered in the literature but nowhere
coordinated. Consequently, there is no single contem
porary paper in which precise diagnostic descriptions
of Calliandra and its component taxa are assembled in
one convenient place, and even the broad outline of its
world dispersal remains controversial. The revision
presented herewith is designed to fill a deficiency that
is keenly felt by botanists working toward an inven
tory of the neotropical flora and a coherent classifica
tion of the family Mimosaceae.
For the past several years I have been engaged, in
collaboration with James Grimes, in revisionary work
on American Mimosaceae of tribe Ingeae, that is, of
MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
the mimosoid genera with indefinitely numerous,
eglandular stamens united at base into a tube. Detailed
surveys have been made of all American Ingeae
(Barneby & Grimes, 1996, 1997) except the follow
ing: a) Inga itself, including pluricarpellate Affonsea,
currently under investigation at Paris, Kew, Mexico,
Maracay, and elsewhere; b) the two small genera
Lysiloma and Enterolobium, capably revised by R. L.
Thompson (1980) and Mesquita (1990), respectively;
c) Calliandra, comprising some 130 species strictly
endemic to the Americas; d) the genus Zapoteca, for
merly included in Calliandra but recently segregated
(Hernandez, 1986); and a few ambiguous species na
tive to continental Africa, Madagascar, and tropical
Asia that, in spite of similar fruits, differ from Ameri
can Calliandra in pollen, leaf-nectaries, and a few
other characters enumerated below. As modern re
search on genus Inga comes to fruition and this review
of Calliandra is published, an organized body of data
bearing on the taxonomy of American Ingeae will be
in reach of systematists, conservationists, biologists,
and foresters, and will contribute to numerous
regional and national floras in preparation.
definition of calliandra,
Past and Present
Prior to 1840, the known species of Calliandra were
referred to Mimosa or Acacia or Inga, which at that
time were categories of convenience rather than bio
logical genera. As an autonomous genus of 18 species,
Calliandra was first characterized (Bentham, 1840:
138) by a syndrome of a) bipinnate leaves; b) globose
units of inflorescence; c) relatively long, basally mon-
adelphous stamens of indefinite number; d) eglandular
anthers; e) pollen shed in polyads; and f) a straight pod
of variable consistence, always elastically dehiscent
through both sutures from apex downward, its valves
simply recurved lengthwise but not horizontally or
obliquely twisted. Bentham later relaxed this generic
character to admit the simply pinnate leaves of C. hy-
menaeodes and the relatively short and few, sometimes
even fewer than ten, stamens of C. parviflora, so that
in practice a Calliandra could be identified technically
only by the fruit. In yet later times, a legume similar to
that of Calliandra in form and dehiscence was found
in monotypic Calliandropsis Hernandez, where it
coincides with the decandrous androecium of tribe
Mimoseae, in some Desmanthus, in Dichrostachys
kirkii Bentham, and even in the caesalpiniaceous
genus Jacqueshuberia Ducke. At this point, no one
macromorphological character stressed by Bentham
remained to distinguish all species of Calliandra from
all other genera of tribe Ingeae. Nevertheless, the orig
inal definition of the genus survived almost intact.
Thus Nielsen (1981: 184) described Calliandra in
Bentham's terms, modified only by lack of spinescent
stipules (except in Asia; but see also Cuban C. pauci-
flora) and by pleurogrammic seeds (in reality, preva
lent in the genus but far from universal). In their revi
sion for North American Flora, Britton and Rose
(1928) adopted for Calliandra the prior synonym
Anneslia Salisbury, but this was a simple nomen-
clatural gambit soon nullified by conservation, and it
had no impact of consequence on the taxonomy of In
geae. The discovery and documentation by Guinet and
Hernandez (1989) of two sorts of pollen in Calliandra
sensu Bentham led to recognition of the American
series Laetevirentes Bentham as an independent genus
Zapoteca, differentiated essentially by pollen alone,
although this criterion is weakly enforced by configu
ration of the stigma, pallid foliage, and relatively thin
texture of the pod's valves. Separation of Zapoteca, of
which two species have petiolar nectaries, confers this
taxonomic benefit: American Calliandra, but not the
Old World taxa referred by Bentham to Calliandra
sensu lato, is now uniformly without nectaries. With
removal of Calliandropsis and Zapoteca (= Anneslia
Portoricenses and Formosae Britton & Rose), the ap
proximately 130 American taxa that have been or
should be referred to Calliandra become a sharply
focused group that can be defined by the generic de
scription that follows. There remain, however, some
so-called calliandras in the Old World that have the
fruit of Calliandra but one or more anomalous fea
tures. The continental African C. redacta (J. H. Ross)
Thulin & Hunde (first described as an Acacia) and C.
gilbertii Thulin & Hunde are microphyll xeromorphic
shrubs, the first of northwestern Cape Province in
South Africa, the second of subequatorial Kenya and
Somalia, and were thought by Thulin and Hunde to be
inseparable from American Calliandra. However, their
acalymmate pollen is discordant. Spinescent stipules
found in C. redacta are encountered in American Cal
liandra only in the rare Cuban C. pauciflora, which
differs in distichous, not spiral, phyllotaxy. It is proba
ble that the pungent stipules of these two remotely al-
lopatric species are independently derived. Calliandra
gilbertii has the distichous phyllotaxy and firm but not
vulnerant stipules that can be matched in American
Calliandra, but some leaflets are alternate along the
rachis, as in the next group. The Madagascan callian
dras are presently under revision by J. F. Villiers (P),
who has generously shared with m e some of his
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING
insights, notably the establishment of the independent
genus Viguieranthus (ined.), differing from American
Calliandra in petiolar nectaries, generally coincident
with leaflets alternate along the pinna-rachis and often
by spiciform units of inflorescence or by genuinely
stipitate, not basally attenuate, fruits. The macrophyll
Indo-Burmese C. geminata Bentham, C. umbrosa
Wallich (with a var. griffithii (Baker) Paul), and C.
cynometroides Beddome also have leaf-nectaries but
are facultatively spiny-stipulate and at least sometimes
cauliflorous. Furthermore, the leaves of C. cyno
metroides are paripinnate bifoliolate. These poorly
known Ingeae have fruits reminiscent of genuine Cal
liandra but are surely not phylogenetically close to the
American core of the genus. Their generic status, es
pecially in relation to Viguieranthus, is unsettled. The
Burmese C. kingii Prain ex Paul & Husain is known to
m e only by the protologue (Feddes Repert. Sp. Nov.
97: 759, fig. 1. 1986), but has been identified by I.
Nielsen (in litt.) as Acacia kingii Prain.
infrageneric classification
of Calliandra
Purged of misplaced Paleotropical elements, Cal
liandra is for the most part rather uniform morpholog
ically, and consequently not readily analyzed into
monophyletic species-groups. A few noteworthy char
acters that occur only once in the genus, or that recur
in few, seemingly random and unrelated taxa, are use
ful specific markers but not significant at higher hier
archic levels. Such are: peculiar trichomes, stellate in
C. setigera or gland-tipped in C. umbellifera; spiral
phyllotaxy in C. chilensis and C. biflora; foliaceous
stipules in C. leptopoda and C. lanata; spinescent stip
ules in C. pauciflora; abruptly pinnate leaves in C. hy-
menaeodes; umbelliform units of inflorescence in C.
sesquipedalis, C. leptopoda, C. houstoniana, C. um
bellifera, C. pedicellata, C. brevicaulis; exaggerated
sutural ribs of pod in C. harrisii; and unexpected mod
ification of some part of the perianth in C. trinervia
var. stenocalyx, C. macrocalyx var. acuta, and C.
physocalyx. Despite anomalous features, the affinity of
these is not in doubt and the idiosyncratic features
themselves are merely distracting. The present classi
fication of Calliandra supposes fundamental diver
gence in a) inflorescence architecture, b) acquisition of
dorsally spiculate stipules, and c) depauperation of the
perianth and androecium. I interpret these characters
as persuasive evidence of real phylogenetic sequences.
The inflorescence of Calliandra takes three forms
characterized by different spatial and temporal rela
tions between the cauline axis of the season and the
complement of individual capitula, racemes, or um
bels. In the most frequent form, the signature of sect.
Androcallis, the units of inflorescence develop either in
primary leaf-axils of the current long-shoot, but below
its foliate apex, or are borne at nodes of short-shoots
and are then commonly delayed until the second year.
In either case, the genesis of the inflorescence-units is
lateral to their axis of origin, and the peduncles are
neither axillary toward apex of the long-shoot nor ter
minally pseudoracemose. In sect. Calliandra and the
small sect. Microcallis, the units of inflorescence arise
directly from axils of coeval leaves on the primary axis
and, beyond these, from efoliate nodes and axillary
stipuliform bracts, to form a terminal pseudoraceme.
N o brachyblasts are formed and no part of the inflo
rescence is postponed until a second season of growth.
As a rule the individual peduncle of sect. Androcallis is
bracteate below the first flower, whereas that of sect.
Calliandra is naked, but this rule is subject to excep
tions (noted in description of the species). In the most
highly specialized third type of inflorescence, encoun
tered only in monotypic sect. Acroscias, the function
ally herbaceous stems of the season, and sometimes
also their few secondary branches, are determinate,
each produced into a single terminal umbel. The stipu-
lar spicules of sect. Acistegis, which consists of two
species endemic to the West Indies, are associated with
the lateral inflorescences of sect. Androcallis and are
duplicated nowhere else in Mimosaceae. The sect.
Microcallis, comprising two Brazilian species very
dissimilar in foliage, resembles sect. Calliandra in
architecture of the inflorescence but differs in a minute
perianth (<3 m m long) and short androecium (3-7 m m
long). Despite congruence in the pollen and the fruit,
the two species of sect. Microcallis are discordant in
Calliandra and are perhaps generically distinct.
Within the two large sections Androcallis and Cal
liandra parallel modifications of leaf and flower are
frequent, and affinities of many species are difficult to
recognize. As will be seen from the conspectus imme
diately following, the species are sorted by phyllotaxy,
leaf-formula, leaflet-venation, and geographical dis
tribution, characters that do not necessarily reflect
phylogenetic relationships.
CALLIANDRA Bentham
Calliandra Bentham, Hooker, J. Bot. (Hooker) 2: 138.
1840, nom. conserv. — Sp. typica (Hernandez &
Nicolson, Taxon 35: 747-748. 1986): C. housto
niana (Miller) Standley.
MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Anneslia Salisbury, Parad. Lond. 64: 1807, nom. rejic. —
Sp. unica: A.falcifolia Salisbury, nom. illegit. = Calliandra
houstoni (L'Heritier) Bentham = C. houstoniana (Miller) Standley.
Clelia Casaretto, Nov. Stirp. Bras. 83. 1845. — Sp. unica:
Clelia ornata Casaretto = Calliandra harrisii (Lindley) Bentham.
Codonandra Karsten, Fl. columb. 2: 43. 1862. — Sp. unica:
Codonandra purpurea Karsten = Calliandra codonandra
Bentham = Calliandra magdalenae (de Candolle) Bentham var. magdalenae.
Calliandra sensu Bentham, Trans. Linn. Soc. London 30:
526-557, exclus. ser. Laetevirentibus, quae = genus Za
poteca Hernandez, necnon spp. paucis paleotropicis.
Trees, shrubs, and subshrubs, some functionally
herbaceous from perennial root-stock or rhizome,
none monocarpic; phyllotaxy distichous or excep
tionally spiral. Indumentum of simple, white, sordid
or brown-black hairs often mixed with short, amor
phously pluricellular, usually red or brown, herein
termed granular trichomes, and in very few spp. with
gland-tipped or stellately branched hairs, exception
ally lacking or almost so. Stipules herbaceous or sub-
coriaceous, commonly striately venulose dorsally and
either deciduous or persistent, exceptionally dilated,
or papery, or spinescent. Leaves bipinnate or (C.
hymenaeodes) simply pinnate; petiolar nectaries 0; lf-
formula diverse and often intraspecifically unstable,
i-many/1-many, the number and size often recipro
cally adjusted; venation of lfts either palmate from
the pulvinule or palmate-pinnate, or in small lfts re
duced to simple midrib. Inflorescence composed of
capitula, or condensed racemes, or umbels, arising
either from contemporary lf-axils or more commonly
from axillary brachyblasts, or forming through sup
pression of distal lvs a terminal psuedoraceme, ex
ceptionally (C. brevicaulis) consisting of a solitary,
truly terminal umbel; peduncles either bracteate or
not; bracts subtending each fl ordinarily persistent
through anthesis; fls of each unit of inflorescence
either homomorphic, or heteromorphic in shape and
the terminal one(s) in some respect larger, and/or the
peripheral ones staminate and the distal one(s) bisex
ual, or the enlarged terminal fl staminate; perianth
normally in most spp. 5-merous, but occasionally 3-,
4-, or 6-merous, or randomly irregular, the outer fls of
densely compact capitula often asymmetric or in
curved, the calyx and corolla often striately veined;
androecium (7-)10(-poly)-merous, the filaments
united into a tube ±lA to twice or more the length of
the corolla, the stemonozone either obscure or form
ing a well-differentiated hypanthium; intrastaminal
nectaries either present or 0, the inner face of hypan
thium then nectariferous; anthers dorsifixed, trans
versely oblong to elliptic; pollen shed in 8-grained ±
pyriform polyads, the basal grain narrowed proxi-
mally, sticky; ovary sessile or almost so, tapering into
the style, the stigma dilated but sometimes scarcely
so, the stigmatic surface either convex, or shallowly
cupular, or obscurely penicellate; ovules often 8,
sometimes fewer, rarely to 11. Pods either erect-
ascending on thickened peduncle or plagio- to geo-
tropic, in profile narrowly oblanceolate or linear-
oblanceolate, straight or slightly falcate, the plane or
low-convex, stiffly coriaceous or ligneous valves re
cessed into a frame formed by the thickened, some
times massive sutural ribs; dehiscence elastic, from
apex downward or from both ends, the valves recurv
ing but not laterally twisted; seeds descending on
short dilated funicle, plumply discoid or compressed-
ovoid or -rhomboid, the testa hard, the pleurogram
either U-shaped or lacking.
A n American genus (see discussion above) of ±135
spp., ranging from s.-w. United States to Uruguay,
warm-temperate Argentina, and n. Chile, most plenti
ful and diverse in monsoon climates and in open brush-
woodland or savanna communities at low and moder
ate elevations, many in riparian forest, a few Hylaean,
some tropical submontane, several adapted to desert.
A Note on the Keys to Species
Because of basic uniformity and repetitious modi
fication in macromorphology, a single key to the
species of Calliandra is difficult to construct and
would be a cumbrous instrument at best. As the dis
persal and ecology of most taxa of the genus are now
known in fair detail, it has become practical to sort
them in the first instance by geographical criteria into
groups of manageable size, the components of which
are then more swiftly and surely keyed. A few culti
vated species, it is true, have acquired erratic patterns
of dispersal and can be expected to turn up, either in
gardens or naturalized from gardens, in any of the
geographic divisions proposed below. These require
repeated entries but are comparatively few in number.
Keys based primarily on dispersal have, of course,
little phylogenetic or systematic significance; their
essential purpose is to link specimen with name and
to afford access to relevant systematic information. A
separate key to some commonly cultivated species of
which the origin is unknown (Key VIII), is provided.
The keys that follow need no further excuse or ex
planation, except for definition of two recurrent terms,
the significance of which must be clearly established:
leaf-formula and inflorescence architecture.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING
Leaf-formula is an abstraction describing the range
of highest numbers of secondary and tertiary divi
sions observed in the larger leaves of a given taxon. A
leaf-formula of ii-vii/5-11 indicates that larger leaves
of any specimen are composed of 2-7 pairs of pinnae,
the longest of which bear five to eleven pairs of
leaflets. Only the maxima of each category are taken
into consideration for each plant, though these max
ima are of course variable with each species.
Terminal and lateral inflorescences, which charac
terize the two principal sections of Calliandra—
namely, sect. Calliandra and sect. Androcallis,
respectively—are defined as follows. In sect. Callian
dra the units of inflorescence (individual capitula or
umbelliform capitula), whether solitary or fascicu
late, arise either directly from leaf-axils of the current
season's long-shoots or from efoliate nodes beyond
the current leaves, or from both. The fertile axes of
the year succeed or surpass the foliage. In the so-
called lateral inflorescence of sect. Androcallis, by
contrast, the peduncles arise either from contempo
rary axils low on the current season's long-shoots,
which continue to bear leaves at further nodes, or
exclusively from coeval or annotinous brachyblasts
generated on long-shoots of the season. In sect. Cal
liandra the flowers follow leaves of the primary stem-
axis, whereas in sect. Androcallis they are borne
either below leaves of the primary axis or on more or
less contracted secondary axes, these often efoliate
but thatched with imbricate stipules.
Conspectus of genus Calliandra Bentham
I. Sectio ANDROCALLIS Barneby
I/A. Series Androcallis * Pinnae of larger lvs 2-14 pairs:
+ North America and n. South America:
1. C. pittieri Standley
a. var. pittieri
b. var. polyphylla (Harms)
Barneby
2. C. purdiaei Bentham
3. C. glomerulata Karsten
a. var. glomerulata
b. var. parvifolia (Bentham)
Barneby
4. C. cruegeri Grisebach
5. C. tolimensis Taubert 6. C. laxa (Willdenow) Barneby
a. var. laxa b. var. stipulacea (Bentham)
Barneby c. var. urimana (Brunner &
Forero, ined.) Barneby
7. C. rubescens (Martens & Galeottii)
Standley 8. C. belizensis (Standley) Standley
9. C. goldmanii Rose ex Barneby
10. C. bijuga Rose
11. C. molinae Standley
12. C. hirsuta (G. Don) Bentham
13. C. peninsularis Rose
14. C. sesquipedalis McVaugh
15. C. californica Bentham
16. C. eriophylla Bentham
a. var. eriophylla
b. var. chamaedrys Isely
17. C. humilis Bentham
a. var. humilis
b. var. gentryana Barneby
c. var. reticulata (A. Gray) L.
Benson
++ Pacific South America:
18. C. tumbeziana Macbride
19. C. taxifolia (Kunth) Bentham
+++ E. Brazil, Uruguay, Argentina, Paraguay:
20. C. parvifolia (Hooker & Arnott)
Spegazzini
20a. C. carrascana Barneby
21. C. foliolosa Bentham
22. C. tweedii Bentham
23. C. bella Bentham 24. C. subspicata Bentham
25. C. dysantha Bentham
a. var. dysantha b. var. macrocephala (Bentham)
Barneby c. var. opulenta Barneby
d. var. turbinata (Bentham)
Barneby 26. C. gardneri Bentham
27. C. macrocalyx Harms a. var. macrocalyx b. var. aucta Barneby
28. C. fernandesii Barneby
29. C. ulei Harms 30. C. pilgerana Harms 31. C. umbellifera Bentham
32. C. imperialis Barneby
33. C. concinna Barneby 34. C. squarrosa Bentham
35. C. glaziovii Taubert
36. C. depauperata Bentham
37. C. silvicola Taubert
** Pinna exactly 1 pair: Amazonian Brazil and n.-ward:
38. C. surinamensis Bentham
39. C. samik Barneby 40. C. purpurea (Linnaeus) Bentham
41. C. riparia Pittier
42. C. magdalenae (de Candolle)
Bentham
a. var. magdalenae
b. var. colombiana (Britton &
Killip) Barneby
43. C. caeciliae Harms
44. C. chulumania Barneby
45. C. carcerea Standley & Steyermark
46. C. pityophila Barneby
47. C. colimae Barneby
48. C. hintonii Barneby
MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
+++++ S.-w. United States (Texas) and adj.
Mexico (Coahuila):
49. C. conferta Bentham
++++++ E. Brazil
50. C. brevipes Bentham
51. C. staminea (Thunberg) Barneby
52. C. sessilis Bentham
53. C. spinosa Ducke
54. C. duckei Barneby
55. C. blanchetii Bentham
56. C. aeschynomenoides Bentham I/B. Series Biflorae Barneby
57. C. biflora Tharp
I/C. Series Chilenses Barneby
58. C. chilensis Bentham
I/D. Series Pauciflorae Barneby
59. C. pauciflora (A. Richard)
Grisebach
60. C. enervis (Britton) Urban
I/E. Series Ambivalentes Barneby
61. C. medellinensis Britton & Killip
62. C. mollissima (Willdenow)
Bentham
63. C. guildingii Bentham
64. C. falcata Bentham
65. C. haematocephala Hasskarl
a. var. haematocephala
b. var. boliviano (Britton) Barneby 66. C. erythrocephala H. Hernandez &
Sousa
67. C. rhodocephala J. D. Smith I/F. Series Macrophyllae Bentham
68. C. trinervia Bentham a. var. trinervia
b. var. carbonaria (Bentham) Barneby
c. var. peruicola Barneby d. var. pilosifolia (Cowan)
Barneby e. var. paniculans Barneby
f. var. stenocylix Barneby
g. var. arborea (Standley) Barneby
69. C. bombycina Spruce ex Bentham 70. C. glyphoxylon Spruce ex Bentham
71. C. jariensis Barneby 72. C. coriacea (Willdenow) Bentham 73. C. antioquiae Barneby
74. C. angustifolia Bentham 75. C. harrisii (Lindley) Bentham
76. C. tergemina (Linnaeus) Bentham
a. var. tergemina b. var. emarginata (Willdenow)
Barneby 77. C. macqueenii Barneby
78. C. brenesii Standley
79. C. laevis Rose
I/G. Series Hymenaeodeae Barneby 80. C. hymenaeodes (Persoon)
Bentham
I/H. Series Longipedes Barneby 81. C. longipes Bentham
II. Sectio ACISTEGIA Barneby 82. C. haematomma (de Candolle)
Bentham
a. var. haematomma b. var. colletioides (Grisebach)
Barneby c. var. correllii Barneby d. var. rivularis (Urban & Ekman)
Barneby e. var. tortuensis (Alain) Barneby
f. var. glabrata Grisebach
g. var. locoensis (Garcia &
Kolterman) Barneby
83. C. pedicellata Bentham
III. Sectio ACROSCIAS Barneby 84. C. brevicaulis M. Micheli
a. var. brevicaulis
b. var. glabra Chodat & Hassler
IV Sectio CALLIANDRA
IV/A. Series Calliandra
+ Brazil: i. Pinnae (4-)5-13-jug.; calyx-teeth
deltate:
85. C. lintea Barneby
86. C. nebulosa Barneby
87. C. bahiana Renvoize
a. var. bahiana
b. var. erythematosa Barneby
88. C. lanata Bentham
89. C. fuscipila Harms
90. C. feioana Renvoize ii. Pinnae 2-5-jug.; calyx-teeth deltate:
91. C. asplenioides (Nees) Renvoize
92. C. viscidula Bentham
93. C. fasciculata Bentham
a. var. fasciculata b. var. bracteosa (Bentham)
Barneby
94. C. linearis Bentham 95. C. elegans Renvoize
96. C. mucugeana Renvoize
iii. Pinnae 1- or 1-2-jug.; calyx-teeth deltate:
97. C. calycina Bentham
98. C.x cumbucana Renvoize
iv. Pinnae 4-8-jug.; calyx-teeth lanceolate: 99. C. hirsuticaulis Harms
100. C. crassipes Bentham 101. C. hirtiflora Bentham
a. var. hirtiflora
b. var. ripicola Barneby
v. Pinnae 2-5-jug.; calyx-teeth lanceolate: 102. C. sincorana Harms
103. C. stelligera Barneby
104. C. coccinea Renvoize
a. var. trimera Barneby b. var. coccinea
105. C. involuta Mackinder & Lewis
vi. Pinnae 1- or 1-2-jug.; calyx-teeth
deltate; lfts 32-88-jug.:
106. C. santosiana Glaziou ex Barneby 107. C. renvoizeana Barneby
108. C. longipinna Bentham
109. C. debilis Renvoize
110. C. iligna Barneby
vii. Pinnae 1- or 1-2-jug.; calyx-teeth
lanceolate; lfts 5-39-jug.:
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING
111. C. paterna Barneby 112. C. ganevii Barneby
viii. Pinnae 1- or 1-2-jug.; lfts 2-11-jug.;
calyx-teeth deltate:
113 C. erubescens Renvoize 114. C. semisepulta Barneby
115. C. germana Barneby
116. C. luetzelburgii Harms
117. C. hygrophila Mackinder & Lewis
North America (cultivated elsewhere):
118. C. houstoniana (Miller) Standley
a. var. anomala (Kunth) Barneby
b. var. colomasensis (Britton &
Rose) Barneby
c. var. acapulcensis (Britton & Rose) Barneby
d. var. calothyrsus (Meisner) Barneby
e. var. houstoniana
119. C. juzepczukii Standley
120. C. palmeri S. Watson
121. C. physocalyx H. Hernandez & Sousa
122. C. wendlandii Bentham
123. C. quetzal (J. D. Smith) J. D. Smith IV/B. Series Virgatae Barneby
124. C. virgata Bentham IV/C. Series Tsugoideae Barneby
125. C. tsugoides Cowan
126. C. vaupesiana Cowan
a. var. vaupesiana
b. var. oligandra Barneby
127. C. pakaraimensis Cowan
128. C. rigida Bentham IV/D. Series Comosae Barneby
129. C. comosa (Swartz) Bentham
130. C. paniculata C. D. Adams V. Sectio MICROCALLIS Barneby
V/A. Series Microcallis
131. C. parviflora Bentham V/B. Series Leptopodae Barneby
132. C. leptopoda Bentham
Regional keys to species of Calliandra
I. Continental North America, n. and w. of Panama Canal (p. 7) II. West Indies ... (p. 10)
III. Northern South America: Panama e. of the Canal, Colombia, Venezuela, Trinidad and Tobago,
and the Guianas (p. 10)
IV Western South America: Ecuador, Peru, and Chile (p. 12)
V Amazonian Brazil: states of Amazonas, Roraima, Amapa, Para e. to the Maranhao line, Acre, Rondonia, and Tocantins s. to lat. 10'S (p. 13)
VI. Northeastern Brazil: states of Maranhao e. to Rio Grande do Norte, s. to the Bahia line.... (p. 14)
VII. Brazilian state of Bahia s. and e. of rio Sao Francisco (p. 14)
VIII. Planaltine and tropical Atlantic Brazil: states of Mato Grosso, Mato Grosso do Sul, s. Tocantins, Goias
with Distrito Federal, transfranciscan Bahia, Minas Gerais, Espirito Santo, Rio de Janeiro (with
Guanabara), and tropical parts of Sao Paulo and Parana (p. 17)
IX. Southern South America: Bolivia, Paraguay, extratropical Brazil, Argentina, and Uruguay (p. 19)
X. Key to some cultivated species (p. 20)
I. K e y to species of Calliandra in continental North America:
s.-w. United States, Mexico, and Central America s.-e. to Panama Canal
1. Inflorescence composed of capitula or of umbelliform capitula borne laterally to homotinous long-shoots,
either from coeval lf-axils or from either foliate or efoliate (then often stipule-thatched) brachyblasts, the
units of inflorescence not assembled into a terminal pseudoraceme or thyrse.
2. Pinnae commonly of all lvs, always of some lvs of any plant, >1 pair.
3. Arborescent shrubs, at maturity 2 m upward (some individual plants and some whole populations
in stressful habitats may flower prolifically when smaller); lft-pulvinules sharply transversely
wrinkled; lowland and submontane tropical Mexico and Central America.
4. Internodes of annotinous and older stems vertically corky-ribbed and sulcate; lfts of longer
pinnae 8-12 pairs; Honduras 11. C. molinae (p. 37) 4. Internodes terete; lfts of longer pinnae mostly 12-29 pairs, in C. bijuga 7-12 pairs; widespread.
5. Capitula (sub)sessile on densely thatched brachyblasts toward apex of long-shoots;
fls at once many and large, the capitula mostly 14—24-fld, the calyx 4.5-7.2 mm, the
corolla 9-13 mm; androecial tassel white; e. Yucatan Peninsula (Quintana Roo),
Belize, and adj. Guatemala (Peten) 8. C. belizensis (p. 34)
5. Capitula evidently pedunculate and fls mostly fewer, or smaller, or both, and the
androecial tassel pink to carmine distally; allopatric.
6. Distributed from tropical Mexico to n.-w. Costa Rica; seed-coat pleurogrammic.
7. Lf-formula ii—v(—vi)/l3—19; s.-e. Mexico (Veracruz to Oaxaca and Chiapas)
to Costa Rica.
8. Stipules 4-7(-l l)-nerved; larger lfts 5-10 x 1.5-3(-3.2) mm; pod 5-10
m m wide. 7. C. rubescens (p. 33)
MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
8. Stipules 11-21-nerved; larger lfts 10-16 x 2.7-4.8 mm; pod 11-16 m m wide 9. C. goldmanii (p. 35)
7. Lf-formula i-ii/8-12; s.-w. Mexico (Jalisco to Guerrero) . . 10. C. bijuga (p. 37)
6. Entering Panama from South America; seed-coat lacking pleurogram 6. C. laxa (p. 29)
3. Shrubs and subshrubs or functional herbs from caudex, seldom >1.2 m tall (but occasionally
attaining 2 m in exceptionally favorable circumstances); lft-pulvinules not transversely wrinkled;
s.-w. United States, extratropical and desert Mexico, and in Mexico extending s. in upland oak-
pine forest and xeric brush-woodland to Jalisco, Colima, and Chiapas.
9. Units of inflorescence umbelliform, the pedicels of all fls 2.5-4 m m ; s.-w. Jalisco.
Lf-formula iv-vii/25-29 14. C. sesquipedalis (p. 41)
9. Units of inflorescence essentially capitulate, the pedicel of peripheral fls <1 m m or obsolete.
10. Distribution in deserts or desert mountains of Baja California Peninsula s.-ward from
Sa. San Pedro Martir; androecium red overall.
11. Lf-formula v-vii/16-21; rachis of longer pinnae 2-5 cm; in pine-oak
woodland at ±1400-1800 m in the Cape mountains of s. Baja California Sur 13. C. peninsularis (p. 40)
11. Lf-formula ii-iii(-iv)/5-12 (exceptionally more); rachis of longer pinnae <3 cm;
in desert below 500 m, from near 30°N in Baja California s. to the Cape in Baja California Sur 15. C. californica (p. 42)
10. Distribution otherwise, one species (C. eriophylla) entering n. Baja California, but its
androecium bicolored, white proximally, pink-tipped.
12. Capitula (1—)2(—3)-fld; phyllotaxy spiral; lf-formula ii-iii/7-10; androecium 14-20-merous; local in s.-centr. Texas and lowland Tamaulipas 57. C. biflora (p. 99)
12. Capitula (3-)5-18-fld; phyllotaxy distichous; either remotely allopatric (from w.
Texas far s. and w.) or, if (C. humilis var. chamaedrys) nearly sympatric in s. Texas,
then the androecium 30-50-merous. 13. Corolla 6-9 m m ; androecium red throughout; in pine-oak woodland of
tropical montane Mexico s. of the Transverse Volcanic Range (Jalisco
to Puebla and Chiapas). 14. Lf-formula iv-xii/19-38; androecium 20-30-merous; range as
given 12. C. hirsuta (p. 38)
14. Lf-formula i-ii/6-9; androecium 14-15-merous; local in
Colima 47. C. colimae (p. 88) 13. Corolla (3.8-)4.3-6.2 m m ; androecium bicolored, pink or carmine
distally; s.-w. United States, Mexican Plateau and Pacific n.-w. Mexico, thence extending s. of the Transverse Volcanic Range in xeric grasslands
to n. Jalisco, Hidalgo, Puebla, Chiapas. 15. Stiffly ramulose shrubs and shrublets, the long-shoots persistent
over winter; androecium 16-24(-27)-merous 16. C. eriophylla (p. 44) 15. Functionally herbaceous from taproot or from terrestrial caudex, or
from a rhizome, the flowering stems arising anew each year from
ground-level; androecium (30-)32-52(-78)-merous 17. C. humilis (p. 45)
2. Pinnae of all lvs exactly 1 pair. 16. Lfts all (in rare C. hintonii of s. Mexico all but the terminal pair) <2 cm; if cultivated, proceed
directly to couplet 25, below. 17. Each pinna exactly 3-foliolate, the rachis bearing a larger distal pair of lfts and one smaller
proximal 1ft on posterior side; microphyll forms of var. emarginata of: . . . . 76. C. tergemina (p. 127) 17. Each pinna at least 6-foliolate, often multifoliolate.
18. Lfts of longer pinnae 12-36 pairs; strong shrubs and trees, mostly 2-6 m or taller;
peduncles mostly from thatched brachyblasts; androecium bicolored, pallid proximally, pink or carmine distally; tropical Mexico and Central America.
19. Largest lfts 11-18 x 3-6 m m ; capitula 16-28-fld; var.
colombiana of: 42. C. magdalenae (p. 82) 19. Largest lfts 5-13 x 0.9-1.9 m m ; capitula 6-14-fld 43. C. caeciliae (p. 84)
18. Lfts of longer pinnae commonly 4-14 pairs, in rare C. pityophila of Guerrero 11-19
pairs; shrubs and subshrubs exceptionally reaching 2 m and most of them native to extratropical Mexico and s.-w. United States but if stature or dispersal as in the
preceding, either the lfts only 4-6 pairs, or the plant <1 m tall, or the androecium red
overall, or some combination of these characters.
20. At once tropical and mesomorphic; local in pine-oak woodland either in Mexico (Guerrero, Mexico) or Guatemala.
21. Shrubs to 3 m; lfts of longer pinnae 4-10 pairs, the longest 11-22 m m .
22. Lfts of longer pinnae 4-6 pairs; pinna-rachises 2^4.5 cm; s.-w.
state of Mexico at ±1100 m 48. C. hintonii (p. 88)
19981 SILK TREE, GUANACASTE, MONKEY'S EARRING 9
22. Lfts of longer pinnae 7-10 pairs; pinna-rachises 4.5-6.5 cm; depto. El Progreso, Guatemala, at ±2100 m 45. C. carcerea (p. 87)
21. Shrublets to 5 d m tall; lfts of longer pinnae 11-19 pairs, the largest
<10 m m ; Mexico (Guerrero) 46. C. pityophila (p. 87)
20. Extratropical and xeromorphic; longer lfts 3-11 m m .
23. Larger 1 fts 3.5-11 x 1.4-4 m m ; capitula 7-13-fld; androecium. bright red;
deserts of peninsular Baja California; unijugate forms of: 15. C. californica (p. 42)
23. Larger lfts 3-5.5 x 0.8-1.4 m m ; capitula 2-8-fld; androecium whitish or
pink-tipped; s. and w. Texas and adj. Mexico 49. C. conferta (p. 89) 16. Terminal pair of lfts (in C. brenesii the solitary terminal 1ft) 2-19 cm.
24. Lfts at least 2 per pinna, all lvs consequently not <4-foliolate.
25. Ornamental shrubs of parks and gardens in warm N. America, variable in lf-formula,
the longer pinnae mostly 10-18-foliolate but sometimes less; androecial tube charged
internally at orifice with a fringe of incurved sterile filaments 65. C. haematocephala (p. 108) 25. Native trees, shrubs, and subshrubs; longer pinnae 1-7-foliolate, >4-foliolate only in
C. rhodocephala of Gulf of Honduras and vicinity; no incurved fringe within orifice
of androecial tube.
26. Venation of distal lfts palmate-pinnate, the primary nerve immediately posterior
to the strongly displaced midrib incurved-ascending beyond mid-blade.
27. Pinnae of all lvs, or at least of all primary lvs, 5-7-foliolate; around
Gulf of Honduras and vicinity in s. Belize, adj. Guatemala, and
Honduras 67. C. rhodocephala (p. Ill)
27. Pinnae of all lvs 1-4-foliolate; widespread.
28. Peduncles 2.5-5 m m and capitula 20-40-fld; local in Guerrero, Mexico 77. C. macqueenii (p. 131)
28. Peduncles 10-40(^4) m m and capitula mostly 12-22-fld; both
sympatric with the last and widely allopatric. 29. Androecial tube 19-35 m m ; pods stiffly ascending from
plagiotropic branches; seed-testa lacking pleurogram. . . 72. C. coriacea (p. 122)
29. Androecial tube <10(—15) m m ; pods either plagiotropic or geotropic, hanging beneath the branches; seed-testa almost
always pleurogrammic. 30. Lft-pulvinules at most 1 m m , commonly less; lft-blades
(variable in size and outline) not acuminate or only
incipiently and very shortly so; widespread from s. Sonora and s. Tamaulipas, Mexico, to centr. Panama; var.
emarginata of: 76. C. tergemina (p. 127)
30. Lft-pulvinules 1.5-3.4 m m ; lft-blades strongly acuminate
in distal XA -lA; s. Mexico interruptedly s.-e. to centr. Costa Rica; var. arborea of: 68. C. trinervia (p. 113)
26. Venation of lfts pinnate, the midrib centric or almost so, subequally branched
on either side. 31. Lfts of each pinna 3^4 pairs; Guerrero and Oaxaca 66. C. erythrocephala (p. 110)
31. Lfts of each pinna 1 pair; s. Sinaloa to Colima 79. C. laevis (p. 133)
24. Lfts 1 per pinna, all lvs consequently 2-foliolate; local in Costa Rica 78. C. brenesii (p. 132)
1. Inflorescence an efoliate or only proximally few-foliate pseudoraceme, either of capitula or of
umbelliform capitula, terminal to stems or branches of current year, wholly or partially exserted
from foliage; brachyblasts 0. 32. Microphyllidious, the lfts of longer pinnae 24 pairs or more per If and the longest blades <1.5 cm;
stipules 2-9 x 1-2.5 m m ; androecium wholly or at least distally pink or red.
33. Pinnae (3-)4-20 pairs per If. 34. Calyx 2.2-9 m m , <lA as long as corolla; pinnae of larger lvs 6-many pairs.
35. Corolla 6.3-12(-l3) m m , if 12 m m or more then brown- or fuscous-pubescent;
widespread in Mexico and Central America 118. C. houstoniana (p. 177)
35. Corolla 13.5-21 m m , white-silky-pilose. 36. Calyx 3.5-6 m m ; larger lfts 4-8 m m ; s.-e. Mexico (Veracruz, Oaxaca,
and Chiapas) 119. C. juzepczukii (p. 182)
36. Calyx 6-9 m m ; larger lfts 9-13.5 m m ; s.-w. Mexico (Nayarit,
Jalisco) 120. C. palmeri (p. 184)
34. Calyx 10-13 m m , at least 2A as long as corolla; pinnae of larger lvs mostly 4—6 pairs; s. Mexico (Oaxaca, Guerrero) 121. C. physocalyx (p. 186)
33. Pinnae 1-2 pairs per If (in random lvs pinnae 3) 122. C. wendlandii (p. 186)
32 MacrophylUdious, the lfts of longer pinnae 6-9 pairs and the larger blades 2-4 cm; stipules
foliaceous 10-24 x 12-36 m m ; androecium white; Guatemala (Baja Verapaz) 123. C. quetzal (p. 187)
!0 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
II. Key to species of Calliandra in the West Indies
1 Lfts of longer pinnae 20-45 pairs; stipules not spurred dorsally; perianth subglabrous; cultivated and potentially naturalized 50. C. brevipes (p. 90)
1. Lfts of longer pinnae mostly <16 pairs, to 18 pairs in rare varieties of C. haematomma but the stipules
then dorsally spurred and the corolla white-silky. 2. Stipules all, or at least those at primary nodes, charged dorsally at base with a conic spur or a spicule
(these often mistaken for spinescent stipules). 3. Units of inflorescence capitate, the pedicels 0 or <0.3 m m 82. C. haematomma (p. 177)
3. Units of inflorescence umbelliform, the pedicels (2-)2.5-8 m m 83. C. pedicellata (p. 144)
2. Stipules never appendaged dorsally, rarely (in Cuba only) indurate and truly spinescent.
4. Longer lfts of larger lvs 2-9 cm. 5. Lfts 7-11 (— 14) pairs per pinna, broad-linear or linear-elliptic, 2.5^.5 m m wide, 3.5-6
times as long as wide; commonly cultivated and sparingly naturalized 41. C. riparia (p. 80) 5. Lfts either fewer {Wi-A pairs) or absolutely and proportionately wider, the amplest at least
7 m m wide. 6. Lfts Wi pairs per pinna, i.e., a terminal pair and one odd posterior; pods plagiotropic
or pendulous on subfiliform peduncle; native in Lesser Antilles, cultivated elsewhere 76. C. tergemina (p. 127)
6. Lfts at least 2 complete pairs per pinna; pods erect on stout lignescent peduncle;
cultivated only. 7. Lfts of longer pinnae 2-5 pairs; capitula 10-26-fld; no retrorse fringe of sterile
filaments within orifice of staminal tube. 8. Larger stipules 0.8-4 x 0.6-1.3 mm; inflorescence immersed in foliage, the
individual capitula axillary (either directly or on very short and obscure
brachyblast) to coeval lvs 63. C. guildingii (p. 104)
8. Larger stipules 2-10 x 1.5-2.8 mm; inflorescence pseudoracemose and
terminal to the year's principal stems and branches, the individual capitula
arising from stipulate but mostly efoliate nodes 64. C.falcata (p. 105) 7. Lfts of longer pinnae 5-10 pairs; capitula 25-80-fld; a retrorse fringe of sterile
filaments within orifice of staminal tube 65. C. haematocephala (p. 108) 4. Largest lfts <2 cm.
9. Units of inflorescence umbelliform, the pedicel of peripheral fls 2-8 mm; Hispaniola only 83. C. pedicellata (p. 144)
9. Units of inflorescence capitulate, the pedicel of peripheral fls <1 m m or imperceptible.
10. Pinnae 2-3 pairs; inflorescence a terminal efoliate panicle of pseudoracemes; Jamaica only.
11. Lf-stk of longer lvs 3-10 mm, and rachis of longer pinnae 4-9.5 cm;
lft-pulvinules subobsolete; lfts lanceolate to lance-oblong obtuse, the longer ones
12-19 x 3.5-5 mm, 3.4-4.8 times as long as wide; inflorescence-axes
subglabrous; calyx ±2 x 2.5 m m . 129. C comosa (p. 196) 11. Lf-stk of longer lvs 1.4-2.2 mm, and rachis of longer pinnae 3.5-4 cm;
lft-pulvinules 0.45-0.6 mm; lfts elliptic to oblong-elliptic, obtuse apiculate, the
longer ones 12-18 x 4.5-6(-7) mm, 2.2-3 times as long as wide; inflorescence-axes
pilosulous; calyx ±1.1 x 1.5 mm. ... 130. C. paniculata (p. 197) 10. Pinnae 1 pair; inflorescence of capitula borne on lateral brachyblasts; allopatric.
12. Larger lfts 7-20 x 4-11 mm; peduncles 8-50 mm; capitula 9-22-fld; Lesser
Antilles 40. C. purpurea (p. 78) 12. Larger lfts 1.3̂ 4 x 0.5-2.2 mm; peduncles 1-4 mm; capitula 2-5-fld; Cuba.
13. Lfts of each pinna (2-)3-7 pairs; stipules at primary nodes of long-shoots
spinescent; lfts plane 59. C. pauciflora (p. 101) 13. Lfts of each pinna exactly 1 pair; stipules subindurate but not spinescent; lfts
biconvex 60. C. enervis (p. 102)
III. Key to species of Calliandra in n. South America: Panama e. of the Canal,
Colombia, Venezuela, Trinidad and Tobago, and the Guianas
1. Lvs bipinnate.
2. Pinnae >1 pair per If. 3. Inflorescence a terminal efoliate pseudoraceme of umbelliform, 3-8-fld capitula, the peduncles
fasciculate by 2-6, the pedicels 2-4.5 mm, and the perianth glabrous; described from Surinam,
where presumably cultivated, known as native, in modern times, only from Mexico and Central
America; var. calothyrsus of: 118. C. houstoniana (p. 177)
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 11
3. Inflorescence of lateral units or, if incipiently pseudoracemose, the units capitate, the peduncles
at most 2 per node, the pedicels <1 m m or obsolete, and the perianth pubescent.
4. Primary nerve of lfts immediately posterior to midrib incurved-ascending through Va-Va
of blade or, in some very small lfts, barely visible, the primary venation thus either conventionally palmate or simple.
5. Peripheral fls of each capitulum relatively small, the calyx 1.1-3.5 m m , the corolla
4.3-9(-9.5) m m .
6. Capitula sessile or almost so, the peduncle not more than 3 m m , often obsolete; lfts at once small (the longest 4-7, exceptionally 9 m m ) and acute; pod 6-8 m m wide 4. C. cruegeri (p. 26)
6. Capitula evidently pedunculate, the peduncle (4-) 15-60 m m ; lfts either longer, or obtuse; pod (10-)11-23 m m wide.
7. Capitula (9-)10-28-fld; seed-coat lacking pleurogram.
8. Pinnae of larger lvs 3-5(-6) pairs and lfts of longer pinnae (8-)9-29
pairs, the longest lfts 5.5-21 x 1.7-8 m m 6. C. laxa (p. 29)
8. Pinnae of larger lvs 6-13 pairs and lfts of longer pinnae 32-70 pairs, the
longest lfts 3-10.5(-13) x 0.6-2.4(-2.8) m m 1. C. pittieri (p. 22)
7. Capitula 5-9(-10)-fld; seed-coat pleurogrammic 3. C. glomerulata (p. 23) 5. Peripheral fls of each capitulum relatively large, the calyx 4-7 m m , the corolla
9.5-14.5 m m .
9. Larger lfts 4.5-8(-10) x 0.7-1.5 m m ; calyx-teeth 1.5-3.2 m m ; pod thinly
puberulent 2. C. purdiaei (p. 23)
9. Larger lfts 8-13 x 1.7-2.3 m m ; calyx-teeth 0.5-1.3 m m ; pod densely pubescent
with either brown or white-silky, either subappressed or erect hairs. . . . 5. C. tolimensis (p. 28)
4. Primary nerve of lfts immediately posterior to midrib produced, parallel to midrib,
almost or quite to blade's apex, the 1ft thus appearing 2-nerved or, by accession of a
similar anterior primary nerve, 3-nerved, its whole length; peduncles arising from primary
lf-axils toward apex of stem, the inflorescence, by suppression of distal lvs, pseudoracemose;
distribution determined by arenitic substrates, in Venezuelan Guayana, adj. Guyana and
Brazil, and Amazonian Colombia.
10. Perianth, or at least the corolla, white-silky externally; androecium (10-)12-57-merous. 11. Larger lfts 13-22 x 2.4—3.6 m m ; Gran Sabana in e. Venezuelan Guayana and in
adj. Guyana 127. C. pakaraimensis (p. 193)
11. Larger lfts 6-12 x 1-2.5 m m ; centr. Venezuelan Guayana to adj. Brazil (Sa. de
Araca), and e. Amazonian Colombia. 12. Primary and secondary lf-axes widely spreading and recurved; lfts of longer
pinnae 46-80 pairs; androecium mostly 36-57-merous, but in rare var.
oligandra only 14-24-merous; some granular trichomes mixed with plain
gray ones in the inflorescence 126. C. vaupesiana (p. 191)
12. Primary and secondary lf-axes mostly straight, ascending; lfts of longer
pinnae 21-52 pairs; androecium 12-20-merous; no granular
trichomes 125. C. tsugoides (p. 190)
10. Perianth glabrous overall; androecium 6-7-merous; e. Venezuelan Guayana and immediately adj. Guyana 128. C. rigida (p. 194)
Pinnae of all lvs exactly 1 pair.
13. Lfts of longer pinnae 13-38 pairs. 14. Filaments of peripheral fls 24-32; pod massively ligneous, the sutural ribs in dorsal
view 2.3-5 m m wide; seed-coat finely pleurogrammic 42. C. magdalenae (p. 82)
14. Filaments of peripheral fls 12-26; pod leathery, the sutural ribs in dorsal view at most 3 m m wide; seed-coat lacking pleurogram 38. C. surinamensis (p. 76)
13. Lfts of longer pinnae 1—12(—13) pairs. 15. Lft-pairs of longer pinnae (2Vfe-)3-13, the whole If at least 10-foliolate.
16. Either lfts of longer pinnae >4 pairs, or the longest lfts <3 c m long, or both.
17. Lfts (disregarding the terminal pair, which may be heteromorphic and longer than
the rest) in outline broad-linear or linear-elliptic, 2.5-6 times as long as wide;
seed-coat pleurogrammic. Native and also planted in Venezuela n. of the Orinoco
and in n.-e. Colombia widely dispersed in cultivation as a street tree or for ornament.
18. Lfts of longer pinnae 7-12 pairs, and the larger lfts ±2.5-4.5 m m wide;
native in n. S. America (Colombia to Guyana), planted in Neo- and
Paleotropics 41. C. riparia (p. 80)
18. Lfts of longer pinnae (3-)4-7 pairs, and the larger lfts 5-9 m m wide; known
for certain only cultivated in inter-Andean Colombia 61. C. medellinensis (p. 103)
17. Lfts (with same reservation) in outline obovate, oblong, rhombic-ovate or -obovate,
or obtusely rhombic, 1.5-2.3 times as long as wide.
12 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
19. In longer pinnae the lfts at least 8 pairs, in outline mostly rhombic or
rhombic-obovate; seed-coat lacking pleurogram; Guianas and Guayana w.
into Colombia, widespread and common 38. C. surinamensis (p. 76)
19. In longer pinnae the lfts 3-6(-7) pairs, in outline mostly obovate; seed-coat
pleurogrammic; Caribbean slope in n. Colombia and Venezuela, disjunct in
s. Guyana and on sandstone cerros of Amazonian Colombia 40. C. purpurea (p. 78)
16. Lfts at once 2V2-5 pairs and the terminal pair (2-)2.5-10.5 x 1.2-3.6 cm.
20. Larger stipules 0.8-4 x 0.6-1.3 mm; inflorescence immersed in foliage, the
individual capitula axillary (either directly, or on very short and obscure brachyblast) to coeval lvs; plants of mesophytic woodland 63. C. guildingii (p. 104)
20. Larger stipules 2-10 x 1.5-2.8 mm, inflorescence pseudoracemose and terminal
to the year's principal stem and branches, the individual capitula arising from
stipulate but mostly efoliate nodes; plants of xeromorphic associations.. 64. C. falcata (p. 105)
15. Lft-pairs of all pinnae exactly 1, or V/i, or 2, the whole If 4-8-foliolate.
21. Both calyx and corolla sharply striate, the corolla its whole length; androecium
38-58-merous; local in n. Colombia (Antioquia) 73. C. antioquiae (p. 123)
21. Only calyx sharply striate, the corolla-tube usually and its lobes always faintly nerved
or externally nerveless; androecium (6-)10-32-merous; widespread.
22. Distal pair of lfts relatively small, the largest blades 1.5-4 cm.
23. Lfts of each pinna 2 pairs; androecium (6-)9-10-merous; pods erect on
stout peduncle; on stream and river banks in Andean Colombia (and widespread outside the range covered by this key, both native and cultivated) 74. C. angustifolia (p. 125)
23. Lfts of each pinna M pairs, the anterior proximal 1ft lacking; androecium
12-18-merous; pods pendulous on subfiliform peduncle; in seasonally dry
brush-woodland of Caribbean slope in Venezuela, discontinuously w. to the Magdalena valley and equatorial Amazonia in Colombia 76. C. tergemina (p. 127)
22. Distal pair of lfts relative ample, the largest blades 4-12 cm.
24. Lfts narrowly or broadly but subequilateraly elliptic or oblance-elliptic, the distal pair 4-10 x 1-3.2 cm; pods stiffly ascending; seeds ±7.5-9 x 5.5-7 mm;
discontinuously dispersed in the Guianas, s.-w. Venezuela (Amazonas), and inter-Andean Colombia 72. C. coriacea (p. 122)
24. Lfts semi-ovate or inequilaterally ovate or lance-elliptic, the distal pair
5.5-16 x 2-6(-7) cm; pods plagio- or geotropic; seeds 11-22 x (6-)7-12 mm;
Colombian and Venezuelan Amazonia (vars. trinervia, pilosifolia) and
Andean Colombia (vars. carbonaria, paniculans) 68. C. trinervia (p. 113) 1. Lvs simply pinnate; lfts either 1 or 2 pairs per pinna, the distal pair ±4-10 x 2.5-4.5 cm;
Guianas 80. C. hymenaeodes (p. 134)
IV. Key to the species of Calliandra in w. South America: Ecuador, Peru, and Chile
1. Pinnae, usually of all lvs, always of most, 2 or more pairs per If.
2. Pinnae (6-)7-13 pairs per If; lfts of longer pinnae 15-70 pairs; arborescent to 15 m: Andean
Ecuador 1. C. pittieri (p. 22) 2. Pinnae 2-5 pairs per If; lfts of longer pinnae 3-30 pairs; if in Ecuador (Azuay, Loja), mostly
diffuse or prostrate shrubs <2 m.
3. Peduncles 0.4-2.2 cm; filament-tassel red or orange; scattered along the Ecuadorean and
Peruvian Andes (in the latitude of Tumbez at 1800 m upward) from s. Ecuador to depto.
Huanuco, Peru, and isolated in lowland Pacific Peru in depto. Arequipa 19. C. taxifolia (p. 50) 3. Peduncles 3.5-4 cm; filament-tassel white; known only from n. Pacific Peru in depto.
Tumbez, alt. 500 m 18. C. tumbeziana (p. 49)
1. Pinnae of all lvs exactly 1 pair. 4. Lfts of each pinna 1, Wi, or 2 pairs, and the blade of the distal lfts 1.5-16 cm.
5. Corolla sharply striate from base to apex of lobes; androecium 44-54-merous, red throughout; local in lowland Amazonian Peru (San Martin).. . 69. C. bombycina (p. 118)
5. Corolla striate only on tube, or not striate; androecium 12-32-merous, variable in color, but in
lowland Amazonian Peru bicolored, the tube white and the tassel carmine (sometimes red on
the Pacific slope). 6. Indumentum of stems and lf-axes thin, or lacking; if upland Andean (in Cauca valley,
Colombia) the stipules at nodes of long-shoots 1.5-2.8 mm; petioles 9-35 mm; capitula
18-27-fld 68. C. trinervia (p. 113) 6. Indumentum of stems and lf-axes spreading; stipules at nodes of long-shoots 3-10 mm;
petioles 1.5-8 mm; capitula 30^5-fld; w. slope of cordillera at 600-2100 m in prov.
Chimborazo, Ecuador 70. C. glyphoxylon (p. 119)
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 13
4. Lfts of each pinna either 4 pairs or more, or the largest of them <1.5 cm, or both.
7. Phyllotaxy distichous; stipules mostly 3-7-nerved (at least when young), never spinescent; rachis of longer pinnae 2.5-11 cm; blade of larger lfts >7.5 x 2 mm.
8. Perianth relatively small, the calyx 1.4-3.4 x 0.8-1.8 mm, the corolla 4.3-10.5 mm.
9. Lfts 3-7 pairs per pinna and the longest 17-55 x (8-)10-18(-25) m m wide. 10. Corolla 4.3-5.4 mm, the tube striate; tassel white; inter-Andean
n. Peru 61. C. mollissima (p. 103)
10. Corolla 5.8-10 mm, the tube not striate; tassel carmine; Amazonian Ecuador and Peru 63. C. guildingii (p. 104)
9. Lfts (in context of this key) 11-17(-25) pairs per pinna and the longest <12(-13) mm; androecium bicolored, the tassel pink-carmine; entering Ecuador from the n. and e 38. C. surinamensis (p. 76)
8. Perianth longer, the calyx 5.5-6 x 3 mm, the corolla 13-14.5 mm; sympatric with
C. mollissima but tassel of C. surinamensis 39. C. samik (p. 77)
7. Phyllotaxy spiral; stipules 1-nerved, subspinose at maturity; rachis of longer pinnae 5-11 mm;
lfts 4-7 pairs per pinna, the blade of larger ones 2.5^.5 x 0.9-1.6 mm; diminutive shrub of Atacama Desert, Chile 58. C. chilensis (p. 100)
V. Key to species of Calliandra in Amazonian Brazil: states of Amazonas, Roraima,
A m a p a , Para e. to the Maranhao line, Acre, Rondonia, and Tocantins n. of lat. 10'S
Pinnae 3-30 pairs per If.
2. Lf-formula iii-v/9-18, and the larger lfts (9-) 10-22 x 3-8 mm; var. stipulacea of: 6. C. laxa (p. 29)
2. Lf-formula in some respect higher than in the last, either the pinnae or the lfts more numerous, and the lfts always narrower, the largest 0.4-2.5 m m wide.
3. Units of inflorescence pseudoracemose and the crimson fls extremely small, the calyx 0.8-1.3 m m and the corolla 1.6-2.4 mm; androecium 6-11-merous and the filaments 3-7 mm. Pinnae of larger lvs 14—36 pairs per If, and larger lfts 2-3.5 mm; weakly entering
the region by way of the Araguaia valley 131. C. parviflora (p. 198) 3. Units of inflorescence not pseudoracemose, and the fls (variously colored) larger, the calyx
1.6-4 m m and the corolla 4.2-8.5 mm; androecium 12-40-merous and the filaments to
17-50 mm. 4. Pinnae of larger lvs 8-18 pairs and lfts only 2.5-5.2 x 0.5-0.9(-l) mm; entering the
region on campirana of lower Rio Tocantins valley 20. C. parvifolia (p. 52)
4. Either the pinnae of larger lvs fewer (3-5 pairs), or the larger lfts 6-12 mm; along the n.
borders of Amazonas and Roraima.
5. Pinnae of larger lvs 4—8 pairs; inner posterior primary nerve of lfts produced, straight
and parallel to midrib, almost or quite to blade's apex; peduncle of capitula 9-26 mm;
near the Venezuelan border in Amazonas (Sas de Tunui and Araca) 125. C. tsugoides (p. 190) 5. Pinnae of larger lvs 3-5 pairs; inner posterior primary nerve of lfts incurved-ascending
and expiring near or below mid-blade; peduncle of capitula 0-3 mm; Rio Surumu in
n. Roraima 4. C. cruegeri (p. 26)
Pinnae exactly 1 pair per If. 6. Lfts of longer pinnae 5-39 pairs, none over 14 m m wide.
7. Peduncle of capitula 7-20 mm; lfts of longer pinnae 5-19, rarely over 17 pairs;
widespread 38. C. surinamensis (p. 76) 7. Peduncle of capitula 0-2 mm; lfts of longer pinnae 18-39 pairs; local on campirana of lower
Rio Tocantins valley 52. C. sessilis (p. 92) 6. Lfts of all pinnae 1 or 2 pairs, the blade of distal pair commonly 2-7 cm wide, rarely narrower.
8. Venation of lfts pinnate, the primary nerve posterior to midrib no longer and no stronger
than the secondary ones. Terminal lfts of each pinna 6-13 cm, the blade inequilaterally
broad-elliptic or ovate-elliptic, not or obscurely acuminate; known only from lower Rio Jari
in Para 71. C. jariensis (p. 120) 8. Venation of lfts palmate-pinnate, the inner primary nerve posterior to midrib manifestly longer
and stronger than succeeding secondary ones, incurved-ascending at least to, commonly well
beyond, mid-blade. 9. Lfts of distal pair inequilaterally ovate or half-ovate, evidently and usually attenuately
acuminate, the larger ones 5.5-16 x 2-6(-7) cm; widespread in Amazonia,
polymorphic 68. C. trinervia (p. 113) 9. Lfts of distal pair elliptic-oblanceolate or less often broad-ovate, but not or very obscurely
and bluntly acuminate, the larger ones mostly 2-5.5 x 1-2 cm (sometimes a bit longer);
in Brazilian Amazonia known only from n.-e. Para (basins of rios Trombetas and
Paru do Oeste) 72. C. coriacea (p. 122)
14 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
VI. Key to species of Calliandra in northeastern Brazil: states of Maranhao e. to Rio Grande do Norte, s. to the Bahia line
1. Stipules foliaceous 1-3.5 cm wide; lf-formula i/3-5, and larger lfts 17-31 x 8-15 mm; pedicel of
peripheral fls 10-20 m m 132. C. leptopoda (p. 199)
1. Stipules <5 m m wide; if lf-formula similar then the larger lft-blades not more than 11 m m and the
fls sessile.
2. Units of inflorescence capitate or only incipiently umbelliform, the pedicel of peripheral
fls <3 mm, commonly almost 0. 3. Pinnae of some larger lvs, usually of all lvs, 2 pairs or more.
4. Pinna-pairs of larger lvs 5-36. 5. Units of inflorescence arising singly from thatched or ± extended brachyblasts;
calyx 1.8-3 m m and corolla 4.2-7 mm; androecium 12-32-merous, the longer
filaments 20-50 mm. 6. Pinnae of larger lvs 4-8 pairs and the longer pinna-rachises 3-5 cm; corolla
densely silky externally; local in n. Piaui 28. C. fernandesii (p. 67)
6. Pinnae of larger lvs 9-20 pairs and the longer pinna-rachises 1.2-2.2 cm; corolla
glabrous, or micropuberulent only at tip of lobes 20. C. parvifolia (p. 52)
5. Units of inflorescence pseudoracemose, the capitula fasciculate by 2-8 at nodes of
terminal and lateral, efoliate axes; androecium 6-11-merous, the longer filaments
3-7 m m 131. C. parviflora (p. 198)
4. Pinna-pairs of larger lvs 2-4. 7. Fls of each capitulum few (4-6) and large, the calyx 7.5-15 m m and the corolla
(10-) 14-21 mm, the whole perianth of thick texture and densely silky-tomentulose
overall 27. C. macrocalyx (p. 65)
7. Fls either more numerous or smaller, often both, the perianth of thin texture and
glabrous or only minutely puberulent.
8. Rachis of longer pinnae 1.4—8.5 cm; capitula 6-24-fld.
9. Lvs relatively small, the longer pinna-rachises 1.4—2 cm, and the larger lfts 3.5-5 mm; capitula mostly 6-10-fld 34. C. squarrosa (p. 72)
9. Lvs relatively large, the longer pinna-rachises 4—8.5 cm, and the larger
lfs 6.5-11.5 mm; capitula 14—24-fld 24. C. subspicata (p. 59)
8. Rachis of longer pinnae <1 cm; capitula 2—4(-6)-fld 36. C. depauperata (p. 73) 3. Pinnae of all lvs exactly 1 pair.
10. Lfts of longer pinnae 4-12 pairs, the larger blades 6.5-18 mm.
11. Larger lfts 12-18 x 6-11 mm; entering Maranhao from the n.-w.; a peripheral
form of:... . 38. C. surinamensis (p. 76)
11. Larger lfts 6.5-10.5 x 2.3-4 mm; Pernambuco 56. C. aeschynomenoides (p. 98)
10. Lfts of longer pinnae (8—)14—39 pairs, if <14 pairs then the blades <5 mm. 12. Longer pinna-rachises 3.5-8 cm, their lfts 20-39 pairs and up to 7-14 mm; capitula
sessile or almost so 52. C. sessilis (p. 92)
12. Longer pinna-rachises 0.3-2.4 cm, their lfts either fewer, or shorter, or both; capitula
usually pedunculate.
13. Capitula 9-30-fld; longest pinna-rachises 1.4—2.4 cm.
14. Pedicel of peripheral fls 1-2.3 mm; perianth thinly silky 54. C. duckei (p. 95) 14. Pedicel of peripheral fls at most 0.4 mm, usually 0; perianth glabrous
or distally micropuberulent 53. C. spinosa (p. 94)
13. Capitula 2-4(-6)-fld; longest pinna rachises 0.3-0.8(-l) cm 36. C. depauperata (p. 73) 2. Units of inflorescence umbelliform, the pedicel of peripheral fls 7-17 mm.
15. Upper stem and inflorescence eglandular; calyx-teeth linear ±4 mm, and nearly 4 times as
long as calyx-tube 32. C. imperialis (p. 70)
15. Upper stem and inflorescence charged with (sometimes minute) gland-capitate trichomes;
calyx-teeth 0.7-1.2 mm.
16. Lf-formula iii—vi/24—31; longer lfts 3-4.5 x 1-1.3 mm; perianth glabrous or minutely
puberulent at apex, eglandular; calyx ±1.2 mm, cleft to base 29. C. ulei (p. 68)
16. Lf-formula i-ii(-iii)/13-17; longer lfts 6-8.5 x 2.2-3 mm; perianth stipitate-glandular
overall; calyx ±1.6-2.4 mm, the teeth 0.7-1.1 m m 31. C. umbellifera (p. 70)
VII. Key to the species of Calliandra in the Brazilian state of Bahia s. and e. of rio
Sao Francisco (the presumed hybrid, C. x cumbucana, p. 162, omitted).
1. Stipules amply foliaceous 12-30 x 10-35 mm. 2. Lf-formula i/3-5; larger lfts 17-31 mm; pedicel of peripheral fls 1-2 cm; corolla <3 mm,
glabrous 132. C. leptopoda (p. 199)
19981 SILK TREE, GUANACASTE, MONKEY'S EARRING 15
2. Lf-formula v-viii/34-50; larger lfts 10-14.5 mm; pedicels all <1 mm; corolla 7.5-10 mm, densely white-silky 88. C. lanata (p. 152)
1. Stipules smaller, at most 12(-17) x 6(-7) mm. 3. Lf-formula i/P/2, each If exactly 6-foliolate; blade of 2 distal lfts 2-5 c m . . 75. C. harrisii (p. 126)
3. Lf-formula otherwise, if pinnae only one pair then lfts at least 2 full pairs and the blade of 2 distal lfts only exceptionally >2 cm.
4. Pinnae of larger lvs 3-13(-l 8) pairs.
5. Calyx-teeth <1.5(-l .9) m m , mostly depressed-deltate, or broad-ovate, or triangular-subulate, evidently shorter than the calyx-tube.
6. Pinnae of larger lvs 7—13 pairs.
7. Inflorescence lateral to primary axis of the season, the peduncles arising from
axillary, either thatched or ± extended brachyblasts; peduncles bracteate (the bract
sometimes lacking in C. parvifolia, but the inflorescence-type then unmistakable).
8. Fls relatively small, the calyx 1.8-4.3 m m , the corolla 4-9 m m ; perianth
either bronze-silky or glabrescent; androecium 20-50-merous.
9. Rachis of longer pinnae 6-9.5 cm; corolla 7-9 m m , bronze-silky 23. C. bella (p. 58)
9. Rachis of longer pinnae 1-2.2 cm; corolla 4-7 m m , thinly puberulent .... 20. C. parvifolia (p. 52)
8. Fls larger, the calyx (4-)4.5-l 1.5 m m , the corolla 10-21 m m ; perianth
densely white-silky; androecium (44-)50-180-merous. 10. Capitula 7-14-fld; calyx 4-8 m m 25. C. dysantha (p. 60)
10. Capitula 4-6-fld; calyx 7.5-11.5 m m 27. C. macrocalyx (p. 65)
7. Inflorescence terminally pseudoracemose, the units arising directly from distal
lf-axils and often, beyond lvs of current season, at bracteate efoliate nodes; peduncles ebracteate.
11. Lfts of longer pinnae 13-25 pairs; perianth densely red-granular and
sometimes in addition gray-pilosulous. 12. Calyx 3-8 m m and corolla 9-11 m m ; filaments ±100-200
per fl 87. C. bahiana (p. 151)
12. Calyx 1.8-2.5 m m and corolla 5-7 m m ; filaments 50-58 per fl.
13. Petiole proper, including pulvinus, 1.5-2 m m ; peduncles 3-10 m m ;
floral bracts ±1 m m ; filaments red 89. C.fuscipila (p. 152) 13. Petiole proper, including pulvinus, 4—7 m m ; peduncles 2-4 cm;
floral bracts 3.5-4.5 m m ; filaments white rubescent. . . 90. C. feioana (p. 154)
11. Lfts of longer pinnae 36-50 pairs; perianth subglabrous 85. C. lintea (p. 149)
6. Pinnae of larger lvs 3-6 pairs. 14. Inflorescence lateral to main axis of the season, the peduncles arising from
axillary, either thatched or ± extended brachyblasts; peduncles bracteate.
15. Units of inflorescence compactly racemose, 14-25-fld, the longer pedicels 1-4.5 m m ; lvs relatively ample, the rachis of longer pinnae 3-8.5 cm.
16. Pedicel of peripheral fls 2-4.5 m m ; larger lfts 3-4.5 m m ; androecium
±12-merous 30. C. pilgerana (p. 69) 16. Pedicel of peripheral fls 1-1.8 m m ; larger lfts 6.5-11.5 m m ;
androecium 28-46-merous 24. C. subspicata (p. 59)
15. Units of inflorescence capitulate, 6-10-fld, the pedicels not over 0.5 m m ;
lvs smaller, the rachis of longer pinnae 1.4—2 c m 34. C. squarrosa (p. 72)
14. Inflorescence terminally pseudoracemose, the units arising directly from
distal lf-axils and often, beyond lvs of current season, at bracteate efoliate
nodes; peduncles (nearly always) ebracteate. 17. Indumentum, especially of peduncles, consisting in part of minute,
pluriradiate stellae 103. C. stelligera (p. 166)
17. N o stelliform trichomes. 18. Petiole proper 10-17 m m ; rachis of longer pinnae 10-14.5 cm; larger
lfts 3-4 m m wide 95. C. elegans (p. 159)
18. Petiole proper (1-) 1.5-7 m m ; rachis of longer pinnae 1.6-7 cm; lfts
mostly 1-3 m m wide, in C. asplenioides to 2-4.5 m m wide but the
petiole then not over 4 m m . 19. Calyx mostly >3.5 m m and corolla 9-13 m m , the corolla densely
red-granular, sometimes in addition white-pilosulous; androecium
±100-200-merous 87. C. bahiana (p. 151)
19.Calyx 1.2-3 m m and corolla 5-7.5(-9) m m ; androecium commonly
30-50-merous, but up to 100-merous in C. nebulosa, which has
minutely ciliolate, non-granular perianth.
16 M E M O I R S OF T H E N E W Y O R K BOTANICAL G A R D E N [VOL. 74(3)
20. Diminutive, thicket-forming subshrubs 2-5(-6) dm;
perianth glabrous or remotely minutely puberulent;
filaments red 96. C. mucugeana (p. 160)
20. Shrubs 5-30 dm; filaments mostly white rubescent, red only
in C. hirsuticaulis, which has hirsutulous perianth.
21. Corolla red-granular and sometimes in addition pilosulous;
floral bracts 2-4.5 m m .
22. Petiole of longer lvs 4-7 m m ; peduncles 2-4 cm;
androecium white rubescent 90. C.feioana (p. 154)
22. Petiole of longer lvs 1-2 m m ; peduncles 0.4-1.6
cm; androecium red 99. C. hirsuticaulis (p. 163)
21. Corolla glabrous or only minutely pilosulous, not
red-granular; floral bracts 0.4-1.2 m m .
23. Pinnae of larger lvs 2-3(^4) pairs; lfts 2.3-4.5 m m
wide, 2.4—3.7 times as long; androecium
24-34-merous 91. C. asplenioides (p. 155)
23. Pinnae of larger lvs 4-6 pairs; lfts 1.2-2.4 m m wide,
mostly 3.7-5.4 times as long; androecium
104-112-merous 86. C. nebulosa (p. 150) 5. Calyx-teeth 2-7 (in rare C. macrocalyx var. aucta to 10) m m , either narrowly ovate, or
subulate, or triangular- to linear-lanceolate, at least as long as calyx-tube, commonly longer. 24. Calyx-teeth ±10 m m , as long as corolla; androecium ±9 cm; Sao Francisco valley
near 9°30'S; var. aucta of: 27. C. macrocalyx (p. 65)
24. Calyx-teeth at most 7 m m , usually surpassed by corolla; androecium 2-7 cm; widespread.
25. Corolla glabrous resinous ... 102. C. sincorana (p. 166)
25. Corolla either rusty-granular, or pilose-pilosulous, or both, or very exceptionally glabrous but then not resinous.
26. Peduncles 1-5-bracteate; larger lfts 11-20 x 3.5-8.3 m m ; calyx 5.6-9.5 (-12) m m and corolla 8-14.3 m m . . 92. C. viscidula (p. 156)
26. Peduncles with rare exceptions ebracteate; larger lfts 3-11 x 1-3.6 m m ; either
calyx or corolla shorter, or if these as long then the lfts decisively smaller. 27. Peduncles 2.5-8.5 cm.
28. Lfts of longer pinnae 17-37 pairs, plane or almost so; calyx
5-9.5 m m , its longest tooth 3-7 m m .
29. Rachis of longer pinnae 2-4 cm; lfts dorsally charged
with rufous granular trichomes, the larger lfts 5-7.5 m m 100. C. crassipes (p. 164)
29. Rachis of longer pinnae 4.5-7.5 cm; lfts smooth and glabrous on both faces, the larger lfts 7-12 m m . 101. C. hirtiflora (p. 165)
28. Lfts of longer pinnae 10-15 pairs, ventrally convex, dorsally concave, the margin revolute; calyx 3-3.5 m m , its longest tooth ±2.5 m m 105. C. involuta (p. 168)
27. Peduncles all, or all but the lowest ones, <2.5 cm.
30. Perianth white-pilose either overall, or in the distal half, not or
inconspicuously granular; longer lfts 4-8 mm.. . . 99. C. hirsuticaulis (p. 163) 30. Perianth densely granular, sometimes in addition minutely
pallid-puberulent but nowhere pilose; longer lfts
3-4.5 m m 104. C. coccinea (p. 167) 4. Pinnae of larger lvs either 1, or 2, or 1-2 pairs.
31. Capitula sessile or almost so 52. C. sessilis (p. 92) 31. Capitula evidently pedunculate.
32. Lvs extremely small, the rachis of longer pinnae 3-10 m m ; capitula 2-6-fld and
corolla ±3-4 m m 36. C. depauperata (p. 73)
32. Lvs larger, the rachis of longer pinnae well over 10 m m ; if fls as few then corolla larger. 33. Lfts of longer pinnae (11-)13 pairs upward.
34. Inflorescence lateral to main axis of the season, the peduncles arising from
axillary, either thatched or ± extended brachyblasts; peduncles bracteate.
35. Pedicel of peripheral fls 3-5.5 m m 55. C. blanchetii (p. 97) 35. Pedicel of peripheral fls <1 m m .
36. Long-shoots and some short-shoots tapering into a stout thorn; lfts
of longer pinnae 14-21 pairs; capitula 25-30-fld; local in middle
Contas valley near 14°40/S (n. into Ceard) . 53. C. spinosa (p. 94) 36. Neither long- nor short-shoots thorny; lfts of longer pinnae
20^t5 pairs; capitula 6-12-fld.
19981 SILK TREE, GUANACASTE, MONKEY'S EARRING 17
37. Calyx 2-4 mm; pinnae of some lvs 2 pairs; corolla striate
5.5-8 mm; rare species of n. Bahia 34. C. squarrosa (p. 72)
37. Calyx 0.9-2.2 mm; pinnae of all lvs exactly one pair; corolla
not striate 3.4-6.6 mm; widespread in Brazil s.-ward from
Minas Gerais, cultivated elsewhere, once recorded from
Bahia 50. C. brevipes (p. 90)
34. Inflorescence terminally pseudoracemose, the peduncles arising directly
from distal lf-axils and often, beyond lvs of current season, at stipulate
efoliate nodes; peduncles ebracteate.
38. Stipules either linear, or subulate, or if ovate then very small, none more that 2 m m wide at base.
39. Lfts of longer pinnae 14—30 pairs.
40. Larger lfts (2-)2.3-4.5 m m wide 91. C. asplenioides (p. 155) 40. Larger lfts 0.5-1.6 m m wide 97. C. calycina (p. 161)
39. Lfts of longer pinnae 37-50 pairs 108. C. longipinna (p. 170)
38. Stipules ovate, the larger ones 3.5-7 m m wide.
41. Calyx 5.5-10 mm; perianth thinly pilose distally.
42. Lfts of longer pinnae (11—)12—39 pairs, up to ±10-16 mm,
palmately 4—5-nerved from pulvinule.
43. Rachis of longer pinnae 8-13 cm, the lfts 32-39 pairs;
calyx 5.5-7 mm, finely multistriate; corolla 8.5-10 mm;
known from near 1000 m on Morro do Pai Inacio, n. fork
of rio Paraguacu near 12°30'S 111. C. paterna (p. 172)
43. Rachis of longer pinnae 4.5-6.5 cm, the lfts 11-17 pairs;
calyx 7.5-10 mm, faintly 5-nerved; corolla 12-12.5 mm;
known from ±1400-1500 m on Sa. da Tromba, near
sources of rio de Contas (mun. Piata and Abaira, 12°58'-13°16'S) and from Mucuge 112. C. ganevii (p. 173)
42. Lfts of longer pinnae 2-12 pairs, up to 5-8.3 mm, inconspicuously 1-nerved 107. C. renvoizeana (p. 169)
41. Calyx ±3 mm; perianth glabrous 109. C. debilis (p. 171)
33. Lfts of longer pinnae 2-12 pairs. 44. Inflorescence lateral to main axis of current season, the peduncles arising
from axillary brachyblasts 56. C. aeschynomenoides (p. 98)
44. Inflorescence terminal, pseudoracemose, the peduncles arising directly from
distal lf-axils and, beyond developed lvs of current season, from stipulate
efoliate nodes; peduncles ebracteate. 45. Lfts of longer pinnae 5-11 pairs.
46. Stems functionally herbaceous, arising yearly from slender rhizomes and <2 dm long. 114. C. semisepulta (p. 174)
46. Stems early frutescent (0.4-)0.5-3 m. 47. Pinnae of larger lvs 2(-3) pairs; lf-stks 10-21 mm; larger lfts
3.5-7 m m wide; androecium 42-44-merous.... 113. C. erubescens (p. 174)
47. Pinnae of all lfs exactly one pair; lf-stks 4.5-12 mm;
androecium 24—30-merous. 48. Larger lfts linear-elliptic 2.8-6 m m wide, 4—5.3 times as long,
remotely resin-dotted but not granular. ... 116. C. luetzelburgii (p. 176)
48. Larger lfts oblong-obovate 9-20 m m wide, less than twice
as long, densely rufous-granular dorsally. ... 115. C. germana (p. 175)
45. Lfts of longer pinnae 2-3 pairs; larger lfts 6-15 m m
wide . . 117. C. hygrophila (p. 176)
VHL Key to species of Calliandra in Planaltine and tropical Atlantic Brazil: states
of M a t o Grosso, M a t o Grosso do Sul, s. Tocantins, Goias with Distrito Federal,
trans-Franciscan Bahia, Minas Gerais, Espirito Santo, Rio de Janeiro (with Guanabara),
and tropical parts of Sao Paulo and Parana
1. Pinnae of some larger lvs, usually of all lvs, >1 pair. 2 Corolla 1.6-2.4 m m and androecium <1 cm; inflorescence composed entirely or in great part
of efoliate or only hysteranthously foliate pseudoracemes, the small capitula fasciculate at each
node on subfiliform peduncles of different lengths; lf-formula x-xxxvi/30-49; larger lfts
2-3.5 x 0.4-0.8 m m 131. C. parviflora (p. 198) 2 Corolla 4—15 mm; inflorescence otherwise; either pinnae or lfts fewer, or lfts larger, or both at once.
18 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
3. Stem-growth determinate, each unit of inflorescence solitary and terminal either to the
primary stem or to 1 to few branches; units of inflorescence umbelliform, the pedicel of outer
peripheral fls 5-26 m m ; barely entering the region in Mato Grosso do Sul, along the Paraguay border. 84. C. brevicaulis (p. 146)
3. Stem-growth indeterminate, each unit of inflorescence lateral, either to a primary stem or to an
axillary branchlet, the units often paired or fasciculate at a node; units of inflorescence either
capitulate or umbelliform, if umbelliform the longer pedicels <5 m m . 4. Stipules (deciduous, but some evident on all flowering stems and at distal nodes of most
fruiting ones) cymbiform, elliptic 4-17 m m , papery, striately many-nerved, glabrous;
peduncular bract of similar texture, 3-9 m m . 5. Each capitulum subtended by a coevally expanding If; androecium bicolored,
pallid proximally, pink-carmine disially 21. C. foliolosa (p. 55)
5. Each capitulum subtended by an efoliate pair of bracteiform stipules; androecium
uniformly red 22. C. tweedii (p. 56)
4. Stipules few-nerved or obscurely nerved, or in outline lanceolate or linear, never papery
nor cymbiform; peduncular bract smaller or lacking. 6. Corolla densely white- or bronze-silky, the surface concealed or almost so by
indumentum.
7. Subshrubs mostly <1.5 m, with stout erect stems arising annually from short
caudex or xylopodium; androecium 44-100-merous; the tassel opening and
remaining scarlet-crimson; pod densely white- or gray-tomentulose overall 25. C. dysantha (p. 60)
7. Arborescent shrubs and treelets, at maturity 2-6 m; with persistent branches; androecium 26^6-merous, the tassel opening white, some-fading pinkish; pod
either glabrate or brown-silky.
8. Rachis of longer pinnae 5.5-9.5 cm, and their lfts 40-64 pairs, the longer
ones to 5-9.5 m m ; pod brown-silky; Atlantic Forest in Bahia, s. perhaps
into n. Espirito Santo 23. C. bella (p. 58) 8. Rachis of longer pinnae 1.3-2 cm, and their lfts 24-29 pairs, the
longer ones 2.5-3.5 m m ; pod glabrate; carrasco of n. Minas Gerais 20a. C. carrascana (p. 55)
6. Corolla either glabrous, or thinly strigulose, or puberulent, but the indumentum
(if any) not concealing the surface. 9. Lfts relatively large, the longer ones (5-)6-15 m m .
10. Arborescent shrubs 2-12 m. 11. Pinnae of larger lvs 3-5 pairs; larger lfts ±10-20 x 3-8 m m ; calyx of
peripheral fls 1.4—2.6 m m and corolla 5.5-8 m m ; androecium (8-)
10-14-merous; feebly entering Mato Grosso from the north; var. stipulacea of: 6. C. laxa (p. 29)
11. Pinnae of larger lvs (l-)2-3 pairs; larger lfts 9-12 x 1.8-2.6(-3) m m ;
androecium 22-36-merous.
12. Lfts of longer pinnae 27-37 pairs; calyx of peripheral fls 4—4.5 m m and corolla 8.5-12.5 m m ; local in centr. Minas
Gerais 35. C. glaziovii (p. 73) 12. Lfts of longer pinnae 12-17 pairs; calyx of peripheral fls
scarcely 2 m m and corolla 4-5 m m ; Goias and Distrito
Federal 37. C. silvicola (p. 75) 10. Either shrubs at most 2 m or functionally herbaceous subshrubs.
13. Lf-formula (i—)ii/43—51; local in centr. Minas Gerais 110. C. iligna (p. 171)
13. Lf-formula variable but, if pinnae as few, then lfts <(35-)25 pairs; both sympatric and allopatric.
14. Dwarf rhizomatous shrublet of Sa. do Cipo in centr. Minas
Gerais, the fertile stems of the season 5-15 cm; larger lfts
0.8-2 m m wide 94. C. linearis (p. 158)
14. Variable in stature, but if resembling the last in that respect the
larger lfts >3 m m wide and dispersal different.
15. Stems frutescent, branched distally, 5 d m upward, densely
multifoliate; corolla at most 9 m m ; mountains of n. and centr. Minas Gerais.
16. Lfts linear or linear-lanceolate from auriculate base,
the larger ones 1-2.3 m m wide, 4.8-8 times as long;
floral bracts usually 2.5-15 m m , but sometimes minute;
lfts not or only sparsely and obscurely resin-dotted
dorsally 93. C. fasciculata (p. 156)
19981 SILK TREE, GUANACASTE, MONKEY'S EARRING 19
16. Lfts narrowly oblong from shallowly cordate or subtruncate base, the larger ones 2.4—4.3 m m wide, 2.4—4 times as long;
floral bracts 0.6-1.2 m m ; lfts evidently and often densely
resin-spotted or papillate dorsally 91. C. asplenioides (p. 155)
15. Stems herbaceous, simple, 2-3 dm, ±3-5-foliate; corolla
13-17 m m ; e.-centr. Goias 26. C. gardneri (p. 64) 9. Lfts minute, the largest 2-5.5 x 0.5-1 m m .
17. Pinnae of larger lvs 5-18 pairs, ascending from rachis; rachis of longer
pinnae 12-22 m m ; fls of most capitula heteromorphic; pedicel of peripheral
fls 0.7-2.4 m m ; widespread 20. C. parvifolia (p. 52)
17. Pinnae of larger lvs 3-5 pairs, divaricate from primary lf-stk at ±90°; rachis
of longer pinnae ±2.5-5 cm; fls homomorphic; pedicel of peripheral fls
1.5-4 m m ; local in centr. Minas Gerais 33. C. concinna (p. 71) 1. Pinnae of all lvs exactly 1 pair.
18. Lfts of each pinna not less than 9 pairs, and blade of longest ones no more than 1.4 cm.
19. In longer pinnae the lfts 70-88 pairs; calyx weakly 5-nerved and corolla nerveless; local near
Diamantina, centr. Minas Gerais 106. C. santosiana (p. 169)
19. In longer pinnae the lfts at most 45 pairs; at least the calyx striate.
20. Capitula sessile or almost so, the peduncle not more than 2 m m , commonly obsolete;
corolla striate like the calyx 52. C. sessilis (p. 92)
20. Capitula evidently pedunculate, the peduncle 5 m m upward; calyx sharply striate but
corolla only faintly venulose. 21. Peduncles 0.5-2.2 cm; no intrastaminal nectary in peripheral fls; shrubs and trees
1.5 m upward. 22. Lf-stk of longer lvs (10-)13-23 m m ; lfts of longer pinnae 12-17 pairs, the
longer ones 9.5-11.5 m m 37. C. silvicola (p. 75)
22. Lf-stk of longer lvs 1.5-7 m m ; lfts of longer pinnae 20-45 pairs, the longer
ones 2.3-9 m m . 23. Stipules 0.6-2 m m ; larger lfts 3.5-6 m m ; calyx of peripheral fls
0.9-1.8 m m 50. C. brevipes (p. 90)
23. Stipules 2.5-6 m m ; larger lfts 7-9 m m ; calyx of peripheral fls
2-2.8 m m 51. C. staminea (p. 92)
21. Peduncles 2.5-11 cm; an intrastaminal nectary in all fls; campo subshrubs
3-7 d m 124. C. virgata (p. 189)
18. Lfts of each pinna \V*-5 pairs, and blade of distal lfts ample, 1.7-10 cm. 24. Stipules ovate-deltate, at most 3 m m ; lfts of all pinnae exactly Wi pairs; pedicel of peripheral
fls 0-1.5 m m ; androecium 1.5-6 cm. 25. Slender shrubs or treelets at least 1 m tall; mature lf-stks 1.5-3.5 cm; capitula arising
from first 1-2 nodes of reactivated brachyblasts, each on peduncle \-A cm. . 75. C. harrisii (p. 126)
25. Functionally herbaceous subshrubs dying back annually to an oblique caudex; mature
lf-stks 4.5-7.5 cm; capitula arising from buds at or near ground-level on scapiform peduncle 6-18 cm, the hysteranthous foliate stems only 1-3 d m 81. C. longipes (p. 137)
24. Stipules foliaceous, reniform-suborbicular 12-26 m m ; lfts of longer pinnae 3-5 pairs; pedicel of peripheral fls 10-20 m m ; androecium ±6-7 m m 132. C. leptopoda (p. 199)
IX. Key to species of Calliandra in s. South America: Bolivia, Paraguay, extratropical
Brazil, Argentina, and U r u g u a y
1. Pinnae of each If (2-)3-many pairs. 2. Units of inflorescence umbelliform (longer pedicels 5-26 m m ) and terminal to the homotinous
stem or to its few branches; functionally herbaceous subshrub in campos of cis-Paraguaian
Paraguay and of adj. Brazil and n.-e. Argentina (Corrientes, Misiones) 84. C. brevicaulis (p. 146)
2. Units of inflorescence either capitulate or umbelliform, but always borne laterally to the
homotinous stem (either in primary lf-axils, or on axillary brachyblasts, or terminally
pseudoracemose), if umbelliform the pedicels exceptionally >5 m m ; habit and dispersal various.
3 Pinnae of larger lvs 14-36 pairs; units of inflorescence pseudoracemose, the crimson fls very
small, the calyx 0.8-1.3 m m , the glabrous corolla 1.6-2.4 m m ; androecium 6-11-merous, the
filaments 3-7 m m 131. C. parviflora (p. 198) 3. Pinnae of larger lvs either 2-9 pairs, or in C. parvifolia to 18 pairs, but the corolla then
>4 m m and the units of inflorescence not pseudoracemose; androecium (12-)14-106-merous,
the filaments 20-55 m m . 4. Stipules 1.5-6 m m ; calyx 1.8-3.3 m m ; corolla glabrous or only thinly puberulent at
tip of lobes 20. C. parvifolia (p. 52)
20 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
4. Stipules (4-)6-17 mm; calyx 3.6-7.5 mm; corolla densely silky-pilose externally.
5. Virgate, functionally herbaceous subshrubs 2-10 dm; stipules lanceolate, firm,
dorsally pubescent at least when young; androecium 90-106-merous; s.-e. Paraguay,
along the Brazilian border; var. opulenta of: 25 C. dysantha (p. 60)
5. Arborescent shrubs 1-8 m, sometimes dwarfed in shallow soils; stipules
(deciduous, but some evident on flowering stems and at distal nodes of fruiting
branches) papery, cymbiform, elliptic 4-17 m m , striately nerved, glabrous; androecium 26-46-merous.
6. Each peduncle subtended by a coevally expanding If; androecium pallid proximally, the tassel pink-carmine 21. C.foliolosa (p. 55)
6. Each peduncle subtended by an efoliolate pair of bracteiform stipules;
androecium bright red throughout 22. C. tweedii (p. 56) 1. Pinnae of all lvs exactly 1 pair.
7. Lfts of each pinna at least 3 pairs.
8. In longer pinnae the lfts 7-45 pairs, the largest 3.5-15 m m ; capitula <20-fld; androecium
bicolored, pallid proximally, carmine distally, the orifice of the androecial tube naked within.
9. Rachis of longer pinnae 3-5 cm and lfts 7-9 pairs, the larger ones 9-15 x 4-6 m m ; submontane Bolivia (La Paz, 980-1300 m ) 44. C. chulumania (p. 85)
9. Rachis of longer pinnae 1.5-3.5 cm and lfts 20-45 pairs, the larger ones 3.5-6.5 x 0.4—1.5 m m ; lowland s. Paraguay e.-ward 50. C. brevipes (p. 90)
8. In longer pinnae the lfts 3-9 pairs, the largest 22-150 m m ; androecium red throughout, the orifice of the androecial tube fringed internally with incurved staminodia 65. C. haematocephala (p. 108)
7. Lfts of each pinna exactly 1, \l/i, or 2 pairs.
10. Arborescent shrubs, not dying back annually to the caudex; units of inflorescence either axillary to primary lvs, or borne on axillary thatched brachyblasts; androecium <50-merous.
11. Venation of lfts palmate-pinnate, the inner primary nerve posterior to midrib manifestly longer and stronger than succeeding secondary ones, incurved-ascending at least to, commonly
well beyond mid-blade; pod bifacial, the recessed valves wider than the sutural ribs. 12. Blade of distal pair of lfts elliptic-oblanceolate ±2-4 x 0.5-1.2 cm; androecium
9-10-merous; pods stiffly erect . . 74. C. angustifolia (p. 125)
12. Blade of distal pair of leaflets obliquely ovate-acuminate 6-16 x 3-6.5 cm;
androecium 12-30-merous; pods either plagiotropic or geotropic 68. C. trinervia (p. 113)
11. Venation of lfts pinnate, the primary nerve posterior to midrib no longer than the
secondary ones; pod obtusely tetragonal, the broad sutural ribs much wider than the
recessed valves ... 75. C harrisii (p. 126) 10. Functionally herbaceous subshrubs from slender rhizome, the inflorescence precocious,
scapiform, arising from lowest nodes of hysteranthously foliate stems, these dying back or burned back annually; androecium 66-82-merous; campos of cis-Paraguaian Paraguay
and adj. Argentina 81. C. longipes (p. 137)
X. Key to some cultivated species of Calliandra
1. Pinnae of larger lvs, commonly of all lvs, 2 pairs or more.
2. Inflorescence-units axillary to a If or to a pair of stipules, lateral to the main stem or to secondary
branches. 3. Lfts of longer pinnae 24-70 pairs.
4. Stipules papery brownish, navicular, (4-)7-17 m m long.
5. Each peduncle subtended by a coevally expanding If; androecium pallid
proximally, the tassel crimson 21. C.foliolosa (p. 55)
5. Each peduncle subtended by a pair of bracteiform stipules; androecium
crimson-carmine throughout 22. C. tweedii (p. 56)
4. Stipules either linear, or subulate, or narrowly ovate, plane or terete, 1.3-8 x 0.8-4 m m . 6. Rachis of longer pinnae (2.5-)3-9.5(-14) cm.
7. Larger lfts 2-8 m m wide 6. C. laxa (p. 29) 7. Larger lfts 0.5-2(-2.4) m m wide.
8. Calyx 5-nerved or almost nerveless; androecium 26-44-merous, the
filaments usually crimson distally 1. C. pittieri (p. 22)
8. Calyx delicately 15-20-nerved; androecium 26-44-merous, the filaments
opening white, sometimes fading pinkish 23. C. bella (p. 58)
6. Rachis of longer pinnae 1-2.2 cm 20. C. parvifolia (p. 52)
3. Lfts of longer pinnae 5-24 pairs.
9. Lf-stk of longer lvs 2.5-9.5(-l 1) cm and rachis of longer pinnae (3-)4-10.5 c m 6. C. laxa (p. 29) 9. Lf-stk of longer lvs <3 cm and rachis of longer pinnae to 2.6 cm.
10. Capitula mostly 7-13-fld; filaments crimson throughout 15. C. californica (p. 42)
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 21
10. Capitula 3-7-fld; filaments pallid proximally, crimson distally 16. C. eriophylla (p. 44)
2. Inflorescence a terminal, efoliate, simple pseudoraceme of capitula or a few-branched panicle.
Lf-formula vi-iv/24-69 118. C. houstoniana (p. 177) 1. Pinnae of all lvs exactly 1 pair.
11. Stipules linear-subulate to lance-ovate, unarmed.
12. Each pinna 4—plurifoliolate, forming at least 2 opposite pairs.
13. A row of sterile filaments forming a retrorse fringe within the androecial tube capitula 20-84-fld 65. C. haematocephala (p. 108)
13. Filaments all erect and exserted from corolla; capitula 7-24(-26)-fld.
14. Lfts of each pinna 20-45 pairs and the largest lfts <6 m m 50. C. brevipes (p. 90)
14. Lfts of each pinnae <21 pairs and the largest lfts at least 8.5 mm. 15. Lfts of longer pinnae 7-19 pairs.
16. Lfts in outline broad-linear or linear-elliptic, 3.5-6 times as long as wide;
seed-coat with pleurogram 41. C. riparia (p. 80)
16. Lfts in outline obovate, oblong, rhombic-ovate or -obovate, 1.5-2.3 times
as long as wide; seed-coat without pleurogram 38. C. surinamensis (p. 76) 15. Lfts of longer pinnae 2-7 pairs.
17. Both calyx and corolla sharply striate-nerved 61. C. medellinensis (p. 103)
17. Calyx sometimes striate, but corolla not or faintly so.
18. Lfts strongly accrescent distally, the longer ones (1.7-)2-9 x 1-4 cm.
19. Larger stipules 0.6-1.3 m m wide; inflorescence immersed in
foliage, the individual capitula axillary (either directly, or on
obscure brachyblast) to coeval lvs 63. C. guildingii (p. 104)
19. Larger stipules 1.5-2.8 m m wide; inflorescence pseudoracemose
and terminal to principal stem and branches, the individual capitula arising from stipulate but efoliate nodes 64. C falcata (p. 105)
18. Lfts not or scarcely accresent distally, the longer ones 0.8-2 x 0.4—1.1 cm 40. C. purpurea (p. 78)
12. Each pinna either 2- or 3-foliolate, the lfts 1 terminal pair with or without a solitary 1ft toward
base of rachis on its proximal side. 20. Lfts subsymmetrically elliptic; both calyx and corolla striate; androecium
38-58-merous 73. C. antioquiae (p. 123) 20. Lfts obliquely obovate or elliptic, or inequilaterally oblanceolate; calyx striate but corolla
not so; androecium 20-28-merous. 21. Pod linear, obtusely 4-angulate, 5-6 m m diam, the sutures dilated and as wide as
the pod itself 75. C. harrisii (p. 126) 21. Pod bifacially compressed, the sutures much narrower than the valves, these
6-14 m m wide 76. C. tergemina (p. 127)
11. Stipules charged dorsally at base with a conic spur or a stiff spinule resembling a spinescent
stipule 82. C. haematomma (p. 140)
A Note o n M e t h o d of Citation
The nomenclature of all taxa is presented in
paragraphs, all elements of which are homotypic
synonyms, cited serially by date of publication. The
authorship of each accepted name is given in full, but
the original author of each epithet is not repeated in
derived combinations. References following the word
sensu are bibliographic, not nomenclatural.
I. Sectio ANDROCALLIS Barneby
Calliandra sect. Androcallis Barneby, sect, no v., in-
florescentiae structura imprimis definienda: florum
capitula seu umbellae cauli contemporaneo lat
erals, aut e foliorum primariorum axillis aut e
brachyblastorum axillis nascentes, nunquam
pseudoracemum terminalem aphyllum efformantes.
Sp. typica sub ser. homonyma indicatur.
Inflorescence lateral to long-shoots of current sea
son, the peduncle of each unit arising either directly
from a contemporary lf-axil or from a (foliate or efo
liate, but stipulate) axil of either a contemporary or an
annotinous brachyblast, the units together never
assembled into a terminal efoliate pseudoraceme.
I/A. Series ANDROCALLIS Barneby
Calliandra sect. Androcallis Barneby ser. Andro
callis, ser. nov. Omnia sectionis suae, sequentia
autem particularia: folia semper bipinnata, micro-
phyllidia; phyllotaxis disticha; stipulae inermes;
capitula haud subscaposa. — Sp. typica: C. laxa
(Willdenow) Bentham.
Calliandra ser. Nitidae Bentham, London J. Bot. 3: 103.
1844. — Sp. lectotypica: C. brevipes Bentham
Calliandra ser. Nitidae sensu Bentham, 1875: 545, max. ex
parte et spp. gerontogaeis exceptis.
22 M E M O I R S OF THE N E W Y O R K BOTANICAL G A R D E N [VOL. 74(3)
Characters of the section; phyllotaxy distichous;
lvs bipinnate microphyllidious; stipules not spines
cent. — Spp. 56, in dispersal almost coextensive with
the genus.
1. Calliandra pittieri Standley, Contr. U.S. Natl.
Herb. 18: 102. 1916.—Typus infra sub var. pittieri
indicatur.
Elaborately microphyll, arborescent shrubs and trees
2—12(—15, -20) m tall with plagiotropic branches and
consequently low-convex or flattened crown (often
leaning over water), the young branches and lf-axes
appressed-puberulent or pilosulous with either gray
or brownish hairs to 0.15-0.7 m m , the crowded, firm,
facially glabrous but often thinly ciliolate lfts bicol
ored, when mature lustrous olivaceous above, paler
dull beneath, the hemispherical capitula arising
singly, or geminate, or rarely 3 together from either a)
nodes of either foliate or efoliate brachyblasts axil
lary to coeval primary lvs, or b) directly from lf-axils
of contemporary long-shoots; phyllotaxy distichous.
Stipules narrowly to broadly lanceolate or narrowly
ovate 2-10 x 0.8^- m m , striately 7-11-nerved when
young, thickened dry in age, either glabrous or pilo
sulous dorsally, deciduous. Lf-formula (vi-)vii-xiii/
(35-)37-62(-70); lf-stks randomly variable in vigor,
(4-)5-14(-15) cm, the petiole, including pulvinus,
(4-)5-30(-33) m m , at middle 0.6-1.2 m m diam, the
longer interpinnal segments (1.5—)2— 14(—25) m m , the
ventral groove either continuous or weakly bridged;
pinnae usually a little accrescent distally, sometimes
subequilong, the rachis of longer ones (2.5-)3-9(-14)
cm, the longer interfoliolar segments 0.3-14(-2.2)
m m ; lft-pulvinules 0.1-0.35 m m ; lft-blades linear or
linear-lanceolate from obtusangulate or shortly ob
tusely auriculate base, straight or incipiently falcate,
acute or subobtuse, the larger ones (2.7—)3—10.5(—13)
x (0.5-)0.6-2.4(-2.8) m m , (4.3-)4.8-6.6 times as
long as wide; midrib subcentric or forwardly dis
placed to divide blade ±1:1.5-2, either simple or (in
larger lfts) giving rise to faint secondary venules. Pe
duncles 1.5-6(-7.5) cm, often appearing ebracteate,
but the peduncular bract inserted close under the ca
pitulum and scarcely distinguishable from floral
bracts; capitula 14-28-fld, the receptacle, including
short terminal pedestal, 2-4 m m diam; bracts oblong
or subulate 0.6-1.4 m m , tardily deciduous; fls (al
ways, so far as known) heteromorphic, the perianth of
all 5-merous, brownish-puberulent or glabrous, the
calyx 5-nerved or almost nerveless, the corolla also
externally nerveless; P E R I P H E R A L FLS: pedicel
broadly turbinate, or drum-shaped, sometimes as wide
as base of calyx but then differentiated by discolora
tion (and solid in longitudinal section) 0.3-2(-2.3) x
0.4-0.9 m m ; calyx 1.3-3.5 m m , the subulate or de
pressed-deltate teeth 0.3-0.6 m m ; corolla (5-)5.5-9
(-9.5) m m , the ovate lobes (1.1—)1.4—3.2 m m ; an
droecium 12-26(-28)-merous, (2.7-)3-6.2 cm, the
tube (2-)3.6-9 m m , the stemonozone 0.9-2.3 m m ,
the tassel usually pallid at base and crimson distally,
rarely crimson throughout; T E R M I N A L FL(s): peri
anth nearly of the peripheral fls but usually broader;
androecial tube broadly cylindric, ± twice as long as
corolla. Pods 7—13(—18) cm, in Colombia mostly
1-1.25 cm, in Venezuela 1-1.8 cm, in Ecuador to 1.6
cm wide, when well fertilized 5-8-seeded, the brown
woody valves coarsely densely cross-venulose,
minutely or densely puberulent; seeds discoid 9-13 x
7-10 m m , the papery testa dull brown, pleurogram 0.
Key to the varieties of C. pittieri
1. Rachis of longer pinnae 3-6.5 cm, and larger
lfts 3-6.5 x 0.6-1.2 mm; s.-e. Panama s. along the w. and centr. Cordilleras to
Ecuador. la. var. pittieri 1. Rachis of longer pinnae 5-9(-14) cm,
and larger lfts 5.5-10.5 x 1.1-2.4 mm;
Venezuela n. of the Orinoco (w.-ward from
Anzoategui) and local in n. Antioquia,
Colombia lb. var. polyphylla
la. Calliandra pittieri Standley var. pittieri. C. pit
tieri Standley, 1916, I.e., sens. str. — "Type . . .
[US] 531146, collected at La Esmeralda, near
Jamundi, Cauca valley, State [sic] of Cauca,
Colombia, at... 1,200 meters, January, 1905, by H.
Pittier (no. 951)." — Holotypus, US!; isotypus
(fragm), NY!. — Anneslia pittieri Britton & Rose,
N. Amer. Fl. 23: 63. 1928.
C. bella var. trianae Bentham, Trans. Linn. Soc. London 30:
556. 1875. — "New Granada, Triana, Jervise, Schlim n.
783." — Lectotypus, Triana 467, K! = N Y Neg. 2000; pos
sible isotypus, Hb. Lehmann. 5367, F!. — Equated with C. pittieri by Britton & Killip, 1936: 136.
C. lehmannii Harms, Repert. Sp. Nov. Regni Veg. 17: 90.
1921. — "Columbia: ... am oberen Rio Bagua, West-
Anden von Cali, 1200-1300 m (Lehmann 5267. . .). —
Campo Alegre (Langlasse no. 26 . . .)." — Lectotypus,
Lehmann 5267 ̂ B = F Neg. 1245V, isotypi, K! (= N Y Neg.
1999\), F (cf. above)!, N Y (fragm)!; isoparatypus, Lan
glasse 26, NY!. — Equated with C. pittieri by Britton & Killip, 1936: 136, pi. 1.
Characterized by relatively short pinnae and lfts, as
given in key to varieties, and distinguished from var.
polyphylla by almost fully allopatric dispersal.
In disturbed woodland, seasonally dry woodland,
and colonial along sunny stream banks, surviving
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 23
deforestation in hedges and pastures, occasionally cul
tivated in its native range, (800-) 1100-3000 m, lo
cally plentiful on both slopes of Cordilleras Occiden
tal and Central in Colombia, n. at lower elevations into
e. Panama (Darien) and s. into Ecuador (Esmeraldas,
Carchi, Imbabura, Pichincha, Cuenca). — Fl. inter
mittently through the year, new fls often coinciding
with old pods. — Map 1. — Carbonero, carbonerito;
pluma de ydtero (Colombia); tura (Ecuador).
lb. Calliandra pittieri Standley var. polyphylla
(Harms) Barneby, stat. nov. Calliandra polyphylla
Harms, Notizbl. Bot. Gart. Berlin-Dahlem 8: 51.
1921. — "Venezuela: Carabobo, Guaremales,
Strasse von P[uerto] Cabello nach San Felipe . . .
10-100 m (PITTIER n. 8861 — V.-VI. 1920)." —
Holotypus, presumably *B; isotypus, NY!.
C. rupicola Pittier, Arb. Arbust. Venez. 4/5: 49. 1923. — "en
tierra caliente, cerca de Curucuti, D.F., (P[ittier] 10.221)."
— Holotypus, presumably V E N (n.v.); isotypus, NY!.
C. porphyrea Pittier, Bol. Soc. Venez. Ci. Nat. 4(30): 81.
1937. — "[Venezuela.] Aragua: Parque Nacional, cerca del
punto culminante de la Carretera a Choroni, a 1600 m. de
altitud; flores Febr. 19, 1937 (Pittier 13925 . .)." — Holo
typus, presumably V E N (n.v.); isotypi, K!, NY!.
Distinguished feebly from var. pittieri by prevail
ingly longer pinna-rachises and mostly longer and
slightly broader lfts, as specified in key to varieties.
In seasonally dry, virgin and second-growth, upland
forest and along sunny riverbanks in more humid for
est, 400-1600 m, on the n. slope of Cordillera Costan-
era in Venezuela descending to Caribbean lowland dry
forest near 10 m, discontinuously dispersed in n.
Venezuela from Anzoategui to Carabobo, Trujillo and
w. Barinas; apparently disjunct in Colombia on the
slope of Cordillera Oriental in s. Boyaca and on trib
utaries of lower rio Cauca in Antioquia. — M a p 1. —
Fl. II-IX, perhaps intermittently through the year. —
Clavellino, a name applied also to related species.
2. Calliandra purdiaei Bentham, London J. Bot. 4:
104. 1846. — "Mountains of Ocana, Columbia,
Purdie." — Holotypus, K (hb. Hook.)! = N Y Neg.
200T, isotypus, K (hb. Benth.)!. — Feuilleea pur
diaei O. Kuntze, Revis. Gen. PI. 1: 188. 1891.
C. clavellina Karsten, Fl. Columb. 1(4): 159, t. LXXIX.
1861. — "Crescit in Cordillera Granatensi, prope oppidum
Ocana, altitudine 1000 metr." — No typus seen, but the
ample description and superlative illustration are decisive.
— Equated with C. purdiaei by Bentham, 1875: 556.
Arborescent shrubs (1.5-)2-8 m, closely resem
bling C. pittieri in habit, phyllotaxy, vesture and inflo
rescence, the peduncles mostly arising from primary
lf-axils. Stipules 4-11 x 1.7-3 m m , 7-13-nerved when
young. Lf-formula vii—xiii/(33—)35—52(—60); lf-stks
(6-)8-17 cm, the petiole 7-25(-30) m m , at middle
0.6-1.4 m m , the longer interpinnal segments (6-)
14-19 m m ; rachis of longer pinnae (2.5-)3.5-6.5 (-8)
cm, the longer interfoliolar segments 0.4—1.4 m m ; lft-
pulvinules 0.1-0.25 m m ; lft-blades linear or linear-
lanceolate from obtusangulate or subauriculate base,
straight or subfalcate, mostly acute, the larger ones
(4.5-)4.7-8(-10) x 0.7-1.5 m m , 4.4-6.6 times as long
as wide; venation of C. pittieri. Peduncles (2-)2.5-5.5
cm, bracteate as C. pittieri; capitula 18—33-fld, the
receptacle including short pedestal 2.5-5.5 m m diam;
bracts subulate or linear 1.3-6 m m , tardily deciduous;
fls heteromorphic; perianth 5-merous, thinly puberu
lent or glabrous, the calyx striate, the corolla exter
nally nerveless; P E R I P H E R A L FLS: pedicel (0.7-)
1.2-2.7 m m ; calyx (4-)5.3-7.2 x 1.3-3.5 m m , the sub
ulate or lanceolate teeth 1.5-3.2 m m ; corolla 10-11.5
m m , the ovate lobes 2.2-3.5 m m ; andreocium 28^-0-
merous, 4.4-6.5 cm, the tube 7-11 m m , the stemon-
ozone (0.8-) 1.2-2 m m , the tassel carmine throughout;
C E N T R A L FL(S): perianth nearly of peripheral fls but
sometimes broader; androecial tube broadly cylindric,
± twice as long as corolla. Pods (few seen) 9-12 x
1-1.3 cm, 5-8-seeded, venulose and thinly puberulent
as in C. pittieri; seeds 9-11 x 6.5-8.5 m m , the testa
papery, dull brown, pleurogram 0.
In seasonally dry, semideciduous and moist ever
green forest, sometimes along streams, surviving
disturbance in hedgerows and in pasture thickets, 50-
980 m, locally plentiful near the w. margin of Mara-
caibo Basin in Zulia, Venezuela, and in valleys of
Cordillera Oriental in Norte de Santander and San-
tander, Colombia; in Colombia known also in culti
vation on the n. slope of Sa. de Sta. Marta, in middle
Magdalena valley (Mariquita, Melgar, Quindio) and
near Villavicencio, probably elsewhere. — M a p 2. —
Fl. XI-V. — Cujicito.
Calliandra purdiaei differs from closely related C.
pittieri in the longer calyx, and in slightly more nu
merous filaments that are red throughout.
3. Calliandra glomerulata Karsten, Fl. Columb. 2: 5.
1862. — T y p u s infra sub var. glomerulata indicatur.
Potentially arborescent, bushy or broad-crowned,
microphyllidious shrubs flowering when (1—)2—8
(once reported -18) m tall, the young stems, lf-axes
and peduncles pilosulous with shorter curved and
long ± spreading, gray or bronze hairs to 0.2-1.2 m m ,
rarely glabrescent, the multifoliolate lvs strongly bi
colored, the lfts above dull lurid gray and papillate or
24 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
M A P 1. Distribution of Calliandra pittieri Standley var. pittieri and var. polyphylla (Harms) Barneby in northwestern South America.
papillate-puberulent or dull dark brown and smooth,
commonly but not invariably pilosulous or puberulent
beneath, the relatively few-fid capitula of ± silky fls at
first solitary or fasciculate in primary lf-axils, later
pseudoracemose along short efoliate (but not thatched)
terminal or lateral axes, the densely silky-pilose pods
erect; phyllotaxy distichous. Stipules firm, ovate or
lanceolate 1-6.5 x 0.6-2 m m , usually dorsally pubes
cent and weakly nerved when young, becoming dry,
pallid, glabrate, tardily deciduous. Lf-formula iii-xi
(-xiii)/16-36; lf-stks of primary lvs 1.2-9(-14) cm, the
petiole including pulvinus 3.5—14(—16) m m , at middle
0.4-1.2 m m diam, the longer interpinnal segments
3—10(—11) m m ; pinnae accrescent from base upward
or the furthest pair sometimes abmptly shorter than
those next below, the rachis of longer ones
(1—)1.2—6(—8) cm, the longer interfoliolar segments
0.4-1.4(-l.7) m m ; lfts usually decrescent near each
end of rachis and subequilong between, the pulvinules
0.1-0.3 x 0.2-0.35 m m ; lft-blades oblong to broad-lin
ear from semicordate base, straight or almost so,
broadly obtuse or depressed-deltate at apex, those near
mid-rachis 3.4-7.5 x l-1.9(-2.4) m m , (2.2-)2.4-4-
(-4.6) times as long as wide; venation of lfts externally
simplified, the midrib scarcely excentric, ventrally
immersed, finely prominulous dorsally, often simple,
sometimes faintly branched on each side, the posterior
primary nerves usually imperceptible. Peduncles 1-3
per node, 4-18 m m , at base discolored quasi-pulvinu-
late, ebracteate, disarticulating if not fertile but com
monly retained on the branch by entangled filaments;
capitula 5-9(-10)-fld, the receptacle including very
short terminal pedestal ±1-2 m m diam; floral bracts
0.3-1.5 m m , persistent; pedicels (sometimes imper-
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 25
M a p 2. Distribution of Calliandra tolimensis Taubert, C. purdiaei Bentham, and C. glomerulata Karsten var. glomerulata
and var. parvifolia (Bentham) Barneby in northern South America.
ceptible in external view) 0.3-1 x 0.5-1.1 m m ; fls
either homomorphic as to androecium and then either
staminate or bisexual, with short androecial tube and
lacking nectary, or dimorphic, one central fl broader
but scarcely longer than the peripheral ones, staminate,
with long-exserted androecial tube and lobed intrasta
minal disc 0.45-1.5 m m , the perianth of either sort of
flower either 4- or 5-merous, the calyx glabrous or pu
berulent distally, the corolla at least thinly, commonly
densely white-silky externally; P E R I P H E R A L FLS:
calyx campanulate or turbinate-campanulate (1.4-)
1.8-3.2 x 1.2-2.3 m m , weakly 8-10-nerved, the subu
late, ± incurved teeth 0.35-0.8 m m ; corolla white,
4.5-7.1 m m , the ovate-lanceolate lobes (1.2-) 1.6-2.7
m m , commonly spreading-recurved at late anthesis;
androecium (8-) 10-15-merous, ±30-39 m m , the tube
2-3.4 m m , the tassel pink-carmine; stemonozone ob
scure; ovary at anthesis silky-barbate; T E R M I N A L FL:
sessile or almost so, the calyx hemispherical, the white
staminal tube 10-12.5 m m , at orifice 3-4 m m diam, the
free filaments ± twice as many as those of peripheral
fls. Pods in broad profile 4.5-12 x 1.1-2.3 cm, silky-
pilose overall with erect, either white or partly bronze
hairs, the sutural ribs ±4.5-6 m m wide in dorsal view,
the sublignescent valves densely subhorizontally venu-
lose with coarse subcontiguous fibers, at maturity con
vex over the 2-5 seeds; seeds (few seen) ±6.5-12 x
4-8 m m , the testa smooth, brown dark-speckled, finely
pleurogrammic.
Key to the varieties of C. glomerulata
1. Lf-formula v-xi(-xiii)/(21-)24-36; lf-stk of
primary lvs (3-)3.5-9(-14) cm; pod (not well
known) 8-12 x (1.3—)1.5—2.3 cm, and seeds
9-12 mm; range of the species: Venezuela,
immediately adj. n. Colombia, and s.
Guyana 3a. var. glomerulata
1. Lf-formula iii—iv(—v)/l 6-21; lf-stk of primary lvs 1.2-3.6(4) cm; pod 4.5-8 x 1-1.2 cm;
seeds 6.5-? mm; within the range of the
preceding, Venezuela only 3b. var. parvifolia
3a. Calliandra glomerulata Karsten var. glomeru
lata. C. glomerulata Karsten, I.e., t. O I L 1862, sens.
str. — "... in montibus Venezuelanis, prope pagum
Sanare inter Quibor et Tucujo [El Tocuyo, edo. Lara]
. . . 1000 metr." — Holotypus, to be expected at L E
or W (n.v.). — Feuilleea glomerulata O. Kuntze,
Revis. Gen. PI. 1: 188. 1891.
26 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
C. glomerulata sensu Bentham, 1875: 554; Britton & Killip,
1936: 137; Steyermark & Huber, Fl. del Avila 560. 1978.
The relatively foliose forms of the species, with lf-
measurements as given in key to varieties.
In seasonally dry brush-woodland, savanna thickets,
and llano, 140-1400 m, scattered around the Orinoco
basin in Venezuela (Monagas w. to Tachira and ZuHa,
s. to n. Bolivar and n. Amazonas), n. in Distrito Federal
to the n. slope of Cord. Costanera, w. just into Co
lombia (Norte de Santander); disjunct in s. Guyana
(Kanuku Mts) and adj. Brazil (Surumu, Sa. da Lua). —
M a p 2. — Fl. (IX-)X-III. — Cansa-caballo; clavel-
lino; quebra-potro (the last two generic in scope).
3b. Calliandra glomerulata Karsten var. parvifolia
(Bentham) Barneby, comb, nov Calliandra laxa
var.(?) parvifolium [sic] Bentham, Trans. Linn. Soc.
London 30: 551. 1875. — "Venezuela [edo. Aragua,
between La Victoria and Biscaina], Fendler, n. 2255."
— Holotypus, K (hb. Benth.)! = photo s.n., NY!.
C. blakeana Pittier, Arb. Arbust. Venez. 6-8: 82. 1927. —
"Distrito Federal: Cerca del Ojo de Agua, carretera antigua
Caracas-La Guaira . . . May 22, 1927 (Pittier 12407 . ..)."
— Holotypus, V E N n.v.; isotypus, NY!. — Provisionally equated by Pittier with the preceding, but mistakenly
referred to ser. Laetevirentes = genus Zapoteca.
C. minutifolia Pittier, Arb. Arbust. Venez. 6-8: 83. 1927. —
"Carabobo: Alrededores de Valencia . . . Agosto 15, 1920 (Pittier 9018 . . .)." — Isotypus, NY!. — Compared by Pit-
tier with C. panlasia = C. laxa var. panlasia.
The relatively microphyll forms of the species,
with lf-measurements as given in key to varieties.
O n dry stony hills and in seasonally deciduous brush-
woodland or chaparral, recorded definitely at 1000-
1200 m, local within the range of var. glomerulata: n.
Venezuela, between w. Distrito Federal and centr. Fal
con, s. interruptedly to Trujillo; cultivated for ornament
in and around Caracas, to be expected elsewhere. —
M a p 2. — Fl. III-V, IX. — Mulato; moromoy.
4. Calliandra cruegeri Grisebach, Fl. Brit. W . I. 224
("Cruegerir). 1860. — " H A B . Trinidad! [Her
mann] Cr[uger], at Chocachocacco [sic, = Cha-
cachacare]." — Holotypus, to be sought at G O E T
(n.v.); isotypi, Cruger s.n., K!, Cruger 1008,
acquired by Britton from herb. Krug. & Urban.,
NY!. — Feuilleea cruegeri O. Kuntze, Revis. Gen.
PL 1: 187. 1891. Fig. 1
C. affinis Pittier, Arb. Legum. 1: 51. 1927. — "Area descono-cida, especie aparentemente propia del Llano." — Lecto
typus (Britton in adnot., NY): Pittier 12529, collected near
Dos Caminos, Guarico, Venezuela, 12 Sep 1927. —Lecto
typus, V E N n.v.; isotypus, NY!. — A second gathering, Pit-
tier 12224 (NY!), collected 10 Oct 1926 between Ortiz and
Guarico bridge, may have contributed to the protologue.
C. cruegerii sensu Bentham, 1875: 553; R. O. Williams, Fl.
Trin. Tob. 1(4): 299. 1931.
Arborescent shrubs (l-)2-6 m with stout stiff, ver
tically fissured, gray or pallid, plagiotropic long-shoots
and densely thatched short-shoots developing from pri
mary lf-axils, the young growth and all lf-axes puberu
lent or finely silky-pilose with white hairs to 0.3-1.4
m m , the narrow imbricate lfts discolorous, lustrous
dark-olivaceous above, paler duller beneath, facially
glabrous but often randomly cili(ol)ate, the sessile or
very shortly pedunculate capitula arising singly from
axils of efoliate stipules on homotinous or older
brachyblasts; phyllotaxy distichous. Stipules papery
stramineous, lanceolate or triangular-lanceolate 3-9
m m , striately nerved, commonly glabrous dorsally but
sometimes micropuberulent, persistent even though
frayed or broken in age. Lf-formula iii-v/ 32-52; lf-
stks of larger lvs 2.54.5 cm, the petiole including
scarcely swollen pulvinus 2-9 m m , at middle
0.45-0.7(-0.9) m m diam, the longer interpinnal seg
ments 5-11 (-12) m m , the shallow ventral groove
bridged; pinnae either regularly accrescent distally or
of randomly unequal lengths, the longer ones
2.5-5(-5.5) cm, the longer interfoliolar segments
0.5-1.1 m m ; lft-pulvinules ±0.1 x 0.2-0.3 m m ; lfts
abruptly decrescent at base of rachis and more gradu
ally so upward from near mid-rachis, the blades linear
or linear-lanceolate from either obtusangulate or
shortly auriculate base, acute, straight or gently falcate,
the longer ones (3-)4.2-7(-9) x 0.6-1.1 (-1.4) m m ,
(4.3-)4.5-6.5(-6.7) times as long as wide; midrib fili
form, displaced to divide blade 1:1.5-2, pinnately few-
branched, the 1-2 posterior primary nerves extremely
slender or barely perceptible, the inner one expiring
well short of mid-blade. Peduncles 0-3 m m bracteate,
the subscarious bracts 1-2, ovate 1-1.5 m m , often con
cealed by stipules or by the lower fls of the capitulum;
capitula ±11-21-fid, the fls ideally (but in fact not
always) dimorphic, the peripheral ones a trifle accres
cent upward, the terminal one (sometimes deformed,
or abortive) larger and with long-exserted trumpet-
shaped androecium, the subglobose or claviform re
ceptacle <2 m m ; floral bracts ovate 1-1.5 m m , persis
tent; pedicels 0-0.45 m m ; perianth submembranous,
the sharply striate calyx glabrous externally, some
times micro-ciliolate, the whitish corolla thinly silky-
pilosulous with wavy hairs; P E R I P H E R A L FLS: calyx
campanulate 1.8-2.9 x 1.1—1.8(—2) m m , the obtuse or
obtusely deltate teeth 0.3-0.55 m m ; corolla 4.3-6.2
m m , the ovate lobes 1.4-2 m m ; androecium 16-25-
merous, 17-24 m m , the stemonozone ±0.5 m m , the
tube 3-6 m m , the tassel whitish proximally, pink
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 27
FIG. 1. Calliandra cruegeri Grisebach.
28 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
distally; no intrastaminal nectary; TERMINAL FL:
calyx scarcely longer but broader than that of periph
eral fls; corolla 6-7 mm; androecial tube white ±10-12
m m , trumpet-shaped, 2-2.5 m m diam at orifice;
intrastaminal nectary cupular 0.5-0.9 mm; ovary at
anthesis glabrous. Pods 5-6.5 x 0.6-0.8 cm, thinly sub-
appressed-pilosulous overall, the dilated rim in dorsal
view 2 m m wide, the recessed lignescent valves low-
bullate over seeds, obliquely venulose; seeds up to 8
per pod, ellipsoid 5.5 x 2.5-3 mm, dusky-mottled, the
testa finely pleurogrammic.
In dry forest, xeromorphic scrub, and savanna,
0^100 m, locally plentiful on the Caribbean coast and
llanos of n. and centr. Venezuela, from n. Falcon and
n. Guarico e to Isla Sta. Margarita and the Paria
peninsula, s. to the Orinoco valley in n.-w. Bolivar
and in adj. Apure and Amazonas, thence e. across the
Dragon's Mouth to Chacachacare and Monos islands
in n.-w. Trinidad; isolated in savanna of n. state of
Roraima, Brazil (rio Surumu). — Map 3. — Fl. III-V,
VIII-XII, perhaps intermittently following rains.
5. Calliandra tolimensis Taubert, Bot. Jahrb. Syst.
21: 314. 1895. — "Columbia: crescit frequenter in
faucibus inter El Hubo et El Gigante sitas et prope
San Augustin civit. Tolima, mense Decembri flo-
rens (coll. columb. [Alphons Stubel] n. 239)." —
Holotypus, ̂ B; clastotypus (fl, lfts), F! = F. Neg.
1263. Fig. 2
C. matisiana Uribe-Uribe, Mutisia 11:2, cum icon, photo.
1952. — "Lorenzo Uribe-Uribe 2326, [Colombia.] Cundi-
namarca, Guaduas, comino antiguo a Honda, a 3 km. al N.O.
de lapoblacion; 1100 m. alt., junio 14,1952 (... Herb. Univ.
Javer. Bogota)." — Holotypus, COL!; isotypus, US!. —
Named in commemoration of Francisco Xavier Matis, artist
with the Real Expedition Botanica under J. C. B. Mutis.
C. tolimensis sensu Britton & Killip, 1936: 136.
Amply microphyllidious, arborescent shrubs 2-8 m,
the young stems, lf-axes and peduncles densely
pilosulous or silky-pilose with gray or brownish hairs
to 0.2-0.75(-1.2) m m , the firm plane lfts bicolored,
lustrous dark green above, paler dull beneath, facially
glabrous but sometimes minutely ciliolate, the massive
dense, technically umbelliform capitula arising either
solitary or 2(-3) together directly from loosely
thatched, efoliate brachyblasts axillary to coeval lvs of
long-shoots or from the primary lf-axils; phyllotaxy
distichous. Stipules firm, ovate, lanceolate, or rarely
depressed-deltate (1.5-)2-8 x 1.5-4 m m , weakly stri
ate dorsally beneath strigose-pilosulous vesture,
glabrous and strongly striate ventrally, deciduous from
broad base. Lf-formula iv-vii/(21-)24-41; lf-stk of
longer lvs (3—)4—9(—12) cm, the petiole including
swollen pulvinus (0.8-) 1-3.5(-4) cm, at middle 1-1.7
m m diam, the longer interpinnal segments (5-)6-14
(-17) m m , the ventral groove weakly bridged or some
times continuous; pinnae a httle accrescent distally, the
rachis of longer ones (4-)4.5-9 cm, the longer interfo-
liolar segments 1.1-2.3 m m ; lft-pulvinules 0.2-0.5
m m ; lfts decrescent toward each end of rachis, linear-
lanceolate from prominently auriculate base, obtuse or
subacute, straight or incipiently falcate, those near
mid-rachis (7.5-)8-13 x (1.5-)1.7-2.3 m m , (3.7-)4.2-
6.1 times as long as wide; midrib straight or nearly so,
forwardly displaced to divide blade 1:2-3, finely pin
nate dorsally, the venulation immersed or almost so
ventrally. Peduncles stout (1.5—)2—8 cm, bracteate
above middle; capitula 18—32-fld, the clavate or sub-
spherical receptacle 3-8 x 3-3.5 m m ; bracts ovate or
lanceolate ±2-2.5 m m , 1-3-nerved, incurved, persis
tent; fls heteromorphic, the peripheral ones at least
shortly pedicellate, the terminal one sessile, the peri
anth of all 5-merous, the faintly 15-nerved calyx thinly
puberulent, the greenish-white corolla appressed-silky,
externally nerveless; P E R I P H E R A L FLS: pedicel 1-
3.3 x 0.8-1.8 m m ; calyx deeply campanulate or nar
rowly um-shaped 4-6 x 2.6-̂ 1.2 m m , the ovate, deltate
subacute, or obtusely semicircular teeth 0.5- 1.3 m m ;
corolla 9.5-14.5 m m , the lanceolate or ovate lobes
(1.4-)3.5-5.5 m m ; androecium 30-42(-46)-merous,
4.4-6.6 cm, the stemonozone 0.8-1.6 m m , the tube
4-7.5(-10) m m , the tassel white proximally, carmine
distally; ovary white-silky beyond middle; TERMI
N A L FL: perianth about as long as that of peripheral
fls, but broader; staminal tube broadly cylindric, ± as
long as corolla; intrastaminal disc present. Pod erect, in
profile 10-15 x 1.1-1.6 cm, 5-8-seeded, the valves
coarsely transverse-venulose, densely pilosulous or
silky-pilosulous overall with mixed shorter brown and
longer white hairs, either type predominant; seeds in
broad view 10-11.5 x 6.5-7.5 m m , the papery testa
dull brown, pleurogram 0.
In seasonally dry woodland, sometimes colonial
along highways, locally plentiful at 600-1250 m in
the upper Magdalena valley between 2°N and 5°N in
Huila, Tolima, and Cundinamarca, Colombia. —
Map 2. — Fl. X-XII, IV.
Uribe recognized the relationship between C. ma
tisiana and C. tolimensis, but distinguished C. mati
siana by the pedicellate flower, more numerous pinnae
and leaflets, and flowers said to be more than three
times larger. The population at Guaduas has indeed on
average slightly more numerous leaflets, but genuine
C. tolimensis, now known by several collections from
Huila, also has pedicellate flowers, and the difference
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 29
M A P 3. The distribution of Calliandra cruegeri Grisebach in Venezuela and northern Brazil.
in flower-size was simply a mistake. The fmits of C.
matisiana are pilosulous with prevailingly short rusty-
brown hairs, whereas those of C. tolimensis elsewhere
are clothed in longer, largely white ones. This visually
arresting difference in pubescence is the one distinctive
character of C. matisiana, which I here regard as a
somewhat aberrant population that does not deserve
taxonomic rank.
6. Calliandra laxa (Willdenow) Bentham, Trans.
Linn. Soc. London 30: 551. 1875. — Typus infra
sub var. laxa indicatur.
Arborescent shrubs, ordinarily fertile at 2-15 m
with trunk to 2 d m dbh but exceptionally attaining
25 m with trunk 4 d m dbh, the stout long-shoots
mostly plagiotropic and the short-shoots usually ±
extended or only loosely thatched, the young stems,
lf-axes (at least along ventral ribs, usually all around)
and peduncles pilosulous with either straight ascend
ing or incurved hairs to 0.25-0.8 m m , the bicolored
lfts either dark olivaceous or (especially at maturity)
dove-gray above, paler brownish beneath, all either
fully glabrous, or glabrous ciliolate, or thinly papil-
late-pilosulous on either face, or softly pilosulous
overall, at maturity chartaceous and either plane or
low-convex ventrally, the incipiently umbelliform ca
pitula either solitary or geminate, arising mostly from
nodes of developing lateral short-shoots but some
times from the axil of primary lvs; phyllotaxy distic
hous. Stipules of firm texture, those subtending pri
mary lvs lanceolate or lance-ovate 2.5-8(-10) x 1-3
m m , either acute or obtuse, at least when young
coarsely 6-13(-15)-nerved but often smooth and
blanched in age, those of short-shoots commonly
smaller, broad-lanceolate, ovate or triangular-deltate,
all persistent but commonly tattered in age. Lf-formula
iii-v(-vi)/(8-)9-29; lf-stk of primary lvs and of larger
lvs on brachyblasts 2.5-9.5(-ll) cm, that of some
brachyblast lvs (no further mentioned) shorter, the
petiole including fuscous pulvinus (8-) 10-34 m m , at
middle 0.6-1.8 m m diam, the longer interpinnal seg
ments 4-18(-24) m m , the ventral groove bridged;
pinnae distally accrescent, the rachis of furthest and
penultimate pairs (3-)4-10.5 cm, the longer interfoli-
olar segments (1—)2—10(—12) m m ; lft-pulvinules 0.4-
1.3 x 0.45-0.9 m m , cross-wrinkled; lfts subequilong
or slightly either ac- or decrescent distally, the blades
(often variable in outline and dimensions between lvs
of one branch) oblong or oblong-elliptic, less often
ovate-oblong or broad-linear, from shallowly semicor-
date or inequilaterally rounded base, either broadly
obtuse or less often deltately subacute, the longer
ones (4-)5.5-21(-23) x 1.7-8 m m , 2-5.3 times as
long as wide; venation of mature blades immersed
or shallowly impressed ventrally, finely prominulous
dorsally, the nearly straight midrib displaced to divide
30 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
FlG. 2. Calliandra tolimensis Taubert.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 31
blade ±1:1.5-2, ±3-6-branched on posterior side,
the inner posterior primary nerve either incurved-
ascending to or even beyond mid-blade or weak and
obscure, tertiary venules faint or imperceptible. Pe
duncles 0.6-6(-6.5) cm, ebracteate; capitula 9-23-
fld, the hemispherical or shortly claviform receptacle
1.5-2 x 1.5-3 m m , the fls either homo- or heteromor-
phic, 1-3 (sub)terminal ones often slightly broader but
not or scarcely longer than the peripheral ones but
their androecium modified into a far-exserted white
trumpet; bract subtending outermost fls minute (<0.9
m m ) , caducous, the inner fls ebracteate; pedicel of pe
ripheral fls sometimes perceptible only in section but
usually externally discolored, 0.3-1.2 x 0.6-1.2 m m ;
perianth submembranous, greenish-white, yellowish,
or red-tinged, the nerveless or weakly 5(-10)-nerved
calyx usually glabrous or only very thinly pilosulous
above middle, 5-merous, the corolla 5-merous or by
adherence of one or two pairs of petals 4- or 3-merous,
usually thinly appressed- or ascending-pilosulous, ex
ceptionally glabrous; P E R I P H E R A L FLS: calyx nar
rowly campanulate or turbinate-campanulate 1.4-2.6
(-3) x 1.1—1.8(—2.2) m m , the obtuse, dorsally convex
teeth 0.3-0.55(-0.8) m m ; corolla 4.5-8.5 m m ; andro
ecium (8-)10-18-merous, 26-41 m m , the stemono
zone 0.8-1.8 m m , the whitish tube 3.3-6.6 m m , the
tassel pink or crimson; intrastaminal nectary 0; ovary
at anthesis either glabrous or villosulous; T E R M I N A L
FL(S): perianth broader than that of peripheral fls but
hardly longer; androecial tube 13-22 m m , at orifice
dilated to 3-4-.5 m m diam; intrastaminal nectary to 0.9
m m tall. Pods in broad view 9—16(—17) x 1.1-1.6
(-1.7) cm, the sutural ribs in dorsal view 4-5 m m
wide, glabrous or early glabrate, the recessed woody
valves coarsely sinuously transverse-venulose, either
glabrous or thinly pallid- or brownish-pilosulous;
seeds in broad view ±10-14 x 5.5-10.5 m m , the loose
testa papery fragile brown, lacking pleurogram.
Key to the varieties of C laxa
1. Petiole of well-developed lvs not more than
1.5 times as long as the first interpinnal
segment of lf-stk; larger lfts 8-21 x 2-8 mm.
2. Longer peduncles, in Venezuela, 0.6-2 cm,
but longer in Panama and n.-w. Colombia;
lfts of longer pinnae 18-29 pairs; Venezuela
n. of the Orinoco and extending weakly into
n.-e. state of Bolivar, thence disjunct in e.
Panama and adj. Colombia 6a. var. laxa
2. Longer peduncles 2-6.5 cm; lfts of longer
pinnae (8—)9—16(—18) pairs; abundant in
Venezuelan Guayana and s., less commonly,
into adj. Guyana and the Amazon Basin in
Brazil and e. Colombia 6b. var. stipulacea
1. Petiole of well-developed lvs ± twice as long
as the first interpinnal segment of lf-stk; larger
lfts 4—6.5 mm, local on Gran Sabana in state
of Bolivar, Venezuela .... 6c. var. urimana
6a. Calliandra laxa (Willdenow) Bentham var. laxa.
Calliandra laxa Bentham, 1875, I.e., sens. str. Acacia
laxa Willdenow, Sp. PI. 4: 1069. 1805. — "Habitat
ad Caracas . . . Bredemeyer." — Holotypus, Brede-
meyer 15 in B-WILLD 19148, seen in Microform!
and F Neg. 7277!. — Mimosa laxa Poiret, Encycl.
suppl. 1: 72. 1810. Feuilleea laxa O. Kuntze, Revis.
Gen. PI. 1: 188. 1891.
C. cumingii Bentham, J. Bot.(Hooker) 2: 140. 1840. —
"Panama. Cuming, n. 1248." — Holotypus, K (hb.
Benth.)!; isotypus, K (hb. Hook.)! = photo s.n., NY!, US! = photo s.n., NY!. — Feuilleea cumingii O. Kuntze, Revis.
Gen. PI. 1: 187. 1891. Anneslia cumingii Britton & Rose, N. Amer. Fl. 23: 57. 1928.
C. panlosia [sic, a mistake for panlasia = hairy overall] J. R.
Johnston, Proc. Amer. Acad. Arts 40(21) [= Contr. Gray
Herb. n. ser. 29]: 686. 1905. — "[VENEZUELA. Nueva
Esparta: Isla Margarita] ... on hills at altitude of 300 to 600 m., El Valle to Juan Griego, Miller & Johnston, n. 58,
July 22, 1901, and Johnston, n. 27, July 2, 1903." — Lec
totypus, J. R. Johnston 27, GH!; isolectotypus, NY!;
paratypi, Miller & Johnston 58, GH!, N Y (fragm)!.
C. trijugata Schery, Fieldiana, Bot. 28(2): 256. 1952. — "[VENEZUELA] ... on rocky savanna slopes between Ciudad Bolivar and Rio Caroni, state of Bolivar, alt. 100 m, Aug. 1, 1944, Julian A. Steyermark 57592." — Holotypus,
F!; isotypus, MO!.
C. laxa sensu Bentham, 1875: 551, exclus. syn. xalapensis
et rubescens; Pittier, 1927: 49, quoad pi. venezol.
C. cumingii sensu Bentham, 1844: 106, var. exclus.; Ben
tham, 1875: 551, quoad typum, caeteris exclusis; Woodson & Schery, 1950: 262.
C. panlosia sensu Pittier, 1927: 51; Hoyos E, 1985: 521, but
ambiguously equated with C. [= Zapoteca] caracasana Benth.
Lf-formula iii-v(-vii)/16-27(-29); lf-stk of longer
lvs 2.5—8(—11) cm, the longer interpinnial segments
6-22(-24) m m ; rachis of longer pinnae 4-9 cm, the
longer interfoliolar segments (1-) 1.3^4 m m ; lfts
facially either glabrous, or papillate, or pilosulous,
the larger ones 7-14 x 1.7-4.5 m m , the inner poste
rior primary nerve short and faint; longer peduncles
0.6-3.5 cm, mostly <2 c m in Venezuela.
In semideciduous and riparian woodland, thorn
scrub, and savanna thickets, surviving disturbance,
near sea level on Caribbean coast and up to 1200 m
inland, locally plentiful in n. Venezuela, from Aragua
and Miranda e. to Nueva Esparta, Sucre (Mochima e.
to Cristobal Colon), Monagas, and n.-e. Bolivar, there
fading into var. stipulacea; disjunct in moist lowlands
of e. Panama (Colon, Canal Zone, San Bias, Darien)
32 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Map 4. The distribution of the varieties of Calliandra laxa (Willdenow) Barneby in South America and adjacent Panama.
and n.-w. Colombia (Choco). — Map 4.
— Clavellino; clavellina serrana.
Fl. I-IX.
6b. Calliandra laxa (Willdenow) Bentham var. stip
ulacea (Bentham) Barneby, stat. nov. C. stipulacea
Bentham in, J. Bot. (Hooker) 2: 137. 1840. —
"[GUYANA.] ... On the Rio Quitaro [= Guidaro,
Kwitaro], [Robert] Schomburgk [1st collection], n.
582." — Holotypus, K (hb. Benth.)! = N Y Neg.
19691; isotypi, +B = F Neg. 7267!, BM!, NY!,
OXF!. — Feuilleea quitaro O. Kuntze, Revis. Gen.
PI. 1: 186. 1891.
C. diversifolia Britton & Killip, Ann. New York Acad. Sci. 35: 136. 1939. — "Columbia, 1760-1808, Mutis 441 . . .
3542." — Holotypus, C. Mutis 441, US!; clastotypus, NY!; paratypus, Mutis 3542, US!, N Y (fragm)!. C. stipulacea sensu Bentham, 1844: 107; 1875: 551; Ducke, 1949: 50; 1958: 49; Cowan, Mem. New York Bot. Gard. 10(1): 143. 1958.
Lf-formula iii-v(-vi)/(8-)9-16(-18); lf-stk of larger
lvs (3-)4-9.5(-10.5) cm, the longer interpinnal seg
ments 8—18(—22) m m ; rachis of longer pinnae
(4-)4.5-10.5 cm, the longer interfoliolar segments
4.5-10(-l2) mm; lfts commonly glabrous facially,
sometimes weakly or remotely pubemlent (but then
fewer and broader than those of pubescent var. laxa),
the larger ones (8-) 10-21 (-23) x (2.5-)3-8 mm; inner
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 33
posterior primary nerve produced to or beyond mid-
blade; longer peduncles 2-6(-6.5) cm.
Along streams in upland savanna, in forest-savanna
ecotone, and in lowland riparian forest, 90-850 m,
locally plentiful in Venezuelan Guayana, s.-w. Guyana,
and adjoining Brazil (Amazonas, Roraima), thence
extending w. into Amazonian Colombia (Cauqueta,
Vaupes) and s. in Brazil to the Madeira basin in n.-w.
Rondonia and n. Mato Grosso and to Sa. do Cachimbo
in s.-w. Para. — Map 4. — Fl. intermittently through
the year.
6c. Calliandra laxa (Willdenow) Bentham var. uri-
mana (Brunner & Forero, ined.) Barneby, var. nov.,
foliorum adultorum petiolo quam petioli communis
segmentum interpinnale proximale subduplo longiori
foliolisque 4-6.5 m m usque longis ab aliis forrnis
diversa. — VENEZUELA. Bolivar: On w. Bank of
no Uriman just above Salto Acarima, 393 m, 8 Jan
1955, /. A. Steyermark & J. J. Wurdack 38. — Holo
typus, NY; isotypus, US. Calliandra urimana Brun-
ner & Forero in sched., nom. nud.
Lf-formula iv-vi/17-22; lf-stks of larger lvs 2.5-5
cm, the petiole 9-17 m m , the longer interpinnal seg
ments 4.5-8 m m ; rachis of longer pinnae 3-5 cm; lfts
thinly pilosulous facially, the larger ones 4-6.5 x
1.8-2.5 m m ; inner posterior primary nerve faint and
short; peduncles 2.5-4 cm.
At savanna edge and on rocky stream banks, ±370-
400 m, localized in the middle Caroni basin in Boli
var, Venezuela, within 5°15'-45TS[, 62°20,-50,W, on
rios Uriman, Aprada and Acanan near the foot of
Auyan- and Uaipan-tepuis. — Map 4. — Fl. I—III.
7. Calliandra rubescens (Martens & Galeotti) Stand-
ley, Publ. Field Mus. Nat. Hist., Bot. Ser. 4(8): 309.
1929. Acacia (?) rubescens Martens & Galeotti,
Enum. PL Galeotti p. 10: 14 [= Bull. Acad. Roy. Sci.
Bruxelles 10(9): 315]. 1843. — "Coll. H. Galeotti
No. 3314 .. . ravins et bois du Puente Nacional, pres
de Vera Cruz [Mexico], et pres de Cantaranas et
Zacuapan." — Holotypus, BR n.v. = isotypus
sequentis. — Anneslia rubescens Britton & Rose, N.
Amer. Fl. 23: 61. 1928. — Mistakenly equated by
Bentham, 1875: 551, and by Standley, 1922: 387,
with C. laxa.
C. xalapensis Bentham, London J. Bot. 3: 106. 1844. — "Mexico; woods of the neighborhood of Xalapa, Galeotti, n. 3314." — Holotypus, K (hb. benth.)! = photo s.n., NY!; isotypus, K (hb. Hook.)!. — Anneslia xalapensis Britton & Rose, N. Amer. Fl. 23: 62. 1928. — Mistakenly equated by Bentham, 1875: 551, and by Standley, 1922: 387, with
C. laxa.
Anneslia compacta Britton & Rose, N. Amer. Fl. 23: 62. 1928. — "[MEXICO.] Chiapas . . . Hacienda Monserrate, September, 1923, [C. A.] Purpus 9063." — Holotypus, NY! = NY Neg. 9372. — Mistakenly equated by Zamora, 1991: 78, with C. bijuga Rose.
A. simulans Britton & Rose, N. Amer. Fl. 23: 63. 1928. — "[MEXICO.] Between La Venta and Niltepec, Oaxaca, July, 1985, E. W. Nelson 2795V — Holotypus, US!; isotypus, NY!.
A. papillosa Britton & Rose, N. Amer. Fl. 23: 63. 1928. — "[MEXICO.] Veracruz. Type from near Cameron [= Ca-mar6n], October, 1925, [C. A.] Purpus 10582." — Holotypus, NY!. — Calliandra papillosa Standley, Publ. Field Columbian Mus., Bot. Ser. 8: 312. 1931. A. pallida Britton & Rose, N. Amer. Fl. 23: 65. 1928. — "Puliorages [= paturages] de la Cruz de Guanacaste, 1890, Costa Rica, Pittier 2727" — Holotypus, US!; clastotypus + photo, NY!. — Equated with C. rubescens by Zamora, 1991: 80. — Calliandra pallida Standley, Field Mus. Nat. Hist. Publ. Bot. 18: 492. 1937. Calliandra yunckeri Standley, Field Mus. Nat. Hist. Bot. Ser. 17(4): 367. 1938. — "[HONDURAS.] . . plain near Siguatepeque, Dept. Comayagua, 1050 meters, July 4, 1936, T. G. Yuncker, R. F. Dawson & H. R. Youse 5622." — Holotypus, F!; isotypi, K!, NY (ex hb. Yuncker.)!. — Equated by Zamora, 1991: 80, with C. rubescens. C. tapirorum Standley, Ceiba 1: 40. 1950. — "Honduras: Dept. Comayagua . . . near Siguatepeque, alt. 1050 m., March-April 1947, Paul C. Standley 6856." — Holotypus, F!; paratypi, Williams & Molina 11209, 11576 F!. — Equated by Zamora, 1991: 80, with C. rubescens. C. rubescens sensu Zamora, 1991: 80.
Microphyll shrubs variable in stature, in exposed
stony sites flowering as a stiffly intricately branched
shrub 0.5-1.5 m, but in riparian or woodland envi
ronment potentially arborescent and attaining 4.5
(-5.5) m, in most respects closely resembling C. laxa,
the young branchlets, lf-axes and peduncles usually
thinly (seldom densely) pilosulous or puberulent with
fine pallid hairs to 0.1-0.5 mm, the bicolored, dorsally
pallid dull, ventrally brown-olivaceous and either dull
or lustrous lfts varying from glabrous facially and ei
ther ciliolate or not to pilosulous on one or both faces
(the hairs of upper face often reduced to a papilliform
base), the solitary or geminate capitula borne mostly
on thatched or shortly caulescent brachyblasts but
some axillary to primary lvs; phyllotaxy distichous.
Stipules triangular to narrowly lanceolate, those sub
tending primary lvs 1-6.5 x 0.4-2.3 mm, weakly 4—7
(-1 l)-nerved, becoming dry fragile, those of brachy
blasts shorter. Lf-formula ii—v(—vi)/( 12—) 14—17; lf-stk
of larger lvs (1.5—)2—5 cm, the petiole 4-16(-19) mm,
at middle 0.5-1.9 m m diam, the longer interpinnal seg
ments 5-14 mm; pinnae either subaccrescent distally
or almost equilong, the rachis of longer ones (2-)
2.5-5.5(-6) cm, the longer interfoliolar segments
1.6- 3.3 mm; lft-pulvinules 0.15-0.5 x 0.2-0.6 mm,
transversely wrinkled; lfts subequilong or gradually
34 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
decrescent distally, the blades inequilaterally lance-
oblong or ovate from shallowly semicordate base,
deltately acute or depressed-deltate-apiculate, the
larger ones (4.5-) 5-10 x (1.4-)1.6-3.2 mm, (2.3-)
2.6-4.1 (-4.4) times as long as wide; venation palmate-
pinnate, prominulous either on both faces or only dor
sally, the midrib straight or incipiently falcate, dis
placed to divide blade ±1:2, 2-3-branched beyond
middle on posterior side, the inner posterior primary
produced either nearly to or beyond mid-blade. Pedun
cles 1.7-3.5(-4) cm, ancipital, ebracteate or exception
ally bracteate near middle; capitula (6-)8-14-fld, the
receptacle at most 2 mm, a terminal pedestal some
times barely differentiated; bracts deltate-ovate or
lanceolate 0.3-1.4 mm, tardily deciduous; fls (sub)
sessile (the pedicel scarcely differentiated externally),
usually heteromorphic, one terminal one with broader
calyx, sometimes longer corolla, and always well-
exserted androecial tube, the perianth of all fls gla
brous or, less often, puberulent distally, the calyx
weakly nerved, the corolla externally nerveless;
PERIPHERAL FLS: pedicel (in vertical section)
0.35-0.6 x 0.55-1.2 mm; perianth (4-)5-merous; calyx
(1.5-)1.7-3.4(-3.6) x (0.8-)l.l-1.9(-2.1) mm, the
broad obtuse teeth 0.15- 0.8 mm; corolla (4.5-)5.4-8.7
mm, the triangular or lanceolate lobes (0.5-)0.7-l .9
mm; androecium (13-)l4-24-merous, 22-32 mm, the
stemonozone 0.5-1.2 mm, the pallid tube 5.5-8.5 mm,
the tassel pink or carmine; ovary at anthesis glabrous;
no intrastaminal nectary; C E N T R A L FL: calyx
scarcely longer but broader than that of peripheral fls;
corolla scarcely longer; androecial tube nearly twice as
long as corolla; intrastaminal nectary to 0.8 m m tall;
gynoecium 0 or rudimentary. Pods erect, in broad pro
file (4-)6-ll x 0.5-0.9(1) cm, the thickened sutures
4—6 m m wide in dorsal view, the densely puberulent or
pilosulous, stiffly woody valves either bluntly, sub-
symmetrically venose or externally almost nerveless;
seeds 9-12.4 x 3-6 mm, the smooth testa fight brown
or putty-colored, dark-speckled, pleurogrammic.
In semideciduous forest, surviving disturbance in
pasture thickets, in thin pinewoods, on stony hillsides
and rocky river banks, 70-1300(-1500-?) m, interrupt
edly dispersed from tropical e. Mexico near 21°N to w.
Costa Rica: in Mexico from lowland Veracruz to s.
Oaxaca and adj. Chiapas; unknown from Guatemala; s.-
centr. Honduras; w. Nicaragua; w. Costa Rica (Guana-
caste). — Map 5. — FL VI-X. — Cabellos de angel.
8. Calliandra belizensis (Standley) Standley, Publ.
Field Columbian Mus., Bot. Ser. 4: 309. 1929.
Anneslia belizensis Britton & Rose ex Standley,
Trop. Woods 11: 19. 1927. — "Collected by H. W.
Winzerling at Hillbank, Orange Walk District,
British Honduras, in 1927 (No. VII.4)." — Holoty
pus, US 12697991; isotypus (on 2 sheets), NY!.
Anneslia belizensis sensu Britton & Rose, 1928: 62. Calliandra belizensis sensu Standley, Publ. Field Columbian Mus., Bot. Ser. 8: 312. 1931; Standley & Steyermark, 1946: 20.
Coarsely microphyllidious, arborescent shrubs fer
tile when 2-9 m tall, with trunk attaining 1 dm dbh,
stout straight terete long-shoots, and dense, conspic
uously bracteate capitula sessile on thatched or incip
iently caulescent brachyblasts axillary to the furthest
or to the two furthest lvs of homotinous long-shoots,
often appearing terminal, the young stems and ventral
face of lf-axes loosely pilosulous with hairs <0.5 mm,
the foliage bicolored, the firm plane, facially gla
brous, sometimes ciliolate lfts dark and lustrous on
upper face, paler dull-olivaceous or cinnamon-brown
beneath. Stipules ovate-triangular or broad-lanceolate
4—12 mm, weakly striate, early stramineous, dry and
eroded, glabrous externally, persistent. Lf-formula
(i-)ii/( 16-) 18-29, some lvs always with bijugate pin
nae; lf-stks of longer lvs (11-) 14-43 mm, the petiole
(5-)7-23 mm, at middle 0.7-1.2 m m diam, the one
interpinnal segment 6-20 mm; rachis of longer pin
nae 6-9 cm, the longer interfoliolar segments 2.2-5
mm; lft-pulvinules 0.4—0.7 x 0.6-0.7 m m , coarsely
wrinkled; lfts subequilong except at extreme ends of
rachis, the blades linear-lanceolate from obtusely au-
riculate base, deltately acute or acuminulate, straight
or almost so, the larger ones 8-18 x 2-3.7 mm, 3.9-
4.6(^.8) times as long as wide; venation externally
faint, the weak midrib displaced to divide blade ±1:2,
the secondary venulation visible only in lfts dried
when young. Peduncles essentially 0, the densely or
loosely imbricate, efoliate stipules of brachyblasts
passing directly into similar floral bracts, the capitula
prior to anthesis conelike; floral bracts resembling
stipules in form and texture, lanceolate or ovate-
elliptic 0.5-9.5 x 1.4-4.2 mm, strongly striate at base,
weakly so distally, thinly pilosulous near apex, per
sistent; fls homomorphic, sessile or almost so, the
perianth 5-merous, the calyx sharply multistriate, the
corolla not or scarcely so, both thinly pilosulous
externally in distal V4—VS; calyx deeply campanulate-
ellipsoid 4.5-7.2 x 2.6-3.3 mm, the broadly obovate-
semicircular teeth (0.6—)1.1—1.8 mm; corolla 9-13
mm, the ovate lobes 2-3.2 m m ; androecium 23-29-
merous, ±4-4.8 cm, the tube (5-)6-10 mm, the in
durate stemonozone 2.4—3.2 m m , the tassel white; in
trastaminal nectary 0; ovary sessile, thinly pilosulous.
Pods (few seen) erect, in profile ±8.5-10 x 1-1.2 cm,
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 35
4—=> ^-4—^l/ tr • j w \~r^—ft-
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-
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MAP 5. Distribution of Calliandra rubescens (Martens & Galeotti) Standley in Mexico and Central America.
4—7-seeded, the sutural ribs in dorsal view 2.3-3 m m
wide, the stiff brittle valves bullate over seed-cavities,
dark brown, gray-pilosulous overall; seeds not seen.
In tropical evergreen forest, either in the under-
story or in sunny openings and on roadsides, mostly
on limestone, below 150 m, best known from about
the common boundary-point of Mexico (s. Quintana
Roo), Guatemala (Peten), and Belize (Corozal and
Orange Walk districts), and collected once on
Cozumel I. in n. Quintana Roo. — Map 6. — Fl.
XI-XII(-?). — Capulin de corona; barba de viejo.
Notable features of C. belizensis are faintly veined
leaflets, capitula subsessile on often subterminal
brachyblasts, large papery floral bracts, and white
androecial tassel.
9. Calliandra goldmanii Rose ex Barneby, sp. nov., C.
rubescenti foliorum formula necnon filamentis parti-
coloribus similis, sed ab ea stipuhs latioribus striatis,
foliolis majoribus, necnon leguminibus latioribus
11-16 (nee 5-10) m m latis abstans. — MEXICO.
Chiapas: Mun. Cintalapa, 23 km w. of Las Cruces,
870 m, 30 Oct 1981 (fl, fr), D. E. Breedlove 54121.
— Holotypus, NY. — Anneslia goldmanii Rose ms,
in sched., E. A. Goldman 850, NY.
Anneslia seleri sensu Britton & Rose, 1928: 62, ex parte, typo excluso.
Stiffly crookedly branched microphyllidious trees
3-9 m with terete plagiotropic or widely ascending
branches and densely thatched florigerous brachy
blasts, the young branches and lf-axes thinly or densely
pilosulous with either straight or incurved, either
loosely ascending or spreading hairs to 0.3-0.7 m m ,
the firm bicolored lfts lustrous and darker green above,
either facially glabrous and cili(ol)ate or erect-pilosu-
lous on lower (rarely also on upper) face, the stout pe
duncles arising singly from stipulate but elaminate
axils of brachyblasts; phyllotaxy distichous. Stipules
broad-lanceolate or triangular (2.5-) 3-8 x 2-4 mm,
the blades firmly chartaceous, when young finely
closely 11-21-nerved, persistent but erose in age. Lf-
formula ii-iii(-iv)/(13-)14-19; lf-stk of primary lvs to
2-4 cm, the petiole 7-17 mm, at middle 0.6-1.4 m m ,
the ventral groove obscurely bridged, the one or the
36 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
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M A P 6. Distribution of Calliandra goldmannii Rose e* Barneby, C. bijuga Rose, C. molinae Standley, and C. belizensis (Standley) Standley in Mexico and Central America.
longest of 2(-3) interpinnal segments commonly ± as
long as petiole, sometimes shorter; pinnae usually a ht
tle accrescent distally, the rachis of furthest pair ±4-7
cm, the longer interfoliolar segments 2-3.2 m m ; lft-
pulvinules 0.2-0.45(-0.6) x 0.4-0.8 m m , wrinkled; lfts
weakly graduated, the blades linear-oblong, rarely
lance- or elliptic-oblong from shortly obtusely auricu-
late base, straight or subincurved beyond middle,
deltately acute or apiculate, the longer ones (8-) 10-16
x 2.7-4.8 m m , (3.4-)3.5-5.4 times as long as wide;
venation palmate-pinnate, the straight or almost
straight midrib displaced to divide blade ±1:1.5-2, on
posterior side branched from middle upward, on
antrorse side branched from base upward, the inner
posterior primary nerve ascending at least to, often
somewhat beyond mid-blade, all these and a fine irreg
ular reticulum of veinlets prominulous on both faces.
Peduncles 9-28 m m , usually bracteate near or below
middle; capitula 12-18-fld, the receptacle 1.5-2 m m
diam; floral bracts ovate or subulate 0.8-2 m m ; fls sub-
sessile, in most capitula heteromorphic, the perianth of
the central-terminal fl wider but scarcely longer than
that of the rest but its androecium scarcely modified;
perianth 4-5-merous, yellowish-green sometimes red-
tinged, externally glabrous overall but the orifice
sometimes microscopically ciliolate, the calyx weakly
striate, the corolla nerveless externally or almost so;
P E R I P H E R A L FLS: pedicel at most 0.4 x 0.8 m m ,
often scarcely perceptible; calyx 3.2-5 x 2-3.2 m m ,
the broad obtuse teeth 0.4—0.8 m m ; corolla 8-9.5 m m ,
the lobes 2-3 m m ; androecium 14— 18-merous, ±3.6-4
cm, the tube 5.5-9.5 m m , the stemonozone 0.7-1.1
m m , the filaments white in lower half, the tassel deep
pink or carmine; ovary at anthesis glabrous; no nectar-
ial disc seen, but probably present in bisexual fls. Pods
stiffly erect-ascending, in broad profile 6.5-11 x
1.1-1.6 cm, the sutural ribs in dorsal view 5-8.5 m m
wide, the inflexibly lignescent valves brown, minutely
pilosulous overall and also thinly or densely granular;
seeds (few seen) 6.3-7.5 x 3.5-5.5 m m , the smooth
brown testa faintly dark-speckled, pleurogrammic.
In tropical deciduous woodland and pine-oak forest,
sometimes riparian and surviving deforestation as a
pasture- or hedge-tree, 250-915 m, locally common,
apparently of bicentric range: within 16°-17°N,
92°-94°W in s. Chiapas, Mexico; and near 10°N,
85°W in prov. Guanacaste and Puntarenas, Costa Rica.
— M a p 6. — Fl.VIII-XII.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 37
Calliandra goldmanii has much in common with C.
rubescens, especially leaf-formula and particolored
androecium, but differs in broader, striately nerved
stipules, larger leaflets, and pods 11-16, not 5-10 m m
wide. Its densely thatched brachyblasts recall C.
belizensis, but this has sessile capitula, ample striate
floral bracts, and more numerous, whitish filaments.
10. Calliandra bijuga Rose, Contr. U.S. Natl. Herb.
5: 135. 1897. — "[MEXICO. Guerrero:] Bottom
lands at Acapulco, Dr. Edward Palmer, November,
1894 (No. 138)." — Holotypus, US!; isotypus, K!;
clastotypus, NY!. — Anneslia bijuga Britton &
Rose, N. Amer. FL 23: 61. 1928.
Anneslia michelii Britton & Rose, N. Amer. Fl. 23: 61. 1928.
— "[MEXICO.]... Monte de las Cortaduras, near Inguaran
[±19°N, 101°40'W]. Michoacan, March 20, 1898, Lan-
glasse 41." — Holotypus, US!; clastotypus, NY!. Calliandra
michelii Standley, Field Mus. Nat. Hist., Bot. Ser. 11: 159.
1936. — Equated with C. bijuga by McVaugh, 1987: 152. Calliandra bijuga sensu M. Micheli, Mem. Soc. Phys.
Geneve 34: 251, t. 3. 1903; Standley, 1922: 387; McVaugh, 1987: 152.
Arborescent shrubs attaining 4—5 m with trunk ±2
d m diam, the annotinous and older branches stiffly di
varicate, terete, gray or fuscous, the young branchlets
and ventral face of lf-axes pilosulous with loosely
ascending or subappressed, pallid hairs to 0.35-0.7
m m , the firmly chartaceous, usually glabrous, rarely
dorsally pilosulous or ciliolate, lfts strongly bicolored,
dark brown or dove-gray and sublustrous above, paler
brown and dull beneath, the capitula arising singly
from lower nodes of shortly extended brachyblasts;
phyllotaxy distichous. Stipules submembranous, lance
olate or lance-attenuate 2.5-6 x 0.7-1.3 m m , weakly or
indistinctly 3-5-nerved, becoming papery fragile. Lf-
formula (i-)ii/7-12; lf-stks of primary lvs 5-23 m m ,
the petiole ±5-14 m m , the one interpinnal segment
(present in all or most lvs) 6-16 m m ; pinnae equilong
or the distal pair slightly longer, the rachis of longer
ones 3.5-6.5 cm, the longer interfoliolar segments
4-7.5 m m ; lft-pulvinules 0.2-0.7 x 0.5-0.8 m m , trans
versely wrinkled; lfts little graduated (but terminal
pair, no further described, sometimes longest), the
blades lanceolate or lance-oblong from shallowly
semicordate base, straight or incipiently falcate above
middle, deltately acute or sometimes apiculate, those
near mid-rachis 9-18 x 3-5.7 m m , 2.9-3.6 times as
long as wide; venation palmate-pinnate, the midrib dis
placed to divide blade ±1:1.5, weakly ±7-12-branched
on each side, the inner posterior primary nerve
incurved-ascending almost to or less often beyond mid-
blade, the (1—)2—3 outer posterior primary veins pro
gressively much shorter, all these and random tertiary
venules prominulous on both faces of blade. Peduncles
1.6-4.3 cm, commonly ebracteate (one bract seen); ca
pitula densely 12-18-fld, the receptacle <2 m m diam,
the fls apparently subhomomorphic, the subterminal
fls sometimes stouter but no longer than the peripheral
ones and their androecium (poorly observed) appar
ently not differentiated, the perianth (4—)5-merous,
glabrous except for microciliolate orifice of calyx and
sometimes for a few random hairs at tip of corolla-
lobes; bracts subulate 0.4-1.2 m m , tardily deciduous;
calyx campanulate (1-)1.3-2.6 x 0.7-2.4 m m , weakly
5-nerved, the depressed-deltate teeth 0.15- 0.3 m m ;
corolla 5.5-7 m m , seemingly ochroleucous reddish-
tinged, the lobes 1.1-2 m m ; androecium 14—18-mer-
ous, the tube 6.5-7 m m , either shortly included or
shortly exserted, the stemonozone 0.7-0.9 m m , the tas
sel either deep crimson-red throughout, or distally
pink; ovary subsessile, glabrous at anthesis, the intrast
aminal nectary ±0.35 m m tall. Pods erect, in broad
profile 7-10.5 x (0.65-)0.8-1.2 cm, the sutural ribs in
dorsal view ±2-3 m m wide, the stiffly coriaceous,
weakly distantly trans verse-venulose valves cinnamon-
brown, like the sutures glabrous throughout; seeds
(few seen) to 13 x 6.5 m m , the brown hard testa indis
tinctly mottled, pleurogrammic.
Described from the Pacific coast of Guerrero (near
Acapulco), where perhaps a waif or cultivated; known
subsequently only from oak-pine forest, sometimes
along streams, at 700-2000 m in mountainous s. trop
ical Mexico, within lat. 18°30'-20°N, 100°-105°W: w.
Jalisco; s. Michoacan; Guerrero, s.-w. Mexico. — M a p
6. — FL XI-II, V, the whole cycle not established.
Calliandra bijuga closely resembles the allopatric
C. rubescens and C. goldmanii, but differs from both
in somewhat lower leaf-formula and dark crimson
androecium, from C. rubescens further in larger leaf
lets and from C. goldmanii further in less condensed
brachyblasts and few-nerved stipules.
The equation C. bijuga = C. michelii, proposed
first by Micheli and lately endorsed by McVaugh, is
here accepted without question.
Collectors have described the androecium as either
uniformly crimson or white with rose-pink tassel.
11. Calliandra molinae Standley, Ceiba 1: 39. 1950.
— " H O N D U R A S : Dept. El Paraiso . . . along Rio
Lizapa at Galeras . . . June 27, 1945, Louis O.
Williams & Antonio Molina R. 14142V — Holotypus,
F!; paratypi, Standley & al. 571, Glassman 571, NY!.
Microphyllidious arborescent shrubs (0.5-)2-6 m
with flattened crown, stout plagiotropic, vertically
38 M E M O I R S OF T H E N E W Y O R K BOTANICAL G A R D E N [VOL. 74(3)
corky-ribbed long-shoots and densely thatched brachy
blasts, the new stems and foliage hirsutulous through
out with fine pallid subvertical hairs to 0.4-1 m m , the
firm plane lfts bicolored, lustrous brown-olivaceous
above, dull rufous beneath, hirsutulous on both faces
(hairs of epiphyllum basally dilated) and ciliolate, the
capitula subsessile in axils of brachyblasts; phyllotaxy
distichous. Stipules lanceolate striate, becoming pa
pery and brittle, pilosulous dorsally but glabrescent,
those at primary nodes 5—9(—11) x 2-3 m m , those of
brachyblasts a httle shorter and broader, densely imbri
cate. Lf-formula ii-iii/(7-)8-12, the pinnae of some
brachyblast lvs 1-jugate; lf-stks of larger lvs 6.5-24
m m , the petiole 3-9 x 0.6-1 m m , shallowly grooved,
the one or the longer of two interpinnal segments 4-13
m m ; rachis of further pinnae ±3-5(-5.5) cm, the longer
interpinnal segments 2.5-4.5 m m ; lfts little graduated,
subsessile, the pulvinules 0.2-0.6 x 0.45-0.65 m m ; lft-
blades oblong or ovate-oblong from broadly obtusely
auriculate base, at apex broadly rounded and minutely
apiculate, the longest ones 7—10.5(—11.3) x (2.6-)
3-4.2(-5.2) m m , 2-2.9(-3) times as long as wide;
venation primarily palmate, of (3-)4 nerves from pulv-
inule, the slender, incipiently sigmoid midrib displaced
to divide blade ±1:2-2.5, on posterior side 1-2-
branched well beyond mid-blade, the inner posterior
primary nerve incurved-ascending well beyond mid-
blade, the outer 1-2 primary nerves much shorter, ter
tiary venulation almost 0, the principal nerves finely
prominulous on both faces. Peduncles <4 m m , mostly
concealed by stipules; capitula 7-12-fld, the axis <2
m m ; bracts papery and striate like stipules, broadly
ovate-triangular ±1.5-2 m m ; fls dimorphic, one centric
one with long androecial tube, an intrastaminal nectary
but no gynoecium, the rest with tube ± as long as
corolla, lacking nectary, bisexual, the perianth of all
5-merous, either glabrous or thinly pilosulous; PE
R I P H E R A L FLS: calyx deeply campanulate 3.4-4.1 x
1.7-2.6 m m , the ovate obtuse teeth 0.6-1.2 m m , often
narrowly imbricate at base; corolla ±7 m m , the oblong
lobes 1.6-2.7 m m ; androecium 20-24-merous, ±25
m m , pallid proximally, pink distally, the tube ±6.5 m m ,
the stemonozone ±1 m m ; ovary sessile, narrowly ellip
soid, glabrous at anthesis but pubescent later; CEN
T R A L FL: perianth slightly longer than that of periph
eral fls; androecial tube to 13 m m , the free filaments
thickened and sigmoid. Pod (not seen fully ripe) erect,
in profile 4-5.5 x 0.5-0.6 cm, gray- or gray-brown-
pilosulous overall, the sutural ribs in dorsal view ±2
m m wide, the leathery valves bullate over ±4—5 seeds.
In rocky matorral, often colonial on stream banks,
600-1100 m, locally plentiful on the Pacific slope of
Honduras between 86° and 91°W, in deptos. Gracias,
Tegucigalpa, and El Paraiso, sometimes cultivated. —
M a p 6. — Fl. mid-VI-IX. — Palo corcho.
A m o n g Central American calliandras of sect. An
drocallis with only two or three pairs of pinnae per
leaf C. molinae is readily recognized by its corky-
ribbed long-shoots, relatively few leaflets (7-12 per
longer pinna), and capitula subsessile on brachyblasts.
12. Calliandra hirsuta (G. Don) Bentham, Trans.
Linn. Soc. London 30: 554. 1875. Inga hirsuta G.
Don, Gen. Hist. 2: 395. 1832. — "Native of N e w
Spain [= Mexico]. Mimosa hirsuta Ruiz et Pav[6n] in
herb. Lamb[ert.]." — Holotypus, OXF!; clastotypus,
sent to Britton by Druce, NY!; presumed isotypus,
mislabeled: "Perou, Pavon," G!. — Feuilleea hirsuta
O. Kuntze, Revis. Gen. PL 1: 188. 1891. Anneslia
hirsuta Britton & Rose, N. Amer. Fl. 23: 58. 1928.
C. cumingiifi ? var. galeottii Bentham, London J. Bot. 3: 306.
1841. "Mexico . .. province of Oaxaca at... 6 to 7000 feet,
Galeotti no. 3148." — Holotypus, K! = isotypus of the next.
Inga speciosa Martens & Galeotti, Enum. pi. Galeotti pt. 10:
19 [320]. 1843. — "(G[aleottii] No. 3148 . . . dans les bois
de chenes de Socorro et de la Sierra de Yavesia (cordillere
orientale d'Oaxaca) de 6,000 a 7,500 pieds." — Holotypus, BR n.v.; isotypus = holotypus of the preceding!. —Annes
lia speciosa Britton & Rose, N. Amer. Fl. 23: 58 1928. Cal
liandra speciosa Standley, Publ. Field Mus., Bot. Ser. 4:
309 1929. Calliandra nitida S. Watson, Proc. Amer. Acad. Arts 22:
410. 1887. — "[Mexico. Jalisco:] Rio Blanco [near
Guadalajara], in deep ravines; July [1886). [E.] Palmer
178." — Holotypus, GH!; isotypi, K!, NY!. — Equated
with Anneslia hirsuta by Britton & Rose, 1928: 58; Mc
Vaugh, 1987: 160.
Anneslia pubiflora Britton & Rose, N. Amer. Fl. 23: 58.
1928. — "[MEXICO.] ... near Fenix, Chiapas, May 1925.
[C. A.]Purpus 10346" — Holotypus, NY!; isotypus, US!.
— Calliandra pubiflora Standley, Publ. Field Columbian Mus., Bot. Ser. 4: 309. 1929.
A. diqueti [sic, the collector's name spelled Diquet on the label] Britton & Rose, N. Amer. Fl. 23: 58. 1928. —
"[MEXICO.] Sierra de Nayarit, Territoire Huichol, Jalisco,
Leon Diguet." — Holotypus, NY!; isotypus, P (fide
McVaugh). — Spelling of the epithet emended to digueti
by McVaugh, 1987: 161.
A. houghiana Britton & Rose, N. Amer. Fl. 23: 59. 1928. —
"[MEXICO.]... type from near Tehuacan, Puebla, August
1, 2,1901, [J. N.] Rose and [R.] Hay 5909." — Holotypus,
NY!; isotypus, US!.
Calliandra matudai Lundell, Lloydia 2(2): 88. 1939. —
"Eizi Matuda 1888, collected at Buena Vista, Escuintla,
Chiapas, Mexico, Jan. 1938." — Holotypus, MICH! = MICH Neg. 3832; isotypus, NY!.
Pithecolobium siltepecense Lundell, Contr. Univ. Michigan
Herb. 7: 16. 1942. — "MEXICO: Chiapas, Barranca
Honda, Siltepec [±15°40'N, 92°20'W], Oct. to Nov., 1940,
Eizi Matuda 4132." — Holotypus, MICH!. — Cojoba sil-
tepecensis L. Rico, Kew Bull. 46: 511. 1991. Non Callian
dra siltepecensis Lundell, Phytologia 2: 1. 1941, quae =
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 39
(H. Hernandez, 1991: 821 ["siltepensis"]) Zapoteca por-toricensis (Jacquin) H. Hernandez. Calliandra hirsuta sensu Standley, 1922: 388 C. nitida sensu Standley, 1922: 386.
Shrubs (l-)2-12(-20) dm with either one or few,
erect or diffuse, terete few-branched stems from a fru-
ticose base or an oblique woody root, variable in pu
bescence of lvs and fls, in lf-formula of primary lvs
(sometimes lacking at anthesis), and in organization of
the inflorescence, the obscurely umbelliform capitula
of bright red fls arising 1-3 together either preco
ciously from efoliate nodes prior to development of
foliage, or seemingly from primary lf-axils, or most
often from axils of short, but not acaulous or thatched,
lateral branchlets, the young stems and lf-axes densely
or thinly pilose or strigulose with fine, wavy or
straight hairs to 0.3-1.1 mm, the bicolored lfts (when
mature and dried) either lustrous dark brown or dull
dove-gray on upper face, usually ± puberulent but
sometimes glabrous dorsally, commonly both densely
and loosely ciliate but sometimes appressed-ciliolate
or almost eciliate; phyllotaxy distichous. Stipules tri
angular, lanceolate or linear-lanceolate (2-) 3-6.5(-8)
x 0.7—1.5(—1.8) mm, the blades firm erect, indistinctly
l-3(-5)-nerved, becoming dry, deciduous. Lf-formula
plastic, that of primary lvs in periods of vigorous
growth iv-xi(-xiii)/(19-)22-35(-38), that of lvs on
depauperate branchlets (no further described) lower;
lf-stk of primary lvs (2-)3-9(-10) cm, the petiole
5-18 mm, at middle 0.5-1.2(-l .4) m m diam, the
longer interpinnal segments (3-)4-16 mm; pinnae
subdecrescent proximally, thence subequilong or
shorter distally, the rachis of longer ones (l-)2-4.5
(-6) cm, or exceptionally (Forero 10107, COL) to
9-12 cm, the longer interfoliolar segments 0.4-2.5
(-3.5) mm; lft-pulvinules 0.1-0.3 x 0.2-0.45 mm, not
wrinkled; lfts narrowly oblong, lance-oblong, nar
rowly ovate-oblong or linear from mostly obtusely
auriculate, sometimes subtmncate base, straight or
almost so, abruptly acute, the longer ones (2.2-)3.3-
7.5(-9.5) x (0.7-)0.9-1.8(-2.2) mm, (2.5-)3.1^.5
(-4.8) times as long as wide; venation except for fine,
dorsally prominulous midrib faint or imperceptible,
the midrib displaced to divide blade ±1:1.5-2, usually
simple, the l(-2) posterior primary venules faint or
barely visible externally. Peduncles slender l^L5(-6)
cm, very exceptionally bracteate; capitula 5—10(—11)-
fld, the receptacle 1-1.5 mm; bracts 0.6-1.9 mm,
tardily deciduous; fls homomorphic, subsessile or dis
tinctly pedicellate, the stout pedicel 0.2-1.2 x 0.3-0.9
mm ; perianth 5-merous, loosely gray-pilosulous over
all with wavy hairs to thinly strigulose with straight
subappressed hairs or glabrous except for scattered
hairs distally; calyx campanulate, deeply campanu
late, turbinate-campanulate or hemispherical 1.3-2.6
(-3.2) x 1.6-2.3 mm, weakly 5-nerved; corolla 6-9
(-9.8) mm, the ovate lobes (l-)1.3-3.2(4) mm; an
droecium red throughout, (17-)20-28(-30)-merous,
the tube (4-)4.4-7.5(-10.5) mm, the stemonozone
(0.5-)0.8-1.4 mm; ovary at anthesis glabrous, sur
rounded at base by a lobed nectary 0.45-0.7 m m tall.
Pods (little known) in broad profile 5-9 x 0.65-1 cm,
the sutural ribs in dorsal view 1.8-2.8 m m wide, the
firm, low-tumid valves densely gray-puberulent over
all; seed (1 seen) in broad view oblong-elliptic 7.2 x
3.6 mm, the hard testa smooth dull, grayish-brown
faintly speckled, pleurogrammic.
In oak-pine- and degraded oak-woodland, 1460-
2250 m, often in stony places but the microhabitats
poorly reported, scattered in the mountains of s. Mex
ico from n.-w. Jalisco to Puebla. s. through Oaxaca into
Pacific Chiapas. — Map 7. — Fl. in all months of the
year, sometimes when efoliate, sometimes with devel
oping foliage, sometimes with full-grown leaves, the
aspect consequently diverse. — Potosina.
As described above, C. hirsuta is uncomfortably
polymorphic, and may consist of several discrete taxa
or geographic races. Features common to these are
relatively small stature, bright red flowers of moder
ate size, red androecia, and habitat in the pine-oak
belt of southern Mexico. Typical C. hirsuta, which
has been collected repeatedly in central Oaxaca, has
leaf-formulae of (iii-)iv-vi(-vii)/17-28, and is
densely loosely pilose throughout. Named segregates
listed in the synonymy above are weakly character
ized as follows:
a. Calliandra nitida S. Wats., from vicinity of
Guadalajara, Jalisco: leaf-formula (v-)vii-xi
(—xiii)/28—35; leaves (except for ventral face of
leaflets) and flowers densely loosely villous-
pilosulous.
b. Anneslia diquetii Britton & Rose, from western
Jalisco: leaf-formula iv-vii/28-32; leaflets dor
sally subappressed-pilosulous.
c. Anneslia pubiflora Britton & Rose, from Chiapas:
leaf-formula vii-viii/34; leaflets thinly sub
appressed-pilosulous, but perianth densely
loosely so.
d. Anneslia houghiana Britton & Rose, described
from Puebla as having leaf-formula of "ii-iv/
6-16," in reality has vi-vii/26; it is not different
in either number or indumentum of leaflets from
the typical form as interpreted above.
40 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
M A P 7. Distribution of Calliandra hirsuta (G. Don) Bentham in southern Mexico.
e. Pithecellobium siltepecense Lundell, from Chiapas:
leaf-formula v-vi/23-25; leaflets glabrous dor
sally, thinly ciliolate; perianth thinly appressed-
pilosulous. The transfer of this taxon to Cojoba,
proposed by L. Rico, is incomprehensible.
A previously undescribed variant from central
Oaxaca (Hughes 1503, N Y ) has leaf-formula of vii-
viii/22—28, but leaflets and perianth only thinly
strigulose, the foliage in consequence bright green
rather than cinereous.
13. Calliandra peninsularis Rose, Contr. U.S. Natl.
Herb. 5: 135. 1897. — "[MEXICO. Baja California
Sur:] ... in a garden at La Paz . . . collected by Dr.
Edward Palmer, January 20 to February 5, 1890 .. .
(n. 22)." — Holotypus, US!; clastotypus + photo,
NY!. — Anneslia peninsularis Britton & Rose, N.
Amer. FL 23: 58. 1928. — Calliandra sp. Rose,
Contr. U.S. Natl. Herb. 1: 69. 1890, a provisional
description, lacking epithet.
Anneslia lagunae Britton & Rose, N. Amer. Fl. 23: 59. 1928.
— "La Laguna, Lower California, [26 Jan,] 1906, [E. W.]
Nelson and [E. A.] Goldman 7455." — Holotypus, NY, on
2 sheets!; isotypus, US!.
A. brandegeei Britton & Rose, N. Amer. Fl. 23: 62. 1928. —
"Sierra de Francisquito [a peak of Sa. La Victoria], Lower
California, September 29,1899, T. S. Brandegee." — Holo
typus, Brandegee s.n., NY!. — Calliandra brandegeei
Gentry, Allan Hancock Pacific Exped. 13(2): 130. 1949,
quoad nom., exclus. Dawson 1165 e San Jose del Cabo.
Calliandra brandegeei sensu Wiggins, Fl. Baja Calif. 703.
1980.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 41
Microphyllidious shrubs attaining ±2 m with slen
der terete fuscous branches, these together with lf-
axes thinly (initially more densely) puberulent and
often in addition granular, but glabrescent, the firm
plane bicolored lfts at maturity dove-gray above, dull
brown on lower face, either glabrous throughout or
(sub)glabrous minutely ciliolate, the capitula arising
singly or paired from axils of contemporary lvs; phyl
lotaxy indecisive, partly distichous, partly spiral or
irregular. Stipules herbaceous, narrowly lanceolate or
lance-attenuate, weakly 3-5-nerved, deciduous. Lf-
formula v-vii/16-21; lf-stk of primary lvs 4-9 cm, the
petiole (6-) 10-24 m m , at middle 0.5-0.8 m m diam,
the ventral groove obscurely bridged between pinna-
pairs, the longer interpinnal segments 4—13 m m ; pin
nae irregularly graduated, the rachis of longer ones
2-5 cm, the longer interfoliolar segments 1-3 m m ; lft-
pulvinules 0.25-0.45 x 0.2-0.3 m m , not wrinkled; lfts
not or scarcely graduated, the rachis of longer ones
2-5 cm, the blades narrowly oblong or narrowly semi-
ovate from obtusely semi-cordate base, deltate-apicu-
late, the longer ones (3.6-)4-9 x (0.9-)1.4-3.5 m m ,
2.6^- times as long as wide; venation weakly promin
ulous ventrally, slenderly raised dorsally, the almost
straight midrib forwardly displaced to divide blade
1:2, feebly 2-3-branched on either side from near
middle or above, the one simple posterior primary
nerve scarcely attaining mid-blade. Peduncles 12-23
m m , ebracteate; capitula 8-12-fld, the receptacle <1.5
m m diam, the obscurely pedicellate fls homomorphic,
the bracts subulate ±1 m m or less, persistent; pedicels
0.5-0.7 x 0.3-0.4 m m ; perianth 5-merous, the sub-
membranous calyx minutely puberulent around ori
fice, the faintly nerved corolla thinly strigulose; calyx
campanulate 1.6-2 x 1.1-1.2 m m , the tube weakly 5-
nerved, the triangular teeth 0.25-0.35 m m ; corolla
subtubular, slightly dilated distally, 6.6-8 m m , the
ovate lobes 1.4-0.9 m m ; androecium 17-26-merous,
17.5-23 m m , the tube ±6 m m , the stemonozone ob
scure, the tassel red; intrastaminal nectary of bisexual
fls 0.25 x 0.4 m m ; ovary subsessile, prior to fertiliza
tion either glabrous or thinly granular. Pods 1-4 per
capitulum, erect, in profile 6-11 x 0.75-1.7 cm, the
sutural keels in dorsal view 1.4-2 m m wide, the stiffly
papery valves reddish-brown, translucent, weakly
cross-nerved, puberulent overall, low-bullate over
seeds; seeds 4-8 per pod, plumply compressed-
obovoid 6.5-7 x 4.7-5.2 m m , the hard testa dull tan,
microscopically speckled, pleurogrammic.
O n open stony hillsides and brushy banks in the
oak-pine belt, 1400-1800 m, best known from Sa. La
Victoria close to the Tropic fine in Baja California Sur;
reported by Wiggins (1980: 703) from Sa. La Giganta
near 26°N; first seen by E. Palmer in cultivation as a
febrifuge in the city of La Paz. — Fl. VII-I. —
Tabardillo, meaning yellow fever, against which its
roots provided medication.
Calliandra peninsularis is notable for indecisive
phyllotaxy, elaborately decompound leaves, and rela
tively thin-textured pod. It differs from C. californica,
which is found in the same latitude but in desert foot
hills below 550 m, in more numerous pinnae and
leaflets.
The glandular indumentum ascribed to C. peninsu
laris by Wiggins consists of small, thickened multi
cellular, so-called granular trichomes common in the
genus, in this case mixed with simple hairs. The seg
regation of Anneslia brandegeei into a monotypic se
ries (Britton & Rose, 1928: 51) characterized by
glabrous flowers has neither fact nor reason to rec
ommend it.
14. Calliandra sesquipedalis McVaugh, Fl. Novo-
Galiciana5: 168. 1987. — " [ M E X I C O . ] W. Jal[isco]
... Sierra de la Campana 12-13 k m N W of Los Vol-
canes, [23-25 Oct 1952, R.] McVaugh 13795." —
Holotypus, M I C H n.v; isotypus, NY!; paratypus,
Breedlove 35769, CAS!.
Erect, densely foliate shrublet ±3-6 d m tall with
terete long-shoots, the growing tips and lf-axes thinly
minutely puberulent and minutely granular, the small
crowded lfts bicolored, facially glabrous ciliate,
adaxially dark and ± lustrous, the few-fid umbelli
form capitula slenderly pedunculate in coeval lf-axils;
phyllotaxy distichous. Stipules linear-lanceolate or
narrowly subulate ±2-3.5 x 0.5 m m , tardily decidu
ous. Lf-formula iv-vii/25-29; lf-stk of primary lvs
±2-5 cm, the petiole 6-12 m m , at middle 0.6-0.9 m m
diam, the ventral groove bridged at insertion of pin
nae, the longer interpinnal segments 5-7 m m ; pinnae
distally subaccrescent, the furthest or the penultimate
pair longest, their rachises 2.2-4.5 cm, the longer in
terfoliolar segments 1-1.8 m m ; lft-pulvinules at most
0.3 x 0.4 m m , not wrinkled, the blades sessile against
the rachis; lft-blades decrescent only at very ends of
rachis, semi-ovate from short obtuse auricle, slightly
curved upward, acute, those near mid-rachis ±3.5-5 x
1.1-1.4 m m , 3-3.5 times as long as wide; venation
indistinct, the midrib displaced to divide blade ±1:2,
faintly 1-2-branched at or beyond middle, the one
posterior primary nerve sometimes immersed, when
visible incurved-ascending to mid-blade at furthest.
Peduncles ±3-4.5 cm, ebracteate; capitula 3-6-fld,
the receptacle 0.5-1 m m ; bracts ovate 0.4-1 m m ,
42 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
deciduous; fls homomorphic; pedicels 2.5^4 x 0.5
m m ; perianth 5-merous, reddish, remotely strigulose
and charged with a few minute, amorphously pluri-
cellular trichomes, the calyx weakly 5-nerved, the
corolla externally nerveless, its lobes granular-
ciliolate; calyx shallowly campanulate ±1.2-1.7 mm,
the deltate teeth ±0.6 m m ; corolla ±4.5 mm, cleft to
middle, the lance-ovate lobes spreading-recurved; an
droecium red-pink, 18- or 32-merous in 2 dissected
fls, ±2 cm, the tube ±2 m m , the stemonozone ±1 mm;
ovary at anthesis glabrous, surrounded by a lobed
disc 0.55 mm. Pods erect, in broad profile ±4.5-6.5 x
0.6 cm, the sutural ribs in dorsal view nearly 2 m m
wide, the brown, stiffly chartaceous valves weakly
randomly nerved, subappressed-puberulent overall;
seeds (1 seen fully ripe) 6.2 x 3.2 mm, the testa
smooth fuscous, pleurogrammic.
In open places on steep mountainsides in dry oak-
pine forest, 1900-2000 m, known only from the type-
locality, near 20°20/N, 104°30'W in s.-w. Jalisco,
Mexico. — Fl. IX-X.
15. Calliandra californica Bentham, Bot. Voy. Sul
phur 14, pi. 11. 1844. — "[MEXICO. Baja Califor
nia Sur:] Bay of Magdalena . .. 24°38'N." — Holo
typus, collected by Barclay & Hinds in Nov. 1839,
K! labeled "Hinds, 1841"; isotypus, Barclay 3102,
BM!. — Feuilleea californica O. Kuntze, Revis.
Gen. PL 1: 187. 1893. Anneslia californica Britton
& Rose, N. Amer. FL 23: 59. 1928.
Anneslia mixta Britton & Rose, N. Amer. Fl. 23: 59. 1928.
— "San Jose del Cabo, Lower California, September 24,
1890, [T. 5.7 Brandegee 194." — Holotypus, NY!; isoty
pus, U C n.v.
A. mucronulata Britton & Rose, N. Amer. Fl. 23: 60. 1928.
— "San Jose del Cabo, Lower California, March 16, 1911,
[J. N.] Rose 16473." — Holotypus, NY!; isotypus, US!.
Calliandra californica sensu Bentham, 1844: 105; 1875:
552; Standley, 1922: 386; Wiggins, 1964: 594; Fl. Baja
Calif. 703. 1980; Lenz, Fl. Cape Region 60. 1992; Anon.,
Desert PL 10(4): 156, 176 (color photo). 1993.
Stiffly repeatedly branched, either erect or diffuse-
depressed, desert shrubs (0.5-)0.8-2 m tall with terete
fuscous annotinous and older stems and lateral short-
shoots variably extended following rains, diverse in
pubescence, the lvs glabrous through thinly strigose to
pilosulous with fine, mostly straight appressed but
sometimes flexuous and spreading hairs to 0.4-0.9
m m , the firm plane lfts becoming dove-gray above,
brownish beneath, either equally pubescent on both
faces, or pubescent beneath only, or glabrous overall,
the capitula of red-crimson-stamened fls solitary or
geminate in lf-axils of actively growing, either termi
nal or lateral branchlets; phyllotaxy indecisive, mostly
spiral, sometimes distichous in part. Stipules linear-
lanceolate or narrowly lance-triangular (1-) 1.5-4.5
(-6.5) x 0.3-1.1 m m , faintly l-3(-5)-nerved, becom
ing dry and tardily deciduous. Lf-formula i-iii(-iv)/
5-12(-21); lf-stk of primary lvs (1.5-)2.5-16 m m , the
petiole (1.5-)2.5-10 m m , at middle 0.25-O.55(-0.8)
m m diam, the one or the longest interpinnal segment
2-7 m m ; rachis of distal (or only) pair of pinnae
6-26(-28) m m , the longer interfoliolar segments
(0.75-)0.9-3.8H0 m m ; lft-pulvinules 0.15-0.4
(-0.45) x 0.2-0.4(-0.5) m m , not wrinkled; lfts either
subequilong or smaller proximally, the blades nar
rowly oblong to ovate-oblong or oblong-elliptic from
shallowly semicordate or obtusangulate base, obtuse
apiculate, straight or rarely subincurved, the larger
ones (3.4-)3.7-8(-11.2) x (l-)1.4-3.2(-5.5) m m ,
(2—)2.1—3.1(—3.4) times as long as wide; venation
weak, prominulous only dorsally, the midrib only a
trifle displaced from mid-blade, 2-3-branched on each
side from near or above middle, the posterior primary
nerve simple, short, faint. Peduncles (4—)6-17(-20)
m m , ebracteate; capitula (5—)7—13-fld, the receptacle
±1-1.5 m m diam; bracts subulate, <1 m m , persistent;
fls homomorphic (except some staminate only), sub
sessile, the pedicel (often obscure externally) 0.15-
0.6(-0.8) x 0.3-0.6 m m ; perianth 5-merous, usually
strig(ul)ose, less often pilosulous overall, or some
times glabrescent in lower half, or merely ciholate,
exceptionally glabrous overall; calyx submembra-
nous, campanulate or hemispherical (1-) 1.2-1.7 x
1.2-1.8 m m , weakly 5-nerved, the depressed-deltate
teeth 0.25 m m or less; corolla (5.5-)5.8-7.2 m m , the
lobes (1.6-)1.8-2.4 m m ; androecium 17-26(-30)-
merous, (12-)17-24(-26) m m , the tube 2.8-4.2 m m ,
the stemonozone 0.4—0.8 m m ; intrastaminal disc of bi
sexual fls 0.45-0.8 m m tall, lobed; ovary at anthesis
glabrous. Pod 1-2 per capitulum, erect, in broad pro
file 4-8.5 x 0.8-1.1 cm, when well fertilized 4-7(-8)-
seeded, the sutural ribs in dorsal view 1.2-1.8 m m
wide, the stiffly chartaceous valves low-bullate over
seeds, almost always densely puberulent overall,
sometimes canescently so, exceptionally glabrous;
ripe seeds not seen.
Along boulder-strewn desert washes and on dry
hillsides with Idria, Pachycormus, and numerous
Cactaceae, near sea level to 550 m, locally abundant,
endemic to the peninsula of Baja California, from
near 30°N in s. state of Baja California s. to the Cape
region in Baja California Sur. — M a p 8. — FL IX-V,
most prolifically following storms, sporadically even
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 43
MAP 8. Distribution of Calliandra californica Bentham, and C. eriophylla Bentham var. eriophylla and var. chamaedrys Isely in the southern United States and Mexico.
in the hot dry summer months. — Chuparosa;
tabardilla, applied also to C. peninsularis.
Calliandra californica is obviously related to C.
eriophylla, from which Britton & Rose (1928: 50,
key) separated it by the feeble and ineffective charac
ter of leaflets 4-6 (not 3^1) m m long. Its imperfectly
distichous phyllotaxy, filaments crimson-scarlet their
whole length, flowers slightly more numerous (7-13)
per capitulum, perianth a bit longer (5.5-7.2, not
3.4-5.5 m m ) , and allopatric dispersal are stronger,
but not very strong supporting differences, and a case
for demoting one or the other to varietal rank could
be made. Unfortunately, if this were proposed, prior
ity would require C. californica, published in mid-
January 1844, to retain specific rank and C. erio
phylla, much more widely used and published on
February 1 of the same year, to be subordinated. I
gladly leave the decision in other hands.
For a species of relatively narrow range (Map 8)
and uniform desert ecology, C. californica is surpris
ingly variable in leaf-formula and indumentum. These
modes of variation, however, are correlated neither
with one another nor with dispersal. Extreme variants,
such as Hughes 1546 (NY), which has relatively few
and ample, glabrous leaflets and completely hairless
perianth, and Nelson & Goldman 7411 (NY), which
has many small, dorsally silky leaflets and hairy peri
anth, are instantly perceived as different. Both are
44 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
from the Cape region and each is connected by inter
mediates with genuine C. californica from Magdalena
Bay and with the taxonomically negligible Anneslia
mixta and A. mucronulata. The former, the closely
similar Wiggins 5675 (NY), both from near Miraflo-
res, and Lehto 19671 (NY) from cultivation in Ari
zona, together suggest passage into the related C.
peninsularis, found close at hand but at greater eleva
tions (in the oak-pine belt) in Sierra de Victoria.
16. Calliandra eriophylla Bentham, London J. Bot.
3: 105. 1844. — Typus infra sub var. eriophylla
indicatur.
Microphyll xeromorphic shrubs variable in stature
and (seasonally) in display of foliage, in desert and
desert-grassland commonly 1.5-6 dm, depressed and
intricately stiffly branched, but in sheltered places or
deeper soils attaining 12 dm and more erect, occa
sionally on the Mexican Plateau fruticose to 2 m tall
but the trunks scarcely 1 cm diam, the terete, annoti-
nous and older stems fuscous or blanched in age, the
young branchlets whitish-puberulent, the lf-axes, dor
sal face of lfts, peduncles and fl-buds finely strigulose
or pilosulous with shining, straight and forwardly
subappressed, or spreading-ascending and then some
times wavy, hairs to 0.2-0.7 mm, the mature lfts
almost always glabrous on upper face, the few-fid ca
pitula borne mostly solitary on short brachyblasts but
sometimes also axillary to primary lvs of new branch-
lets; phyllotaxy distichous. Stipules linear-lanceolate
or narrowly subulate, those associated with primary
lvs 1^4(-4.8) x 0.15-0.5 mm, all smooth dorsally or
faintly l(-3)-nerved, deciduous. Lf-formula (i-)ii-iv/
8-17(-20), the pinnae of at least some primary lvs 2
or more pairs, but those of many brachyblast-lvs
geminate; lf-stks of primary lvs 7-30 mm, the petiole
2-9.5 m m , at middle 0.2-0.55 m m diam, the inter
pinnal segments a little longer or shorter; pinnae
scarcely graduated, the rachis of longer ones (6.5-)
8-20 mm, the longer interfoliolar segments (0.6-)
0.7-1.8(-2) m m ; lft-pulvinules 0.1-0.25 x 0.15-0.25
m m , not wrinkled; lfts decrescent only at very ends of
rachis, otherwise subequiform, the blades narrowly
ovate-oblong, broad-linear or linear from bluntly au-
riculate base, deltately apiculate or obtuse, the larger
ones 2.5-6(-8) x (0.6-)0.7-1.8(-2.1) mm, (2.3-)2.5-
5.2(-5.7) times as long as wide; midrib of lfts a trifle
forwardly displaced from mid-blade, straight or
almost so, simple or sometimes faintly 2-3-branched
on each side, rarely accompanied by 1-2 short and
faint incurved primary posterior venules, the venation
immersed on upper face of blade, sometimes on both
faces. Peduncles subfiliform 3-16(-24, seldom over
12) mm, ebracteate; capitula (2-)3-7-fld, the fls
homomorphic, the receptacle scarcely more than 1
m m diam; bracts 0.6-1.1 mm, tardily deciduous;
pedicels 0, or very short, then often visible only in
section, at most 0.4-0.7 mm; perianth 5-merous, the
submembranous calyx 5-nerved, the corolla reddish-
pink or carmine, both usually silky-pilosulous but
sometimes thinly so and occasionally glabrous; calyx
shallowly campanulate or patelliform 0.6-2.1 (-2.4)
mm, the teeth depressed-ovate or -deltate 0.3-0.6
mm, ± incurved; corolla (2.4-)3.4-5.5 mm, the lobes
(1.5-)1.8-2.4(-2.8) mm, commonly recurved in age;
androecium (21-)24-34-merous, (13-)16-24(-27)
mm, the internally thickened tube 1.7-3.2 mm, the
stemonozone to 1.2 m m but sometimes obscure, the
tassel distally pink or crimson, an intrastaminal nec
tary in most fls 0.4—1 m m tall; ovary subsessile,
at anthesis glabrous, becoming sericeous after fertil
ization. Pods erect, 4-10 x 0.5-0.75(-l) cm, the
sutural ribs in dorsal view 1.6-2.4 m m wide, the
recessed valves stiffly papery, bullate over 3-8 seeds,
nearly always densely pilosulous with either spread
ing or retrorse hairs, rarely densely minutely pilosu
lous, the ribs either equally pubescent or glabrescent;
seeds 4.8-6 x 3-4.6 mm, oblong-ellisoid to pyriform
or plumply discoid, the hard smooth testa light
brown, nearly always mottled, the pleurogram slen
derly engraved.
Calliandra eriophylla, the one species of its genus
common in the northern Sonoran and Chihuahuan
deserts, with disjunct populations further south, is
difficult to characterize neatly, because of marked
seasonal dimorphism of the foliage. Plants bearing
many, relatively large, primary leaves, in vigorous
spring growth, are deceptively different from those
flowering from brachyblasts after the ephemeral pri
mary leaves have been shed. The species is closely
akin to the vicariant, but less widely dispersed C. con-
ferta, C. californica, and C. biflora, which see for
commentary on differential characters.
According to Benson and Darrow (1954: I.e.) the
plants furnish excellent browse and can regenerate by
suckering.
Despite distracting random variation in leaf-formula
and size of flowers, only one morphologically weak
variety is taxonomically recognizable. With misgiving
I refer to var. eriophylla two collections from Sonora
(VanDevender 92-227; H. S. Gentry 1281, both NY)
that differ in relatively ample leaves (lf-stks to 3-5.5
cm, pinna-rachises to 2-3.5 cm) and more numerous
(28-38, not 16-27) filaments. The specimens, both
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 45
gathered in February, are in poor condition and may
represent a distinct taxon.
Key to the varieties of C. eriophylla
1. Pod 4-7(-7.5) cm long; s.-e. California and n. Baja California e. to s.-w. New Mexico, thence s. to Sonora and over the Meseta Central to Jalisco and Chiapas 16a. var. eriophylla
1. Pod 8-10 cm long; s. Edwards Plateau, Texas 16b. var. chamaedrys
16a. Calliandra eriophylla Bentham var. erio
phylla. C. eriophylla Bentham, 1844, I.e., sens. str.
— "Mexico; Chila in the district of Pueblo [= state
of Puebla], Andrieux, n. 405." — Holotypus, K! = K
Neg. 15532. — Feuilleea eriophylla O. Kuntze,
Revis. Gen. PL 1: 187. 1890. Anneslea [sic] erio
phylla Britton in Britton & Kearney, Trans. N e w
York Acad. Sci. 14: 32. 1894.
C. chamaedrys Engelmann in A. Gray, PI. Fendler. 1: 39. 1848. — "Chihuahua, Dr. Wislizenus [from "Cachimba"], Dr. Gregg [from "Canyon of Ojito"]." — Syntypi, M O . — Equated with C. eriophylla by Bentham, 1875: 552. C. eriophylla sensu S. Watson, Proc. Amer. Acad. Arts 17: 351. 1882; Coulter, Contr. U.S. Natl. Herb. 2: 100, exclus. syn. 1891; Wooton & Standley, Contr. U.S. Natl. Herb. 13(6): 328, exclus. syn. 1910; Standley, 1922: Benson & Darrow, Trees and Shrubs Southw. Deserts ed. 2, 164, fig 34(d), pi. XIX(A). 1954; McVaugh, 1983: 155; Estrada, Legum. Centro-Sur de Nuevo Leon 53. 1992; McClintock in The Jepson Man. 605, 607(fig.). 1993. Anneslia eriophylla sensu Britton & Rose, 1928: 59, exclus. syn. C. conferta. C. chamaedrys sensu A. Gray, PL Wright. 1: 63. 1852.
As described for the species, but the pod relatively
short, mostly 4—6, exceptionally to 7(-7.5) c m long.
In sandy or rocky, well-drained soils derived from
either granitic, calcareous, or recent volcanic
bedrock, in low desert, desert grassland, and mator-
ral, on the Mexican Plateau entering open pine forest,
in the Sonoran Desert mostly below 1000 m, but in
Mexico mostly between 900 and 1950 m, locally
plentiful in the Sonoran and Chihuahuan deserts of s.
United States and Mexico, discontinuously s. to
Jalisco and Chiapas: in United States from s.-e. San
Diego Co. in California e. across Arizona s. of the
Mogollon Escarpment to the s.-w. corner of N e w
Mexico; in Mexico from n. Baja California (Matias
Pass, ±900-1000 m ) and lowland Sonora to w.
Tamaulipas, s. to n. Jalisco and n.-e. Oaxaca; disjunct
s. of the Meseta Central in Jalisco (near Autian,
±1000 m ) and in Chiapas (La Trinitaria, ±1080 m ) . —
M a p g — FL in U.S. II-V, and following rains in
IX-XI, in Mexico mostly in late summer or in winter.
— Fairyduster; mock mesquite.
16b. Calliandra eriophylla Bentham var. cha
maedrys Isely, Madrono 21: 276. 1972. — "[B. L.]
Turner 3642. 15 miles north of Uvalde, Uvalde Co.,
Texas. June 26, 1954." — Holotypus, SMU!. —
Non C. chamaedrys Engelmann, which is a taxo-
nomic synonym of C. eriophylla var. eriophylla, the
varietal epithet in the circumstances deplorable.
C. chamaedrys var. A. Gray, PL Wright. 2: 52. 1853. — "... between the copper mines, New Mexico and the Chiricahui Mountains; July ([C. Wright] 1043). Between the Leona and Rio Frio, Western Texas; July ([C. Wright] 1367)." — The three sheets (NY) mentioned by Isely (1972: 276) correspond to this entity, which Gray provisionally distinguished from C. chamaedrys Engelmann before Bentham had recognized the latter as a taxonomic synonym of C. eriophylla. The three plants are suspiciously alike in age, discoloration, and details of pubescence, as well as in the long pod, such as is unknown in southwestern New Mexico. Probably all are parts of one gathering made by Wright in the first week of July 1849, between the present Uvalde and Brackettville in Uvalde or Kinney County, Texas. C. eriophylla var. chamaedrys sensu Isely, 1973: 79.
In habit, foliage, and flowers exactly duplicating
the more amply foliate states of var. eriophylla found
in the Sonoran Desert, but notable for the relatively
long pod (8-10 cm) and remotely allopatric dispersal.
O n caliche soils of dry hillsides below 450 m, very
local, known with certainty only from the valley of
upper Leona River in Uvalde County, Texas, perhaps
also shortly to the w. in adj. Kinney County. — M a p 8.
— Fl. V-VI.
Characters other than the long pod listed by Isely
(1972: 276) as supporting the status of var. chamaedrys
are ineffective: in Arizona the petiole of larger leaves
reaches 9 m m , and the flowers vary from 3 to 7,
exceptionally 9 per capitulum. Isely's count of 2-5
flowers per capitulum, modified (1973: 79) to 2-4, is
incompatible with a 7-flowered capitulum evident on
Wright 1367 (NY).
17. Calliandra humilis Bentham, London J. Bot.
5: 103. 1846. — Holotypus infra sub var. humili
indicatur.
Functionally herbaceous, microphyll subshrubs
with slender, loosely tufted or weakly assurgent to
humifuse, mostly simple stems (0.1-)0.5-2.5(-3) d m
from woody root and superficial or shallowly subter
ranean rhizomatous caudex, strigulose or pilosulous
nearly overall with fine white hairs to 1(—1.4) m m to
nearly glabrous, the subconcolorous lfts glabrous on
upper face, sometimes (especially in early lvs, some
times in all) strigulose or pilosulous beneath, ciholate
46 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
or not, the small capitula either pedunculate or sessile
and either solitary or paired in a succession of lf-axils
(no brachyblasts), the lf-formula and lft-size also
highly variable; phyllotaxy distichous. Stipules firmly
herbaceous, linear-lanceolate or ovate-elliptic and
straight to falcately lance-elliptic, 2-7 x 0.4-2.3 m m ,
coarsely 1-5-nerved, green or brown, turning dry per
sistent. Lf-formula (i-)ii/(6-)7 to x(-xi)/23(-27), most
often within the extremes of ii-v/8-17, in n.-w. Mex
ico attaining vii-x/27-31, the number and size com
monly mutually adjusted, the lvs often heteromorphic,
the early ones (no further described) smaller simpler,
the lf-stk of those associated with capitula 2-7(-8)
cm, its ventral groove usually continuous between
pinna-pairs, the petiole ±1-4 cm, the longer interpin
nal segments 4—16(-20) m m ; pinnae subequilong or
distally accrescent, the rachis of longer ones
(10-) 13-38 m m , the longer interfoliolar segments
(0.45-)0.7-7(-9) m m ; lft-pulvinules 0.2-0.45 m m
diam, not wrinkled; lft-blades inequilaterally broad-
elliptic, ovate-elliptic, lanceolate, or linear-lanceolate
from shallowly obtusely auriculate base, obtuse when
relatively broad but when narrow sometimes triangu
lar-acute, the largest of a plant 3—10.5(—12.5) x
0.65-4(-6) m m , 2-3.8X4.2) times as long as wide;
venation of lfts primarily palmate, the primary nerves
(l-)3 in smaller lfts, 5 in larger ones, the straight
midrib forwardly displaced to divide blade 1:1.5-2,
either 1- or 2-forked only well above mid-blade, the
inner posterior primary nerve almost as strong, pro
duced almost to tip of blade and enclosing with the
midrib a nerveless elliptic areole, in ample lfts 1 ante
rior and 1-2 weak and short outer posterior primary
nerves, all these immersed on blade's upper face, ob
tusely prominulous beneath. Peduncles 0-32(-40)
m m , either ebracteate or 1-bracteate close below
capitulum, the capitula in some plants all sessile, in
others all pedunculate, in yet others some of each, and
then not rarely 1 sessile and 1 pedunculate from the
same axil; capitula (2-)5-12-fld, the fls homo
morphic, the receptacle not over 1.5 m m ; bracts subu
late or linear-attenuate 0.6-1.6 m m ; pedicels mostly
obsolete, but rarely 1 m m , exceptionally 1.5-2.5 m m ;
perianth 4- or 5-merous, gray-strigulose or -pilosulous
overall to glabrous except for a few fine hairs at tip of
lobes, the calyx 5-nerved or weakly 10-15-nerved, the
corolla externally nerveless; calyx campanulate 1.9-3
x (0.75-)1.2-1.8(-2.4) m m , the orifice often asym
metrical, the longer teeth 0.3-0.7 m m ; corolla nar
rowly campanulate (4.4-)4.8-6.4(-7.5) m m , the erect,
often unequal lobes 0.7-1.3 m m ; androecium (30-)
32-52(-78)-merous, (10-)ll-15(-28) m m , the tube
1.8-4.4 m m , the stemonozone 0.55-1 m m , the tassel
either pink or white; intrastaminal nectary in some bi
sexual fls to 0.4 m m ; ovary subsessile, glabrous at
anthesis. Pods subvertically erect, in profile (2.3-)
2.6-5.5(-5.8) x 0.5-0.7 cm, mostly 3-7-seeded, the
sutural ribs in dorsal view 1.1-1.8 m m wide, the stiffly
chartaceous valves becoming brown or stramineous,
distantly transverse-venulose, either pubemlent over
all, or glabrous, or minutely granular; seeds plumply
rhombic-oblong in broad view, 3.6-5.5 x 2.6-3.6 m m ,
the hard smooth testa gray, variably mottled or speck
led, the pleurogram finely engraved.
In the short span of time between 1838 and 1853,
botanists independently described as species four taxa
that in recent years have been combined with good
reason into one, taxonomically complex C. humilis,
sens. lat. Two of these were encountered by European
travelers at the south margin of the Mexican Plateau,
in Hidalgo and Zacatecas, and were described in
Europe. The two others were based by Engelmann and
Asa Gray on collections of August Fendler and
Charles Wright from N e w Mexico. Bentham (1875)
perceived that only two substantial taxa, C. humilis
and C. reticulata could be maintained, but as new
material accumulated, from both southwestern United
States and Mexico, these also proved to be geograph
ically and morphologically confluent. M o d e m ana
lysts (Benson, Isely, McVaugh) of the complex are
unanimous in recognizing one species composed of
two varieties differing ideally in leaf-formula, ampli
tude of leaflets, and some more trifling characters, but
propose somewhat different definitions of these. Isely
(1973) proposed for var. reticulata in United States
a leaf-formula of i-ii(-iii)/5-6(-8) coinciding with
leaflets 5—10(—12) m m long, in contrast to leaf-
formula iii—vi/6— 14(—20) in var. humilis, coinciding
with leaflets 3-5 m m long. It is disconcerting to find
in an isotype of C. reticulata (NY) pinnae bearing 11,
12, even 14 pairs of leaflets. In fact, leaf-formulae
overlap between plants with relatively ample and dis
tant, as opposed to small and crowded, leaflets. Sup
porting differential characters that have been sought in
density of pubescence, prominence of venulation, and
length of peduncles are all now known to vary inde
pendently and continuously, without any strong corre
lation with dispersal, for the range of var. reticulata is
almost fully contained within that of var. humilis.
Nevertheless the extreme forms of var. reticulata
are instantly recognizable. With their relatively low
leaf-formula and especially with their larger, more
prominently venulose leaflets are generally associ
ated a condensed, even subacaulescent growth-habit,
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 47
indumentum sparse or almost lacking, and capitula
consistently sessile. The taxon reticulata may per
haps consist of neotenous variants of var. humilis that
have arisen independently from the matrix of the
species. Its strange, partly sympatric, partly vicariant
dispersal within the whole range of C. humilis would
harmonize with such a hypothesis.
Granted the significance of leaf-formula in separat
ing vars. humilis and reticulata, it becomes necessary
to recognize further a new variety, gentryana, which
differs in yet more elaborately decompound leaf-
blades and a vicariant dispersal on the west slope of
Sa. Madre Occidental.
Calliandra humilis ranges further north than any
other member of the genus, in Arizona north to the
Coconino Plateau and in N e w Mexico to the upper
Pecos valley in latitude 35°30,N, where it must sur
vive frigid winter temperatures.
The key that follows will discriminate between
most collections of var. humilis and var. reticulata, but
obstinately individual specimens must be expected.
Key to the varieties of C. humilis
1. Lf-formula (ii-)iii-x/10-31 and lfts relatively
small, the larger ones 3-6 mm. 2. Pinnae of larger lvs mostly 3-6, rarely to
9(—11) pairs but lfts <22 pairs; range of the species in s.-w. United States and
upland interior Mexico, in Mexico wholly
e. of Sa. Madre Occidental 17a. var. humilis
2. Pinnae of larger lvs 7-10 pairs and lfts
(23-)26-31 pairs; local on w. slope of Sa.
Madre Occidental, lat. 26o-28°20'N in
Sonora and Sinaloa 17b. var. gentryana
1. Lf-formula i—iii/(5—)6—12 and lfts relatively
ample, the larger ones 6 m m upward; sympatric
with var. humilis .... 17c. var. reticulata
17a. Calliandra humilis Bentham var. humilis. C. hu
milis Bentham, 1846, I.e., sens. str. — "[MEXICO.]
Zacatecas, Coulter." — Holotypus, Coulter 511, K!;
isotypus (fragm), GH!; phototypus, NY!. — Equated
with Anneslia herbacea by Britton & Rose, 1928:
57; non Anneslia humilis (Schlechtendal) Britton
& Rose. Fig. 3
Acacia humilis Schlechtendal, Linnaea 12: 567. 1838. —
"Pr[ope] Reglam [estado Hidalgo, Mexico] . . . (C. Ehren-
berg)." — Holotypus, to be sought at H A L (n.v.); photo
typus, Ehrenberg 563, NY!. — Feuilleea humilis O.
Kuntze, Revis. Gen. PL 1: 188. 1891. Anneslia humilis
Britton & Rose, N. Amer. Fl. 23: 57. 1928. Calliandra hu
milis L. Benson ex B. L. Turner, Leg. Tex. 31 (lapsu pro C.
humilis Bentham). 1959. — Non Calliandra humilis Ben
tham, 1846. Calliandra (?) herbacea Engelmann ex A. Gray, PL Fendler.
39. 1849. — "[August Fendler] 180 . . . Between San
Miguel and Las Vegas [San Miguel County, New Mexico]
. . in 1847." — Lectotypus (Isely, 1972: 179), GH!. —
Anneslia herbacea Britton & Rose, N. Amer. Fl. 23: 57.
1928. — Equated with Calliandra humilis by A. Gray, PL
Wright. 2:53. 1853.
C. humilis sensu Bentham, 1875: 552; Kearney & Peebles,
1951: 396; Correll & Johnston, 1979: 770; C. humilis var.
humilis sensu Isely, 1972: 278; 1973: 80, map 17 (U.S.);
McVaugh, 1987: 163-164.
C. herbacea sensu Turner, Leg. Tex. 31: 1959; Correll &
Johnston, 1979: 770.
Variable in stature, density of indumentum, and
development of peduncles, but lf-formula and lft-size
as given in the key to varieties; lfts commonly imbri
cate or contiguous along pinna-rachis; androecial tas
sel commonly pink in United States, often white s.-
ward from Durango, but the color in most populations
not known.
In desert grassland, open pinyon-juniper and oak-
juniper woodland, and thin yellow-pine forest, n.-e.-
ward in short-grass prairie transitional to forest,
(95O-)1200-2250(-2400) m, locally plentiful but the
populations widely scattered, interior s.-w. United
States and parts of the Mexican Plateau: in Arizona
and N e w Mexico common around the sources of the
Gila River, n. and n.-w. to Hualpai Mts. in Mohave
County and the Coconino Plateau s. of Grand Canyon;
disjunct in N e w Mexico on the upper Rio Grande in
Sandoval County and the sources of Pecos River in
San Miguel Co.; trans-Pecos Texas (Davis, Tierra
Vieja and Chinati Mts.) and adjoining n. Coahuila;
and in Mexico e. of the Continental Divide from
Chihuahua to Zacatecas and n. Jalisco. — M a p 9. —
Fl.(lateV-)VI-VIII.
17b. Calliandra humilis Bentham var. gentryana
Bameby, var. nov., inter speciei suae formas folio-
rum foliolisque valde numerosis, illis 7-10- his
27-31-jugis usque, ulterius patria transmontana
praestans. — M E X I C O . Sinaloa: Ocurahui, Sa.
Surutato, ±26°N, 107°40'W, 1-10 Sep 1941 (fr), H.
S. Gentry 6360. — Holotypus, NY. — Sonora-
Chihuahua line: 56 mi. n.-e. of Alamos, 7.7 mi. w.
of Chinipas, ±27°20'N, 108°40'W, 11 Aug 1980
(fl), E. Lehto 24845. — Paratypus, NY.
Resembling multifoliolate forms of var. humilis in
everything but yet higher lf-formula of vii-x/27-31;
larger lfts 3^.5 x 0.7-1.3 m m .
Open places in pine-oak forest, ±1800-2100 m,
and descending along rio Mayo to the lowlands; local
on the w. slope of Sa. Madre Occidental within lat.
26°-28°N in s.-e. Sonora, adj. Chihuahua, and n.-e.
Sinaloa, Mexico. — M a p 9. — Fl. VIII-X.
48 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
FIG. 3. Calliandra humilis var. reticulata (A. Gray) L. Benson (left) and var. humilis (right).
17c. Calliandra humilis Bentham var. reticulata (A.
Gray) L. Benson, Amer. J. Bot. 30: 630. 1943. C.
reticulata A. Gray, PL Wright. 2: 53. 1853. —
"Stony hills at the copper mines [= Sta. Rita del
Cobre, Grant County], N e w Mexico, Aug. [1851] . .
. [C. Wright] 1045." — Holotypus, GH!; isotypi,
K!, ny!. — Feuilleea reticulata O. Kuntze, Revis.
Gen. PL 1: 189. 1891. Anneslea [sic] reticulata
Britton in Britton & Kearney, Trans. N e w York
Acad. Sci. 14: 32. 1894. FIG. 3
C. reticulata sensu Kearney & Peebles, 1951: 396. C. humilis
var. reticulata sensu Isely, 1972: 279; 1973: 81, map 17
(US); McVaugh, 1987: 164.
Thinly pubescent or glabrate, the stems mostly <1
dm, the peduncles most often 0, rarely well developed
at some nodes; lf-formula and lft-size as given in the
key to varieties; lfts often well separated along pinna-
rachis; color of androecial tassel scarcely known.
In habitats of var. humilis but of narrower range in
and around the margin of the Gila Basin in Arizona
and s.-w. N e w Mexico, thence s. along the w. and s.
margins of the Mexican Plateau to Jalisco, n. Mexico,
w. Hidalgo, and (reportedly, Standley, 1922: 387)
Puebla. — M a p 10. — FL VI-IX.
The vars. humilis and reticulata are known to
occur in close proximity and in a c o m m o n habitat,
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 49
115 no 105 100
C A L L I A N D R A H U M I L I S
• var. H U M I L I S
y var. G E N T R Y A N A
MAP 9. Distribution of the varieties of Calliandra humilis Bentham in southern United States and Mexico.
but no mixed populations or mixed collections are
on record.
18. Calliandra tumbeziana Macbride, Field Mus.
Nat. Hist., Bot. Ser. 8: 89. 1930. — "Pern ... moun
tains east of Hacienda Chicama, Prov and Dept. of
Tumbez, Feb. 19-24, 1927, Weberbauer 7677." —
Holotypus, F!; isotypi, GH!, K!, NY!.
C. tumbeziana sensu Macbride, 1943: 73.
Stiffly branched, microphyll shrubs attaining 3 m,
in habit and lvs resembhng C. taxifolia of the same
latitude on the w. slope of the Andes, the terete long-
shoots gray or blanched, glabrate, the bifariously stip
ule-thatched brachyblasts efohate except at tip, where
bearing 1-3 lvs and a fihform peduncle, the lf-axes
pilosulous with erect pallid hairs to 0.3-0.4 mm, the
crowded, thin-textured lfts facially glabrous, thinly
ciholate; phyllotaxy distichous. Stipules triangular-
lanceolate 2-3 mm, striate persistent. Lf-formula (ii-)
iii-v/18-30; lf-stks 1-2.5 cm, the petiole 1-5 mm, at
middle 0.4-0.5 m m diam, the longer interpinnal seg
ments 5-8 mm; rachis of longer pinnae 12-22 mm, the
interfoliolar segments 0.7-1 m m ; lft-pulvinules
scarcely 0.1 x 0.2 mm; lfts equilong except at further
end of rachis, the blades linear-lanceolate from obtus-
angulate base, subacute, often gently falcate, the larger
ones 2.5-4 x 0.5-0.6 mm, 5-8 times as long as wide;
midrib fihform, a little forwardly displaced, simple,
weakly prominulous dorsally, further venulation not
50 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
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'< X. I A. \
J N V 1 ° \ ̂
^
3^=^-—=llL - -̂ i V J • n/ ^ T̂T —
/N \ ' ) !
1 V o \ •> V)
S. pi t %-i. \ ^ y\ ° \ - m -j\ \
' / h + i F ^ X i \ , o • . v.
\ , t . \ K n ^ •>/>:;
MAP 10. Distribution of Calliandra humilis Bentham var. reticulata (A. Gray) L. Benson in Mexico and the southwestern United States.
externally perceptible. Peduncles ±3.5-4 cm, 0.3-0.5
m m diam, bracteate beyond middle; capitula hemi
spherical ±10—12-fld, the receptacle ±1 m m diam;
floral bracts linear 0.7-1.3 mm; fls homomorphic,
subsessile, the pedicels not more than 0.3 mm; peri
anth striate, seemingly pallid (dried), glabrous prox
imally, the calyx-teeth and corolla-lobes thinly pilo
sulous; calyx narrowly campanulate 2.2-2.4 x 0.9-1
m m , 15-20-nerved, the obtuse teeth mostly ±0.4
m m , but one sinus often much deeper; corolla tubu-
lar-campanulate, little dilated upward, ±5 mm, the
ovate lobes ±1 mm; androecium 26-merous, ±21-23
mm, the stemonozone 0.6 mm, the tube 5-6 m m , the
tassel white; ovary glabrous at anthesis; no intrasta
minal nectary. Pod unknown.
In seasonally dry brush-woodland, near 500 m,
known only from the type-locality on the w. slope of
Cerros de la Brea, near 4° 1 0 % 80°45,W, in depart
ment of Tumbez, Pern. — Map 11. — Fl. II—III(—?).
19. Calliandra taxifolia (Kunth) Bentham, London
J. Bot. 3: 104. 1844. Inga taxifolia Kunth, Mimoses
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 51
MAP 11. Distribution of Calliandra taxifolia (Kunth) Bentham, C. chilensis Bentham, and C. tumbeziana Macbride in South America.
64, t. 20. 1820. — "Crescit in Andibus Quitensium,
Ofiam inter et Loxam, in Paramo de Saraguru
[Zaraguru], alt. 1300 hexapodamm [±3°30'-4°S,
49°10'W]." — Holotypus, P-HBK!; isotypus, +B =
F Neg. 12621. — Feuilleea taxifolia O. Kuntze,
Revis. Gen. PI. 1: 189. 1891.
C. expansa Bentham, Trans. Linn. Soc. London 30: 540. 1875. "Peru near Huanuco, etc., Ruiz and Pavon, Lobb." — Lectotypus, Ruiz & Pavon s.n., K (hb. Benth.)!; isotypi, tB ex G = F Neg. 7229!, G (fragm)!, OXF!. — Mimosa expansa Ruiz & Pavon ex Bentham, I.e. in syn. Feuilleea expansa O. Kuntze, Revis. Gen. PI. 1: 187. 1891.
C. prostrata Bentham, Trans. Linn. Soc. London 30: 554. 1875. — "Peru, McLean in Herb. Hook." — Holotypus, K! = N Y Neg. 1998. — Feuilleea prostrata O. Kuntze, Revis. Gen. PI. 1: 188. 1891. Inga taxifolia sensu Humboldt, Bonpland & Kunth, Nov. Gen. Sp. 6 (qu): 301. 1824; de Candolle, Prodr. 2: 439. 1825. Calliandra prostrata sensu Bentham, 1875: 546. Calliandra expansa sensu Macbride, 1943: 71. C. prostrata sensu Macbride, 1943: 73.
Xeromorphic microphyll, stiffly awkwardly
branched, diffuse shrubs and bushy, even subarbores-
cent shrubs (2—)3—18 dm, with straight long-shoots,
caducous primary lvs, and small capitula of red
(orange) fls arising singly from densely stipule-
thatched axillary brachyblasts, except for glabrous
or facially glabrous but ciholate lfts and glabrate
annotinous stems inconspicuously fuscous- or white-
pilosulous throughout and often in addition randomly
reddish-granular, the firm plane lfts lustrous dark
green above, paler beneath, either 1-nerved or pin-
nately venulose dorsally or on both faces. Stipules
erect, narrowly lance-attenuate or triangular-acumi
nate 2-7 x 0.3-1.4 mm, striately 3-9(-15)-nerved,
becoming stiff and dry, persistent. Lf-formula season
ally variable, that of primary lvs (absent from many
flowering spms) (i—)ii—iv(—v)/15—32(—3 8), that of
brachyblast lvs i-ii/11-20, the lf-stk of plurijugate lvs
to 35 mm, of simpler lvs at most 1-5 x 0.5 mm; rachis
of longer pinnae 1.1-3.2 cm, in random primary lvs
attaining 5 cm, the longer interfoliolar segments
(0.5-)0.6-1.6 mm; lft-pulvinules 0.1-0.25 mm; lfts
decrescent near each end of rachis, otherwise sube-
quilong, the blades narrowly oblong or linear-lanceo
late from shallowly semicordate base, obtuse or
deltately subacute, those near mid-rachis 3-6 x
(0.7—)1—1.5 mm, 3-5.2 times as long as wide; vena
tion most commonly palmate-pinnate, the straight,
either simple or branched midrib forwardly displaced
to divide blade 1:1.5-2, the 2 posterior primary
nerves shortly incurved-ascending, the 5-7 secondary
venules (when externally visible) from each side of
midrib divergent at wide angles to anastomosis with a
slender elevated marginal nerve, the whole venation
either equally prominulous on both faces or im
mersed on upper one. Peduncles 4—22 m m , bracteate
1-2.5 m m below apex, the ovate bract scarcely 1 m m ;
capitula compactly umbelliform, 4—13-fld, the nearly
erect fls homomorphic, the receptacle ±1-2.5 x 1
mm; bracts ovate acute 0.6-1.2 mm, incurved, persis
tent; pedicels 0.2-1.2 x 0.3-0.8 mm; perianth com
monly thinly pilosulous overall but sometimes only
distally, the calyx-tube 15-20-nerved, usually sharply
striate but sometimes faintly so, the crimson (orange,
52 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
yellow) corolla externally nerveless; calyx shallow-
campanulate 1—1.9(—2) x 1-1.4 mm, the tube some
times deeply split at one sinus (hence spathiform), the
deltate or ovate teeth 0.3-0.5 mm; corolla tubular-
campanulate (4.5—)5—9.5(—11) m m , slightly expanded
distally, the ovate lobes 0.5-2.8 mm; androecium
crimson (orange) throughout, 19-32-merous, (14.5-)
16-25 m m , the stemonozone 1-1.4 mm, the tube 3-5
(-6) m m ; intrastaminal disc sometimes well differen
tiated 0.35-0.5 mm, but in some fls reduced to a
thickened lining of the stemonozone; ovary shortly
stipitate, at anthesis glabrous, becoming pilosulous
after fertilization. Pods (poorly known) 4.5-9 x ±0.75-
0.9 cm, 5-7-seeded, the sutural ribs in dorsal view 2-3
m m wide, the recessed, stiffly chartaceous valves
thinly pilosulous, low-bullate on each face over alter
nate seeds; seeds in profile 6-7.5 x 4 mm, the smooth
testa dull brown-olivaceous, randomly dark-speckled,
the deeply U-shaped pleurogram ±4.5 x 2 mm.
In xeromorphic brush-thickets and on barren hill
sides at (1250-) 1850-2800 m in w.-draining interan-
dean valleys of s. Ecuador (prov. Azuay and Loja)
and n. Peru, in lat. 3°20'-10°S, in deptos. Amazonas
and Huanuco crossing the Andean crest to the head
waters of rios Maranon and Huallaga; and apparently
disjunct at 350-510 m on the w. foothills of the Andes
in back of Mollendo, near 16°30/S in depto. Are-
quipa. — Map 11. — Fl. II, V-IX, XI, perhaps inter
mittently through the year.
Because of seasonal dimorphism of the foliage and
the appearance of flowers associated either with pri
mary leaves or, after their fall, on brachyblasts lateral
to defoliate stems, the specimens here referred to C.
taxifolia are superficially diverse. In addition to sea
sonal variation in foliage, they demonstrate a clinal
north-south variation in venulation of leaflets and in
prominence of nerves on the calyx. However, I have
found no discrete taxa. The typus of C. taxifolia has
only conjugately pinnate leaves and distinctly venu-
lose leaflets; but Hitchcock 21324 (NY), from a few
kilometers west of Loja, has the same leaflets on pri
mary leaves with up to three pairs of pinnae. The col
lections from Mollendo, interpreted by Macbride
(1943) as C. prostrata, seem morphologically insepa
rable, except perhaps for flower color, said (Dillon
3936, N Y ) to be orange or (Worth & Morison 15748,
G H ) orange-red, rather than simply red as in Ecuador
and most of Peru. Macbride (1943: 71) described the
flower of his collection from A m b o in Huanuco, iden
tified as C. expansa, as "very deep and bright yellow,"
but this may perhaps refer only to the perianth, and
not to the freshy expanded androecium.
20. Calliandra parvifolia (Hooker & Arnott)
Spegazzini, Revista Argent. Bot. 1: 193. 1926. Inga
parvifolia Hooker & Arnott, Bot. Misc. 3: 202.
1833. — "Banda Orientale, Baird. Uruguay,
Tweedie." — Lectoholotypus, Baird s.n., K!. — A n -
nesleya parvifolia Britton, Ann. N e w York Acad.
Sci. 7: 101. 1892, quoad nom., exclus. pi. cit.
(Morong 412), quae = C. brevicaulis M . Micheli.
C. bicolor Bentham, J. Bot. (Hooker) 2: 139. 1840. —
"Uruguay, Tweedie." — Holotypus, Tweedie s.n. from rio
Uruguay, K (hb. Hook.) = N Y Neg. 7957 (2 plants at bot
tom of sheet)!. C. microphylla Bentham, J. Bot. (Hooker) 2: 139. 1840. —
"Minas Gerais. P. Claussen." — Holotypus, Claussen 24 e
Cachoeira do Campo, K (hb. Benth.)! = N Y Neg. 7979. —
Feuilleea multifoliolata O. Kuntze, Revis. Gen. PI. 1: 190.
1891. C. myriophylla Bentham, London J. Bot. 3: 111. 1844. —
"Brazil (Minas Geraes?), Sello, Pohl, Miers." — Lecto
holotypus, Sello s.n., K! = N Y Neg. 1980. — Feuilleea
myriophylla O. Kuntze, Revis. Gen. PI. 1: 188. 1891.
C. peckoltii Bentham, Trans. Linn. Soc. London 30: 555. 1875; & in Martius, Fl. Bras. 15(2): 425. 1876. — "Habitat
ad Canta Gallo [= Cantagallo, near 22°S, 42°30'W], prov. Rio de Janeiro, Peckolt n. 399." — Holotypus not found in
hb. Mart. (BR). — Feuilleea peckoltii O. Kuntze, Revis.
Gen. PI. 1: 188. 1891. C. microcalyx Harms, Feddes Repert. Sp. Nov. Regni Veg. 17: 91. 1921. — "Brasilien: Minas Geraes, Caraca
(Glaziou no. 14648)." — Holotypus, fB = F Neg. 1246;
isotypi, K! = N Y Neg. 7987, P!.
C. falcifera Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 71. 1922. — "[BRAZIL. Para:] . ad stationem Arumateua
viae ferreae alcobacensis prope cataractas inferiores fluvii
Tocantins, l[egit] A. Ducke 4-1-1915 n. 15.650, fruct, 15-7-1916 n. 16.256." — Syntypi, RB n.v.; isosyntypus,
Ducke 15650, M G = F 602743, photo + fragm!.
C. parvifolia sensu Burkart, 1952: 111; Cabrera, Man. Fl.
Aired. Buenos Aires 246, fig. 82(A-C). 1953; Burkart in Parodi, Encicl. Argent. Agri. Jard. 455, fig. 126(B). 1959; Fl.
Buenos Aires 4(3): 421, fig. 133. 1967; Jozami & Munoz,
Arbust. Prov. Entre Rios 118, 120. 1982; Bernardi 1984: 175; Hoc, 1992: 212, fig. 4 + map 2; Stannard, 1995: 382.
C. bicolor sensu Bentham, 1844: 107; 1875: 555, exclus. syn.; 1876: 425.
C. microphylla sensu Bentham, 1844: 110; 1875: 555; 1876: 424.
C. myriophylla sensu Bentham, 1844: 111; 1875: 555; 1876:
425; Glaziou, 1906: 188 (n. 14648, P! exclus. n. 10620, P!).
C. microcalyx sensu Renvoize, 1981: 67; G. P. Lewis, 1987: 175.
Slender arborescent shrubs or subshrubs (0.5-)l-4
m with fuscous, stiffly virgate long-shoots and narrow
multifoliolate primary lvs, the young growth and
especially the dorsal face of lf-axes finely pilosulous
with spreading or forwardly subappressed, white hairs
to 0.25-0.9 m m , the tiny, closely imbricate, either
glabrous, or microscopically cioliolate, or loosely
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 53
ciliate (cilia to 0.4-1.1 m m ) lfts subconcolorous, the
umbelliform capitula arising singly from brachyblasts,
subtended either by a developed coeval If or by a pair
of annotinous stipules, the older brachyblasts thatched
with imbricate stipules; phyllotaxy distichous. Stip
ules lanceolate or triangular-lanceolate 1.5-6(-7) mm,
at first soft green, early stiff, dorsally 1-9-nerved, per
sistent but becoming dry fragile, often blanched in
age. Lf-formula variable, even on one plant, between
primary lvs and those of brachyblasts, (v-)viii-xvii
(-xx)/(20-)24-52; lf-stks of primary lvs 2-8(-9.5)
cm, the petiole 3-12 mm, the longer interpinnal seg
ments 2.5-6 m m , the stalk little swollen at insertion of
pinnae, the ventral groove continuous or weakly
bridged; rachis of longer pinnae (8-) 12-22 mm, the
interfoliolar segments 0.15-0.5 m m ; pulvinules
0.1-0.2 m m ; lfts linear from obtusely auriculate base,
obtuse or apiculate, straight, the longer ones (2-)2.2-
5.2(-6) x 0.5-0.9(-l) mm, 4-6.5(-6.9) times as long
as wide; midrib simple or very faintly pinnate, sub-
centric, immersed on upper face. Peduncles slender or
subfiliform 1-3.4 cm, either bracteate or not, the bract
when present ±1 m m , inserted either above or below
mid-peduncle; capitula 6-16-fld, the fls ordinarily
heteromorphic, the peripheral ones subsessile to slen
derly pedicellate, the terminal one (sub)sessile, not or
scarcely longer than the rest but broader, the floral
receptacle 1-1.5 mm, sometimes produced as a short
terminal pedestal; bracts of outer fls submembranous,
caducous, those of further fls wanting; perianth either
reddish or greenish-yellow with rubescent teeth and
lobes, thinly pubemlent distally or almost glabrous;
PERIPHERAL FLS: longer pedicels (0.4-)0.7-4
(-4.5) m m ; calyx vase-shaped 1.8-2.6(-3) x 1.4-2.2
mm, weakly 5-nerved, the obtuse teeth 0.25-1 mm;
corolla vase-shaped 4.2-7 mm, the ovate lobes
1.1-1.6 m m ; androecium 12-32-merous, (19-)21-50
(-54) m m , the stemonozone 0.45-0.8 mm, the tube
1.5-3.1 m m , the tassel of filaments bicolored, white or
pallid in lower half, carmine distally, the color deep
ening with age; ovary subsessile, at anthesis either
glabrous or distally pubemlent; intrastaminal nectary
0; T E R M I N A L FL (in some capitula abortive):
broadly campanulate, with 5-lobed intrastaminal nec
tary 0.6-0.8 m m tall, the stamens sometimes more
numerous than those of outer fls. Pods in profile (4-)
5-10.5 x 0.6-1.2 cm, straight or gently retro-arcuate,
the dilated sutural ribs ±2-3.5 m m wide in dorsal
view, the plane, recessed valves stiffly coriaceous
or lignescent, transversely sinuously venulose, the
whole pod either pilosulous overall, or glabrous
except for puberulent margin, or glabrous except for
red-granular faces; seed-funicles basally dilated; seeds
(few seen) ±6.5-7.5 x 5 mm, the U-shaped pleuro
gram 6 x 3 mm.
On river banks, rocky shores, and in gallery-mar
gins, n.-ward around outcrops in campo cerrado, from
near sea level on dunes of coastal n.-e. Bahia and on
the Plata estuary in Argentina to 950-1150 m on the
Brazilian Planalto and 1250 m on Pico de Almas in
Bahia, discontinuously dispersed over e. Brazil from
s. Maranhao (Grajau) and Ceara to interior Rio de
Janeiro, w. to s. Goias and w. Parana, thence w. into s.-
e. Paraguay and along river banks s. to the w. Uruguay
and the Plata estuary in Buenos Aires, Argentina; dis
junct, in Amazonian campina, on lower rio Tocantins
in Para (near Camet Arumateua) Brazil. — Map 12.
— Fl. (VIII—)X-I(-II). — Angiquinho; plumerillo;
borbas de obispo; niho azote; flor de seda; chicote de
niho (Argentina, Hoc).
As might be foreseen from the synonymy cited
above, C. parvifolia varies considerably in features
of which the importance has been exaggerated: leaf-
formula, pubescence (especially of the fruit), length of
peripheral pedicels, flower-size, and number of sta
mens. Bentham acknowledged close similarity
between C. bicolor (an admitted synonym, now
untenable, of C. parvifolia), C. myriophylla, C. peck
oltii, and C. microphylla, but noted these particulari
ties (see his conspectus of Calliandra, Bentham,
1876: 408) of each: in C. bicolor only 3-6 pairs of
pinnae; in the rest 10-20 pinna pairs coupled in C.
myriophylla with densely pubescent young foliage
and scarcely pedicelled flowers about 6 m m long, in
C. microphylla with glabrescent leaves and sessile
flowers about 4 m m long, and in C. peckoltii with gla
brous leaves and pedicellate flowers 4 m m long. The
fruit of C. bicolor was pubescent, that of C. myrio
phylla glabrous, and that of C. microphylla facially
pubescent but marginally glabrous. In material now
available for comparison, the leaf-formula, leaf-
pubescence, and flower-size are independently and
continuously variable, and I have found no credible
geographical or morphological patterns in them. Ves
ture of the fruits needs further study, for few have been
collected, and the ovary at anthesis is always glabrous,
acquiring vesture only after fertilization. There are
trends toward higher leaf-formula and longer pedicels
in Brazil northward from state of Rio de Janeiro, and
these are accompanied by slightly shorter and some
times narrower calyx; whereas from central Goias to
Paraguay and Argentina the pedicels are often (but not
in fact always) shorter and the corolla a trifle longer,
but not in concert with pinna- or leaflet-number. I
54 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
M A P 12. Distribution of Calliandra parvifolia (Hooker & Arnott) Spegazzini in South America.
evaluate C. microcalyx, a name taken up by Renvoize of short-shoots, which may be the only leaves present
(1981) for the Bahian forms of C. parvifolia, as essen- at flowering time. I have found, in addition, that the
tially equivalent to C. peckoltii though the leaflets are androecium in plants that approximate the C. peck-
a little fewer. Leaflet-number is, however, quite vari- oltii-microcalyx form is commonly less than 20-mer-
able between leaves of primary long-shoots and those ous as contrasted with 20-32-merous elsewhere in the
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 55
species. It might in consequence be possible to recog
nize in C. parviflora two varieties divided approxi
mately by the 45th meridian (see M a p 12), but cer
tainly not more than one independent species. The
altitudinal range of C. parvifolia is notable. It is as
sumed that the riparian populations along the lower
Uruguai and Paraguai rivers are colonists from further
north, where precisely similar forms occur at eleva
tions up to 650 m. In Bahia C. parvifolia is well
known from elevations of 900 m upward in Chapada
Diamantina and recurs, somewhat surprisingly, on
dunes of the Atlantic coast north of Salvador. The ped
icels in this coastal form are exceptionally long (to 4.5
mm, Queiroz 2534, HUEFS).
20a. Calliandra carrascana Barneby, sp. nov, folio-
rum formula foliolisque minimis imbricatis C. parv-
ifoliam Sprengel et C. bellam Bentham revocans, ab
ambabus floribus maribus (calyce 7.5-9.5, nee 1.8-
4.3 m m , corolla 11-14, nee 4.5-9 m m longis) et
legumine latiusculo (12-) 13-16, nee 8-12 m m lato,
ulterius a priori filamentis ineunte anthesi albidis, ab
hac pinnarum longiorum axi 1.3-2.7, nee 5.5-9.5
m m longo necnon foliolis minus numerosis 24—29-,
nee 40-64-jugis parvis ±2.5-3.5, nee 5-9.5 m m
usque longis diversa. — BRAZIL. Minas Gerais,
Mun. Januaria: Distrito Fabiao, in carrasco, 2 k m na
estrada partindo do abrigo do Malhado, ±15°7'S,
44°15,W, 23.V. 1997 (fl), J. A. Lombardi 1656 & A.
Salino in herb. B H C B 37138. — Holotypus, B H C B ;
isotypus, NY. — Ibidem, eadem die, J. A. Lombardi
1707 (fr). — Paratypi, B H C B 37759, B H C B , NY. —
Minas Gerais: Vale do rio Peracu, ±15°7'S, 44°15'W,
24. V. 1997 (fr), A. Salino 3068 in herb. B H C B 37071,
B H C B , NY.
Stiffly branched but unarmed, elaborately micro
phyllidious shrubs and treelets 2.5-3 m, with terete
brown older stems, thinly pubemlent lf-axes, and
facially glabrous, microciliolate lfts, the phyllotaxy
distichous, the capitula arising singly from thatched
brachyblasts. Stipules firmly papery, finely striate,
ovate or lanceolate 2-7 m m , the blades subulate or
linear-lanceolate glabrous, becoming dry and brittle.
Lf-formula vi-xi/24-29; lf-stks 2.5-6 cm, the petiole
5-10 m m , the interpinnal segments 3.5-7 m m ; rachis
of longer pinnae 1.3-2.7 cm, the interfoliolar seg
ments 0.3-0.6 m m ; lfts scarcely graduated, subses
sile, linear-lanceolate or linear, obtuse or acute,
straight or slightly incurved, the longer ones (2-)2.5-
3.5 x 0.45-0.75 m m , the very slender midrib scarcely
excentric, simple or almost so. Peduncles 14—24 m m ,
bracteolate, the bracteoles ±4 m m ; capitula (few
seen) ±9-flowered, the axis scarcely 2 m m ; fls 5-mer
ous, seemingly homomorphic; pedicels obconic,
scarcely 2 m m ; calyx campanulate, a trifle tumid,
7.5-9.8 m m , thinly pilosulous, the faintly striate tube
4.5-7 m m , the ovate lobes 2.5-3 m m ; corolla pale
green, white-strigulose, 11-14 m m , the campanulate
tube ±4.5 m m , the ovate lobes 5-6 m m ; androecium
31-33-merous, 3.5-6 cm, the tube 5-6 m m , the tassel
at first white, faintly pink-tinged distally in age; ovary
at anthesis densely minutely pubemlent, glabrescent.
Pods subsessile, apparently ascending, in profile
oblanceolate straight, obtuse or barely apiculate, 7-
8.5 x (1.2-) 1.3-1.6 cm, planocompressed, the sutures
in dorsal view ±3-4 m m wide, the stiff plane, almost
glabrous valves transversely venulose; dehiscence of
the genus; seeds not seen ripe.
In carrasco, at elevations not recorded, known only
from the upper S. Francisco valley in mun. Januaria,
near 15°07'S, 44°15'W, in n. Minas Gerais, Brazil. —
Fl. IV; fr. V-VI.
In shrubby stature and relatively small plurifolio-
late leaves C. carrascana closely resembles C. parvi
folia, but is abmptly distinguished by large gray-silky
flowers, a campanulate rather than obconic calyx, and
broader pod. For comparative measurements, see the
Latin diagnosis. It may be related also to C. bella,
which is characteristic of Atlantic forest immediately
to the northeast, but differs not only in flower-size but
in much longer pinnae and more numerous leaflets,
and thereby a distinct facies. The androecial tassel of
the one known flowering collection of C. carrascana
is recorded as white, like that of C. bella. The sym
patric C. dysantha is suffruticose, and has subsessile
capitula with crimson-scarlet filaments.
21. Calliandra foliolosa Bentham, London J. Bot. 3:
110. 1844. — "Brazil, Sello, near Formigas in Minas
Geraes, Gardner, n. 4525." — Lectotypus, Gardner
4525, collected 11-13 Jul 1839 (fl) near the present
Montes Claros, Minas Gerais (±17°S, 44°W), K (hb.
Benth.)! = N Y Neg. 7977; isotypus, K (hb. Hook.)!;
paratypus, Sello s.n., K! = N Y Neg. 7976. —
Feuilleea foliolosa O. Kuntze, Revis. Gen. PL 1:
187. 1891.
C. diademata Lemaire, Jard. Fleur. 3: t. 305, p. 3-6. 1853. —
Described from living plants cultivated in the Netherlands
from seeds collected in 1842 by Libon on river banks near
Vila Franca, Sao Paulo. — No typus seen. — Equated by
Bentham (1875: 555) with C. bicolor, but the perules of
squames naviculaires glabres scarieuses brunes described
by Lemaire, in conjunction with bicolored filaments, are
diagnostic of C. foliolosa.
C. Sancti Pauli Hasskarl, Retzia 214. 1855. — "St. Paul,
Americae meridionalis nescio an sit provincia Brasiliae
56 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
aut insula Oceani atlantici." — Described from plants that
flowered at Bogor, Java, in 1855, grown from stock origi
nating from American seed cultivated at Utrecht; authen
tic specimens, perhaps isotypic, Hasskarl s.n., e. Java
mis it Teysmann n. 7568, K!, NY!. — C. tweedii var. Sancti
Pauli [sic] Bentham, Trans. Linn. Soc. London 30; 553.
1875. C. hirsuta var. Sanct-Pauli [sic] Macbride, Contr.
Gray Herb., ser. 59: 5. 1959. — Mistakenly equated with
C. tweedii by Parodi, Encicl. Argent. Agri. Jard. 455. 1959.
C.foliolosa sensu Bentham, 1875: 553; 1876: 423; Glaziou,
1905: 188, lapsu "foliosa"; Burkart, 1952: 111; 1979: 98;
Bernardi, 1984: 173; Hoc, 1992: 217, fig. 7 + map 2.
C. tweediei var. sancti-pauli sensu Bentham, 1876: 424;
Burkart, 1979: 103 + map.
C. hirsuta sensu Spegazzini, Rev. Argent. Bot. 1: 195. 1926;
non C. hirsuta (G. Don) Bentham; synonymy confirmed by Hoc, 1992: 220.
Arborescent shrubs 2-7(-8) m with pallid glabrous
annotinous and older branches, the new stems and lf-
axes silky-pilose with mostly straight, in age more
flexuous, slender lustrous hairs to 0.7-1.8 m m , the
narrow crowded subconcolorous, facially glabrous
lfts silky-ciliate or finally glabrate, the umbelliform
capitula arising singly and geminate from the axil of
coeval lvs low on homotinous long-shoots. Stipules
papery brown striate, elliptic obtuse or acute 7-17
m m , loosely involute, mostly deciduous at maturity
of associated If. Lf-formula v-ix/30-57(-60); lf-stks
and pinnae pliantly sinuous when expanding, when
mature stiff straight, the former 4.5-9.5(-l 1) cm, the
petiole 8-22 m m , the ventral groove continuous be
tween pinna-pairs, the longer interpinnal segments
5-14 m m ; pinnae proximally decrescent, thence sub-
equilong, the longer distal ones becoming 3-6(-6.5)
cm, the longer interfoliolar segments 0.4—1.1 m m ; lfts
decrescent at each end of rachis or only distally, either
contiguous or narrowly imbricate, sessile or nearly so
against rachis, the pulvinule 0.1-0.2 m m , the blade
linear from obtusangulate or obtusely auriculate base,
acute or apiculate, straight or almost so, the larger
ones 4.5-9.5 x 0.7-14(-l.7) m m , (5-)5.5-8 times as
long as wide; midrib slightly displaced forward from
middle of blade, simple or weakly pinnate, the sec
ondary venules widely divergent, the venation pro
minulous only beneath. Peduncles 2.5-5.5 cm; bract
resembhng stipules but smaller, caducous; capitula
(3—)5—13-fld, the fls usually dimorphic, the periph
eral ones cuneate in outline and ± pedicellate, 1-2 ter
minal ones (sub)sessile, broadly campanulate, the
receptacle 1.5-3 m m ; bracts subtending 1-3 outer
most fls linear-oblanceolate 4-8.5 m m , deciduous;
P E R I P H E R A L FLS: pedicel (0.6-)l-4 m m ; perianth
thinly pilose, especially distally, with white or partly
ferruginous hairs, weakly arborescently venulose, not
striate; calyx 4.5-7 m m , the ovate or lanceolate teeth
1.6-3.5 m m ; corolla vase-shaped (7.5—)8—11(—13.5)
m m , the ovate lobes 2.2-3.5 m m ; androecium 26-AA-
merous, 43-55 m m , the stemonozone 0.7-1.6 m m ,
the tube 1.5-4.5(-5.5) m m , the tassel white or pale
pink proximally, pink or crimson distally; ovary at
anthesis either glabrous or barbellate distally; C E N
T R A L FL(S): perianth scarcely longer than that of
peripheral fls, but broader and calyx rounded at base;
androecium to 64-merous; intrastaminal nectary 5-
lobed, to 2 m m . Pods 1—2(—3) per capitulum, erect, in
profile (4.5-)6-9 x 0.7-11 cm, the thickened sutural
ribs and the recessed valves woody throughout, verti
cally venulose, densely pilose-tomentulose overall
with silvery-gray or rufescent hairs; seeds (httle
known) oblong-obovoid, plumply compressed, 8-9.5
(-10) x 4.5-5 m m , the testa brown, sometimes fus
cous-spotted, the U-shaped pleurogram 5-5.5 m m .
In dense moist lowland forest and ascending in
gallery woodland to 1000 m, surviving in second-
growth woodland on moist soils, locally plentiful in
the valleys of middle Paraguai, lower Parana, and
upper Umguay rivers in s.-e. Paraguay, n.-e. Argentina
(Misiones), and adj. Brazil, thence n. in Brazil in scat
tered stations to s.-w. Goias (Caiaponia) and centr.
Minas Gerais (Monte Claros, the type-locality). —
M a p 13. — Fl. VII-IX(-?). — Maricd, cabela de
anjo, angico, sarandi (Brazil); nino azote (Argentina).
Calliandra foliolosa resembles C. tweedii in boat-
shaped, brown, striate stipules, and in coeval expan
sion of leaves and flowers, but differs in prevailingly
higher leaf-formula, capitula arising directly from
axils of new leaves rather than from the axils of efoli
ate stipules, and especially in the white-and-pink, not
completely red tassel of filaments. In few examples
dissected, the thick distal flower of C. foliolosa had a
pronounced five-lobed disc around the base of the
ovary, whereas the corolla of C. tweedii was merely
thickened internally in the region of the stemonozone.
22. Calliandra tweedii Bentham, J. Bot. (Hooker) 2:
140. 1840. — "Mountains of the Rio Jaqury [=
Jacuhy, Rio Grande do Sul, Brazil], Tweedie." —
Holotypus, K (hb. Hook.)! = N Y Neg. 1978; isoty
pus, K (hb. Benth.)!. — Feuilleea tweedii O. Kuntze,
Revis. Gen. PI. 1: 189. 1891. Anneslia tweediei
Lindman, Bihang Kongl. Svenska Vetensk.-Akad.
Handl. 24,Afd. 3(7): 51. 1898.
Inga pulcherrima Sweet [Hort. brit. 1: 483. 1827, nom.
nud.] ex Paxton, Mag. Bot. 11: 147 + fig. 1844. — De
scribed from plants cultivated at Chatsworth. — Lectoty
pus, the cited figure!. — A plant from "Chelsea, 1848," ex
herb. Thomas Moore (K) confirms its identity.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 57
M A P 13. Distribution of Calliandra foliolosa Bentham in South America.
Mimosa yaguaronensis Larranaga, Inst. Hist. Geograf. Uruguay 1: 176. 1922. — "Se cultiva . . . traido de
Yaguaron." — No specimen seen; equated with C. tweedii by Burkart, Darwiniana 8: 226. 1948.
Calliandra tweedii sensu Bentham, 1844: 107; 1875: 553;
Burkart, 1952: 111; Parodi, Encicl. Argent. Agri. Jard. 455, fig. 126C. 1959; Hoc, 1992: 215, fig. 6 + map 2.
C. tweediei sensu Hooker, Bot. Mag. 71, t. 4: 88. 1845; Ben
tham, 1876: 424; Glaziou, 1905: 188, exclus. no. 10608,
which = Chloroleucon dumosum (Bentham) G. Lewis; Spegazzini, 1926: 196.
Anneslia tweediei sensu Boynton, Addisonia 1: 75, t. 78 1917.
Arborescent shrubs and tree lets 1-3 (-5) m with
widely spreading branches, dwarfed and diffuse in
rocky places, the young stems, lf-axes, and peduncles
variably pilose or pilosulous with either straight or
entangled, gray or whitish hairs but the branchlets
quickly glabrate, often blanched in age, the plane nar
row lfts either facially glabrous or on dorsal face
thinly pilosulous, commonly ciliate with fine white
hairs to 0.6-1.4 m m , less often glabrous overall, the
compactly umbelliform capitula arising singly, sub
tended by papery striate efoliate stipules, from near
base of short-shoots axillary to coeval lvs, the soft new
foliage and the showy red capitula expanding ±
coevally at annual renewal of growth; phyllotaxy dis
tichous. Stipules papery brown multi-striate, mostly
broadly to narrowly lanceolate or elliptic 4—16 m m ,
loosely involute, deciduous. Lf-formula (ii-)iii-v
(-vi)/25-42(-45); lf-stks when fully grown (2-)2.5-
6.5(-8) cm, the petiole 0.5-2 cm, the longer interpin
nal segments 0.6-1.5 cm, the ventral groove inter
rupted between pinna-pairs; pinnae subequilong, the
longer ones 2-5(-6.5) cm, the longer interfoliolar seg
ments 0.5-1.3 m m ; lfts sessile against rachis and
decrescent at each end of it, the pulvinules 0.15-0.25
m m , the blades linear from shallowly auriculate-semi-
cordate base, acute or deltate-apiculate, straight or
nearly so, those near mid-rachis 5.5-10 x 1-1.8 m m ,
5-6.4 times as long as wide; venation simple or faintly
pinnate, the subcentric midrib usually immersed on
upper face, pallid and finely prominulous beneath.
Peduncles 2-4(-5) cm, charged below middle with a
lanceolate or narrow-elliptic, striate bract 3-9 m m ;
capitula ±8-16-fld, the fls moderately heteromorphic,
the peripheral ones subsessile to distinctly pedicellate,
one or more subterminal ones scarcely longer but ses
sile and distinctly broader, its androecium scarcely
modified; bract of 1-3 outermost fls linear 2-4- m m ;
pedicel of outer fls 0.5^ m m ; perianth of all fls usu
ally pilosulous overall but not so thickly as to conceal
the form or surface, the calyx sometimes glabrate
proximally, always faintly striate, the corolla exter
nally nerveless; P E R I P H E R A L FLS: calyx turbinate
(3-)3.6-6.8(-7.5) m m , the lanceolate or rarely ovate
teeth (1.3-) 1.5-3 m m ; corolla turbinate (5-)6-9(-9.5)
m m , the ovate lobes 1.6-3(-3.6) m m ; androecium
26-46-merous, 32-48 m m , the stemonozone 0.5-1.8
m m , the tube 2-4.5(-5) m m , not thickened within, the
tassel red throughout; DISTAL FL(s): sessile or nearly
58 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
so, the calyx broadly campanulate ±2.5-3 m m diam;
corolla 8.5-11.5 m m ; androecium to 52-merous, the
stemonozone 1-2 m m , thickened (and presumably
nectariferous) internally; ovary subsessile, at anthe
sis glabrous or micropapillate. Pod in profile linear-
oblanceolate 5.5-9 x 0.6-0.7 cm, pilosulous overall,
the massive sutural ribs shallowly sulcate length
wise, the deeply recessed valves coarsely vertically
venulose; seeds not seen.
In scrub woodland, riparian forest, about outcrops,
occasional in restinga, below 800 m (but to 1600 m on
Sa. dos Orgaos), native in s.-e. Brazil, from s. Minas
Gerais to Rio Grande do Sul, and extending into
Umguay and Argentina (Misiones), often cultivated
for ornament in its native range and, since early 19th
century, in warm temperate and tropical regions of
both hemispheres. — Map 14. — Fl. IX-II, IV, VII,
perhaps at intervals through the year. — Diadema,
mandavart, quebra-foice, topete de cardenal (Brazil);
pincel del aire, palode pincel, plumerillo, borlas de
obispo (Argentina).
23. Calliandra bella Bentham, London J. Bot. 3:
110. 1844. — "Brazil, Sello: Villa nova de Almeida,
Martius." — Lectoholotypus (Renvoize, 1981: 72),
Sello s.n., K! = N Y Neg. 7952; probable isotypi,
Sello 822, ̂ B = F Neg. 1230; Sello s.n., G!;
paratypi, collected by Maximilian Wied, not by
Martius, BR (3 sheets, 2 annotated by Bentham, the
third bearing handwritten ticket dated March
1816)!. Vila Nova do Almeida, according to Boker-
man (1957: 219) is in coastal Espirito Santo, near
21°10,S, and Maximilian was in southern Bahia,
near Caravelas, in early 1816. Both Vila Nova and
Caravelas are far south of the certainly known range
of C. bella. — Note that the name C. bella has been
attributed to (Sprengel) Bentham, as though based
on Acacia bella Sprengel, under which name Ben
tham found Martius's specimen at M; but in the pro-
tologue Sprengel is quoted with interrogation, a
mark that disappeared only in the Revision of Mi
moseae (1875). — Feuilleea bella O. Kuntze,
Revis. Gen. PI. 1: 187. 1891.
C. bella sensu Bentham, 1875: 555; 1876: 426; Lewis, 1987: 171.
Slender, amply multifoliolate, arborescent shrubs
flowering when 2-6 m tall, the trunk attaining 8(-?)
cm diam, the young stems and the If- and inflores
cence-axes variably pilosulous with brownish or sor
did-gray hairs to 0.3-0.7 mm, the narrow imbricate
lfts glabrous except for random cilia, lustrous dark
green above, pallid dull beneath, the long-pedunculate
capituliform racemes of usually bronze-, rarely pallid-
silky fls arising solitary or geminate, either a) directly
from distal lf-axils, or b) from short axillary, bracteate
but efoliate lateral branchlets, these often by suppres
sion of distal primary lvs forming a shortly exserted
compound panicle; phyllotaxy distichous. Stipules of
primary lvs linear or linear-lanceolate 3-7.5 x 0.7-1.7
mm, indistinctly ±5-nerved, deciduous, those of florif-
erous lateral axes triangular-acuminate and propor
tionately broader, or lanceolate, exceptionally to 18 x
3 mm, persistent. Lf-formula (v-)vii-xiii(-xiv)/40-64;
lf-stks of larger lvs (7—)8—15 cm, the petiole including
pulvinus (8-) 12-26 mm, the longer interpinnal seg
ments 8-19 mm, the ventral groove bridged at inser
tion of each pinna-pair; pinnae commonly decrescent
proximally, the further ones subequilong or erratically
longer and shorter, the rachis of longer ones (5.5-)
6-9.5 cm, the longer interfoliolar segments 0.7-1.6
mm; lft-pulvinules either pallid or nigrescent, in dor
sal view 0.2-0.5 mm; lfts subequilong except near
base of rachis, the blades linear from deltately auricu-
late base, straight or incipiently sigmoid, porrectly
deltate-apiculate, the larger ones 5-9.5 x 0.8-1.9 mm,
5-6.3 times as long as wide; venation simple or nearly
so, the slender, dorsally prominulous midrib forwardly
displaced to divide blade ±1:2, further venulation
immersed or barely perceptible, but the low-convex
ventral face of blade sometimes finely rugulose. Pe
duncles (when paired of unequal length) to
(2-)2.5-7.5 cm, all charged, mostly above middle,
with an elliptic bracteole 1.5-3(-4) mm; capitula 14-
28-fld, prior to anthesis subglobose becoming at full
anthesis oblong-ellipsoid, the receptacle at and after
full anthesis 4-11 mm; bracts subtending proximal fls
linear or linear-spatulate 1.2-2 mm, caducous, those
of further fls minute or wanting; fls essentially homo
morphic, the terminal one sometimes a trifle larger but
otherwise unmodified; pedicel of lowest fls 0.7-2.4 x
0.6-1.5 mm, when very short drum-shaped, not well
differentiated externally from calyx proper; perianth
appressed-silky or the calyx tube only minutely so or
glabrate, the corolla densely so, the calyx delicately
15-20-nerved, the vesture usually golden-bronze,
sometimes whitish; calyx campanulate 2-4.3 x 2-3.6
mm, the teeth either depressed-deltate or ovate 0.4-1
mm; corolla 7-9 mm, 2.1-3.5 times as long as calyx,
the ovate lobes 2.8-3.7 mm; androecium 26-44-mer
ous, opening white, sometimes fading pinkish, when
fully expanded 3.5-5.8 cm, the thickened stemono
zone 1.8-2.5 mm, the tube 3.5-6 mm; nectarial disc 0;
ovary at anthesis either glabrous, or puberulent at base
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 59
MAP 14. Distribution of Calliandra tweedii Bentham in southern South America.
of style. Pods (little known) ±7-10 x 0.8-0.9 cm, the
sutural ribs in dorsal view 2-3 m m wide, the recessed
valves transversely venulose, the whole densely
brown-pilosulous overall; ripe seeds unknown.
In moist lowland forest and restinga, locally plenti
ful, often in secondary growth and along roadsides,
collected frequently along and near the coast of Bahia
in lat. 13°30'-16°30'S, but known by one collection
from mata de cipo in middle Pardo valley near 15°30/S
and by three from campos gerais at 600-1000 m on
upper forks of rio Paraguacu in lat. 12°45'-13°45'S.
— Map 15. — Esponja; pincel. — Visited by hum
mingbirds (Lewis 2025, NY).
24. Calliandra subspicata Bentham, Trans. Linn.
Soc. London 30: 556. 1875; & in Martius, Fl. bras.
15(2): 427. 1876. — "Habitat ad ripas fluminis
Coanara [? Cachoeira] prope Ilheus provinciae
Bahiensis: Luschnath? (in herb. Martii)." — Holo
typus, BR!= K Neg. 194291. — Feuilleea subspi
cata O. Kuntze, Revis. Gen. PI. 1: 189. 1891.
C. bracteosa sensu Ducke, Mem. Inst. Oswaldo Cruz 51: 426. 1953.
C. bella sensu Renvoize, 1971: 72, fig. 2(16); Lewis, 1987-177.
Amply leafy microphyllidious shrubs 1-2 m with
flexuous long-shoots, the young stems, lf-axes, and
peduncles variably pubemlent or more densely pilose
with fine whitish hairs to 0.2-1.2 m m (interspersed
sometimes with minute reddish granules), the striate
stipules glabrous or glabrescent, the lvs bicolored,
lustrous dark brown-olivaceous above, paler dull be
neath, facially glabrous, ciholate, the compactly rece-
mose capitula arising singly from a stipulate node of
efoliate brachyblasts coeval with primary ivs); phyl
lotaxy distichous. Stipules of primary lvs broadly
lanceolate or linear-attenuate 8-12 x 2-4 m m , decur-
rent on lf-spur ±1-3 mm, the blades stiffly papery,
stramineous or finally blanched, striately many-
nerved, persistent, those of brachyblasts a little
smaller, loosely imbricate. Lf-formula ii-iv/28-^3; lf-
stks 1.5̂ 1.5 cm, the petiole including pulvinus 4.5-
13 x 0.6-1.2 mm, the one or the longer of 2-3
interpinnal segments 5-11 mm, the lf-stk dilated
under each pinna, the ventral groove bridged; pinnae
60 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
M A P 15. Distribution of Calliandra bella in Bahia, Brazil.
often a httle accrescent distally, but the furthest pair
sometimes shorter, the rachis of longer ones 4-8.5
cm, the longer interfoliolar segments 0.9-1.5 m m ; lft-
pulvinules 0.1-0.25 x 0.4-0.6 m m ; lft-blades linear
or linear-lanceolate from obtusangulate or shortly ob
tusely auriculate base, deltately (sub)acute, straight or
nearly so, the longer ones 6.5-11.5 x 1.2—1.9(—2)
m m ; midrib of lfts forwardly displaced to divide
blade ±1:2, on posterior side delicately 3-44oranched,
the secondary nerves camptodrome well within the
plane margin, the inner posterior primary nerve in
curved-ascending nearly to mid-blade, these all finely
prominulous on both faces, a weak random tertiary
venulation perceptible on upper face only. Peduncles
12-21 m m , 1-2-bracteate, the lance-ovate bract(s)
4—6.5 m m , resembling stipules in texture and vena
tion, persistent; capitula 14—24-fld, at full anthesis
plumply ellipsoid (discounting androecia), the linear
or linear-clavate receptacle 5-9 m m ; bracts of a few
lower fls resembling bracts of peduncle but smaller,
deciduous, the further fls ebracteate; pedicels 0.8-1.8
x 0.4—1.1 m m ; fls homomorphic, the 5-merous peri
anth ochroleucous glabrous, the firm calyx prominu-
lously 18-25-nerved, the corolla submembranous,
faintly nerved but the nerves immersed; calyx 2.4-4 x
1.5-3 m m , the triangular acute or depressed-triangu
lar teeth (0.3-)0.6-1.2 m m ; corolla 6.8-10 m m , the
ovate acute lobes 2.2-4 m m ; androecium 24~46-mer-
ous, when fully expanded 3.8-7 cm, the stemonozone
M A P 16. Distribution of Calliandra gardneri Bentham
and C. subspicata Bentham in eastern Brazil.
(l-)1.3-4 m m , the tube 3-5 m m , the tassel white;
ovary subsessile, above middle distended and pubem
lent. Pods pilosulous when young, not seen fully
formed.
O n stream-banks and in stony stream-beds, mostly
Below 100 m, and in low caatinga perhaps somewhat
above 100 m, apparently uncommon, in Pemambuco
near Tapera and on Sa. das Varas, and in lowland e.-
centr. Bahia inland on rio de Contas to Jequie reported
by Lewis (1987: 177) s. to valley of rio Pardo. — M a p
16. — Fl. II-IV, VII-X, the full cycle not established.
25. Calliandra dysantha Bentham, J. Bot. (Hooker) 2:
138. 1840. — T y p u s infra sub var. dysantha indicatur.
Micro- or coarsely polyphyllidious, mostly virgate
subshrubs from xylopodium or oblique woody, fire-
resistent rootstock, the stiff, simple or distally few-
branched stems commonly (1.5-)2-14(-20) d m and
dying back annually to the ground, but sometimes per
sistent and fmticose or even subarborescent (report
edly attaining 4 m ) , at anthesis only distally foliate,
the dense, sometimes umbelliform capitula of large,
externally white- or bronze-silky-barbate fls borne
generally at 1^1 furthest lf-axils, either directly or on
condensed brachyblasts of few efoliate internodes, on
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 61
peduncle varying from almost 0 to 5.5 cm, the tassel
of filaments either blood-red or pink-and-white; ves
ture variable in composition, density, and distribution
on the plant, that of young stems and all lf-axes usu
ally composed of longer straight, spreading-ascending
needle-like hairs to 0.8-2.1 m m and shorter, incurved
or partly uncinate ones mixed or not with reddish-
granular trichomes, the long-ciliate or minutely ciho
late lfts bicolored, the ± lustrous upper face of leaden
hue, varying from glabrous to papillate (and then
sometimes microscopically hirtellous), the lower face
cinnamon-brown, dull, either appressed-pilose or
glabrous; phyllotaxy distichous. Stipules of primary
(laminate) lvs erect, symmetrically or asymmetrically
lanceolate, lance-attenuate, or lance-ovate, mostly 6-
18 m m , rarely 2-6 m m long, at base (1-) 1.3-4 m m
wide, striately nerved, dorsally pubescent or glabrate
in age, becoming dry papery and persistent (but ulti
mately fragile), the bracteiform (elaminate) stipules of
short-shoots, when present, sometimes shorter and
wider, commonly glabrate. Lf-formula iii-xi(-xii)/
(16-)18-35(-38); lf-stks (2.5-)4.5-10.5(-15.5) cm,
the obscurely pulvinate petiole 0.5-2.5(-3) cm, the
longer interpinnal segments (0.5-)0.8-2.1(-2.6) cm,
the ventral groove bridged at insertion of pinnae;
rachis of longer pinnae (4.5-)5-14 cm, the longer in-
terfoliolar segments (1.4-)2-6 m m ; lfts subequilong
except at each end of rachis, subsessile in a shallowly
dilated socket, the cross-wrinkled pulvinule (0.2-)
0.3-0.8 m m , the blades oblong or lance-oblong or nar
rowly ovate from shallowly semicordate, obhquely
truncate, or bluntly auriculate base, those near mid-
rachis commonly 7-18(-21) x (1.8-)2-6.5(-7) m m ,
but in var. turbinata 3.5-8 x 0.8-1.8 m m , in any case
2.2-4.3(-5) times as long as wide; venation palmate,
the straight or distally incurved midrib displaced to di
vide blade 1:2-3.4, the inner posterior primary nerve
incurved-ascending to or far beyond mid-blade, the
outer posterior one(s) much shorter, the whole vena
tion either immersed or raised on ventral face, sharply
prominulous beneath. Peduncles often almost 0, or the
lowest, or all, raised on stout, bracteate, white-pilosu-
lous peduncle up to 1.5-5.5 cm; bract sessile, broadly
deltate-ovate or semiorbicular 3-7.5 m m , in texture,
venation and external pubescence resembhng the stip
ules; floral receptacle claviform or subspherical 2.5-4
m m diam; bract of peripheral fls lanceolate or elliptic-
obovate or obovate-tmncate 3-5.5 m m , deciduous,
that of inner fls wanting; capitula prior to anthesis
forming a hairy ball, becoming at anthesis 2-3.5 c m
diam (without filaments), the (7-)8-14 fls contiguous,
either subsessile or contracted at base into a sohd
pedicel 0.8-5.2 m m , all homomorphic or the inner
most broader (but not longer) than outer ones and
these often staminate; perianth firm, externally
densely silky-pilose overall or proximally glabrescent,
brown glabrous within, normally 5-merous (random
exceptions); calyx either broadly or narrowly
turbinate, disregarding vesture 4—7.5(-8) m m , the
(often unequal) lobes usually depressed-deltate or
semicircular 0.4—3 m m , or the orifice merely undu
late; corolla (8.5-)10-14(-15) m m , the spreading-
ascending lance-ovate lobes (2.6-)4—6.5 m m at anthe
sis, sometimes more deeply split in age; androecium
(44-)52-92(-106)-merous, 3.3-5.7(-6.5) cm, the
stemonozone (1.4—) 1.6-4 m m , comeously thickened
internally, the tube 3-6.5 m m , intrastminal nectary 0;
ovary subsessile, at anthesis varying from papillate-
puberulent to white-tomentulose either overall or be
yond middle. Pods 1-3 per capitulum, erect, in profile
oblanceolate 7-13 x 1.1-1.6 cm, 3-7-seeded, the mas
sive sutural frame and the recessed ligneous valves
ahke densely white-, gray-, or yellowish-tomentose;
seeds in broad profile 9-13 x 5.5-9 m m , the hard brit
tle testa smooth, dull-ochraceous, the narrowly U-
shaped pleurogram ±7-9 m m .
A polymorphic species of wide dispersal over the
Brazilian Planalto, in campo cerrado and campo ru-
pestre upward of 500 m, extending from extreme s.
Piauf s. to extreme n. Parana, w. just into Paraguay,
the range more exactly described under each variety.
The foregoing description of C. dysantha records
the presently known macromorphological variation in
a complex species that includes not only the original
C. dysantha as conceived by Bentham but also his
species abbreviata, macrocephala, and turbinata, and
furthermore several minor variants that were not yet
known in 1875-1876. Bentham's four major species
were placed close together in a subordinate group
within ser. Nitidae, within which they were separated
by combinations of three characters: number of pinnae
per leaf; length of peduncles, and mutual proportions
of calyx and corolla. These formulae are no longer ap
plicable to much of the material that has accumulated
in herbaria subsequently.
The commonest, most widely dispersed, and conse
quently best known member of this group, C. dysantha
sens, str., was defined by 4-6 pairs of pinnae, subses
sile capitula, and corolla more than twice as long
as calyx. Copious collections from the relatively re
stricted territory of the Brazilian Distrito Federal now
show pinnae in three to seven, even nine pairs, pedun
cles indeed often obsolete or almost so but the first of
each stem not seldom developed (to 1.5, exceptionally
62 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
2 cm), and calyx often half or more than half as long
as corolla. The free filaments are constantly blood-red.
Calliandra abbreviata was described from a depau
perate specimen with only two pairs of pinnae in the
few remaining leaves, though the description was
soon modified to three pairs seen on the isotype in
Hooker's herbarium (K!) and on a second collection
(Gardner 2835, K!) from northwestern Bahia. A mod
e m collection from nearly the same place (Coradin
5775, K!), otherwise identical, has four pairs, and is
indistinguishable from some C. dysantha of Goias and
Minas Gerais. Calliandra abbreviata is confidently
equated taxonomically with typical C. dysantha.
Most capitula of C. dysantha var. dysantha arise
from brachyblasts on 1-3 efoliate nodes axillary to a
primary leaf—that is, on axes of the second order—
and are huddled together toward the stem's apex on
obsolescent but bracteolate peduncles; the filament-
tassel is uniformly and consistently red. By contrast,
the capitula of C. macrocephala are borne on ex
tended ebracteolate peduncles that arise directly from
well-spaced primary leaves, from mid-stem upward;
and the tassel, so far as known, is never red, but pink
white-tipped, or pink at each end and whitish in be
tween. Against these rather striking differences are
marshalled similarity in leaves, capitula, individual
flowers, and tomentose pod, pointing to a very close
relationship, expressed at the varietal rank.
Calliandra turbinata, also here interpreted as a
variety of C. dysantha, was at first assigned 8-12
pairs of pinnae, short but evident peduncles, and a
corolla scarcely half as long as the calyx. It is now
certain that leaf-formula alone does not separate C
turbinata from var. dysantha, but the notably short
and narrow leaflets, the distinctly pedicelled periph
eral flowers, and the whitish androecium constitute a
strong diagnostic syndrome. Calliandra turbinata
(which includes the inconsiderable C. chapadae)
might be maintained as in independent species except
that w e now have plants, described below as C.
dysantha var. opulenta, in which the foliage of nor
mal var. dysantha coincides not only with the pedun
culate capitula and pink-white tassel of var. macro
cephala but with the pedicellate peripheral flowers of
C. turbinata. A practical advantage of amalgamating
these taxa into one specific unit is that C. dysantha
sens. lat. becomes definable by wide discontinuities
on all sides from related species.
Here must be mentioned an ambiguous specimen
from the eastern edge of the range of C. dysantha in
Bahia (5 k m s. of Caietete, Harley 21096A, K). It
closely resembles some small-leaflet forms of var.
dysantha in all respects but two: the calyx (disregard
ing pedicel) is only about 2.5 m m long, and the fila
ments are described as white pink-tipped. Possibly
this may represent a hybrid derivative of C. dysantha
and C. nebulosa, which has been collected within two
kilometers along the road south of Caietete.
Key to the varieties of C. dysantha
1. Pedicel of peripheral fls of the capitulum 0-1.
8(-2) mm; filament color variable.
2. Peduncles subobsolete, or the lowest
attaining 1.5(—2) cm; filament-tassel
blood-red; extreme s. Piaui s. through w.
Bahia to s. Minas Gerais, w. to centr. and
s. Goias 25a. var. dysantha 2. Peduncles 1.5-5 cm; filament-tassel
either pink white-tipped, or pink at
base and tip but whitish medially; of
bicentric range, sympatric with var.
dysantha in centr. Goias, disjunct around
the S. Paulo-Parana boundary near
23-24°S 25b. var. macrocephala
1. Pedicel of peripheral fls of the capitulum 2-5 mm; filament-tassel colored as that of
var. macrocephala; e. Paraguay, Mato Grosso,
Mato Grosso do Sul. 3. Larger lfts 10-16 x 3-5 mm; range as
given 25c. var. opulenta
3. Larger lfts 3.5-8 x 0.8-1.8 mm; range as
given, except Paraguay 25d. var. turbinata
25a. Calliandra dysantha Bentham var. dysantha. C.
dysantha Bentham, 1840, I.e., sens. str. — "Minas
Geraes, P. Claussen." — Syntypi, Claussen 41, 122
[sent to G from Cachoeira do Campo by Claussen,
and from G to Bentham], K (hb. Benth., mounted on
one sheet)! = N Y Neg. 1970; isosyntypus, Claussen
s.n. K (hb. Hook.)!. — Feuilleea dysantha O. Kuntze,
Revis. Gen. PI. 1: 187. 1891.
C. abbreviata Bentham, London J. Bot. 3: 108. 1844. —
"Province of Piauhy [expanded in Martius, 1876: 422, to 'in
districtu Paranagoa', near 10°20'S, 44°40'W], Brazil, Gard
ner, n. 2556." — Holotypus, K (hb. Benth., pinnae 2-jug.)!
= IPA Neg. 1473; isotypus, K (hb. Hook., pinnae 2-3-jug.,
not accounted for in the protologue)!. — Note: 2 sheets of
Gardner 2835 (K) from Fda. Sta. Rosa on rio Preto in n.-w.
Bahia (not Pernambuco) were mistakenly photographed as isotypes (NY Neg. 7972, 7975!); they are, however, con-
specific. — Feuilleea abbreviata O. Kuntze, Revis. Gen. PI. 1: 187. 1891.
C. dysantha fi pilosa Bentham in Martius, Fl. bras. 15(2):
421. 1876. — "Locis Brasiliae meridionalis non indicatis: St.-Hilaire, Sello." — Lectotypus, Sello 68, K (hb. Hook.)! = N Y Neg. 7977.
C. dysantha sensu Bentham, 1844: 109; 1875: 552; 1876: 421, var. beta et gamma exceptis.
C. abbreviata sensu Bentham, 1875: 553; 1876: 188, exclus. loco pernambucensi.
C. macrocephala sensu Glaziou, 1905: 188; non Bentham.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 63
Stems mostly virgate, simple or few-branched dis
tally (3-)4-14(-20) dm and dying (or burned) back
annually to the rootstock, but occasionally persistent
and fruticose (even arborescent to 4 m), variable in
indumentum, that of young stems and lf-axes com
posed mostly of mixed longer straight needle-like
trichomes to 0.9-2.1 m m mixed with shorter incurved
ones, the lft-blades usually pilose (sometimes
glabrous) beneath and glabrous, papillate, minutely
hirtellous, rarely thinly pilose above, cili(ol)ate; stip
ule-blades 7-18 x (1-) 1.3-4 mm, those of brachy
blasts sometimes shorter; lf-formula (ii-)iii-vii(-ix)/
(16-)18-33, the longer lfts 7-18(-21) x (1.8-)2-6
(-7) m m , 2.2-4.3 times as long as wide; peduncles
often almost 0, but that of the first capitulum often de
veloped and to 1.5(—2.5) cm; calyx of peripheral fls
5-7.5 mm; corolla 9.5—14(—15) mm; androecium
(46-)60-92-merous, the tassel blood-red.
In campo cerrado and campo mpestre, 530-1250 m,
widespread and locally plentiful on the Brazilian
Planalto from far s. Piaui (Paranagoa) s. through w.
Bahia and upland Minas Gerais to ±19°30'S, thence w.
abundantly into centr. and s. Goias and Distrito Fed
eral. — Map 17. — Fl. all months of the year, but most
prolifically XII-IV. — Flor do cao; quebra-foice.
The var. dysantha is variable in width of leaflets:
the narrow and broad extremes appear very different,
and are to some degree correlated with dispersal. In
Goias and Distrito Federal the larger leaflets are al
most consistently (3.5-)4—6.5(-7) m m wide, in Bahia
and Minas Gerais they are mostly commonly 2-3.5
m m wide. However, the apparent gap between the ex
tremes is bridged in Minas Gerais by leaflets 2.5-6
m m wide (in mun. Gouveia) and 2-4 m m wide (Sa.
do Cipo), and on Chapada dos Veadieros in Goias by
leaflets 3-5 m m wide. Frutescent forms, in which the
new stem regenerates from apex of the past year's
growth rather than from the rootstock, are found most
frequently in trans-Franciscan Bahia and may be ge
netically determined, but their flowering branches are
not distinguishable from the whole annual growth of
the more common forms. In the same general region,
the flowers tend to be a trifle smaller than on the
Goias highlands, but not discontinuously so.
25b. Calliandra dysantha Bentham var. macro
cephala (Bentham) Barneby, stat. nov. C. macro
cephala Bentham, J. Bot. (Hooker) 2: 140. 1840. —
"Brasilia, Pohl." — Holotypus, Pohl 1362, K (hb.
Benth.)!; isotypus, K (hb. Hook.)!; presumed isoty
pus, Pohl 605 from "San Pedro," perhaps = Sa. S.
Pedro near Cavalcante, Goias, visited by Pohl in
October 1819, NY!. — Feuilleea macrocephala O.
Kuntze, Revis. Gen. PI. 1: 188. 1891.
C. macrocephala sensu Bentham, 1844: 108; 1875: 553; 1876:423, t.CIX.
Stems mostly 1.5-3 dm, functionally herbaceous
from fmtescent caudex or knotty xylopodium, to
gether with lf-axes and peduncles pilose with fine
hairs to 1-1.6 m m mixed with minute puberulence and
sometimes with scattered granular trichomes, the lfts
all or almost all facially glabrous, cihate, the pedun
cles arising directly from 1-4 distal lf-axils; stipule-
blades 3.5-9 x 1.5-2.5 mm; lf-formula iv-viii/27-35,
the longer lfts 7-11 x 1.8^ mm, 2.8-3.8 times as long
as wide, the venation sharply defined on both faces;
peduncles 1.5-5.5 cm; pedicel of peripheral fls subob-
solete or up to 2 x 2.8 mm; calyx ±4.5-5.5 mm; corolla
±13-14 mm; androecium 44—52-merous, the tassel
pink proximally, white distally.
In campo and campo mpestre, 700-1100 m, of bi-
centric range in centr. and s. planaltine Brazil: between
13°30,S and 17°S on and near the Tocantins-Parana
watershed in centr. Goias (Cavalcante, Chapada dos
Veadeiros, Sa. Dourada); and in lat. 23°20'-24°20'S
astride the Sao Paulo-Parana state line (Tatui, Itarare
Senges). — Map 18. — Fl. IV, IX-XI.
25c. Calliandra dysantha Bentham var. opulenta
Barneby, var. nov, a var. dysantha, quacum foliis
pubeque congruit, capitulorum pedunculo elongato
(22-50 m m longo), florum periphericorum pedi-
cello evoluto ±2-5 m m longo, necnon filamentis
basi roseatis dein albidis (nee omnino sanguineis), a
var. macrocephala (proxime affini) praesertim flo
rum periphericorum pedicello (2-)3-5 m m longo, a
var. turbinato, quoad flores simillimo, foliolis am-
phusculis 10-16 (nee 3.5-8) m m usque longis di-
versa. — PARAGUAY. Depto Amambay: [Esper-
anza,] Sa. de Amambay, 1907/1908 (fl), T. Rojas in
Hassler 10818. — Holotypus, A; isotypi, B M , NY.
— Ibid., 10818a (fr). — Paratypus, A.
C. macrocephala sensu Hassler, Bull. Herb. Boiss. II.
Stems 2-10 dm, functionally herbaceous from
oblique rootstock, the young parts silky-pilose, the fa
cially glabrous lfts cihate, the peduncles and individual
fls white-silky-tomentose; stipule-blades 6-16 x 2-4
mm; lf-formula iv-vii/18-33, the lfts lance-oblong,
deltately subacute, the longer ones 10-16 x 3-5.5 mm,
2.8-4.2 times as long as wide; peduncles (15-)22-50
mm; pedicel of peripheral fls (2-)3-5.2 mm; calyx
4-5.5 x 6-9 mm; corolla 12-14 mm; androecium
90-106-merous, the tassel pink at base, white distally.
64 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Map 17. Distribution of Calliandra dysantha Bentham var. dysantha in eastern Brazil.
In campo subject to fire, ±450-950(-?) m, scat
tered over the s.-w. quarter of the Brazilian Planalto,
from the sources of rio Araguaia in s.-e. Mato Grosso
s. to the rio Pardo in Mato Grosso do Sul, w. into Sas
A m a m b a y and Maracayu in cis-Paraguaian Paraguay.
— M a p 18. — F1.VI-XI.
25d. Calliandra dysantha Bentham var. turbinata
(Bentham) Barneby, stat. nov. C. turbinata Ben
tham, London J. Bot. 3: 109. 1844. — "[Brazil.]
Matto grosso, Manso, Lhotzky," the locality more
precisely stated by Bentham, 1876: 422: "in Serra
Santa provinciae Matto Grosso." — Holotypus, col
lected by Silva Manso and distributed by L'Hotsky,
K!; isotypus, ^B = F Neg. 7265. — Feuilleea
turbinata O. Kuntze, Revis. Gen. PI. 1: 189. 1891.
C. chapadae S. Moore, Trans. Linn. Soc. London 11, Bot. 4:
349. 1895. — "Hab. Serra de Chapada, circa 1000 ped.
supra mare . . . ([S. Moore] N. 160.)" — Holotypus, la
beled: "chapada plateau, frequent at about 1800 ft [540
m]," BM!. C. turbinata sensu Bentham, 1875: 552.
C. chapadae sensu Hoehne, Com. Lin. Telegr., Bot. 5(8): 21.
1919.
Shrubs 1-3 m, except for facially glabrous lfts
finely pubemlent or hirsutulous throughout, the stems
distally, the peduncles, and the individual fls white-
pilose-tomentose; stipule-blades (2-)3-9 x 1.2-2.5
m m ; lf-formula (vi-)vii-xi(-xii)/26-38, the lfts lin
ear-oblong, the longer ones 3.5-8 x 0.8-1.8 m m ,
3.5-5 times as long as wide; peduncles 1.5-5.5 cm;
pedicel of peripheral fls 2-5 m m ; calyx 6-8.5 m m ;
corolla 9-13 m m ; androecium 58-80-merous, 46-52
m m , the tassel white.
In campo cerrado, often in rocky places, ±550-830
m, localized on the Brazilian Planalto in s. Mato
Grosso and centr. Mato Grosso do Sul, there sympatric
with var. opulenta. — M a p 18. — Fl. Ill, V, VIII-X.
26. Calliandra gardneri Bentham, London J. Bot. 3:
108. 1844. — "Province of Goyaz, Brazil, Gardner,
n. 3703." — Holotypus, collected at Conceicao,
near 11°20'S, 47°W, in Feb 1840, K (hb. Hooker.)!
= N Y Neg. 1968; isotypi, K (hb. Benth.)!, OXF!. —
Feuilleea gardneri O. Kuntze, Revis. Gen. PI. 1:
187.1891.
C. gardneri sensu Bentham, 1875: 550; 1876: 421.
Microphylhdious, functionally herbaceous sub shrub
from obhque woody rootstock, the simple ascending
stems 2-3 d m bearing ±3-5 lvs and 1-2, proportion
ately large, axillary capitula, the whole plant glabrous
except for traces of puberulence on lf-axes and mi
nutely strigulose perianth, the firm plane lfts bicolored,
dark ohvaceous sublustrous above, pale dull green be
neath, the furthest (or only) peduncle often appearing
terminal and continuous with primary cauline axis
due to abrupt inhibition of terminal meristem beyond
the furthest If. Stipules lanceolate 2-5 x 0.7-2 m m ,
weakly several-nerved, becoming dry fragile, but only
very tardily deciduous. Lf-formula (ii-)iii-v/(14—)
18-29; lf-stks of longer lvs 5-9 cm, the petiole 8-28
m m , the longer interpinnal segments 13-24 m m , the
rachis dilated into a cupule under each pinna, the ven
tral groove continuous but constricted at insertion of
pinnae; lft-pulvinules 0.4-0.7 m m ; lfts subequilong
except at each end of pinna-rachis, the blade narrowly
oblong or lance-oblong from obtusangulate base, at
apex either rounded or depressed-deltate, those near
mid-rachis (9-)10-15 x 3.3-4.7 m m , 2.7-3.6 times as
long as wide; venation palmate-pinnate, the straight
midrib forwardly displaced to divide blade 1:1.8-3
and giving rise on its posterior side, near and beyond
mid-blade, to 3-6 secondary nerves ascending to
anastomosis well within the margin, the inner poste
rior primary nerve incurved-ascending ± to mid-
blade, the outer 1-2 much shorter, tertiary venulation
0 or faint, the whole venation prominulous on hy-
pophyllum, scarcely raised above. Peduncles solitary
or exceptionally geminate, erect stout 6-12 cm, 1-
bracteate below middle; capitula densely 5-11-fid,
the ascending fls homomorphic or almost so (the ter
minal fl sometimes stouter than the rest but neither
longer nor otherwise modified), the clavate receptacle
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 65
M A P 18. Distribution of Calliandra dysantha Bentham vars. turbinata (Bentham) Barneby, opulenta Barneby, and macrocephala (Bentham) Barneby.
2-4 m m diam; floral bracts 0 or reduced to vestigial
caducous scales; pedicels claviform or stoutly
turbinate 1-2.5 x 1.4—2 m m ; perianth pallid pink-
tipped, the calyx either pubemlent or glabrous ciho
late, faintly nerved, the corolla minutely strigulose;
calyx campanulate 3-5 x 1-\ m m , the broad obtuse
teeth 1.4—2.2 m m ; corolla 13-17 m m , the broadly
ovate lobes 4.5-7.5 m m ; androecium 46-92-merous,
6-7.4 cm, the inwardly thickened stemonozone 3.5-4
m m , the tube to 12 m m , the tassel carmine proxi
mally, whitish distally; no intrastaminal disc; ovary
subsessile, at anthesis glabrous. Pods 1-2 per capitu
lum, in profile linear-oblanceolate 9.5-14 x 1.2-1.6
cm, the sutural ribs in dorsal view 3-4 m m wide, the
recessed valves castaneous sublustrous, transversely
venulose, minutely pubemlent overall; seeds in broad
profile obovate-elliptic 10.5-14 x 6-8 m m , the testa
light brown, smooth but dull, the pleurogram lacking.
In sandy and rocky places in campo cerrado,
600-1000 m, localized on the w. slope of Sa. Geral de
Goias, on right effluents of upper rio Tocantins, be
tween 11°S and 13°20'S, in e.-centr. state of Goias.
— M a p 16. — Fl. I-III.
Calliandra gardneri is notable for low stature, few
cauline leaves, a pseudoterminal capitulum (not truly
terminal, as in C. brevicaulis), large flowers, filament-
tassel white distally but pink proximally (the reverse of
the usual pattern) and seed-coat lacking pleurogram.
27. Calliandra macrocalyx Harms, Bot. Jahrb. Syst.
42: 203. 1908. — Holotypus infra sub var. macro-
calyce indicatur.
Subarborescent shrubs 1-5 m with terete, almost
straight branches, pilose nearly throughout with as
cending and subappressed gray hairs to 0.6-1.5 m m
but the annotinous and older stems glabrescent, the
66 M E M O I R S OF THE N E W Y O R K BOTANICAL G A R D E N [VOL. 74(3)
lvs bicolored, dull dark olivaceous and either smooth
glabrous, or microscopically red-brown-papillate, or
thinly pilosulous above, paler and either subap-
pressed-pilose or exceptionally glabrous beneath, the
stout peduncles and perianth of fls more densely pal
lid- or brown-pilose-tomentulose overall, the capitate
units of inflorescence arising singly either from the
axil of few furthest primary lvs, or terminally very
shortly pseudoracemose, or borne exceptionally at
nodes of axillary brachyblasts, the individual fls pro
portionally massive and thick-textured. Stipules firm,
becoming stiff-papery, brownish, finely striate, the
blades (linear-)lanceolate or triangular-ovate-acumi
nate, including lf-spur 4.5-10 x (1-)1.3-3.4 m m , per
sistent or tardily deciduous. Lf-formula ii-iv(-v)/15-
23(-25); lf-stk of longer lvs (0.7-)l^t cm, the petiole
including discolored pulvinus 3-16 m m , at middle
0.5-1 m m diam, the ventral groove bridged at inser
tion of pinnae, the longer interpinnal segments 3-9
m m ; pinnae scarcely graduated, the longer ones
(1.8-)2^(-4.5) cm, the longer interfoliolar segments
±1-1.5 m m ; lft-pulvinules 0.3-0.5 x 0.3-0.7 m m ; lfts
distally decrescent or subequilong, the blades nar
rowly oblong or rhombic-oblong from shallowly
semicordate or obliquely subtruncate base, deltately
acute, the longer ones 4—11.5 x 1—1.2(—3.7) m m ,
2.7-4.1 times as long as wide; midrib straight or gen
tly incurved near apex, simple or weakly 1-2-
branched above middle on posterior side, forwardly
displaced to divide blade ±1:1.5-2.5, 1-2 posterior
primary nerves sometimes perceptible, all venation
prominulous only dorsally, immersed or nearly so
ventrally. Peduncles 12-29 m m , bracteate above mid
dle, the bract deltate-ovate or broad-lanceolate 2-6 x
3-6 m m , resembling a stipule in texture and striation;
capitula 4—6-fld, the fls appearing subsessile, homo
morphic, the hemispherical or depressed-pyramidal
receptacle <2 m m ; bract of outermost fls resembling
peduncular bract but smaller and caducous, that of
terminal fl wanting; pedicel (often differentiated only
in vertical dissection of fl) turbinate 1.5-3 x 2-5 m m ;
perianth either 4- or 5-merous, densely brownish- or
pallid-silky-tomentulose overall; calyx at full anthe
sis 7.5-15 x 7-12 m m , the coriaceous walls nearly
1.5 m m thick in section, the teeth usually deltate or
depressed-ovate ±1.5^.2 m m , but in var. aucta lance-
acuminate 9-10 m m ; corolla (10-) 14-21 m m , the
lobes (disregarding indumentum) 3.5-9 m m ; androe
cium 5-9 cm, 64-185-merous, the coriaceous stem
onozone 3-8 m m , the tube 6-9 m m , the tassel at first
white, rubescent; no intrastaminal nectary; ovary ses
sile, at anthesis densely white-villous-tomentulose.
Pods ±8-ll(-16) x (0.75-)l-1.9 cm, the lignescent
valves densely tomentose; seeds (few seen) in broad
profile obtusely rhombic, to ±15 x 11 m m , the testa
light brown, smooth, pleurogrammic.
Systematically important features of C. macroca
lyx are bracteate peduncles, very large silky-tomen-
tose flowers, and coriaceous perianth and stemono
zone. The inflorescence-type and bracteate peduncles
suggest much closer affinity to sect. Androcallis than
to compatriot species of sect. Calliandra. A mature
fruiting specimen (Ganev 548, H U E F S ) from 1220 m
near Catoles in Chapada Diamantina has the massive,
densely velutinous perianth of C. macrocalyx com
bined with foliage suggesting C. bahiana; it could
perhaps be of hybrid origin.
Key to the varieties of C. macrocalyx
1. Calyx 7.5-13 mm, the deltate or depressed-
ovate teeth 1.5-4.2 mm, 4-8 m m shorter than corolla; range of the species. . . 26a. var. macrocalyx
1. Calyx ±16 mm, the lance-acuminate teeth
±10 mm, about as long as corolla; known
only from the type-locality, on rio S. Francisco
near 9°30'S 26b. var. aucta
27a. Calliandra macrocalyx Harms var. macro
calyx. C. macrocalyx Harms, 1908, I.e. sens. str. —
"Brasilien: Bahia, Campo der Serra do Sao Ignacio
[= n. end of Sa. Acurua near 12°10'S, 42°40'W]
(ULE n. 7586. — Dez. 1906)." — Holotypus, fB =
F Neg. 7249; isotypus, H B G ! = K Neg. 75755.
C. villosiflora Harms, Bot. Jahrb. Syst. 42: 205. 1908.
"Bahia: Catinga bei Remanso (ULE n. 7586. — Dez
1906)." — Holotypus, +B = F Neg. 7267; isotypus, HBG!. — Equated with C. macrocalyx by Renvoize 1981: 77.
C. macrocalyx sensu Renvoize, 1981: fig. 3(28); Lewis, 1987: 77.
Calyx as described in key to varieties; lfts pubes
cent dorsally; androecium 5-7 cm.
In caatinga scrub and about outcrops in cerrado,
near 750-1100 m, perhaps lower at its n. limit, scat
tered along right affluents of rio S. Francisco in inte
rior Bahia, from Remanso s. to Caietete thence n.
across river into Pemambuco (Petrolina) and s.-w.
Piaui (Pagehu), in lat. 9°40'-14°S. — M a p 19. — Fl.
XII, II-VI.
27b. Calliandra macrocalyx Harms var. aucta
Barneby, var. nov, a var. macrocalyce calycis den-
tibus lanceolato-caudatis ±10 m m (nee depresse
deltatis ovatisve ±1.5-4 m m ) longis, ulterius foholis
dorso glabris (nee pilosulis) androecioque longis-
simo 9 cm usque (nee 5-7 cm) longo diversa. —
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 67
CALLIANDRA MACROCALYX
T var MACROCALYX
* var. AUCTA
MAP 19. Distribution of Calliandra macrocalyx Harms var. macrocalyx and var. aucta Barneby in eastern Brazil.
BRAZIL. Bahia: Mun. Casa Nova, 9°31'S, 41°23'W,
8 Oct 1981 (fl), Lucia M. C. Gongalves 209 = Herb.
R A D A M B R A S I L 4825. — Holotypus, K.
Calyx as described in key to varieties; lfts dorsally
glabrous; androecium ±9 mm.
In sandy soil of estepe arborea aberta, at un
recorded elevation, known only from the type-locality
on the left bank of rio Sao Francisco at 41°23/W in n.-
e. Bahia. — Map 19.
This remarkable plant differs substantially from var.
macrocalyx only in the modified calyx-teeth, which
are prolonged to equal the corolla. The dorsally gla
brous leaflets and longer androecium mentioned in the
Latin diagnosis are perhaps only individual aberra
tions. The collector of the type described var. aucta as
a subshrub, whereas other material of C. macrocalyx
is definitely fruticose or arborescent.
28. Calliandra fernandesii Barneby, sp. nov., habitu,
foliis aliisque notulis C. ulei revocans, sed ab ea
foliorum primariorum axi principali 3-7 (nee 1-2)
cm longo, flosculis obscure pedicellatis (pedicello
vix 1, nee 8-12 m m usque longo) ante anthesin
argenteo-sericeis (nee subglabris), calycis dentibus
deltatis tubo brevioribus (nee lanceolatis fere basin
usque liberis), androecioque rubro ±2.5 cm (nee
albido fere 5 cm) usque longo diversissima. —
BRAZIL. Piaui. entre Altos e Campo Maior, ±5°S,
42°20'W, 29 Jul 1979 (fl, fr), A. Fernandes & P
Martins 6839. — Holotypus, EAC; isotypus, NY.
— Piaui: Sete Cidades-Piracuruca, 16 Oct 1977 &
5 Jul 1991 (fl, fr), A. Fernandes 3502, 17884,
paratypi, NY. FlG. 4
Slender microphyllidious shrubs of unknown po
tential stature with terete defoliate annotinous and
older stem, the young stems, lf-axes, and peduncles
gray-pilosulous with spreading or incurved-ascend
ing grayish hairs to ±0.5-0.7 mm, the lvs bicolored,
the firm, contiguous or imbricate, facially glabrous but
finely cihate or granular-ciliolate lfts lustrous dark
brown above, paler dull beneath, the capitula arising
singly from efohate axils of either short and thatched
or ± extended brachyblasts; phyllotaxy distichous. Stip
ules firm, lanceolate or narrowly triangular-acuminate
±1.4—5 x 1-1.5 mm, dorsally not or indistinctly venu
lose, pubemlent but glabrate in age, persistent. Lf-
formula iv-viii/32-40; lf-stk of longer lvs 3-7 cm, the
petiole including pulvinus 5-11 mm, at middle 0.7-1
m m diam, the longer interpinnal segments 5—10(—12)
mm, the shallow sulcus bridged; pinnae a little, some
times randomly, accrescent distally, the rachis of fur
ther ones 3-5 cm, the longer interfoholar segments
0.45-1 mm; lft-pulvinules ±0.2 x 0.3 mm, scarcely
wrinkled; lfts subequilong or gradually decrescent
from near mid-rachis upward, the blades linear or
linear-oblong from obtusangulate base, obtuse or
deltately subacute, the longer ones 3.6-5 x 0.9-14
mm, (3.2-)3.5-4.8 times as long as wide; midrib
straight, finely prominulous only dorsally, displaced to
divide blade ±1:2, weakly 2-3-branched distally, the
posterior primary nerve very short or imperceptible.
Peduncles X.5-4 cm, bracteate above middle, the bract
lance-elliptic 1.5-2 mm; capitula incipiently recemi-
form ±15-20-fld, the ellipsoid-claviform pitted recep
tacle 2-3 x 1.5-2 mm; floral bracts minute fugacious;
pedicels 0.3-1 x 0.5-0.9 mm, perianth submembra-
nous 5-merous, the calyx thinly pubemlent, brownish,
faintly 5-nerved (not striate), the corolla densely sub-
appressed-silky, externally nerveless; calyx campanu
late ±2 x 2.5 mm, the deltate teeth 0.5-0.7 mm; corolla
6.5-7.5 mm, the erect ovate-lanceolate lobes 2.2-3.8
mm; androecium (one exactly observed) 28-merous,
2.5 cm, the stemonozone 1.3-1.5 mm, the tube 2.6-3.7
mm, the tassel red. Pods 3.5-9.5 cm, in broad profile
6-8 m m wide, minutely erect-pilosulous overall, the
ribs in dorsal view 1.3-2 m m wide, the recessed valves
openly trans verse-venulose; seeds not seen.
In campo cerrado at elevations not recorded, known
from two stations in hill country between 4°S and 5°S
in n. Piaui and from Chapada da Ibiapaba in adj. n.-w.
Ceara, Brazil. — Map 20. — Fl. VI-X, or following
rains.
Calliandra fernandesii resembles C. ulei in habit
and leaf-formula, but differs in longer leaf-stalks, ob
scurely pedicellate flowers, silvery-silky perianth,
68 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
FIG. 4. Calliandra fernandesii Barneby.
deltate calyx-teeth, and relatively short perianth-tassel
red throughout. This distinctive species is named in
honor of Professor Afranio Gomes Fernandes of Uni-
versidade Federal do Ceara, Fortaleza, to w h o m The
N e w York Botanical Garden is indebted for gifts of rare
and critical Leguminosae from northeastern Brazil.
29. Calliandra ulei Harms in Engler, Bot. Jahrb.
Syst. 42; 205. 1908. — "Piauhy: Catinga der Serra
Branca [8°10'S, 41°15'W] (ULE n. 7440 — Jan.
1907)." — Holotypus, fB = F. Neg. 7266!; isotypi
H B G ! , K ! = I P A N e g . 7476.
Microphyll shrubs reportedly (protologue) 1-3 m
tall with virgate long-shoots and crowded foliate
brachyblasts, the primary internodes ±5-14 m m , the
new stems, lf-axes and peduncles pilosulous with fine
straight subvertical hairs to ±0.6 m m mixed, in inflo
rescence, with few minute capitellate trichomes, the
foliage bicolored, the small imbricate lfts dark lustrous
green (when dry brown) above, pallid dull beneath,
cihate with straight ascending or distally hooked hairs,
the stout peduncles arising singly from primary lf-
axils toward top of long-shoots, the capitula umbelli
form; phyllotaxy distichous. Stipules narrowly lanceo
late, those of primary lvs, including lf-spur, 5-8 x 1-
1.5 m m , those of brachyblasts imbricate and some
what shorter, all weakly nerved, pubescent dorsally,
marcescent. Lf-formula iii—v(—vi)/24—31, the pinnae of
most primary lvs 3-5-, of brachyblast lvs 2-3-jug.; lf-
stk of primary lvs ±1-2 cm, the petiole 1.5̂ 4- m m , the
interpinnal segments 2-4 m m , the ventral groove nar
row, bridged at insertion of pinnae; pinnae a little
accrescent distally, the axis of furthest ones 22-27 m m ,
the interfoholar segments 0.3-0.6 m m ; lft-pulvinules
0.15-0.2 m m ; lfts equilong except at very ends of
rachis, the blades oblong or lance-elliptic from ob
tusely auriculate base, obtuse, the longer ones 3-4.5 x
1-1.3 m m , 3-3.4 times as long as wide; midrib either
subcentric or forwardly displaced to divide blade 1:2,
simple or faintly 2-4-branched on each side, the pos
terior primary nerve not or scarcely perceptible exter
nally. Peduncles 1-3.3 cm, 1-bracteate near middle,
the broad-lanceolate bract 2-2.5 m m , the clavate axis
of capitulum 2-3 m m ; floral bracts 0.4 m m or less, ca
ducous; umbels 15-22-fld, the fls heteromorphic, the
peripheral ones slenderly pedicellate, the 2-4 furthest
ones shorter and stouter, shortly pedicellate, all gla
brous except for minutely puberulent tip of calyx- and
corolla-lobes; P E R I P H E R A L FLS: pedicel 8-12 x
0.3-0.35 m m , turbinately dilated at base of fl; perianth
5-merous; calyx ±1.2 m m , cleft to base into narrowly
lanceolate lobes; corolla whitish or perhaps reddish-
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 69
** CALLIANDRA FERNANDESII
•* C. ULEI
A C. PILGERANA
¥ C. UMBELLIFERA
* C. IMPERIALIS
Map 20. Distribution of Calliandra fernandesii Barneby, C. ulei Harms, C. pilgerana Harms, C. umbellifera Bentham, and C. imperialis Barneby in eastern Brazil.
tinged, ±7 m m , the tube much dilated at the limb, the
lobes ±2.6 x 2 m m ; androecium white (Ule), to 53
mm, 22-merous, the thickened stemonozone 2.5 mm,
the tube ±4.5 m m , some filaments irregularly fascicu
late beyond orifice; ovary glabrous; TERMINAL FLS:
pedicel ±2 x 0.8; calyx as in peripheral fls but teeth to
3 mm; corolla ±9 mm, the stoutly cylindric stemono
zone ±3.5 m m , in section 0.4 m m thick, the broadly
ovate lobes ± 4 x 2 mm; androecium to 66 mm, 26-
merous, the tube 5 m m ; no ovary and no intrastaminal
disc. Pod unknown.
In caatinga, at unrecorded elevation, known only
from the type-locality near 8°S in s.-e. Piaui, Brazil.
— Map 20. — Fl. I—II.
30. Calliandra pilgerana Harms, Bot. Jahrb. Syst.
42: 204 ("Pilgeriana"). 1908. — "Bahia: Serra do
Sao Ignacio [= Sa. Acurua] (ULE n. 7530. — Febr.
1907)." — Holotypus, +B = F Neg. 12531; clastoty
pus (fragm), F!; isotypus, HBG! = K Neg. 75752.
C. pilgerana sensu Renvoize, 1981: 67, fig. 1(2); Lewis, 1987: 175 ("pilgeriana").
Microphyllidious arborescent shrubs attaining 1.5-
4 m, with stout terete glabrate annotinous long-shoots
and rather loosely thatched short-shoots, the new
stems, young stipules, all lf-axes, and peduncles silky-
villosulous with extremely fine, ascending and sub
appressed, gray hairs 0.1-0.5 m m mixed with a few,
sometimes colored granular trichomes, the lvs strongly
bicolored, the firm plane imbricate lfts fuscous brown
and lustrous (dry) on upper face, beneath palhd dull
and strigulose with palhd hairs arising from a fuscous
papilla, randomly ciholate, the short racemes of "pur
ple" fls arising singly from efoliate brachyblasts; phyl
lotaxy distichous. Stipules broadly lanceolate or lance-
attenuate, including lf-spur 4—7 mm, several-nerved
but not sharply so, becoming stiffly papery, gray
glabrous, persistent. Lf-formula iii—vii/28-̂ J4(—52) the
pinnae of brachyblasts not more than 3 pairs; lf-stks of
primary (early deciduous) lvs ±2.5-4.5 cm, the petiole
including pulvinus and the longer interpinnal segments
±4—8.5 mm, the shallow ventral groove bridged at in
sertion of pinnae; pinnae a httle accrescent distally, the
rachis of longer ones (of both long- and short-shoots)
±3-4 cm, the longer interfoholar segments 0.6-1 mm;
lft-pulvinules 0.2-0.3 mm, nearly as wide; lfts decres
cent at each end of rachis, the blades narrowly oblong
from obtusely auriculate base, obtuse, straight or al
most so, the larger ones 3^-4 x 0.85-1.2 mm, 3.3^-
times as long as wide; externally perceptible venation
simple, the midrib only a httle forwardly displaced
from mid-blade, unbranched, shallowly depressed on
upper face of 1ft, fihform dorsally. Peduncles 3-4.5 cm,
bracteate near middle, the lanceolate bract 2-2.5 mm;
racemes closely 18-25-fld, the receptacular axis nar
rowly clavate 6-8 mm; floral bracts obsolete; pedicels
2-4.5 mm, slightly shorter distally, at middle ±0.4—
0.65 m m diam, thickened and discolored at base; peri
anth 5-merous, carnosulous only at the stemonozone,
the calyx glabrescent, the corolla finely minutely
strigulose overall with lutescent hairs; calyx shallowly
campanulate, weakly 15-nerved, 1.8-2.2 mm, the del
tate teeth ±0.5 m m (but one sinus often deeper);
corolla tubular-infundibuhform 10.5-11 mm, the erect,
lance-ovate lobes ±3-4 mm, below middle sometimes
densely white-fimbriolate; androecium 12-merous,
±44 mm, the thickened stemonozone 3.5 mm, the
tube 6.5 mm; ovary subsessile, villosulous around
the top, elsewhere glabrous; no intrastaminal disc. Pod
unknown.
In rocky places above 500 m, in campo mpestre,
known only from Sa. Acurua, near 11°30'S, 42°30'W,
70 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
in n.-centr. Bahia, Brazil. — Map 20. — Fl. II—III,
IX-X, the full season probably greater.
Calliandra pilgerana has the relatively many flow
ers on shortly elongated floral receptacle, the bracteate
peduncles, and the lutescent perianth vesture of C.
bella and C. subspicata, but differs from these in the
well-developed pedicels and remarkably few stamens.
31. Calliandra umbellifera Bentham, J. Bot.
(Hooker) 2: 141. 1840. — "Ceara, Brazil, Gardner,
n. 1581," the locality more precisely stated in
Hooker, London J. Bot. 3: 102. 1844. — "Dry hills
near Crato in Ceara." — Holotypus, K (hb. Hook.)!
= IPA Neg. 7466 = N Y Neg. 7956; isotypi, GH!,
OXF!. — Feuilleea cearana O. Kuntze, Revis. Gen.
PI. 1: 185. 1891.
C. umbellifera sensu Bentham, 1875: 544; 1876: 413;
Ducke, 1953: 427.
Drought-deciduous microphyll shrubs 1.3 m and
probably taller with stiff, simply virgate or few-
branched long-shoots but no brachyblasts or resting-
buds, the new stems at once minutely pubemlent,
thinly pilose with straight white hairs to 0.4—1.1 m m ,
and, with the inflorescence, in addition minutely cap
itate-glandular and resinous, the bicolored lfts ciliate,
glabrous facially but dorsally micropapillate, the um
bellate units of inflorescence borne singly in the axil
of distal lvs of current year; phyllotaxy distichous.
Stipules erect, lanceolate or narrowly ovate 3-9 x
0.7-1.4 m m , the stiffly papery blade 4—6-nerved, per
sistent. Lf-formula i-ii(-iii)/13-17; lf-stks 2-16 m m ,
the one (or longer) interpinnal segment about as long
as petiole proper, the ventral groove narrow or ob
scure; rachis of longer pinnae 1.6-4.2 cm, the longer
interfoliolar segments 1.2-2.3 m m ; lft-pulvinules
0.25-0.35 m m ; lfts subequilong except at very ends
of rachis, the blade oblong from obtusely auriculate
base, obtuse or obscurely apiculate, the longer ones
6-8.5 x 2.2-3 m m , 2.5-3 times as long as wide; vena
tion palmate-pinnate, the straight midrib only slightly
eccentric, giving rise on each side to 3-5 divaricate
secondary nerves brochidodrome well within the plane
margin, the inner of 2 primary posterior nerves pro
duced no further than xh length of blade, tertiary
venulation imperceptible, the whole venation im
mersed or almost so above, finely prominulous be
neath. Peduncles stout 0.9-3 cm, bracteolate below
middle, the bracteole resembling a stipule but smaller,
the hemispherical or narrowly clavate receptacle
0.15-0.25 m m ; bracts of lowest firs linear-lanceolate
0.6-1 m m , caducous, the upper fls ebracteate; umbels
10-17-fld, the fls heteromorphic, the peripheral ones
slenderly long-pedicellate, the 1-3 innermost ones
shortly stoutly pedicellate and the perianth stouter
and longer (these occasionally abortive); PERIPH
E R A L FLS: pedicel 10-17 x 0.25-0.4 m m ; perianth
stipitate-glandular overall, sometimes in addition
minutely pubemlent, the corolla prior to anthesis
plumply pyriform, the venation fine, subimmersed;
calyx campanulate or campanulate-patelliform 1.6-
2.4 x 1.5-2 m m , the subulate obtuse teeth 0.7-1.1
m m ; corolla 8.5-9.5 m m , the broadly ovate lobes
2.5-3.5 m m ; androecium 26-38-merous, ±5 cm, the
thickened, externally ribbed stemonozone 2-3 m m ,
the tube 4—5 m m , the filaments whitish; intrastaminal
disc 0; ovary at early anthesis glabrous, subsessile,
becoming densely glandular after fertilization; C E N
T R A L FLS (not well known): calyx almost of the pe
ripheral fls, sometimes slightly longer; corolla
broadly rounded at base, 9-11 x 2-6 m m ; ovary (al
ways?) 0. Pod unknown.
In unrecorded habitats, to be expected in caatinga
thickets, apparently local, known only from s. Ceara
and s.-w. Piaui, Brazil. — M a p 20. — Fl. VII-VIII(-?).
A m o n g the members of sect. Androcallis distin
guished by umbelliform capitula that are known from
eastern Brazil north of Bahia, C. umbellifera is emi
nently singular in the stipitate-glandular perianth.
Lacking flowers, the species differs from the probably
related C. ulei in somewhat fewer (13-17, not 24—31)
pairs of leaflets on the longer pinnae, and in leaflets
nearly twice as large (to 8-8.5, not 3^.5, m m long),
dorsally micropapillate, and distinctly pinnate-veined.
32. Calliandra imperialis Bameby, sp. nov, habitu
toto et imprimis umbellae flosculis periphericis
longe pedicellatis C. umbelliferam et C. ulei simu-
lans, ab amabus calycis dentibus linearibus tubo suo
subquadruplo longioribus, ulterius a priori flosculis
parce puberulis eglandulosis, ab altera bracteis
florahbus 4 m m usque longis (nee minimis) necnon
florum periphericorum androecio 10-12 (nee ±22)-
mero diversa. — BRAZIL. Piaui: pr. Pedro Se-
gundo, near 4°25'S, 41°25'W, anno 1935, S. E.
Dahlgren 875. — Holotypus, F.
Drought-deciduous microphyll shrubs of unknown
stature, pilosulous with fine white hairs to ±0.4-0.5
m m , the young stems and lf-axes densely, the lfts and
fls thinly so, eglandular, the umbels of heteromorphic
fls solitary, pedunculate in the efoliate axils of short
brachyblasts; phyllotaxy distichous. Stipules of pri-
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 71
mary lvs lanceolate ±4-6 x 1-1.5 mm, firm, weakly
3-6-nerved, persistent, those of brachyblasts similar
but somewhat shorter. Lf-formula iii-iv/20-23; lf-stks
4—11 m m , the longer interpinnal segments 1-2.5 mm;
pinnae subequilong, the rachis of longer ones 10-18
m m , the longer interfoliolar segments 0.3-0.5 mm;
lfts subsessile, imbricate, the blades oblong from
shallowly semicordate base, obtuse, the longer ones
1.8-2.7 x 0.7-1 m m , 2.7-2.8 times as long as wide;
venation immersed. Peduncles 7-14 mm, bracteate
above middle; umbels 12-20-fld, the subglobose re
ceptacle 1.5-2 m m diam; bracts linear-lanceolate ±4
mm, caducous; pedicels graduated, those of outer
most bisexual fls to 7-9 x 0.3-0.4 mm, those of 3-4
subterminal staminate ones ±1.5-2 x 0.9 mm; peri
anth of all fls 5-merous, thinly minutely pubemlent
eglandular; PERIPHERAL FLS: calyx ±5.5 mm, the
campanulate tube ±1 mm, bluntly 5-angulate, the lin
ear teeth ±4 x 0.2-0.4 mm; corolla 6.5 mm, the ovate
lobes 2.8 m m ; androecium 10-12-merous, ±4.5 cm,
the stemonozone thickened internally ±1.4 mm, the
tube 3.5 m m , disc 0; ovary glabrous, grooved later
ally; SUBTERMINAL FLS: calyx ±7 mm, the tube
1.8 m m , the linear teeth 5.2 mm; androecium 20-mer-
ous, the tube 3.5 m m , the tassel of unknown color
(probably white brunnescent); ovary 0. Pod unknown.
In unrecorded habitat, but to be expected in caatinga,
known only from slopes of Sa. do Pedro Segundo in
n.-w. Piaui, Brazil. — Map 20. — Fl. probably after
rains.
Calliandra imperialis resembles C. umbellifera
and C. ulei in general habit and particularly in long-
pedicellate peripheral flowers of the capitulum, but
differs from both in linear calyx-teeth about four
times as long as the tube. Its flowers lack the gland-
tipped trichomes peculiar to C. umbellifera, and its
androecium has only about half as many filaments
(10-12, not ±22) as that of C. ulei.
33. Calliandra concinna Barneby, sp. nov. affinitatis
intra sect. Androcallin incertae, C. parvifoliae sub-
similis sed notulis sequentibus diversa: pinnae 3-5
(nee 5-18)-jugis, a rachi communi subhorizontal-
iter patulae; pinnarum longiorum axis 2.5-5 (nee
1.2-2.2) cm longa; capitulorum manifeste umbelli-
formium flosculi homo-, nee saepissime hetero-
morphi, peripherici pedicello 4 m m usque elevati.
— BRAZIL. Minas Gerais: 3.5 km by road s.-w. of
Rio Jequiti and Mendanha, 14 Apr 1973 (fl), W. R.
Anderson 8902. — Holotypus, UB; isotypi,
CEPEC, F, K, M B M , MICH, N Y (2 sheets), RB,
US. FIG. 5 FIG. 5. Calliandra concinna Barneby.
72 M E M O I R S OF T H E N E W Y O R K BOTANICAL G A R D E N [VOL. 74(3)
Slender but stiffly branching, microphyllidious
shrubs 1-2 m, appearing glabrous throughout but
microscopically pubemlent on some new growth and
some lfts microscopically ciholate, the burnished red-
brown homotinous branchlets becoming fuscous
lamellate, the narrow multifoliolate pinnae stiffly
widely divaricate from lf-stks, the closely imbricate,
thick-textured lfts lustrous dark green above, paler
dull beneath, the proportionately large, umbelliform
capitula borne singly or rarely geminate on lateral
brachyblasts, subtended either by a developed If or by
efoliate stipules. Stipules firm, subulate from triangu
lar base, mostly 1.5-3.5 m m , 3-11-nerved, persistent
but the tips often blanched and fragile in age. Lf-for
mula iii-v/42-64; lf-stks 1.5-̂ 4.5 cm, the petiole 5-14
m m , the longer interpinnal segments 5-11 m m , the
shallow ventral groove interrupted at insertion of pin
nae by dilated, shallow-cupulate sockets; pinnae ran
domly graduated or sub-equilong, bulbously dilated
at base, the longer ones 2.5-4.5(-5.5) cm, the longer
interfoliolar segments 0.35-0.6 m m ; lfts subequilong,
subsessile, the pulvinule ±0.15 m m , the blades linear
or linear-oblong from bluntly auriculate base, obtuse,
the longer ones 3-5 x 0.5-0.7 m m , 4—7 times as long
as wide; midrib subcentric, simple or faintly pinnate,
a posterior primary vein sometimes barely percepti
ble dorsally. Peduncles very slender 20-38 x 0.4-0.5
m m , ebracteate; capitula 11-20-fld, the clavate recep
tacle 1.5-2.5 m m ; floral bracts membranous oblance
olate 1 m m or less, absent from some distal fls;
pedicels 1.5-4 m m ; fls homomorphic, the reddish pe
rianth glabrous except for sometimes microscopically
ciholate calyx-teeth; calyx campanulate, bluntly 5-
angulate 5-nerved, 1.8-2.3 m m , the broadly ovate
teeth 0.4-0.9 m m ; corolla 5-6.5 m m , the lobes 1.9-
2.3 m m , recurved at tip; androecium 28-44-merous,
18-28 m m , the filaments uniformly red, the stemono
zone 1-1.3 m m , the tube 3.2^-.5 m m , scarcely thick
ened internally at base; ovary substipitate, glabrous.
Pod (one seen) in profile 6.5 x 0.7 cm, 7-seeded, the
sutural ribs in dorsal view ±1.2 m m wide, the re
cessed valves lustrous castaneous, finely transverse-
venulose, pilosulous overall (but more densely over
seeds) with straight white hairs ±0.5-1 m m ; ripe
seeds not seen.
O n quartzite outcrops, 800-880 m, along rio
Jequiti e. and n.-e. of Diamantina in centr. Minas
Gerais, Brazil. — Fl. III-V, VIII-IX.
34. Calliandra squarrosa Bentham, London J. Bot.
3: 104. 1844. — "Acacia squarrosa Mart, in Herb.
Hook. . . . Brazil, Campo Serral, Martius," the data
emended by Bentham (1876: 418) to: "in C a m p o
Geral provinciae Bahiensis, Luschnath." — Holo
typus, K (hb. Hook.)!; isotypus, B R (hb. Mart.)!.
The isotypus is dated March 1817, 14 years prior to
Luschnath's arrival in Brazil, and was probably col
lected by Prince Maximilian. According to Urban's
itinerary (1906: 144), Maximilian passed through a
Campos Gerais, northwest of Vitoria da Conquista,
in the first quarter of 1817, but the term is perhaps
topographic rather than toponymic. In any case, the
type of C. squarrosa must have been collected on
Maximilian's journey across interior Bahia some
where between Vitoria and Salvador. — Feuilleea
squarrosa O. Kuntze, Revis. Gen. PI. 1: 189. 1891.
C. catingae Harms in Engler, Bot. Jahrb. Syst. 42: 202.
1908. — "Bahia: Catinga bei Remanso (ULE n. 7573 —
Jan. 1907)." — Holotypus, +B = F Neg. 12341; isotypus,
HBG! = K Neg. 18753. — Equated with C. squarrosa by
Renvoize, 1981:73. C. squarrosa sensu Bentham, 1875: 549; 1876: 418, quoad
typum, planta Martiusiana exclusa; Renvoize, 1981: 73,
fig. 2; Lewis, 1987: 176.
Microphyll shrubs 3-20 dm with virgate long-
shoots and crowded acaulous brachyblasts, the older
stems terete glabrate, the new branches, lf-axes, and
peduncles thinly pilosulous with fine silky hairs to
±0.4—0.6 m m , the lvs subconcolorous but the small
plane imbricate lfts a little darker and more lustrous
above, facially glabrous but microscopically scabrous-
ciliolate, the capitula solitary in axils of first 1-2 lvs
of brachyblasts subtended by a coeval If; phyllotaxy
distichous. Stipules erect, narrowly lance-attenuate,
those of primary lvs 2-5.5 m m , 5-7-nerved, those of
secondary lvs scarcely shorter, all in age papery, pal
lid, persistent. Lf-formula i-iii/24—34, the pinnae of
most lvs or of some brachyblast-lvs only 1 pair; lf-stk
of primary lvs 1-6 m m , the petiole (disregarding lf-
spur) ±1-1.5 m m , the one or the longer interpinnal
segments at most 2 m m , the ventral groove bridged;
rachis of longer pinnae 14-20 m m , the longer inter
foliolar segments 0.3-0.6 m m ; lft-pulvinules 0.1-
0.15 m m ; lfts gradually decrescent toward each end
of rachis, the straight blades linear from auriculate
base, obtuse, marginally thickened dorsally, the larger
ones 3.5-5 x 0.6-0.9 m m , 5.5-7 times as long as
wide; midrib subcentric simple, prominulous only
dorsally, a very short obscure posterior nerve some
times barely perceptible. Peduncles very slender 0.8-
2 cm, usually 1-bracteate distally, the bract like those
of individual fls, narrowly ovate, at most 1.5 m m , and
like them persistent; capitula (4-)6-10-fld, the fls
heteromorphic, the perianth glabrous, striately nerved;
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 73
PERIPHERAL FLS: pedicel 0.3-0.5 x 0.6 mm; calyx
5-merous, turbinate-campanulate 2-3.8 mm, the trian
gular acute teeth 0.35-1.5 mm; corolla 4-merous
5.5-8 m m , the broadly ovate lobes 1.5-2 mm, each 7-
nerved; androecium ±17-merous, nearly 4 cm, the ste
monozone 1 m m , the tube exserted 1-2 m m from
corolla and dilated at orifice, the tassel red distally;
ovary not seen; C E N T R A L FL (sometimes lacking):
calyx scarcely longer but broader than that of periph
eral fls; corolla 8.5-9.5 mm; tube of androecium
16-20 mm, trumpet-shaped distally. Pods (two seen)
in profile 5-7 x ±0.9 cm, densely gray-pilosulous
overall, the sutural ribs ±4 m m wide in dorsal view, the
recessed valves transversely venulose; seeds not seen.
In unreported habitats, but to be expected in
caatinga at moderate elevations, known precisely
only from the lower Rio S. Francisco in n. Bahia (Re
manso., Cachoeira Paulo Afonso, near 9°20'-30'S,
38°-42°W), to be sought in adj. Piaui and Pernam-
buco. — Fl. I—III.
This is a poorly known species, perhaps related
to C. brevipes. The androecium of all flowers in the-
capitulum is dilated distally, but the tube of central
flowers is further exserted, about twice the length of
the corolla.
The foregoing description does not include C.
squarrosa var. crassifolia Benth., referred herein to
C. nebulosa.
35. CalHandra glaziovii Taubert, Flora 71: 75. 1892.
— "Habitat in Brasilia austro-orientali loco non
indicato [supplied by Glaziou, 1905: 188: 'Serra do
Campanema [sic, properly Capanema, near 20°10'S,
43°40'W, MINAS.'], Glaziou n. 23640." — Holoty
pus, collected 18 Jun 1882 (fl), fB = F Neg. 12431;
isotypi, K! = N Y Neg. 7956!, P (2 sheets)!.
C. bracteosa sensu Glaziou. 1905: 188 (Glaziou 13787, P!); non Bentham, 1846.
Shrubs of unknown stature with stout lignescent
long-shoots and in each primary lf-axil a short-shoot
thatched in dry striate stipules, the young stems, lf-
axes, and peduncles pilose-pilosulous with fine white
hairs to 0.4-1.1 m m , the lvs strongly bicolored, the
firm plane lfts glossy dark-olivaceous and glabrous
above, paler and appressed-pilose beneath, the dense
many-fid capituliform racemes arising singly from a
succession of brachyblasts, each subtended by a pair
of efoliate stipules; phyllotaxy distichous. Stipules of
primary lvs (excluding prominent lf-spur) lanceolate
or triangular-acuminate 6-12 x 1.7-3.5(-4) mm,
early dry papery, striately nerved, long persistent,
those of brachyblasts similar but a trifle narrower. Lf-
formula (i-)ii(-iii)/27-37; lf-stk of primary lvs 13-33
mm, the petiole 8-18 mm, at middle 0.9-1.5 m m
diam, the one interpinnal segment (or the longer of 2)
8-15 mm, the ventral groove bridged at insertion of
pinnae; rachis of longer pinnae 5.5-7.5 cm, the longer
interfoliolar segments 1.2-2.1 mm; lft-pulvinules
0.2-0.25 x 0.5-0.6 mm; lfts gently decrescent along
upper lA of rachis and close to its base, otherwise sub
equilong, the blades hnear from slightly dilated obtu-
sangulate base, either obtuse or apiculate, straight or
nearly so, those near mid-rachis 9-12 x 1.8-2.6 m m ,
4.3-5 times as long as wide; midrib forwardly dis
placed to divide blade 1:1.5-2, weakly pinnately
branched, the 2 posterior primary nerves very short.
Peduncles stout 8-25 mm, 1-2-bracteate near or
above middle, the bract(s) lanceolate striate 2.5-4.5
mm, persistent; capitula ±20-40-fld, prior to anthesis
globose, disregarding androecium becoming plumply
ellipsoid, the calyces contiguous, the lowest fls sub
tended by a bract similar to that on peduncle, the rest
ebracteate, all homomorphic; pedicels stout 0.9-1.5 x
0.9-1.2 mm; perianth 5-merous, the calyx striately
±20-nerved, minutely pubemlent, the corolla exter
nally nerveless, thinly appressed-silky, the tip of lobes
barbellate; calyx deeply campanulate 4-4.5 x 2.4—3.2
mm, the broadly obtuse, apically convex teeth 0.8-
1.1 mm; corolla 8.5-12.5 mm, the ovate lobes 2.5-4.2
x 1.7-3 mm; androecium 22-36-merous, (28-)32-42
mm, the stemonozone 2.4—3 mm, internally thick
ened, the tube 5-7.5 mm, the tassel red-purple; intra
staminal nectary 0; ovary sessile, at anthesis glabrous.
Pod unknown.
In habitats as yet unknown, to be expected in scrub-
woodland near 500 m, collected twice by Glaziou
in 1882 in s.-e. Minas Gerais (near Ouro Preto and
Mar d'Hespanha), not since seen. — Fl. I, VI, all
capitula and all fls of each capitulum simultaneously
expanding.
In the protologue Taubert compared C. glaziovii
with C. abbreviata Bentham, but it appears more
closely related to C. subspicata Bentham, which is
similar in the thatched brachyblasts, bracteate pedun
cles, and many-flowered capituliform racemes. The
distantly allopatric C. subspicata differs, however, in
dorsally glabrous, not appressed-silky, leaflets and in
white, not red-purple, androecium.
36. Calliandra depauperata Bentham, Trans. Linn.
Soc. London 30: 546. 1875; & in Martius, Fl. Bras.
15(2): 415. 1876. — "... in provincia Bahia:
Blanchet n. 3900 (specimina pauca vidi in Herb.
74 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Martii)." — Holotypus, BR n.v; presumed isotypus,
Blanchet 148, BM!. — Feuilleea depauperata O.
Kuntze, Revis. Gen. PI. 1: 187. 1891.
Fig. 6
C. depauperata sensu Renvoize, 1981: 71, fig. 2(12); Lewis, 1987: 172.
Small or diminutive, stiffly repeatedly branched,
microphyllous shrubs mostly 2-9 dm, occasionally
attaining 2 m in thickets, with smooth blanched older
stems, the widely divergent or horizontal short-shoots
thom-tipped at maturity, the young stems and back of
lf-axes thinly pilosulous with fine white hairs or sub
glabrous, the minute crowded lfts facially glabrous,
cili(ol)ate or not, the few-fid capitula arising singly or
geminate from coeval lf-axils toward apex of lateral
branchlets or from stemless brachyblasts; resting-buds
not present at anthesis, sometimes developed later;
phyllotaxy distichous. Stipules erect, narrowly lanceo
late 0.8-3 m m , pallidly (1-)3-5-nerved dorsally,
becoming dry brittle, deciduous. Lf-formula either i or
i-iii/8-16(-17), the pinnae of most lvs, in some plants
of all lvs, conjugate, the petiole deflexed from pulvi
nus; lf-stks (including small pulvinus) 1-4 m m , the
petiole ±1-1.5 m m , at middle 0.2-0.3 m m diam, the
interpinnal segment(s), when present, 1-2.3 m m ;
rachis of longer pinnae 3-8(-10) m m , the longer
interfoliolar segments 0.15-0.7 m m ; lft-pulvinules
minute, not over 0.1 m m ; lfts ± decrescent proximally,
the blades narrowly oblong-elliptic or hnear-oblong
from shortly auriculate base, obtuse, the longer ones
1.5-3.6 x 0.45-0.9 m m , (3-)3.4-4.4 times as long as
wide; venation of upper face of lfts imperceptible, the
very slender, simple, subcentric midrib faintly
prominulous beneath. Peduncles mostly 1-3 m m ,
sometimes obsolescent, rarely attaining 14(—16) m m ,
when well developed obviously 1-bracteate, the re
ceptacle <1 m m ; bracts triangular-lanceolate 0.5-1.7
m m , incurved, persistent; capitula (l-)2^-(-6)-fld,
the fls sessile, homomorphic; perianth greenish or
dark reddish, glabrous or remotely pubemlent distally,
both calyx and corolla pallidly 5-ribbed, obtusangu-
late, the androecium crimson; calyx deeply campanu
late 1.7-2.1 x 0.7-0.8 m m , the teeth 0.4-0.65 m m ;
corolla tubular 3-4.2 m m , 0.75-0.9 m m diam, the
nearly erect, subulate lobes 0.5-0.6 m m ; androecium
8-15-merous, 8.5-12 m m , the stemonozone ±0.5 m m ,
the tube 2-3 m m ; a thin nectary ±0.7 m m tall around
stipe of ovary; ovary glabrous at anthesis; style well
exserted, the stigma dilated to ±0.25 m m diam. Pods
solitary, erect, linear-oblanceolate, when well fertil
ized 30-45 x 0.5-0.6 m m , 4-8-seeded, the sutural ribs FlG. 6. Calliandra depauperata Bentham.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 75
1.2-1.7 m m wide in dorsal view, the recessed valves
thinly coriaceous, transversely venulose, the whole
minutely remotely pubemlent; ripe seeds unknown.
In caatinga and caatinga transitional to cerrado,
320-420 m, locally subdominant, often associated
with cactaceae, localized in e. Brazil: n.-e. quarter of
state of Bahia and immediately adjoining Piaui and
Pemambuco, mostly within the lower S. Francisco
valley, but extending n. just into s.-w. Ceara (±6°
30'N) and s. in the e. foothills of Chapada Diaman-
tina to 14°S. — Map 21. — Fl. I-V. — Carqueija;
alecrim-do-campo.
Calliandra depauperata is instantly recognized by
stiff, often repeatedly and intricately branched stems,
minute leaves loosely deflexed in dried specimens,
small, commonly 2-4-flowered capitula, and 8-15-
merous androecium.
37. Calliandra silvicola Taubert in Engler, Bot.
Jahrb. 21: 429. 1896, — "[BRAZIL Goias:] ... in
ditione fluminis Maranhao superioris [= rio das
Almas] in silva ad Monro do Salto: U L E n. 2834." —
Holotypus,^B; isotypus, HBG!.
C. mertensioides var. debilis Bentham in Martius, Fl. Bras.
15(2): 420, in obs. 1876. — ". . . legit Burchell ad Forna prov. Goyaz [ace. Smith & Smith (1967: 500) near
Pirenopolis, 15°58'S, 48°56,W, 11 Oct 1827] sub n. 6232."
— Holotypus, K (hb. Hooker.)! = N Y Neg. 1966.
C. surinamensis sensu Bentham, 1876: 417, minore ex parte
(Burchell 7526, NY!).
Microphyllidious, widely branching, arborescent
shrubs attaining 3 m, with slender smooth, gray and
in age blanched annotinous branches, appearing
glabrous but the lf-axes pilosulous with fine erect or
incurved hairs to 0.4—0.7 m m , the ohvaceous subcon
colorous, moderately lustrous lfts glabrous, the capit
ula arising either singly or geminate from incipiently
elongating, efoliate brachyblasts axillary to contem
porary or lately shed lvs of long-shoots; phyllotaxy
distichous. Stipules inequilaterally lanceolate 2.5-5 x
1-1.3 m m , ±5-7-nerved, persistent. Lf-formula
i-ii/12-17, the pinnae 1 pair in most brachyblast-lvs,
sometimes in all; lf-stk of primary lvs (1-)1.3-3.5
cm, bridged and bicupulate at insertion of each pinna-
pair, the petiole mostly (0.7-) 1.3-2.3 cm, the inter
pinnal segment to 8-15 m m ; pinnae equilong or the
proximal pair a trifle shorter, the rachises of distal
pair 2.5-5.6 cm, the longer interfoholar segments
2-3.5 m m ; lft-pulvinules transversely elliptic 0.2-0.3
x 0.5-0.6 m m ; lfts subequilong except at far ends of
rachis, the blades linear-oblong from proximally ob-
tusangulate base, deltately acute, those near mid-
M A P 21. Distribution of Calliandra depauperata Bentham and C. silvicola Taubert in eastern Brazil.
rachis 9.5-11.5 x 2.2-3 m m , 3.4-4.5 times as long as
wide; venation slenderly prominulous on both faces of
young lfts, less pronounced in age, primarily pinnate,
the straight subcentric midrib ±5-9-branched on each
side, a weak posterior primary produced barely to,
more often far short of, mid-blade, the tertiary venula
tion random. Peduncles slender 10-35 m m , 1-bracteate
near middle, the bract ovate 1-1.5 m m ; capitula 7-12-
fld, the receptacle 1.5 m m or less; floral bracts minute;
fls (so far as known) homomorphic, the calyx 5- and
the corolla 5-6-merous; pedicels at most 0.3 m m ; pe
rianth submembranous, glabrous except for micro
scopically ciholate calyx-teeth, "purple" ace. to proto-
logue; calyx campanulate ±1.2-1.8 x 1.5 m m , bluntly
5-angulate 5-ribbed, the deltate or depressed-deltate
teeth 0.3-0.7 m m ; corolla ±4—5 m m , cleft to middle or
below, the lobes recurved at anthesis; androecium
26-28-merous, 15-20 m m , the continuous tube only
±1.5-2 m m , the filaments thence united in fascicles to
8-9 m m , the tassel apparently pink or carmine; ovary
subsessile glabrous. Pod unknown.
Of poorly known ecology and dispersal, reported
from "woods" and "gallery-woodland," first collected
in 1827 by Burchell near C o m m b a and Pirenopolis,
then in 1892 by Ule on the upper Maranhao fork of
the Tocantins, in 1964 by Irwin between Brasilia and
Planaltina, and in 1968 by Irwin between C o m m b a
and Niquelandia at 750 m, all within 15°-16°S and
47°45'^9°W in Distrito Federal and state of Goias,
Brazil. — M a p 21. — Fl. IX-X(-?).
W h e n first discovered, C. silvicola was mis
interpreted by Bentham partly as a variety of C.
76 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
mertensioides, which is C. asplenioides of this ac
count and a member of sect. Calliandra, not of sect.
Androcallis, and partly as the Amazonian C. surina
mensis, in which the number of pinnae is stabilized at
exactly one pair per leaf. None of the collections of C.
silvicola presently available for study are of good
quality, and the affinities of the species are obscure.
38. Calliandra surinamensis Bentham, London J.
Bot. 3: 105. 1844. — "Surinam, Hostmann,n. 171."
— Holotypus, K! = photo s.n., NY!; isotypi, BM!,
NY!, OXF!.
Inga fasciculata Willdenow, Sp. PI. 4(2): 1022. 1806. — "Habitat in provincia Para Brasiliae . . . [F. W. Sieber com-
mun.] Com[es] de Hoffmannsegg." — Holotypus, B-
WILLD 19048, seen in Microform! = F. Neg. 7240!. —
Acacia fasciculata Poiret, Encycl. Suppl. 1: 46. 1810.
Feuilleea fasciculata O. Kuntze, Revis. Gen. PI. 1: 185.
1891. Anneslia fasciculata Kleinhoonte in Pulle, Fl. Suri-
name 2(2): 322. 1940. — Non Calliandra fasciculata Bentham, 1840. — Equated with C. surinamensis by Bentham, 1875: 547.
C. tenuiflora Bentham, Trans. Linn. Soc. London 30: 547.
1875; & in Martius, Fl. Bras. 15(2): 416. 1876. — "Habitat ad Rio Tapajos prope Santarem Brasiliae boreahs
[Para]: Spruce n. 389." — Holotypus, K!; isotypus, NY!.
— Feuilleea tenuiflora O. Kuntze, Revis. Gen. PI. 1: 189. 1891.
C. angustidens Britton & Killip, Ann. New York Acad. Sci.
35: 134. 1936. — "[cult, in a plaza at] Villavicencio, Meta,
Colombia, E. Perez Arbeldez 195." — Holotypus, US!; clastotypus (fragm.), NY!; isotypus, COL!. C. surinamensis sensu Bentham, 1876: 417 (excl. Burchell
7526, quae = C. silvicola Taubert); Ducke, 1949: 50; Maas
& Westra., Neotrop. PI. Fam. fig. 41. 1993. C. tenuiflora sensu Ducke, 1949: 50; Irwin, 1966: 97;
Jansen-Jacobs, 1976: 469.
Low bushy but potentially arborescent shrubs with
plagiotropic long-shoots, flowering at (0.7-) 1.5-6
(-10, -12) m, at maturity with flat or asymmetrically
tilted crown, heteromorphic in number, size and out
line of lfts and in composition of capitula, the young
branchlets, lf-axes (either all around or only ven
trally) and peduncles either thinly or densely pilosu
lous with pallid or sordid, either erect, incurved-
ascending or subappressed hairs to 0.1-0.5(-0.7)
m m , the annotinous and older branches glabrate
blanched, the chartaceous (when young submembra-
nous) lfts ± bicolored, lustrous and commonly gla
brous (exceptionally papillate or thinly minutely
pubemlent) on both faces, ciholate or not, the capit
ula arising singly from nearly always efoliate nodes
of brachyblasts axillary to coeval or lately shed pri
mary lvs of long-shoots, the androecia bicolored, pal
lid proximally, rose-carmine distally; phyllotaxy dis
tichous. Stipules lanceolate from ± dilated base or
less often ovate-triangular, mostly glabrous dorsally,
those subtending primary lvs (1.5—)3—11(—16) x
(0.8-)l-2.7(-3) m m , 8-19-nerved when young, be
coming dry, pallid, brittle in age, those of brachy
blasts mostly shorter. Lf-formula i/5-7, 6-9, 8-19
(-25), the lfts when relatively numerous tending to be
more crowded but whether few or many equally vari
able in outline; petioles including pulvinus and
dilated apex 2—9(—11) m m , at middle 0.5-1.3 m m
diam, shallowly grooved ventrally; rachis of longer
pinnae (3.5—)4—9(—11) cm, the longer interfoholar
segments (2—)2.3—12(—21) m m ; lft-pulvinules
0.2-0.6 x 0.5-1 m m ; lfts either decrescent toward
base of rachis and thence equilong, or decrescent also
distally, or accrescent from base upward, the furthest
pair often longer and proportionately narrower than
the penultimate one, the blades varying from linear-
lanceolate and straight or almost so to lance-oblong,
oblong, or rhombic-oblong (and often subsigmoid or
subfalcate) from semicordate or shortly auriculate
base, at apex obtuse, obtuse apiculate, or deltately
acute, the penultimate pair (7.5-)8.5-25 x (1.8-)2-14
(-18) m m , (1.5—)1.8—5(—5.3) times as long as wide;
venation palmate-pinnate, the midrib of narrower lfts
subcentric, of broader ones displaced to divide blade
±1:1.5, or that of relatively short and broad ones sub-
diagonal, the inner of 3-4 posterior primary nerves
incurved-ascending nearly to or shortly beyond mid-
blade, the secondary and reticular venules finely
sharply prominulous on both faces or sometimes
nearly immersed on upper. Peduncles 0.7-2 cm, ran
domly to 2-4(-4.5) cm, 1 (-2)-bracteate, the bract
near or below middle; capitula (6-)8-21(-26)-fld, the
receptacle 1.5-3.5 m m diam; bracts narrow-ovate,
subulate, or narrowly lanceolate, 0.5-2.3 m m , 1^1-
nerved, persistent; fls of capitulum subhomomorphic
as to perianth (that of subterminal fls sometimes a
little wider) but either homomorphic or heteromor
phic as to androecia, these potentially of 3 sorts, with
tubes: a) scarcely exserted, b) long-exserted and nar
rowly tubular, or c) long-exserted and dilated into a
funnel up to ±4 m m diam at orifice; pedicels (some
times scarcely differentiated externally) 0.2-0.8 x
0.4-1 m m ; perianth (3-)4-5-merous, most often
glabrous but sometimes minutely pubemlent or thinly
strigulose, the calyx sharply striate, the membranous,
greenish-white or reddish corolla not so; calyx (1.5-)
1.8-3.2(-3.4) x 0.9-14(-l.7) m m , the depressed-
ovate obtuse or triangular-subulate acute teeth 0.15-1
m m ; corolla (5-)5.4-10.5(-11.4) m m , the ovate or
lance-ovate lobes (0.6-)0.8-2.4(-3.3) m m ; androe
cium 24-48(-52) m m , (ll-)12-26(-34)-merous, the
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 77
tube 7.5-17.5(-21) m m ; peripheral fls either stami
nate or bisexual, lacking disc, some differentiated
distal fls staminate or neuter, with disc; ovary at
anthesis glabrous, often pubescent after fertilization.
Pods stiffly ascending on thickened peduncles, in
broad profile 4.5-11(-11.5) x 0.7-1.3(-1.4) cm, the
sutures to 3 m m wide in dorsal view, the stiffly leath
ery, obliquely (subvertically) venulose, dark brown
nigrescent valves either glabrous or brownish-
puberulent, the inner face of the sutures revealed at
dehiscence 0.85-1.25 m m wide; seeds in broad view
7.5-10.5 x 0.45-0.7 m m , the smooth brown testa
sometimes darker-speckled, lacking pleurogram.
Open places in savanna, bmsh-woodland, and dis
turbed moist forest, especially abundant and colonial
along rocky riverbanks and on islands in river rapids,
mostly between 5 and 450 m but attaining 1200 m on
the Gran Sabana in Venezuelan Guayana, 650 m in
pre-Andean Ecuador, and 1440 m in n. Colombia,
widely dispersed in n. S. America: most abundant in
the Guianas, s. Venezuela, and the central and lower
Amazon basin in Brazil, w. to the e. Cordillera and n.
Antioquia in Colombia, Amazonian Ecuador, the
Ucayali valley in Pern, and Rondonia, s.-e. in Brazil
into Maranhao. — Map 22. — Fl. through the year,
unless drought-inhibited. — Salsa (Brazil).
The broadly drawn definition of C. surinamensis
proposed herewith may come as a surprise to those
familiar with the traditional taxonomy, in which C.
tenuiflora has invariably been accepted as a distinct
taxon. As first known to Bentham (1875: 547), C.
tenuiflora differed from C. surinamensis, as then
known, in having 5-6 rather than 8-12 pairs of leaflets
per pinna, the largest of them >2 cm, as opposed to
<1.5 cm, in length. No substantial difference in the
inflorescence or the individual flower was then
known, and both species were described from Para.
Subsequently C. tenuiflora (sens, str.) has been con
sidered (Irwin, 1966: 97) closer to C. purpurea, and its
affinity to C. surinamensis has been lost sight of.
Specimens from the whole vast range of C. surina
mensis now demonstrate a continuous series of vari
ants in number and size of leaflets that has erased the
supposed discontinuity between it and C. tenuiflora.
As a rule, relatively numerous leaflets are relatively
small and fewer leaflets larger, and the furthest pair
either is or may be largest of all in both cases. Whether
few or many, the leaflets vary in outhne from narrowly
lanceolate to obtusely rhombic. Plurifoliolate forms of
C. surinamensis are distributed over the species range,
from the Guianas and lower Amazonian Brazil west to
Colombia and Pern, whereas the paucifoholate ones
are scattered along the Amazon and tributaries from
Para almost to the Peruvian border and north through
the Guianas to Venezuelan Guyana, without establish
ing any independent area of dispersal. The variable
development of the androecial tube, as described
above, is linked neither with leaf-formula nor with
geographic dispersal.
39. Calliandra samik Barneby, sp. nov., C. surina-
mensi habitu, indumento foliisque simillimo haud
nisi perianthio insigniter ampliori, calyce 5.5-6 x 3
(nee 1.5-3.4 x 0.9-1.7) m m et corolla 13-14.5 (nee
5-10.5, rarissime 11.4) m m diversa. — PERU.
Amazonas, prov. Bagua: Rio Cenepa, 5 minutes
downstream from Chavez Valdivia, 16 Dec 1972
(fl), Brent Berlin 554. — Holotypus, NY.
Arborescent shrubs, closely resembling plurifolio
late forms of C. surinamensis in habit, indumentum,
and inflorescence but markedly different in ampler
perianth. Stipules of primary lvs ±5-6 x 2 mm, stri
ate. Lf-formula i/17-19; lf-stks of primary lvs 4—6 x
0.7-1.1 mm, of brachyblast lvs scarcely shorter;
pinna-rachis in primary lvs 6-7.5 cm, the longer
interfoholar segments ±3^4.5 mm; lft-pulvinules 0.5-
0.7-0.8 mm; lfts a little decrescent at each end of
rachis, otherwise subequilong, the blades lance-el-
lipitic from semicordate base, deltately acute apicu-
late, those near mid-rachis 11-12.5 x 4-4.5 m m ,
±2.7-2.8 times as long as wide; midrib only a trifle
excentric, the inner of 3(-4) posterior primary nerves
produced ± to mid-blade. Peduncles stout 14—16 m m ,
bracteate below middle; capitula ±10—16-fld, the re
ceptacle 2 m m diam; bracts lanceolate 1.5-2 m m ;
pedicels 0.6 x 0.7-1 mm; fls homomorphic, the peri
anth thinly sordid-strigulose, the calyx weakly striate,
the membranous corolla not so; calyx ±6x3 mm, the
ovate-triangular teeth 2-2.8 mm; corolla 13-14.5
mm, the lobes 3.4-4 mm; androecium of all fls 34-
merous, ±41 mm, the far-exserted tube 26 m m ,
slightly dilated distally; no disc observed; ovary and
pod unknown.
On banks of rapidly flowing stream, ±225 m,
known only from the type-locality in the middle
Marafion basin, near 4°30/S, 78°15,W in prov. Bagua
of depto. Amazonas, Pern. — Fl. XII—I, probably at
other times. — Samik.
Calliandra samik closely resembles the polymor
phic C. surinamensis in foliage and indumentum and
may prove to be merely another extreme variant of it.
However, the much longer perianth (for measurements
see the Latin diagnosis above) can hardly be accom
modated in my present concept of C surinamensis.
78 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
M a p 22. Distribution of Calliandra surinamensis Bentham in northern South America.
40. Calliandra purpurea (Linnaeus) Bentham, Lon
don J. Bot. 3: 104. 1844, based directly on Inga pur
purea (Linnaeus) Willdenow, Sp. PL 4(2): 1021.1806
— "Habitat in Martinica," which = Mimosa purpurea
Linnaeus, Sp. PL 517. 1753. — "Habitat in America
meridionali." — Based wholly on Acacia frutescens
non aculeata, flore purpurascente Plumier ed. Bur-
mann, PL Amer. t. X, fig. 2 (bottom right + misplaced
caption bottom left). — Holotypus (Howard, Fl.
Lesser Antilles 4: 350. 1988), Plumier's figure, cited
above!. — Feuilleea purpurea O. Kuntze, Revis.
Gen. PL 1: 188. 1891. Anneslia purpurea Britton,
Brooklyn Bot. Gard. M e m . 1: 50. 1918.
Inga obtusifolia Willdenow, Sp. PL 4(2): 1022. 1806. —
"Habitat in Cumana [Sucre, Venezuela]." — Holotypus,
Humboldt 260 labeled "Cumana, an 8," B-WILLD (seen in
Microform)!. —Mimosa obtusifolia Poiret, Encycl. Suppl.
1: 46. 1810. — Calliandra obtusifolia Karsten, Fl.
Columb. 2: 41, t. 121 (left, the type-locality elaborated to
Rio Manzanare, 300 ft elev.). 1863. — Equated by Kunth
(1824: 301) and Bentham (1875: 547) with C. purpurea.
Calliandra coroensis Karsten, Fl. Columb. 2: 41, t. 121
(right). 1863. — "Habitat regiones aridas calidasque
provinciae Venezuelensis Coro [state of Falcon]." — Holo
typus to be sought at W — Equated with C. purpurea by
Bentham (1875: 547). C. purpurea var. dussiana Stehl,, Bull. Mus. Hist. Nat.
(Paris) II, 8(2): 192. 1946. — Said to correspond exactly
with C. purpurea (Linnaeus) Bentham, so best considered
= C. purpurea var. purpurea, the autonym generated by the next.
C. purpurea var. quentiniana Stehle Bull. Mus. Hist. Nat.
(Paris) II, 8(2): 192. 1946. — ". . . Stehle et Quentin n.
5546 . . . [Guadeloupe] Pointe Noire ... Deshaies, par
Ferry ... 3 septembre 1944." — Holotypus, P n.v.; isotypus, NY!.
C. slaneae Howard, Phytologia 61: 3. 1986. — "St. Lucia, 3
miles northeast of Dennery, May 15, 1985, V[erna] Slane 541." — Holotypus, A!.
Inga purpurea sensu de Candolle, 1825: 439; Calliandra
purpurea sensu Bentham, 1875: 546; Britton & Killip,
1936: 193. Anneslia purpurea sensu Britton & Rose, 1923: 60.
Calliandra purpurea + vars. dussiana et quentiniana sensu
Stehle & Quentin, Fl. Guadeloupe Martinique 2(3): 41. 1949; Foumet, Fl. Guadeloupe Martinique 694, 695. 1978.
Stiffly branched, arborescent shrubs flowering
when 1.5-6 m tall, with pallid, either straight or flexu-
ous, usually densely foliate long-shoots and epidermis
soon exfoliating in strips, either glabrous except for
minutely pubemlent lf-axes and peduncles or the new
growth and lfts finely pilosulous with erect-incurved,
palhd or sordid hairs to 0.15-0.35 m m , the mature lfts
firm, facially lustrous, dark above, paler beneath, the
capitula arising, mostly singly, from efohate nodes of
condensed or sometimes longer and loosely thatched
brachyblasts; phyllotaxy distichous. Stipules obtusely
deltate or broad-lanceolate 1.2-4.5(-6) x 0.7-2 m m ,
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 79
when young striately 6-11-nerved, becoming dry
pallescent, mostly persistent. Lf-formula i/3-1 (on
Montserrat and Guadeloupe -9); lf-stks (l-)2-12(-30)
m m , at middle 0.3-0.8 m m diam, narrowly grooved
ventrally, dilated at tip; pinna-rachises (11—)15—38
(-62) m m , the longer interfoholar segments (2.5-)3.5-
9 m m ; lft-pulvinules 0.3-1 x 0.3-0.8 m m ; lfts usually
± accrescent distally, sometimes subequilong, sub-
equiform (the furthest pair then not obviously longer
and narrower), the blades oblong or inequilaterally
(ob)ovate (rarely oblong-rhombic) from semi-cordate
base, broadly obtuse mucronulate, the penultimate
ones (7-)9-20 x (3.5-)4-ll m m , 1.5-2.3(-2.5) times
as long as wide; venation palmate-pinnate, the almost
straight midrib displaced to divide blade ±1:1.5, the
strong inner posterior primary nerve incurved-ascend
ing beyond mid-blade, the outer one much shorter, the
secondary and reticular venulation prominulous on
both faces. Peduncles 0.8-5 cm, bracteate near or
below middle or ebracteate, thickened in fruit; capitula
9-22-fld, the fls (sub)sessile, homomorphic as to peri
anth but either staminate or bisexual and the staminal
tube varying in length between populations; perianth
4- or 5-merous, glabrous, pallid; calyx campanulate or
deeply campanulate 1—2.3(—3.3) x 1-1.8 m m , the ob
tuse teeth 0.15-0.3 m m ; corolla 5-7(-7.8) m m , the
lobes 1.4—2 m m ; androecium 10-16(-18)-merous, red
throughout, 22-34(-39) m m , the tube (4.5-)5.5-14.5
m m , 1 m m shorter to 7 m m longer than the corolla, the
disc in bisexual fls 0.4 m m tall, in staminate fls obso
lete; ovary glabrous. Pods glabrous, stiffly erect, 5.5-9
x 0.8-1.1 m m , the thickened sutural ribs 2.5-3 m m
wide in dorsal view, the recessed plane valves faintly
obliquely venulose; seeds 6-8.6 x 4-6.2 m m , the
smooth brown testa dark-speckled, pleurogrammic.
In semideciduous scmb woodland and chaparral, on
seasonally dry hills in shallow soil, locally plentiful
below 500 m in the Lesser Antilles, on the Caribbean
slope in n. Venezuela and n.-e. Colombia, and
remotely disjunct in s. Guyana and e. equatorial
Colombia: in the Antilles native on almost all the
islands s.-ward from St. Kitts and Antigua, cultivated
in its native range and further n.; in Venezuela local in
Sucre, and in Falcon and Lara; in n. Colombia local in
Atlantico and Magdalena (to be expected in Guajira);
in Amazonian Colombia (introduced "chiribiquete");
in Guyana, perhaps varietally distinct in shghtiy
longer perianth (calyx to 3.3 m m , corolla to 7.8 m m ) ,
at 500 m on the Kanuku Mts. — M a p 23. — Flower
ing throughout the year, but most prolifically follow
ing rains. — Buisson ardent, bois patate (French
Antilles, in reference to red filaments and tuberiferous
M a p 23. Distribution of Calliandra purpurea (Linnaeus)
Bentham in the Lesser Antilles and northern Venezuela
(location in the Kanuku Mts. omitted).
roots); soldier bush (former British West Indies, the
vivid red capitula recalhng uniformed Redcoats).
Leaflet size and leaflet number vary considerably
between populations of C. purpurea on the Lesser An
tilles and between those scattered along the Caribbean
slope in Venezuela and Colombia. The following num
bers of leaflet-pairs (per pinna of primary leaves) are
on record: Antigua: 5; Montserrat and Guadeloupe:
6-8(-9); Martinique, St. Lucia, St. Vincent, and the
Grenadines: mostly 3-4, rarely 6; northern Venezuela:
(2-)3, 4, or 5; northern Colombia: 4-7. Plumier's
drawing, which furnished the Linnaean protologue,
shows 3 and 4 pairs; the model very hkely came from
Martinique, as did that of var. dussiana, which is es
sentially typical C. purpurea. The populations with
6-8 pairs of leaflets on Montserrat and Guadeloupe
correspond with var. quentiniana, and are mildly dis
tinctive; however, 6-7 pairs recur in Magdalena,
Colombia. Calliandra slaneae, with 3-4 pairs, is inter
preted as the xeromorphic extreme from arid insular
habitats. Venezuelan C. obtusifolia (from Sucre) and
C. coroensis (from Falcon), with respectively 4—6 and
(2-)3 pairs of leaflets per pinna, were long ago equated
with C. purpurea by Bentham. The geographically re
mote population in Guyana (A. C. Smith 3168, K, N Y )
was identified as C. purpurea by Sandwith and appears
at very most varietally distinct from Caribbean C. pur
purea in shghtiy larger flowers. Finally, size of leaflets
and pubescence of leaflets are randomly correlated
with dispersal and offer no firm basis for segregation.
80 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
41. Calliandra riparia Pittier, Arb. Arbust Venez. 6-8:
80.1927 (Aug-Sep); & Arb. Legum. 1 (Mimosaceae):
10. 1927 (Dec). — "[VENEZUELA.] Riberas del rio
San Juan de los Morros, Aragua; flores y fmtos Abril
9,1927 (Pittier 12309...)." — Holotypus, V E N n.v;
isotypi, G!, NY!. — Mistakenly equated by Woodson
and Schery (1950: 259) with C. magdalenae.
Fig. 7
C. schultzei Harms, Repert. Spec. Nov. Regni Veg. 24: 209.
1928. — "Colombia: Santa Marta, Rio Pedras, Nordfuss
der Sierra Nevada . . . (Arnold Schultze no. 553;
7.IX. 1926)." — Holotypus, tB = F Neg. 7255 + clastotypus
(If, fls), F!, duplicate photo, NY!. — Mistakenly equated by
Woodson and Schery (1950: 259) with C. magdalenae.
C. schultzei sensu Britton & Killip, 1936: 135.
C. surinamensis sensu Prance & Silva, Arvores de Manaus
156, fotos 73,74. 1975; Howard, 1988: 351; Pauwels, Nza-
yiluN'tit. 135(1-3). 1993.
Arborescent shrubs 1.5-6 m with palhd plagiotropic
branches and virgate long-shoots, at maturity flat-
topped or leaning, the young branches, peduncles and
especially the lf-axes pilosulous with either erect or
incurved, gray or sordid hairs 0.2-0.7 m m , the firm
green lfts not strongly bicolored, facially glabrous but
often thinly ciholate, the lvs relatively insensitive
(often fully expanded in pressed specimens), the capit
ula arising singly or exceptionally geminate either
directly from primary lf-axils or from very short, often
fohate axillary brachyblasts; phyllotaxy distichous.
Stipules subtending primary lvs triangular-lanceolate
to lance-acuminate or -caudate (3-)4—11 x 1.2-2.7
m m , striately 9-17-nerved, becoming dry, palhd, and
brittle but not spontaneously disjointing. Lf-formula
i/7—11(—13); petioles 4-15(-18) m m , at middle 0.4-
0.85(-l) m m diam; pinna-rachises (2.5-)3-6(-6.5)
cm, the longer interfoholar segments (2.5-)3-7 m m ;
lft-pulvinules 0.4-0.8(-0.9) x 0.5-0.8 m m , wrinkled;
lfts broad-linear or linear-elliptic from semicordate
base, straight or almost so, at apex triangular-apiculate
or mucronulate, the longer ones 12-22 x (2.2-)2.5-
4.6(-6) m m , (3.5-)3.8-5.7(-6.5) times as long as wide,
the terminal pair no longer than the penultimate; vena
tion palmate-pinnate, the straight midrib dividing blade
±1:1.5-1.75, the inner posterior primary nerve pro
duced nearly to or shortly beyond mid-blade, the 1-2
(-3) outer ones progressively much shorter, the sec
ondary and reticular venules sharply finely prominu
lous on both faces. Peduncles (9-)10-32(-36) m m ,
1-2-bracteate near or below middle or sometimes
ebracteate; capitula 12-14(-?)-fld, the clavate recepta
cle ±2-3 m m (sometimes one fl downwardly displaced
onto peduncle), bracts subulate or linear 0.8-2.2 m m ,
3-4-nerved, persistent; fls at full anthesis all strongly
ascending, subhomomorphic as to perianth but the an
droecia varying from almost homomorphic to strongly
differentiated in length and amplitude of staminal tube;
pedicels externally discolored 0.1-0.8 x (0.3-)0.4-
0.8 m m ; perianth glabrous, the calyx sharply finely
striate, the corolla not or faintly so; androecial tube of
peripheral fls as long as or ± twice as long as the the
corolla, that of 1-several central fls often but not
always much longer than that of peripheral fls and
expanded at apex, the tube of all palhd or faintly pink-
tinged, the tassel pink or carmine; calyx 2-2.6(-2.8) x
0.9-1.5 m m , the teeth (often unequal) 0.25-0.6 m m ;
corolla either 4- or 5-merous, greenish or whitish,
(5-)5.5-8 m m , the ovate-deltate lobes 1.1-2.3 m m ; an
droecium 12-20-merous, 31^42 m m , the tube of pe
ripheral fls 6.5-11 m m , that of central fls 14-31 m m ,
expanded at orifice to 3-5 m m diam; ovary at anthesis
glabrous. Pods stiffly ascending toward vertical, (5-)
6-8 x 0.8-1.1 cm, glabrous or ciholate along sutures,
the valves stiffly leathery, obhquely or subhorizontally
venulose, the inner face of the sutures exposed by de
hiscence 0.9-1.1 m m wide; seeds in broad view
7.3-9.8 x 4.5-5.4 m m , the pale brown, often dusky-
speckled testa lacking pleurogram.
Native in semideciduous or thin evergreen wood
land, often colonial on rocky river banks, 50-750 m,
scattered in the Orinoco valley and Maracaibo basin
in Venezuela e. to the Pakaraima mountains in adj.
Guyana, w. into n. Colombia and adj. Panama; culti
vated and locally naturalized up to 1330 m in Central
America, Amazonian Brazil and Bolivia, e. Brazil,
the Bahamas and Antilles, and in gardens and botan
ical gardens in s.-e. Asia and Hawai'i. — M a p 24. —
FL nearly yearlong, except when drought-stricken. —
Canasta mexicana (Puerto Rico, the adjective signi
fying foreign rather than Mexican).
In 70 years since its discovery C. riparia has ac
quired no substantial identity in the literature, but
is nevertheless one of the commoner unijugate
calliandras in seasonally dry parts of northern South
America; moreover it is widely cultivated both in its
native range and elsewhere, but where cultivated
passing as C. surinamensis. It does indeed resemble
forms of polymorphic C. surinamensis that have
about 7-11 pairs of leaflets per pinna, but these
leaflets are linear or linear-elliptic rather than rhom
bic or oblong. The hard seed-coat, lacking pleuro
gram, provides the decisive differential character.
The pre-colonial distribution of C. riparia, before
it was taken into parks and gardens, seems to have
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 81
FIG. 7. Calliandra riparia Pittier.
82 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
M A P 24. Distribution of Calliandra riparia Pittier in Central and South America and the West Indies.
centered in the Orinoco valley between Zulia and Boli
var, reaching the Pakaraima mountains in Guyana and
extending westward through the Caribbean lowlands
of Colombia to Choco and to eastern Panama. Beyond
this range there are records from Trinidad, Atlantic
Guyana, one each from Honduras and Costa Rica,
many from inter-Andean Colombia, and from further
afield, all probably or certainly taken from ornamental
plantings. As native status is seldom noted on herbar
ium labels, it is not possible to distinguish between
aboriginal and subsequent status, even in Venezuela.
The accompanying map records all known occurrences
of C. riparia in the Americas.
The remarks by Skutch (Brenesia 37: 141-143, fig.
1) on floral dimorphism and phenology of C. surina
mensis cultivated at El General, Costa Rica, are per
haps applicable to C. riparia.
42. Calliandra magdalenae (de Candolle) Bentham,
London J. Bot. 102. 1846. — Typus infra sub var.
magdalenae indicatur.
Trees 2-8 (-10, once reported 20) m tall with habit,
indumentum and bicolored androecia of Cc. surina
mensis and riparia, variable in size and number of
lfts, these lustrous dark green above, paler duller be
neath, facially glabrous or rarely pilosulous, ran
domly ciholate, the peduncles arising singly or less
often geminate from bracteate axils of either loosely
or densely thatched brachyblasts; phyllotaxy distic
hous. Stipules subtending primary lvs of long-shoots
lanceolate, lance-attenuate or narrowly triangular
2.5-9 x 0.7-2.7 m m , when young striately 7-15-
nerved, becoming dry and fragile, those of brachy
blasts similar but a little shorter. Lf-formula i/(12-)
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 83
13-36; petioles 2-10(-12) x 0.35-1.2(-l.4) m m ;
pinna-rachises of primary lvs 4-10 cm, of brachyblast
lvs often shorter and in foliose saplings reaching 12
cm, the longer interfoholar segments 1.3-6.5 m m ; lft-
pulvinules 0.1-04 x 0.35-O.8(-0.9) m m ; lft-blades
usually longest at mid-rachis but only a little shorter at
base and apex, the terminal pair no (scarcely) longer
than the penultimate, all in outhne varying from linear
to oblong from semicordate or angulately auriculate
base, obtuse mucronulate or apiculate, straight or gen
tly incurved beyond middle, the longer ones (6-)7-18
x 1.4-6 m m , 2.2-5.6(-6.4) times as long as wide;
venation of narrower lfts almost simply pinnate, of
broader lfts clearly palmate-pinnate, the midrib only
shghtiy excentric, the inner posterior primary nerve
incurved-ascending to or beyond mid-blade of broader
lfts but weak and short in narrow ones, the secondary
and reticular venules all finely sharply prominulous
on both faces. Peduncles (2-)5-12 m m , mostly
bracteate below middle; capitula 16-28-fld, the recep
tacle 1.5^4 m m ; bracts subulate 0.6-2.2 m m , 1-3-
nerved, persistent; pedicels (sometimes scarcely dif
ferentiated externally) 0.25-0.55(-0.7) x (0.25-)0.3-
0.9 m m ; perianth either 4- or 5-merous, commonly
glabrous but sometimes minutely pubemlent or strigu
lose, the calyx striate, the corolla not so; flowers
within the capitulum either subhomomorphic or
strongly heteromorphic, the perianths subequal in all
but the androecial tube of the peripheral ones, some
times of all, as long as or up to 4.5 m m longer than
corolla, that of 1-3 subterminal fls cylindric 20-32
m m , expanded at orifice to 4—5 m m diam; calyx cam
panulate 1.2-2.2(-2.7) x 0.9-1.3(-l.5) m m , the teeth
0.1-0.4 m m ; corolla (3.8-)4.2-6.6(-7.5) m m , the
lobes 0.9-1.8 m m ; androecium 25-^2 m m , (9-)10-
15-merous, pallid proximally, the tassel carmine;
ovary at anthesis glabrous, becoming pubemlent after
fertilization. Pods stiffly erect from the plagiotropic
branches, in profile 7-11 x 0.9-1.35 cm, massively
woody and inflexible in texture, brown nigrescent, the
longitudinally ribbed sutures 3.5-5 m m wide in ex
ternal view, 1.5-2 m m thick at the plane of dehis
cence, the deeply recessed, coarsely obliquely venu
lose valves 2-5 m m wide, often narrower than either
suture, the whole sordid- or brown-puberulent in
youth but sometimes glabrate at maturity; seeds fawn,
fuscous-speckled, in broad view 8-9.6 x 5-6.2 m m ,
the smooth testa finely pleurogrammic.
Calliandra magdalenae is closely related to C.
surinamensis and C. riparia, differing, however, from
both in the thickly woody fmits and in pleurogram
mic seeds, and from the second further in more
numerous leaflets. Like C. surinamensis the species
has differentiated out into geographic races charac
terized by fewer larger and more numerous, crowded,
and narrower leaflets separable as follows:
Key to the varieties of C. magdalenae
1. Lfts of longer pinnae (12-) 13-19(-20) pairs
spaced along the pinna-rachis at intervals
to 3-6.5 mm, the larger blades 11-18 x
3-6 mm, 2.2-3.7(-4) times as long as wide;
s. Mexico (Oaxaca, Chiapas) s.-e. through
Centr. America to the Magdalena Valley in
Colombia 42a. var. colombiana
1. Lfts of longer pinnae 20-36 pairs spaced along
the pinna-rachis at intervals to 1.3-2.6(-3) mm, the larger blades (6-)7-12 x 1.4-2.6
(-2.8) mm, (3.7-)4-5.6(-6.4) times as long
as wide; n.-w. Venezuela (Falcon, Zulia, Merida), n. Colombia (La Guajira, Magdalena,
Bolivar) and adj. Panama 42b. var. magdalenae
42a. Calliandra magdalenae (de Candolle) Ben
tham var. colombiana (Britton & Kilhp) Barneby,
stat. nov. C. colombiana Britton & Rose ex Britton
& Kilhp, Ann. N e w York Acad. Sci. 35: 135. 1936.
— "Colombia. Natagaima, Huila . . . 450-500 m.,
August 12, 1917, [F W.] Pennell 1153." — Holo
typus, NY!; isotypus, NY!.
Anneslia chiapensis Britton & Rose, N. Amer. Fl. 23: 61.
1928. "Type from near Tapachula, Chiapas, 1896, [E. W.] Nelson 3838" — Holotypus, US!; isotypus, NY!. — Calliandra chiapensis Lundell, Phytologia 1: 369. 1940. —
Equated with C. magdalenae by Woodson and Schery (1950: 259).
A. tonduzii Britton & Rose, N. Amer. Fl. 23: 61. 1928. —
"Type collected at Bonica [= Burica], Costa Rica, 1891,
Tonduz 4544" — Holotypus, US!; clastotypus, NY!. —
Calliandra tonduzii Standley, Publ. Field Columbian Mus.,
Bot. ser. 4: 309. 1929. — Equated with C. magdalenae by Woodson and Schery (1950: 259).
Calliandra santanderensis Britton & Rose ex Britton & Kil
lip, Ann. New York Acad. Sci. 35: 135. 1936. — "Rio
Surata Valley, near Bucaramanga, Santander, Colombia,
400-600 m ... December 28, 1926, Killip & Smith 16205."
— Holotypus, NY!. — Mistakenly described as having pinnae bijugae, this derived from a short branchlet bearing two
distinct but partly overlapping leaves with geminate pinnae.
As described for the species and modified by the
key to varieties.
Colonial along streams in open woodland, surviv
ing and sometimes invasive in cafetal and pastureland,
200-900 m, discontinuously dispersed in Central
America and Colombia: from e. Oaxaca and Chiapas
in Mexico s. to Panama; Magdalena valley, Colombia,
in deptos. Santander, Cundinamarca, Caldas, Tolima,
Huila. — Fl. VI-I. (Central America), VI-VIII
84 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
(Colombia) — Map 25.
(Colombia).
Aroma (Panama); rayado
42b. Calliandra magdalenae (de Candolle) Ben
tham var. magdalenae. C. magdalenae Bentham,
1846, I.e., sens. str. Acacia magdalenae de Can
dolle, Prodr. 2: 455. 1825. — "ad Sanctam Martham
legit cl. Bertero." — Holotypus, G-DC, seen in Mi
croform 25, box 13!. — Feuilleea magdalenae O.
Kuntze, Revis. Gen. PL 1: 188. 1891. Anneslia mag
dalenae Britton & Rose, N. Amer. Fl. 23: 60. 1928.
Acacia magdalenae Sprengel, Syst. Veg. 3: 137. 1826. —
"Ad fl. Magdalenae [sic], Bertero." — Typus, presumably a duplicate of the specimen described by de Candolle
under the same epithet, not known to survive.
Codonandra purpurea Karsten, Fl. Columb. 2: 43, t. CXXII.
1863. — "Habitat planities aridas siccas provinciae
Venezuelanae Coro, in valle Uvedal collecta." — Holotypus
not seen, but the protologue decisive. — Calliandra
codonandra Bentham, Trans. Linn. Soc. London 30: 547.
1875; non C. purpurea (Linnaeus) Bentham, 1844. Feuilleea
codonandra O. Kuntze, Revis. Gen. PL 1: 187. 1891.
Calliandra magdalenae sensu Britton & Killip, 1936: 136;
Woodson & Schery, 1950: 259, sens, lat., max. pro parte
(exclus. syn. C. riparia, schultzei, angustidens).
As described for the species, and modified by the
key to varieties.
Colonial along riverbanks in semideciduous forest-
climax, in seasonally dry thickets, and surviving in
pastures, 50-650 m, discontinuously dispersed in n.-w.
Venezuela (Falcon, Zuha, Merida), n. Colombia (La
Guajira, Santa Marta, Bolivar), and adj. e. Panama, in
ecology and dispersal suggestive of a xeromorphic de
rivative of var. colombiana. — M a p 25. — Fl.
(V-)VI-XII. —Anda-arriba (Zulia).
43. Calliandra caeciliae Harms, Repert. Spec. Nov.
Regni Veg. 17: 89. 1921. —"Guatemala: Dep. Hue
huetenango, Uaxackanal [= Guaxacana] . . .
1300-1400 m... (Caec[ilie] u. E. Seler no. 3006 —
Aug. 1896)." — Holotypus. +B = F Neg. 7257!;
clastotypus + photo, NY!. — Anneslia caeciliae
Britton & Rose, N. Amer. Fl. 23: 61. 1928.
C. densifolia Rose ex Harms, Repert. Spec. Nov. Regni Veg.
17: 89. 1921, nom. subnud., only inadvertently published in
discussion of the preceding. — "Pringle no. 8671, 1902;
Michoacan, Monte Leon Stat." — Lectoholotypus (Mc
Vaugh, 1987: 153), US!; isotypus, K!. —Anneslia densi
folia Britton & Rose, N. Amer. Fl. 23: 60. 1928. — Equated with C. caeciliae by Standley and Steyermark (1946: 20).
Microphyllidious trees 2-6 m with plagiotropic
long-shoots, habitally resembling C. magdalenae and
C. cruegeri, the young stems, lf-axes, and peduncles
thinly (densely) strigulose-pilosulous with gray hairs
to 0.2-0.6 m m , the crowded narrow lfts olivaceous,
scarcely paler beneath, (sub)glabrous facially, often
thinly subappressed-ciliolate, the often very short
peduncles arising singly from condensed, densely
thatched, mostly efohate brachyblasts axillary to coe
val or lately fallen primary lvs; phyllotaxy distichous.
Stipules lance- or ovate-acuminate, those subtending
primary lvs 2.5-5.5 x 0.7-1.8 m m , striately 7-14-
nerved when young, early dry deciduous, those of
brachyblasts shorter, persistent. Lf-formula i/23-37,
but often seemingly fewer when primary lvs have
fallen, the lfts fewer in many brachyblast lvs; petioles
1-3.5 m m , at middle 0.4—1 m m diam; rachis of longer
pinnae 3-7 cm, the longer interfoholar segments 0.8-
2.5 m m , the lfts decrescent toward each end of rachis;
lft-pulvinules 0.1-0.3 x 0.3-0.5 m m ; lft-blades hnear
or linear-lanceolate from semicordate or obtusangu-
late base, triangular-acute, straight or almost so, those
near mid-rachis 5-13 x 0.9-1.9(-2.2) m m , 5-8.2
times as long as wide; venation weakly palmate-
pinnate, the midrib scarcely excentric, the inner pos
terior primary nerve produced at most to mid-blade,
often shorter, the 1-2 outer ones much shorter, the sec
ondary venules either finely prominulous or almost
immersed ventrally. Peduncles 1—15(—17) m m ,
ebracteate; capitula 6-14-fld, the receptacle ±1-1.5
m m ; bracts ovate or subulate 0.45-1 m m , persistent;
perianth either 4- or 5-merous, either greenish-white
or red-tinged, glabrous except for sometimes minutely
ciholate calyx-rim, the calyx sharply striate, the
corolla not so, the androecium pallid proximally, its
tassel carmine; fls of the capitulum variably homo- or
heteromorphic, the androecial tube of the peripheral
(either staminate or bisexual) fls ± as long as corolla
or exserted to 2 m m , that of one or more central (ex
ceptionally of all) fls cylindric or trumpet-shaped and
up to ±15 m m , its orifice expanded to 4 m m diam;
P E R I P H E R A L FLS: pedicels 0.1-0.5 x 0.4-0.6 m m ;
calyx campanulate or deeply campanulate 1.2-2.1
(-2.8) x 0.9-1.2 m m , the obtuse teeth 0.1-0.25 m m ;
corolla (3.4-)4-6(-7.7) m m , the ovate lobes 0.8-1.5
m m ; androecium 9-15-merous, 22-29 m m , the tube
3-6.5(-8) m m . Pods erect on thickened peduncle, in
profile 5-7 x 0.5-0.8 cm, the sutures in dorsal view
±2.5 m m wide, their planes of dehiscence ±0.6 m m ,
the leathery valves obliquely (sometimes reticulately)
venulose, thinly pilosulous glabrescent; seeds (few
seen) in broad view 5.5 x 4 m m , pleurogrammic.
In semideciduous woodland and on riverbanks, in
moderately dry sites in the uplands, in moist ravines in
the lowlands, 150-1080 m, scattered in s. Mexico and
Central America: Mexico (Jalisco to Chiapas);
Guatemala (Huehuetenango), Honduras (El Paraiso
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 85
CALLIANDRA MAGDALENAE
• var. MAGDALENAE
f var. COLOMBIANA
MAP 25. Distribution of Calliandra magdalenae (de Candolle) Bentham var. magdalenae and var. colombiana (Britton & Killip) Barneby in southern Mexico, Central America, and northwestern South America.
Ocotepeque); Nicaragua (Esteli). — Map 26. — Fl.
VI-XI. — Cinchote, guaricho (Guerrero).
Calliandra caeciliae differs from other unijugate
calhandras of tropical Mexico and Central America in
high numbers and small size of the leaflets.
44. Calliandra chulumania Barneby, sp. nov, inter
sectionis Androcallidis species austro-americanas
geminato-pinnatas foliolis paucis (7-9-jugis usque)
majusculis (±9-15 x 4—6 m m ) pedunculisque
abbreviatis (5-12 m m longis) praestans, a C. suri
namensis formis paucifoliolatis seminibus pleuro-
grammate ornatis diversa; a C. (ser. Ambivalentes)
mollissima, in Andibus Peruviae boreahs obvia,
petiolis abbreviatis 2-6 (nee 5-20) m m longis, foli
olis minoribus 9-15 x 4-6 (nee 1742 x 10-20) m m
usque, androecioque saturate sanguineo (nee al-
bido) distat. — BOLIVIA. La Paz, Sud-Yungas:
Boopi valley, 980 m, at km 52 on Chulumani-
Asunta road, 8 Aug 1983 (fl), St. G Beck 8597. —
Holotypus, NY; isotypus, COL.
Arborescent shrubs 1.54 m with virgate plagio
tropic homotinous long-shoots, probably drought-
deciduous, the young stems and foliage densely
softly pilosulous with fine straight, erect and ascend
ing hairs to 0.45-0.7 mm, the ventrally low-convex
lfts dull-olivaceous slightly paler dorsally, the small
dense capitula of carmine fls shortly pedunculate,
arising from the first, or first and second, bracteate
but efohate nodes of depauperate brachyblasts axil
lary to homotinous primary lvs; phyllotaxy distic
hous. Stipules herbaceous, triangular-lanceolate 1.5-
5 x 0.6-2 mm, faintly nerved, deciduous. Lf-formula
ill-9; lf-stks 2-6 mm, at middle 0.45-0.8 m m diam,
shallowly grooved; rachis of longer pinnae 34.5(-5)
cm, the longer interfoholar segments 2-5.5(-6.5) mm;
lft-pulvinules 0.25-0.5 mm; lfts usually subdecres-
cent proximally and otherwise subequilong, the
blades obtusely rhombic-oblong from shallowly semi-
cordate base, broadly obtuse apiculate, at tip straight
or incipiently incurved, the larger ones 9-15 x 4-6
mm, 2.2-2.7 times as long as wide; venation primarily
palmate, the midrib forwardly displaced to divide
blade ±1:2, weakly 1-2-branched above middle, the
inner posterior primary nerve incurved to or a little
beyond mid-blade, the outer one shorter, these slen
derly prominulous dorsally, the tertiary venulation
86 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
• C A L L I A N D R A C O N F E R T A
Y C. CAECILIAE
• C. BIFLORA
M a p 26. Distribution of Calliandra biflora Tharp, C. caeciliae Harms, and C. conferta Bentham in southern U.S., Mexico, and Central America.
faint. Peduncles sohtary 5-12 m m , 1-bracteate below
middle, the receptacle ±1.5-2 m m diam; bracts nar
rowly triangular 0.4—0.9 m m , persistent; fls sessile,
the calyx glabrous, the corolla thinly or remotely
appressed- or ascending-pilosulous; calyx deeply
campanulate 1.2-1.5 x 0.6-0.8 m m , finely striate, the
teeth minute; corolla slenderly tubular 5-6.4 m m , the
lobes 0.6-1.4 m m ; androecium 8-14-merous, dark
red-brown when dried, ±2-2.3 cm, the tube 6.5-12
m m ; no disc seen. Pods oblanceolate 6-8 x 1-1.1 cm,
5-6-seeded, the sutural ribs in dorsal view ±3 m m
wide, the pale brown hgnescent valves low-convex
over each seed, closely cross-venulose, minutely
densely pubemlent overall; seeds (not seen quite ripe)
±7-7.5 x 5.5 m m , the U-shaped pleurogram sharply
engraved.
In drought-deciduous submontane woodland,
980-1300 m, known only from the South Yungas of
La Paz, Bolivia, at points 26, 30, and 52 k m from
Chulumani on the road to Asunta. — FL VI-VIII.
In the context of the Bolivian flora and the sect. An
drocallis, C. chulumania is quickly identified by a
leaf-formula of i/7-9, softly pilosulous leaflets, rela
tively short peduncles, and deep red tassel of fila
ments. The similar C. mollissima of northern P e m has
rather fewer (4-7) pairs of leaflets per pinna and larger
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 87
blades about 2-4 c m long, prevailingly longer pedun
cles 5-20 (not 2-6) c m long, and white (not crimson)
filament-tassel. The largely Hylaean C. surinamensis
can have similar leaf-formula, but is not known from
Bolivia and has no pleurogram on the seed-coat.
At kilometer 30 on the Chulumani-Asunta road,
Beck collected both typical C. chulumania (no.
72766, N Y ) and an anomalous plant (no. 72762, N Y )
different in longer petioles (to 10-16 m m ) , leaflets to
10 pairs per pinna, and facially glabrous (but ciliate)
leaflets; its status is not determined, but it could be
either juvenile or a shade-form.
45. Calliandra carcerea Standley & Steyermark,
Publ. Field Mus. Nat. Hist., Bot. Ser. 23(4): 161.
1944. — "Guatemala: Dept. El Progreso ... between
Calera and middle slopes of Volcan Siglo, alt.
2,000-2,200 meters, January 20, 1942, Julian A.
Stevermark 42985." — Holotypus (mounted on 2
sheets), F! = F Neg. 53831, 53832.
Arborescent shrubs of unrecorded stature with
dense hard wood, the terete gray virgate long-shoots
hirsutulous when young but early glabrate, the lf-axes
and peduncles more densely gray-hirsutulous but the
firm, prominently venulose lfts facially glabrous lus
trous, ciholate, the peduncles arising singly from con
densed axillary brachyblasts; phyllotaxy distichous.
Stipules narrowly triangular ±1.2-2.5 m m , weakly
striate, persistent. Lf-formula i/7-10; petioles 6-17
m m , at middle 0.7-1.1 m m diam; rachis of longer
pinnae 4.5-6.5 cm, the longer interfoholar segments
5-9 m m ; lft-pulvinules ±0.45-0.6 x 0.5 m m , cross-
wrinkled; lfts proximally decrescent, inequilaterally
ovate or elliptic-ovate from semicordate base, ob
tusely or acutely deltate at apex, the distal ones ±11-
22 x 5-8 m m , 2.2-2.8 times as long as wide; venation
pinnate and obscurely palmate-pinnate, one weak
posterior primary nerve in some lfts produced ± to
mid-blade. Peduncles 15-23 m m , apparently ebract
eate; capitula ±12-20-fld, the receptacle not more
than 1.5 m m diam, the fls homomorphic; floral bracts
linear-attenuate ±1.3-2.4 m m , tardily deciduous;
pedicels ±0.3 m m ; perianth glabrous except for few
loose hairs about orifice of calyx and tip of corolla-
lobes, the calyx-tube weakly striate, the corolla not;
calyx campanulate ±2.5-2.8 x 1.4 m m , the triangular
obtuse teeth ±0.5 m m ; corolla ±6.5 m m ; androecium
±20-merous, vivid red, ±20 m m , the tube 9-10 m m ,
distinctly exserted. Pods in profile linear-oblanceo-
late, straight, including attenuate base 9-11 c m x 8-9
m m , 5-6-seeded, the sutural keels in dorsal view ±1.5
m m wide, thinly hirsute, the valves leathery, glabrous,
micropapillate; dehiscence and seeds not known.
In unreported habitat, known by one collection from
quebradas of Volcan Siglo at ±2100 m in Sa. de las
Minas, depto. El Progreso, Guatemala. — Fl. XII-I.
— Tamarindo de montaha.
In the protologue C. carcerea was described as re
sembling Antillean C. purpurea, but more densely
pubescent. This is not universally true, and it is likely
that C. carcerea, when better known, will prove to be
an outlying disjunct form of C. purpurea sens, lat.,
different if at all in shghtiy longer pinnae and slightly
more numerous leaflets.
The epithet carcerea (of prison) is unexplained. I
conjecture it may be a misreading of calcarea (of
limestone) and allude to the town of Calera (lime pit)
near which the species was discovered.
46. Calliandra pityophila Barneby, sp. nov, fruticu-
losa microphyllidia vix semimetralis, parcissime
puberula, foliorum omnium pinnis 1-jugis foliolisque
11-19-jugis, necnon flosculis saturate sanguineis ut
videtur C. colimae affinis, sed foliolis magis numer-
osis 11-19 (nee 6-9)-jugis usque pinnisque exacte
geminatis (nee 1-2-jugis) diversa. " M E X I C O .
Guerrero, distr. Mina [= mun. Ajuchitlan]: Chilcay-
ote, 1500 m, 7 Jan 1939 (fl), G. B. Hinton 143771'
— Holotypus, NY.
Slender microphyllidious shrublets to 5 dm tall
with erect terete lenticellate stems, appearing glabrous
but the young branchlets, lf-axes, and peduncles thinly
minutely appressed-pilosulous, the moderately bicol
ored lfts brown-olivaceous when dry, dull and gla
brous on both faces, a little paler beneath, thinly
appressed-ciliolate, the few-flowered capitula arising
singly or geminate on subfiliform peduncle from low
est nodes of shortly caulescent brachyblasts; phyl
lotaxy distichous. Stipules firm, narrowly triangular or
subulate ±1-2.4 m m , evenulose dorsally, tardily
deciduous. Lf-formula i/11-19; lf-stks 2.5-7 x 0.3-0.6
m m ; pinna-rachises 2-A cm, the longer interfoholar
segments 1.4-3.3 m m ; lft-pulvinules 0.2-0.4 x 0.3-
0.5 m m , not wrinkled; lfts a little decrescent at each
end of rachis, otherwise subequiform, the blades
oblong-elliptic from semicordate base, obtuse apicu-
late, those near mid-rachis 4.5-8.2 x 1.5-2.7 m m ,
2.4-3.1 times as long as wide; venation faint, the
midrib displaced to divide blade ±1:1.5-2, finely 2-3-
branched on each side, the inner of 2(-3) posterior pri
mary nerves scarcely attaining mid-blade. Peduncles
±2-3 cm, ebracteate; capitula 4-9-fld, the receptacle
88 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
0.5-1 m m diam, one terminal fl sometimes broader
than the rest but hardly longer and its androecium not
differentiated; pedicels 0.7-1 x 0.4 mm; perianth 5-
merous, almost glabrous; calyx of peripheral fls 1-
1.15 x 1.3 m m , weakly 5-nerved, the deltate-ovate
teeth 0.2-0.3 mm; corolla 6 mm, the ovate lobes ±1.4
m m (one pair sometimes united almost to tip); andro
ecium 14—15-merous, red throughout, 26 mm, the
tube ±5 mm, the stemonozone ±1 mm; intrastaminal
nectary 0.6 m m ; ovary substipitate, glabrous at anthe
sis. Pod unknown.
In pine forest near 1500 m, known only by the
type-collection from the n. slope of Sa. Madre del Sur
near 17°40'N, 10°38'W in Guerrero, Mexico. — Fl.
XII-II(-?).
This is one of several, as yet poorly known calhan
dras of low slender growth found in pine forest of
tropical Mexico, but is the only one of them with only
one pair of pinnae per leaf. It has no obvious close
kindred and is arbitrarily compared in the diagnosis
with C. colimae, which differs in leaf-formula.
47. Calliandra colimae Barneby, sp. nov, C. hirsu-
tae et forsan propius ut videtur foliorum pinnis 1-2
(nee 4-13)-jugis C. pityophilae affinis, ab hac prae-
cipue foholis paucis 6-9 (nee 11—19)-jugis usque
diversa. — MEXICO. Colima: mountain summits
near pass ca. 11 miles s.-s.-w. of Colima on Man-
zanillo road, 18 Jul 1957 (fl), 7?. McVaugh 15550.
— Holotypus, NY.
Slender, repeatedly branched, microphyllidious
shrubs ±1 m tall with terete gray glabrate annotinous
and older stems, the young stems, lf-axes, and dorsal
face of lfts subappressed-pilosulous with fine straight
lustrous hairs to 0.65-1 mm, the lfts bicolored, dull
dark brown-olivaceous on upper face, paler dull
beneath, the capitula of crimson-orange fls arising
singly on fihform peduncle from 1-2 nodes of at least
shortly extended lateral branchlets; phyllotaxy dis
tichous. Stipules firm, deltate to lanceolate ±1-2.5
m m , 1-nerved or externally nerveless, persistent. Lf-
formula i(—ii)/6—9, the lvs with 2 pairs of pinnae few
and scattered; lf-stks 3-10 mm, the petiole 3-7 x
0.3-0.4 m m , the one interpinnal segment (seldom
present) to 3 mm; rachis of longer pinnae ±14—19
m m , the longer interfoholar segments ±1.7-2.4 mm;
lft-pulvinules 0.3 x 0.3 mm, not wrinkled; lfts a trifle
smaller at each end of rachis, in outline oblong-
elliptic from broadly shortly semicordate base, trian-
gular-apiculate, those near mid-rachis 6-7.4 x
1.7-2.5 m m , 2.4-3 times as long as wide; venation
palmate-pinnate, the slender midrib displaced to
divide blade 1:1.3-1.6, faintly 2-3-branched from
middle upward, the inner of 2-3 posterior primary
nerves scarcely attaining mid-blade, the whole vena
tion pallid and weakly prominulous dorsally, im
mersed or nearly so on upper face. Peduncles ±2-2.5
cm, strigulose and remotely granular, either ebract
eate or 1-bracteate above middle; capitula 11-18-fld,
the receptacle 1-1.5 mm; bracts subulate obtuse 0.6-
1 mm, deciduous; fls homomorphic, the perianth
either 4- or 5-merous, the calyx glabrous or almost
so, the corolla thinly appressed-pilose; pedicels
almost 0 to 0.6 x 0.6 mm; calyx campanulate 1.5-1.8
x 1.2-1.5 mm, delicately 5-nerved, the triangular or
deltate teeth 0.3-0.9 mm; corolla 6-7 mm, apparently
red overall, the lobes separated by very unequal si
nuses 0.3-1 m m deep; androecium 14— 15-merous,
±29 mm, the tube 5-7 mm, the stemonozone 1.3-1.6
mm, the tassel crimson-orange overall; intrastaminal
nectary 0.4—0.5 m m tall; ovary at anthesis glabrous,
incipiently stipitate. Pod not seen.
On grassy mountain summits in deciduous wood
land near 500 m, known only from the type-locality
in Colima, Mexico. — Fl. VII-VIII.
McVaugh (1987: 161) interpreted this attractive
small Calliandra as a variant of the polymorphic C.
hirsuta, and there is little doubt that the relationship
between them, expressed in the similar stature, in
plane leaflet-pulvinules, and in form and color of the
flowers is a real one. The extremely reduced leaf-
formula and fewer stamens are, however, abmptly
different. In the geminate or randomly bijugate pin
nae C. colimae is more like C. pityophila, but differs
in notably few leaflets.
48. Calliandra hintonii Barneby, sp. nov, inter
species macro- et microphyllidias ambigens sed
affinitatis verae ignotae, his signis recognoscenda:
foliorum pinnae 1-jugae et foliola cujusque pinnae
4-6-juga; foliola (ultimis majoribus exceptis)
14-19 x 4.5-8 mm; flores in capitulo homomorphi;
calyx fere 2 mm, corolla usque 7 m m longa; an
droecium ±16-merum vivide ut videtur rubrum;
statura fruticosa arborescens 3-metralis et ultra. —
MEXICO. Estado de Mexico, distr. Temascaltepec:
Nanchititla, 6-9-1933 (fl), G. B. Hinton 4099. —
Holotypus, NY.
Shrubs 3 m with slender terete branchlets, the new
stems and lf-stks densely gray-pubemlent with in
curved hairs to 0.4 m m but the relatively ample, ven
ulose lfts facially (sub)glabrous ciholate, dark brown-
olivaceous sublustrous above, dull pale brown beneath,
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 89
the capitula of glabrous fls borne singly on loosely
thatched lateral brachyblasts; phyllotaxy distichous.
Stipules subtending primary lvs linear-lanceolate ± 2 ^
x 0.4-0.6 m m , weakly ±5-nerved, becoming papery,
those of thatched brachyblasts broader and shorter. Lf-
formula i/4-6; lf-stks ±5-8 m m , at middle 0.5-0.8 m m
diam; pinna-rachises (1.5-)2-4.5 cm, the narrow ven
tral groove interrupted at insertion of lfts, the longer
interfoholar segments 5-8 m m ; lft-pulvinules 0.4-0.5
x 0.6-0.8 m m , indistinctly cross-wrinkled; lfts distally
accrescent, the blades oblong-, ovate- or incipiently
rhombic-elliptic from broadly obtusely semicordate
base, abmptly deltate-apiculate at rounded or shal
lowly emarginate apex, the penultimate pair 14-19 x
4.5-8 m m , 2.3-3 times as long as wide, the terminal
pair often somewhat longer; venation palmate-pinnate,
the midrib scarcely excentric, straight or almost so,
4—6-branched on anterior side, the inner of 3 posterior
primary nerves incurved-ascending beyond mid-blade,
the outer ones much shorter, all these together with ter
tiary venules pallid and slenderly prominulous on both
faces. Peduncles 2-2.5 cm, ebracteate; capitula ±12-
16-fld, the receptacle 1-1.5 m m diam; bracts ovate or
subulate 0.4—0.8 m m , microciliolate, persistent; fls
homomorphic, the 5-merous perianth glabrous except
for microscopic cihola, the calyx weakly ±15-striate,
the corolla not so; pedicels 0.2-0.6 x 0.3-0.5 m m ,
essentially 0 in furthest fls; calyx campanulate 1.6-2 x
1-1.6 m m , the depressed-deltate teeth 0.2-0.3 m m ;
corolla 6.6-7 m m , the ovate, broadly obtuse lobes to 2
m m ; androecium 16-merous, 16-17.5 m m , the tube ±5
m m , the stemonozone 1.1 m m , the tassel (not seen
fresh) apparently red; ovary at anthesis glabrous; in
trastaminal nectary (of 1 bisexual fl dissected) 0.6 m m
tall. Pod not known.
In a wet barranca, probably in the oak-pine belt
near or above 1100 m, known only from the type-lo
cality at 15°53'N, 100°28'W in district of Temascal-
tepec, state of Mexico. — Fl. VI, and probably in
other months. — Cabello de angel, a generic name
for red-stamened Calliandra in Mexico.
The type-collection of C. hintonii has now lain for
sixty years among unidentified calhandras, unmatched
by new material and anonymous. It represents an un
described species of uncertain affinity, characterized
by the low leaf-formula of i/4-6, leaflets of moderate
size, and a red 16-merous androecium. In technical
characters it approaches the Guatemalan C. carcerea,
but has fewer leaflets per pinna.
49. Calliandra conferta Bentham in A. Gray, PL
Wright. 1: 63. 1852. — "[C. Wright] 166, 167.. .
Hills at the head of the San Felipe, in flower; and on
Zacate Creek, July, in fmit; also on the Rio Grande,
Texas." — Syntypi (Isely, 1972: 276), Wright 166
and 767, NY!; isosyntypi, K! = K. Neg. 75575. —
Feuilleea texana O. Kuntze, Revis. Gen. PL 1: 187.
1891 (non F. conferta (Bentham) O. Kuntze). —
Mistakenly equated with Anneslia eriophylla by Brit
ton and Rose (1928: 59). FIG. 8
C. conferta sensu A. Gray, PL Wright. 2: 53.1853; Bentham,
1875: 546; Turner, 1959: 32, map 4; Isely, 1972: 275; 1973: 78, map 18; Correll & Johnston, 1979: 770.
Intricately branched, mounded or straggling, micro
phyll shrubs 1-3 dm, the stiffly flexuous branches
brown or blanched in age, the new branchlets gray-
pubemlent or -pilosulous, glabrescent, the lf-axes, pe
duncles and dorsal face of the imbricate lfts thinly
strigulose or ascending-pilosulous with fine lustrous
hairs to 0.2-0.7 m m , the small firm lfts glabrous on
upper face, the few-fid capitula solitary or less often
geminate in axils of contemporary lvs; phyllotaxy dis
tichous. Stipules firm, linear-lanceolate or narrowly
triangular 1-2.5 x 0.2-0.5 m m , l-6(-8)-nerved dor
sally, tardily deciduous by weathering. Lf-formula i/8-
12; lf-stks 0.6-2.8 m m , at middle 0.25-0.5 m m diam,
shallowly grooved ventrally; rachis of longer pinnae
(5-)6-12.5 m m , the longer interfoholar segments
0.45-1.2 m m ; lft-pulvinules 0.1-0.2 m m diam, not
wrinkled; lfts often decrescent near top of rachis,
otherwise subequilong, the blades linear, linear-
lanceolate or ovate-elhptic from shallowly auriculate
base, deltately acute or apiculate, those near mid-
rachis (2.3-)3-5.5 x 0.8-1.4 m m , (2.6-)3.4-4.6 times
as long as wide; venation variable in strength, usually
immersed on upper face and obtusely prominulous
dorsally, the straight midrib displaced to divide blade
1:2-3, either simple or faintly 1-2-branched, the inner
posterior primary nerve produced to or commonly be
yond mid-blade. Peduncles 2.5-20 m m , usually
ebracteate; capitula 2-8-fld, the receptacle not over 1
m m ; bracts ovate or subulate 0.45-1 m m ; pedicels 0
or obscure, not over 0.2 m m ; perianth 5-merous,
loosely pilosulous overall to glabrous except at apex;
calyx campanulate, obtusely 5-nerved, 1-1.8 x
1.1-1.8 m m , the deltate-ovate teeth 0.35-0.6 m m ;
corolla narrowly campanulate (3.5-)3.8-5.2 m m , the
tube 5- or weakly 10-15-nerved, the lobes 0.9-1.7
m m ; androecium 20-28(-30)-merous, 7.5-14 m m ,
the tube 1.8-2.1 m m , the stemonozone 0.6-1.1 m m ,
the tassel pale pink or whitish; ovary glabrous at an
thesis, silky-strigulose following fertilization; intrast
aminal nectary 0. Pods erect, in profile (2-)2.6-4 x
90 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
FIG. 8. Calliandra conferta Bentham.
0.4-0.6 cm, the sutural ribs in dorsal view 0.9-1.2 m m
wide, the stiffly papery, when ripe stramineous valves
densely white-silky-strigulose, low-convex over (1-)
2-5(-6) seeds, the ribs similarly strigulose or glabres
cent; seeds plumply ovoid-discoid ±4-4.3 x 3-3.6
m m , the testa pale brown or putty-colored flecked with
darker brown, the small pleurogram dehcately incised.
Bare stony hilltops and thinly vegetated slopes or
rocky flats, mostly on caliche soils, 30-1200 m, lo
cally plentiful, scattered over s.-centr. and s.-w. Texas
(Trans-Pecos, Edwards Plateau and South Texas
Plains n.-ward to Travis County), crossing the Rio
Grande very locally into n. Coahuila (Sa. de Jardm),
Mexico. — M a p 26. — FL IV-VI, sporadically later,
following rains.
Calliandra conferta closely resembles C. eriophylla
in most resepcts, and is easily mistaken for drought-
stressed aspects of C. eriophylla which have lost the
primary leaves and retain only the simpler brachy
blast leaves, many or most of which may consist of
only one pair of pinnae. Even these, however, have a
longer primary axis. Further differential characters of
C. conferta are the shorter androecium (7.5-14, not
16-27 m m long), paler pink or whitish filaments and
on the average shorter, thinner-textured pods with
narrower sutural ribs. A n intrastaminal nectary is
found at the base of almost every flower of C. erio
phylla, but has not been seen in any flower of C. con
ferta. These relatively weak differences, in context of
an almost perfectly vicariant dispersal, would seem to
justify the reduction of C. conferta to varietal status,
as evidently contemplated (in annotation of speci
mens, N Y ) by Isely. The complete subordination of
C. conferta, in North American Flora, is, on the other
hand, clearly mistaken.
50. Calliandra brevipes Bentham, J. Bot. London 2:
140. 1840. — "Brasilia: Pohl." — Holotypus, Pohl
1455, K (2 sheets, one ex hb. Hook. = IPA Neg. 7472
= K Neg. 12040 = N Y Neg. 1960, the second ex hb.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 91
FIG. 9. Calliandra brevipes Bentham. Reproduced from Bentham in Matrius, Fl. bras. 15(2): t. 107. 1876.
Benth. = K Neg. 12041)\. — Feuilleea brevipes O.
Kuntze, Revis. Gen. PL 1: 187. 1891. Fig. 9
C. selloi sensu Macbride, Contrib. Gray Herb., n.s. 59: 5.
1919, exclus. basionym. Acacia selloi Sprengel, Syst. Veg.
3: 137. 1826, — "Brasil. Sello!' — No typus known to survive, but, contrary to Macbride, the specified lf-formula of
i/12 is far outside that known for C. brevipes to which Ben
tham (1876: 416) only doubtfully reduced Acacia selloi
Sprengel. (?) C. yucunensis N. Mattos, Loefgrenia 71: 3, fig. 1. 1977.
— "[Brasil.] Estado do Rio Grande do Sul: Tenente
Portela, Parque Estadual do Turvo, no Sal to Yucuna, 10-1-
1977 (fl), J. Mattos 16411 & N. Mattos." — Holotypus,
IPRN n.v. C brevipes sensu Bentham, 1844: 144; 1875: 416, t. 107;
Glaziou, 1905: 187 ("5787" = 7587\).
C. selloi sensu Burkart, 1952: 111; 1979: 94, est. 11 (copied
from Bentham, 1876); Parodi, Encicl. Argent. Agri. Jard.
455: fig. 126A, 1959; Hoc, 1992: 211, fig. 4 + map 2.
Freely, stiffly branched microphyllidious shrubs
±1.5-2 m, with fuscous annotinous and older (defoli
ate) stems, the young branchlets, lf-stks and pedun
cles minutely, sometimes densely brown-puberulent,
the crowded bicolored lfts glabrous facially, some
times minutely ciholate, lustrously dark green above,
paler dull beneath, the dense capitula solitary, pedun
culate in lf-axils of annotinous long-shoots or (later)
of axillary foliate short-shoots; phyllotaxy distichous.
Stipules triangular-lanceolate or deltate 0.6-2 m m ,
dorsally glabrous, 1-7-nerved, becoming dry persis
tent. Lf-formula i/(20-)23^5; lf-stks slender (1-)
1.5^-.5(-5) x 0.3-0.45 m m , the appendage ovate-
triangular 0.4—1 m m ; rachis of longer pinnae (12-)
15—35(—38) m m , the longer interfoholar segments
0.3-1 (-1.2) m m ; lft-pulvinules 0.1-0.2 m m , the
blade sessile against rachis; lfts decrescent at each
end of rachis, otherwise subequilong, in outline linear
or linear-oblong from obhquely subtmncate base,
straight or gently incurved, obtuse or subapiculate,
the larger ones 3.5-5.5(-6.5) x 0.4-1.4(-l.6) m m ,
(3.2-)4—6 times as long as wide; midrib subcentric,
commonly (in superficial view) unbranched, other
venulation immersed or only faintly perceptible.
Peduncles (4—)5-14(-22) m m , 1-bracteate beyond
middle; capitula (technically umbelliform, but often
obscurely so) 7-12-fld, the claviform or subglobose
receptacle 1-2 m m ; bracts ovate-triangular 0.3-0.9
m m , incurved, persistent; fls of each capitulum ordi
narily heteromorphic, the peripheral ones shortly
pedicellate, the terminal one sessile, broader but no
longer than the rest and its androecium not modified;
P E R I P H E R A L FLS: pedicel 0.1-0.7 m m ; perianth
glabrous or very thinly minutely pubemlent, the
pallid calyx ±15-striate, the pink-reddish corolla
evenulose or almost so; calyx campanulate or
turbinate-campanulate 0.9-1.8 x 1-1.6 m m , the broad
subincurved teeth 0.2-0.5 m m ; corolla 4—6.6 m m , the
ovate lobes 1.4—2.3 m m ; androecium 18-25-merous,
(22-)25-32 m m , the stemonozone 0.35-0.6 m m , the
tube 2.2-4 m m , the tassel whitish proximally, pink-
or crimson-tipped (exceptionally all white); ovary
glabrous; style of peripheral fls linear tmncate, that of
terminal fl(s) dilate at apex to 0.22-0.3 m m . Pods in
profile 4.5-6 x 0.7-0.8 cm, the coarsely framed
valves stiffly coriaceous, glabrous, weakly cross-
venulose; seeds unknown.
Native on rocky stream banks and at margin of
gallery woodland, 90-1200 m, locally c o m m o n in e.
Brazil s.-ward from centr. Minas Gerais to Umguay,
w. into n.-e. Argentina (Misiones) and adj. Paraguay;
one record from Salvador, Bahia; cultivated in its
92 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
native range and far afield. — Map 27. — Fl. in
Brazil mostly XII-I, IV-VII, but in cultivation spo
radically through the year. — Munduruva (Brazil);
sarandi, esponginha (Argentina).
Calliandra brevipes, commonly passing under the
misnomer C. selloi, might stand as the prototype of
the conjugately pinnate, plurifoholate species of sect.
Androcallis. It was cultivated at K e w in 1850 (K,
N Y ) , on Martinique in 1857 (Belanger 297, 1046,
both G ) , at Hortus Bogoriensis in 1903 (NY) and is
currently planted in gardens of Dominican Republic.
Its many small, almost veinless leaflets and its star-
bursts of bicolored filaments arising from parallel
ranks of brachyblasts are unquestionably ornamental.
For purposes of nomenclature I have ignored Aca
cia selloi Sprengel, which Macbride interpreted as
the basionym of Calliandra selloi. N o typus of A. sel
loi is known to survive, and the leaflet number
ascribed to the species is much lower than seen in any
collection of C. brevipes. The typus of C. yucunensis
has not been seen, but there is nothing in the proto-
logue incompatible with other examples of C. bre
vipes from Rio Grande do Sul.
51. Calliandra staminea (Thunberg) Barneby, comb.
nov. Mimosa staminea Thunberg, PL Bras. Decas
Secunda 22. 1818. — "Crescit ad villam Ricam [=
Ouro Preto, Minas Gerais]." — Holotypus, col
lected by G. W . Freyreiss in 1814-1815, U P S (hb.
Thunberg. 24189 ex hb. Westin.), seen in Hb.
Thunb. Microfiche 7075!.
C. cinerea Taubert, Flora 75: 70. 1892. — "Habitat in Brasilia loco non indicato [supplied by Glaziou, 1905: 188:
'Serra de Caraca, pres de Alegria, MINAS'], Glaziou n.
12639." — Holotypus, *B = F Neg. 7236!; isotypi, K! = N Y
Neg. 1984, P (2 sheets)!.
Microphyllidious shrubs of unknown potential
stature, with stout terete glabrate long-shoots and
shortly caulescent short-shoots, the young stems, lf-
axes, and peduncles thinly pilosulous with fine loose
hairs to ±0.6 m m , the crowded conjugate-pinnate lvs
bicolored, when dry dark above, pallid beneath, dull
and glabrous on both faces, thinly ascending-ciliolate,
the umbelliform capitula arising singly from either fo
liate or efoliate nodes of developing brachyblasts,
nestling in foliage; phyllotaxy distichous. Stipules
papery, striately nerved, those of primary lvs lance-
attenuate ±2.5-6 x 1-1.5 m m , those of brachyblasts
similar but a little shorter. Lf-formula i/26-30; lf-stk of
longer lvs 4-7 m m , at middle 0.4-0.45 m m diam;
rachis of longer pinnae 30-38 m m , the longer interfo
holar segments ±1 m m ; lft-pulvinules 0.2-0.3 x 0.2
m m , faintly cross-wrinkled; lfts backwardly decres
cent toward base of rachis, then subequilong, the
blades linear-oblong from obtusangulate base,
deltately subacute, the longer ones 7-9 x 1.5-1.8 m m ,
4.4—5.3 times as long as wide; primary venation of 2
nerves, the scarcely displaced midrib giving rise on
each side to 4—6 weak, divaricate or obliquely reflexed
secondary venules expiring short of the plane margin,
the posterior primary nerve very short, all nerves im
mersed on upper face of blade, faintly prominulous
beneath. Peduncles 7-13 m m , bracteate well above
middle, the bract 2-2.4 m m , persistent; capitula 10-
16-fld, the clavate receptacle ±1-1.5 x 1-2 m m , the fls
heteromorphic, the peripheral ones shortly pedicel
late, the terminal one (sometimes abortive) subsessile,
longer, and with modified androecium; floral bracts
rudimentary caducous or 0; P E R I P H E R A L FLS:
pedicel 1-1.6 x 0.4 m m ; perianth (either 4- or 5-mer
ous) of thin texture, remotely pubemlent, especially
toward tip of teeth and lobes; calyx campanulate 2-2.8
x 1.7-1.9 m m , faintly (12-) 15-nerved, the broad ob
tuse teeth 0.3-04 m m ; corolla 6-6.5 m m , the ovate
lobes 1.5-1.8 m m ; androecium 12-18-merous, 3-3.3
cm, described by Glaziou as pink, stemonozone
scarcely 1.5 m m , tube 3-4.8 m m ; ovary (sometimes
abortive) glabrous at anthesis; T E R M I N A L FL: calyx
±3.6 x 3.5 m m ; corolla stoutly cylindric 9.5 m m , the
erect lobes ±2 m m ; androecium ±22-merous, 3 cm,
the tube 16 m m , a httle dilated at orifice; a lobed
intrastaminal disc 0.3 m m . Pods (one seen, immature)
±3 x 0.8 cm, the strong sutural keels 3(-?) m m wide
in dorsal view, glabrate, the recessed valves densely
pilosulous and strongly transverse-venulose.
In unrecorded habitat, to be expected on open hill
sides or on outcrops near and above 1000 m, appar
ently very local in Sa. do Espinhaco of s.-e. Minas
Gerais in lat. 20°00'-30'S near 43°30'W (Sa. da
Caraca; Ouro Preto). — Fl. IX-X(-?).
This rare species, not collected in the past century,
has much in common with C. brevipes, but it seems
nevertheless distinct in larger stipules and leaflets, in
deeper calyx, in tmmpetlike androecium of the termi
nal flower of capitula, and in densely pilosulous, not
glabrous pod. The type-collections of Mimosa sta
minea and Calliandra cinerea were collected in the
same segment of Sa. do Espinhaco, a region noted for
local endemism, and their protologues are in close
agreement. Whatever the status of the species, these
two names are sure to remain taxonomic synonyms.
52. Calliandra sessilis Bentham, J. Bot. (Hooker) 2:
141. 1840. — "[BRAZIL. Bahia:] Sierra Acurua,
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 93
MAP 27. Distribution of Calliandra brevipes Bentham and C. spinosa Ducke in eastern South America.
Blanchet, n. 2816." — Holotypus, K (hb. Benth.)! = C. axillaris Bentham, Trans. Linn. Soc. London 30: 546.
NY Neg. 1958; isotypi, fB = F Neg. 7225 BM!, G!, 1875' & in Martius, Fl. Bras. 15(2): 415. 1876. — "Habi-
K (hb. Hook.)!, NY!. - Feuilleea mutica O. JlT*^ r r \ r> n dii iq*iqow zr •#• (hb. Benth.)! = NY Neg. 7959; isotypi, ̂ B = F Neg. 7228!, Kuntze, Revis. Gen. PL 1: 186. 1891 (non £ sessilis GH!, NY (the locality given as "Igreja Velha")!, OXF!. — (Vellozo) O. Kuntze). Feuilleea axillaris O. Kuntze, Revis. Gen. PL 1:187. 1891.
94 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
C. tocantina Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 71.
1922. — "[BRAZIL. Para:]... prope stationem Arumateua
viae ferreae alcobacensis in regione fluvii Tocantins
cataractarum inferiorum, l[egit] A. Ducke 3-1-1915, n.
15.607." — Holotypus, RB n.v.; isotypus, M G = photo + clastotypus, F 6023191.
C. sessilis sensu Bentham, 1844: 103; 1875: 545; 1876: 414; Lewis, 1987: 176.
C. axillaris sensu Harley & Simmons, 1986: 114; Lewis, 1987: 171, pi. 10, fig. B.
C. tocantina sensu Ducke, 1949: 50.
Dimorphic in habit and stature, most commonly a)
suffmticose, with diffuse-ascending stems ±2-5 d m
dying back annually to a palhd creeping rhizome
(hence forming dwarf thickets), less often (but sym-
patrically) b) a spindly shmb 1—2(—3) m tall, with old
defohate stems regenerating (in sheltered places, at
edge of woods) through two or more seasons, the ses
sile or subsessile, ± hemispherical capitula borne
singly or two together at top of short-shoots axillary to
primary lvs of long-shoots, the short-shoots often
densely thatched with bifariously imbricate stipules
but sometimes reduced to as few as two nodes, the
capitulum then appearing axillary to a primary If but in
reahty separated from the leaf by two or more bracts;
vesture variable, the stems often glabrous, sometimes
pubemlent or thinly hirsutulous, the lf-axes always
ventrally pubemlent or hisutulous, the firm plane bi
colored, ventrally lustrous lfts always glabrous on
upper face and nearly so on lower face, their margin
either smooth or ciliate with stiff spreading-ascending
hairs to 0.2-0.7 m m . Lf-formula i/(17—)21—39; lf-spurs
1—2.5(—3) m m ; stipules lanceolate, narrowly ovate, or
subhnear 1.5-9 x 0.7-2.2 m m , striate, commonly gla
brous dorsally but occasionally hirsutulous, some
times ciliate; lf-stks including swollen base and apex
1-7 m m , the terminal appendage 1.5-5 m m ; rachis of
longer pinnae (3.5—)4—8(—10) cm, the longer inter
foholar segments (1 —) 1.3—2.6(—3) m m ; lft-pulvinules
0.2-0.4 m m , nearly twice as wide; lfts a trifle shorter
at each end of rachis, elsewhere equilong or nearly so,
in outhne lanceolate (narrow-ovate) from shallowly
semicordate base, abmptly acuminate-mucronate,
7_13(_14) x 1.7-3 m m , (3-)3.4-4.5 times as long as
wide; venation palmate-pinnate, the shghtiy displaced
midrib giving rise on each side to ±7-11 short brochi-
dodrome secondary nerves, the 1-2 posterior and
sometimes one anterior primary nerve not attaining
mid-blade, the venulation prominulous on both faces.
Peduncles <2 m m , often obsolete; capitula ±9-24-fld,
the fls sessile or raised on a sohd pediment to 0.2 m m ,
almost always heteromorphic, the androecial tube of
the peripheral ones included or exserted to 2.5(-3)
m m , 1-4 distal ones a trifle larger but scarcely broader,
their androecial tube exserted 4—9 m m ; bracts striate,
shorter than calyx; perianth sharply striate from base
to apex, often glabrous, less often white-hirsutulous
distally; P E R I P H E R A L FLS: calyx campanulate or
turbinate-campanulate, sometimes deeply so, 1.9-3.1 x
1.2-1.8 m m , the depressed-deltate, often unequal teeth
0.15-0.5 m m ; corolla (4.3-)4.5-6.8 m m , the ovate,
dorsally convex lobes 1.2-2.5 x 0.8-1.6 m m ; androe
cium (8-)10-16-merous, to 2.4—3.7 cm, the stemono
zone 0.8-1.5 m m , the tube 4—7.5(-9) m m , the tassel
pink or crimson; ovary glabrous; style fihform, the
stigma truncate-poriform; intrastaminal nectary 0. Pod
not seen.
Eastern Brazil in lat. 3°30/-18°S: best known from
Chapada Diamantina in Bahia and the upper S. Fran
cisco Basin in Minas Gerais, where found in campo
cerrado and campo mpestre, mostly between 600 and
1200 m; recorded once from the Una de Balsas in
s.-e. Maranhao, in chapadao at 300 m; once from
Piaui (Sa. da Lagoa, not located, not mapped) and
collected several times in the lower Tocantins Valley
near Tucurui Reservoir, on white sand campirana at
70 m. — M a p 28. — FL (XI-)I-III(-V).
Calliandra sessilis can be recognized, in the context
of sect. Androcallis, by unijugate pinnae, sessile or
subsessile capitula, and obhque rootstocks which give
rise to stoloniferous thickets. Flowering stems com
monly arise directly from the rhizome, but sometimes
persist as slender woody trunks. The relatively uncom
mon fruticose form has been found in Bahia (Harley
19978, N Y ) and in Para (Lisboa 1252, N Y ) , in both
cases close to the diffuse rhizomatous, functionally
herbaceous form. N o differences in leaf or in individ
ual flower are correlated with the variations in stature,
and the growth-forms seem to be no more than aspects
of one variable species. There is some variation among
specimens in length of the floriferous brachyblasts, C.
axillaris sens. str. being the state in which one or two
capitula are borne together on a very short or even ob
scure branchlet axillary to a coeval leaf, whereas C.
sessilis sens. str. is the state in which the capitula arise
from near apex of an older brachyblast thatched with
closely imbricate, defohate stipules. Harley recorded
the flowers of his no. 75975 as intensely fragrant.
53. Calliandra spinosa Ducke, Anais Acad. Brasil. Ci.
32(2): 289. 1959. — "In Brasihae boreali-orientalis
regione sicca 'Sertao' appellata locis nonnullis fre-
quens. Ceara: circa Caridade (mun. Caninde), A.
Ducke, Herb. Mus. Paraensis 2777." — Holotypus,
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 95
MAP 28. Distribution of Calliandra sessilis Bentham in eastern Brazil.
M G n.v; isotypus, RB 56691; paratypus e loco typico:
W. Franga, May 1958 (fl), NY!. FIG. 10
C. suberifera Rizzini, Rodriguesia 41: 175. 1976. — "Provenit in caatinga, Paulistana, Piaui, collegit D. P. Lima
173.307 (6-XI-1974)." — Holotypus, RB n.v.
C. brevipes auct. brasil. bor.-or., fide Ducke, I.e. 1959; non
Bentham.
Stiffly intricately branched, microphyll, drought-
deciduous shrubs and treelets attaining ±3 m but often
smaller, with sinuous terete long-shoots tapering at
apex into a stout vulnerant thorn, and at each primary
node a condensed, closely thatched, subacaulous 1- to
few-lvd brachyblast, the annotinous and older stems
blanched, becoming suberous, flaking, and shallowly
sulcate, the young stems, lf-axes and peduncles thinly
pilosulous with straight palhd and often some twisted
or granular orange-brown hairs, the narrow imbricate
plane lfts nearly concolorous but more lustrous on
ventral face, finely ciholate, the dense capitula soli
tary, subtended by elaminate stipules of brachyblasts;
phyllotaxy distichous. Stipules papery lustrous brown
ish, sharply striate, narrowly lanceolate or hnear 3-6
x 0.6-1.1 m m , often frayed or tattered in age but not
disarticulating. Lf-formula i/14-21; lf-stks beyond lf-
spur 0.5-2.5 m m ; rachis of pinnae 14—26 m m , the
longer interfoholar segments 0.4—1.4 m m ; lft-pulvin
ules 0.15-0.4 x 0.2-0.5 m m ; lfts decrescent near each
end of rachis, the blades linear from angulate base,
acute, those near mid-rachis 4-10 x 0.7-2 m m , 5-5.8
times as long as wide; venation essentially pinnate, the
straight midrib dividing blade 1:1.5-2, 5-8-branched
on each side, the secondary venules brochidodrome,
the posterior primary nerve not or scarcely longer, the
venulation delicately prominulous on both faces. Pe
duncles 4—13 m m , bracteate below middle, the bract
lanceolate ±1-1.5 m m ; capitula densely 25-30-fld,
the receptacle 1.5-2.5 m m ; floral bracts lanceolate
incurved 1-1.5 m m , persistent; fls heteromorphic, one
or more (sub)terminal ones broader and longer, all
subsessile, the pedicels not over 0.4 m m ; perianth
either 4- or 5-merous, glabrous, the calyx finely striate
but the corolla not so; P E R I P H E R A L FLS: calyx
campanulate or cuneate-campanulate (1.1—)1.4—2.2 x
(0.7-)1.2 m m , the obtuse, dorsally convex teeth
0.2-0.3 m m ; corolla 3.6-5.8 m m , the ovate lobes
0.8-1.2 m m ; androecium 10-20-merous, 18-24 m m ,
the stemonozone 0.5-0.9 m m , the tube 2-3 m m , the
tassel white-mbescent; nectary 0; ovary minutely stip-
itate, at anthesis glabrous; T E R M I N A L FL(S): calyx
2.3-2.6 x 2-2.5 m m ; corolla 5.5-7 m m ; androecium
22-36-merous, the tube 6.5-10 m m , the intrastaminal
nectary ±0.7 m m . Pod not seen, described by Ducke
(protologue) as 6 x 0.7 cm, glabrous.
In caatinga, at elevations not recorded, locally fre
quent within 3°30'-8°30'S and 37°-41°W in n.-e.
Brazil, from n.-e. Ceara. to centr. Pemambuco and
extreme s.-e. Piaui; and disjunct near 14°S, 4 1 ° W in
middle Contas Valley in Bahia (mun. Tanhacu). —
M a p 27. — Fl. intermittently after rains. — Umari
bravo; m a n bravo; espinheiro bronco.
In the context of sect. Androcallis and geminate
pinnae, C. spinosa is readily recognized by its xero-
morphic habit, stiff, early blanched, thom-tipped
branches, relatively numerous glabrous flowers, and
white but early rubescent androecium.
54. Calliandra duckei Barneby, sp. nov., hucusque
cum C. blanchetii, cui foliorum formula ac capitulis
unbelliformibus accedit, confusa, sed ab ea im
primis florum periphericorum perianthio subduplo
minori, calyce ±1.4 (nee 3^.3) m m et corolla 4
(nee 6.5-8) m m longis necnon flosculi terminalis
tubo androeciali longe exserto diversa. — BRAZIL.
96 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
FIG. 10. Calliandra spinosa Ducke.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 97
Pemambuco: Russinha, 4 Feb 1933 (fl), D. Bento
Pickel 3213. — Holotypus, GH.
C. blanchetii sensu Ducke, 1953: 51; non Bentham.
Stiffly awkwardly branched, microphyll, drought-
deciduous shrubs or treelets of unknown stature, the
new stems, lf-axes, and units of inflorescence thinly
pilosulous with extremely fine white ascending hairs
to ±0.5 m m , the lvs bicolored, the facially glabrous
ciholate lfts dull olivaceous above, paler beneath, the
umbelliform capitula borne single in the first efohate
axil of brachyblasts axillary to primary lvs. Stipules
of primary lvs (few seen) lance-attenuate ± 3 ^ mm,
striately 3-5-nerved, becoming dry fragile, those of
brachyblasts similar but smaller, closely imbricate.
Lf-formula i/20-24; petioles 3-4 x 0.2-0.4 mm;
rachis of pinnae ±1.5-2 cm, the longer interfoholar
segments 0.6-0.8 m m ; lft-pulvinules 0.2 mm; lfts
abmptly smaller at each end of rachis, otherwise
subequilong, the blades linear-oblong from bluntly
auriculate base, obtuse, nearly straight, the longer
ones ±3.6-4 x 0.9-1.1 mm; venation pinnate, the
midrib only a trifle displaced from mid-blade, divari
cately 5-6-branched on either side. Peduncles 5-18
m m , 1-bracteate well above middle, the papery bract
<1.5 m m , the receptacle 1-1.5 mm; floral bracts ±1
m m , deciduous; fls dimorphic, the peripheral ones
pedicellate, the central one sessile or nearly so, larger,
and its androecium trumpet-shaped, its tube far-
exserted; perianth of all fls 5-merous, thinly finely
pilosulous; PERIPHERAL FLS: pedicel 1-2.3 x 0.2-
0.3 m m ; calyx turbinate-campanulate ±1.4 mm, the
tube bluntly 5-angulate, the obtuse teeth 0.25 mm;
corolla 4 m m , the ovate lobes 1.7-2 mm; androecium
16-18-merous, ±17 m m , the stemonozone <1 mm,
the tube ±2.5 m m ; intrastaminal disc 0; TERMINAL
FL: calyx broadly campanulate ±2.5 x 2.5 mm; co
rolla 7.5 m m , the lobes 2.2 mm; androecium 18-mer-
ous, ±20 m m , the tube 14 mm, the intrastaminal disc
±0.6 m m ; ovary subsessile, distally pilosulous. Pod
unknown.
In unreported habitat, but to be expected in season
ally dry scrub thickets at low elevations, known at
present only from the type-locality at 8°09'S,
35°30'W in w. Pemambuco, Brazil. — FL I-H(-?).
Calliandra duckei has much in common with C.
blanchetii and could easily be mistaken for it in ab
sence of authentic material of the latter, which Ducke
may well not have seen. It differs from C. blanchetii
primarily in much shorter peripheral flowers; see
Latin diagnosis for measurements. Calliandra bre
vipes, which ranges from Salvador, Bahia, far south
ward, has similar leaves, but broad short stipules,
truly capitate, not umbelliform units of inflorescence,
and prominently striate-venulose calyx.
55. Calliandra blanchetii Bentham, London J. Bot.
3: 102 ("Blanched"). 1844. — "Sierra Jacobina,
Brazil, Blanchet, n. 2584." — Holotypus, K (hb.
Benth.)! = N Y Neg. 7957; isotypi, BM!, K (hb.
Hook.)!, NY!, OXF!. — Feuilleea blanchetii O.
Kuntze, Revis. Gen. PL 1: 187. 1891.
C. blanchetii sensu Bentham, 1875: 544 ("Blancheti"); 1876: 414; Renvoize, 1981: 67, fig. 1(4); Lewis, 1987: 171.
C. umbellifera sensu Lewis, 1987: 177; non Bentham.
Subshrubs or shrubs of unknown height, with de
foliate older stems and densely leafy annotinous
branches, the young stems and ventral face of lf-axes
pilosulous with fine weak whitish hairs <0.6 m m , the
lvs strongly bicolored, the small firm, facially gla
brous but microscopically ciliolate lfts brown-
olivaceous above, paler dull beneath, the umbelliform
capitula arising singly and geminate from efoliate
brachyblasts axillary to several distal primary lvs, and
beyond these forming a shortly exserted efohate
pseudoraceme, the fls glabrous except for microscop
ically silky-strigulose limb of corolla. Stipules of pri
mary lvs erect, narrowly lanceolate 4.5-8 x 0.9-1.4
m m , striately ±9-nerved, becoming dry, deciduous,
those of brachyblasts densely imbricate, similar in
form but much shorter. Lf-formula i/21—28(—32); lf-
stk of primary lvs, including the pulvinus but disre
garding the prominent lf-spur, 1.5-5 m m , at middle
0.3-0.5 m m diam, that of brachyblast lvs mostly
shorter and reflexed; pinna-rachis of primary lvs 17-
30(-33) m m , the longer interfoholar segments 0.6-
1.3 m m ; lfts a little decrescent at each end of rachis,
otherwise equilong, the blades linear-lanceolate from
bluntly auriculate base, deltately subacute, those near
mid-rachis 3.5-5.5 x 1-1.2 m m , 3.5-5 times as long
as wide; midrib simple, nearly straight, forwardly dis
placed to divide blade ±1:1.5-2, prominulous only
on dorsal face of 1ft, a very short and faint posterior
primary nerve sometimes perceptible. Peduncles
stout, compressed, 1.2-2.5 cm, bracteate above mid
dle, the lanceolate bract 1-1.5 m m , sometimes decid
uous; capitula ±10-20-fld, the receptacle, including a
sometimes well-defined terminal pedestal, 1.5-3 x
1.5-2 m m ; floral bracts ephemeral or 0; peripheral
pedicels 3-5.5 x 0.3-0.5 m m , the terminal one a httle
shorter; fl-buds broadly pyriform; perianth 4- or 5-
merous, pallid, externally nerveless; P E R I P H E R A L
98 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
FLS: calyx turbinate-campanulate 2-4.3 mm, gla
brous, the deltate-obovate obtuse teeth unequal, the
deepest sinus 0.5-2 m m ; corolla 6.3-8 mm, the
broadly ovate obtuse unequal lobes 2.2-3 mm; andro
ecium 16-40-merous, 2.8-5 cm, the thickened, exter
nally fuscous stemonozone 0.7-1.6 mm, the tube
±3.5 m m , the tassel pink; intrastaminal nectary 0;
T E R M I N A L FL: scarcely known, sometimes early
deciduous, a httle wider than peripheral ones but
scarcely longer, the androecium apparently little
modified; ovary of peripheral fls subsessile, glabrous
at anthesis. Pods (two seen) 6-6.5 x 0.75-0.8 cm, the
sutural ribs in dorsal view scarcely 2 m m wide, the
dark brown valves densely cross-venulose, micropu
berulent; seeds unknown.
In stony ground, in habitats and at elevations not
recorded, known only from interior Bahia, Brazil: at
Jacobina, near 11°S, 40°30/W; confluence of rios
Jacuipe and Paraguacu, 12°32'S, 39°05'W. — FL
VII-?.
Calliandra blanchetii is well characterized by the
combination of geminate pinnae and umbelliform ca
pitula arising from efohate axillary brachyblasts. Cal
liandra umbellifera, similar in the umbellate flowers,
has a second pair of pinnae in some or all leaves, and
capitate-glandular perianth.
56. Calliandra aeschynomenoides Bentham, Trans.
Linn. Soc. London 30: 546. 1875; & in Martius, Fl.
Bras. 15(2): 415. 1876. — ". . . in provincia Bahia:
Blanchet n. 3896 ... in herbario Martiano." —
Holotypus, BR! = K Neg. 19427; isotypi, BM!, G!,
GH!, P!. — Feuilleea aeschynomenoides O.
Kuntze, Revis. Gen. PL 1: 187. 1891.
C. aeschynomenoides sensu Renvoize, 1981: 71, fig. 2(13); Lewis, 1987: 170.
Slender ("esguio") shrubs with phant branches,
attaining 2 m, the young stems, lf-axes, and units
of inflorescence thinly loosely pilose or strigulose
with fine flexuous white hairs to ±0.7-1 mm, the
small, relatively simple lvs bicolored, the plane thin-
textured lfts dull olivaceous glabrous above, paler
pilose beneath, the capitula shortly pedunculate in
the lower lf-axils of shortly caulescent branchlets lat
eral to and coeval with homotinous long-shoots;
phyllotaxy distichous. Stipules lance-ovate (1.5-)
2.5-5.5 m m , early dry stramineous, striately nerved,
persistent. Lf-formula i(—iv)/5—12(—13); petioles
1.5-5 x 0.4-0.5 m m ; rachis of longer pinnae
12-28(-35) m m , the longer interfoholar segments
2-3 m m ; lft-pulvinules 0.2-0.4 mm; lfts accrescent
distally, the blades obliquely oblong-elliptic from
broadly shortly auriculate base, either obtuse or ob
tuse apiculate, the larger ones 6.5-10.5 x 2.3-4 m m ,
2.5-3 times as long as wide; venation of 3 primary
nerves, the gently incurved midrib forwardly dis
placed to divide blade ±1:2, 3-5-branched on each
side upward from mid-blade, the inner posterior
nerve incurved-ascending to near mid-blade, the
much shorter outer one subhorizontally divaricate,
the secondary nerves Y-forked within the margin,
scarcely or not camptodrome. Peduncles 2-5 cm, 1 -
bracteate close under first fl, the bract ovate ±2-3
mm, striate like stipules; capitula hemispherical
3-15-fld, the receptacle ±1 mm; floral bracts nar
rowly ovate 2-3 mm, faintly striate, tardily decidu
ous; fls homomorphic, sessile, the 5-merous perianth
thinly minutely pilosulous; calyx campanulate 2-2.2
x 1.2-1.5 mm, the tube faintly striate-nerved, the
teeth 0.5-0.8 mm; corolla 4.2-4.4 m m , the narrowly
ovate lobes 1.2-1.4 m m ; androecium 12-16-merous,
to 23 mm, the stemonozone 0.8 m m , the very slender
tube ±9 mm, the tassel united into fascicles, the ste
monozone not internally thickened, intrastaminal
nectary 0. Pod (one seen) narrowly oblanceolate 6
x 0.7 cm, 7-seeded, thinly finely pilosulous, the
sutures in dorsal view <2 m m wide, the leathery
valves bullately distended over seed-locules, finely
venulose; seeds unknown.
In caatinga on sandy substrates, known precisely
only from centr. Pemambuco (Ibimirim) and n.-e.
Bahia (Reserva Ecologica do Raso da Catarina). —
Fl. VI-?. — Forrageira.
Among the calhandras of eastern tropical Brazil
with geminate pinnae C. aeschynomenoides is no
table for relatively few and large leaflets. Its "flow
ers" have been described by collectors as either red or
pink. The species was described from specimens col
lected somewhere in Bahia by (or for) J. S. Blanchet,
before 1856, and not seen again until 1982, when re
discovered in lower Sao Francisco valley by L. S.
Guedes. The one record from Pemambuco (Ataide
587, COL) dates to 1985.
I/B. Series BIFLORAE Barneby
Calliandra sect. Androcallis ser. Biflorae Barneby,
ser. nov. Caules e caudice subterraneo herbacei; phyl-
lotaxis spiralis; fohomm formula ii-iii/7-10; capitula
axillaria subbiflora. — Sp. unica: C. biflora Tharp.
Herbaceous from subterranean caudex; phyllotaxy
spiral; lf-formula ii-iii/7-10; peduncles axillary to
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 99
contemporary lvs, mostly 2-fld. — Sp. 1, of s. Texas
and adj. Tamauhpas.
The combination of erect, functionally herbaceous
stems, mostly two-flowered capitula axillary to con
temporary leaves, and spiral phyllotaxy makes the
one species of ser. Biflorae unique in sect. Androcal
lis. Each of the three diagnostic characters seems
individually derived, as is the habitat in the warm
temperate zone.
57. Calliandra biflora Tharp, Rhodora 56: 132. 1954.
— "TEXAS. DeWitt Co.: . . . southwestern part of
the county, July 5, 1942, Maguerite Ridel and B. C.
Tharp 44419." — Holotypus, TEX!. Fig. 11
C. biflora sensu Turner, Leg. Tex. 32, map 4. 1959; Isely,
1973: 78, map 18; Correll & Johnston, 1979: 770.
Functionally herbaceous subshrubs with few erect,
simple or strictly few-branched, closely foliate stems
3-6 d m that die back annually to a shortly buried
caudex, except for upper face of lfts stigulose or sub-
appressed-pilosulous throughout with fine, appressed
or partly ascending, straight white hairs to 0.3-0.9
m m , the fohage dull green subconcolorous, the mostly
2-fld capitula borne solitary or 2-4 together in axils
of primary lvs from mid-stem upward, sometimes
incipiently pseudoracemose distally; phyllotaxy spi
ral. Stipules lanceolate or linear-attenuate 2-4 x 0.2-
0.8 m m , 1-nerved, tardily deciduous. Lf-formula
ii-iii/7-10, the pinnae of some early and of random
distal lvs (no further described) 1-jug.; lf-stk of larger
lvs 1-1.8 cm, the petiole 3-9 m m , at middle 0.4-0.5
m m diam, the one or the longer of 2 interpinnal seg
ments (2-)2.5-7 m m ; rachis of longer pinnae 8-14
m m , the longer interfoholar segments 0.8-2.2 m m ;
lft-pulvinules 0.15-0.3 x 0.2-0.4 m m ; lfts subequi
long except at very ends of rachis, the blades nar
rowly oblong or inequilaterally lanceolate from shal
lowly semicordate base, acute, the larger ones 4-6.3
x 1.3-2.2 m m , 2.7-3.4 times as long as wide; venula
tion palmate, weakly raised on dorsal face only, the
midrib only a little forwardly displaced from mid-
blade, simple, the 1-2 posterior primary venules
much shorter. Peduncles 5-9.5 m m , bracteate close
under the fls, the bract resembling those subtending
the fls, these subulate 1.2-1.8 m m , persistent; capit
ula (l-)2(-3)-fld, the receptacle <0.5 m m ; fls homo
morphic subsessile, the pedicel at most 0.25 x 0.4
m m ; perianth 5-merous, thinly strigose overall, the
corolla brownish; calyx campanulate 2-2.3 x 2 m m ,
obtusely 5-angulate, the narrowly triangular teeth
0.85-1 m m ; corolla 5-5.6 m m , the lobes ±1.8 m m ; Fig. 11. Calliandra biflora Tharp.
100 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
androecium 14-20-merous, ±18-21 mm, the tube 4
m m , the stemonozone 0.8-1 mm, the tassel whitish.
Pods vertically erect, in profile 4.5-6.5(-"8") x 0.8-1
cm, the sutural keels in dorsal view scarcely 2 m m
wide, the reddish-brown, stiffly chartaceous (but
translucent) valves low-convex over 5-8 seeds, strig
ose overall; seeds obtusely rhomboid, compressed but
plump, in broad view 6.2-7.8 mm, the smooth hard
putty-colored testa fuscous-mottled, the pleurogram
long and narrow.
In xeromorphic brush-woodland or thom-forest on
either sandy or stony soils below 350 m, surviving
grazing and clearing in roadside thickets, apparently
rare and local, known only from lowland s. Texas
(Goliad and DeWitt Counties) and Tamaulipas, Mex
ico (Aldama), in lat. 23°-28°30'N. — Map 26. — Fl.
IV-VII.
I/C. Series CHILENSES Barneby
Calliandra sect. Androcallis ser. Chilenses Barneby,
ser. nov. monotypica a sectionis suae speciebus micro-
et ohgophyllidiis (foliorum formula i/4—7) compara-
bihbus praesertim phyllotaxi spirah, ulterius stipuhs
uninerviis subpungentibus et patria atacamensi
diversa. — Sp. unica: C. chilensis Bentham.
Dwarf microphyll shmblets; phyllotaxy spiral; stip
ules 1-nerved, subpungent; lf-formula i/4—7. — Sp. 1,
of fog-desert in Atacama Desert, near 27°45'S in
n. Chile.
In the context of sect. Androcallis, the one species
of ser. Chilenses is taxonomically isolated by spiral
phyllotaxy and incipiently spinescent one-nerved
stipules. Britton in manuscript (Morong 143, N Y )
provisionally transferred it to a monotypic genus, per
haps on account of the relatively thin-textured, few-
seeded fmit; but in general aspect and in form of the
flower, C. chilensis seems generically inseparable
from xeromorphic microphyll species such as Peru
vian C. taxifolia. Calliandra chilensis is the only
member of the genus native in Chile and thus marks
the southwestern hmit of the genus. The spiral phyl
lotaxy, like that seen in C. biflora at the northeastern
periphery of the generic range, is presumably of
independent origin. In one dissected flower I found
two ovaries, a random abnormality.
58. Calliandra chilensis Bentham, London J. Bot. 3:
103. 1844. — "Chili, Bridges, n. 1291." — Holoty
pus, K (hb. Hook.)!. — Feuilleea chilensis O.
Kuntze, Revis. Gen. PL 1: 187. 1891.
Acacia nigra Clos in Gay, Fl. Chil. 2: 253. 1847 — "[Chile.
Coquimbo:] . . en sitios muy descubiertos del camino de
Arqueros." — Holotypus, P (not verified); isotypus, K!. —
Equated with C. chilensis by Bentham, 1875: 545.
Calliandra chilensis sensu Gay, Fl. Chil. 2: 253. 1847; Ben
tham, 1875: 545; Reiche, Fl. Chil. 2: 28. 1897.
Diffuse, stiffly branched, xeromorphic microphyll
shmblets 3-10 dm, with straight divaricate long-
shoots, the young growth thinly strigulose with white
and scattered reddish granular trichomes, the ephem
eral lvs almost concolorous, the plane thin-textured
lfts either pubemlent overall or glabrous on upper
face, the small, compactly umbelliform capitula
borne singly at the lowest nodes of new growth, either
directly from a primary lf-axil or from a short axillary
brachyblast; phyllotaxy spiral. Stipules firm, nar
rowly lanceolate acute 1.5-4.5 m m , strongly 1-
nerved, persistent and subpungent in age. Lf-formula
i/4—7; lf-stks 1.5-5 m m , at middle 0.3-0.4 m m diam;
rachis of pinnae 5-11 m m , the longer interfoholar
segments 0.8-2.2 m m ; lft-pulvinules 0.1-0.15 m m ;
lfts scarcely graduated, the blades oblong-elliptic or
ovate from obtusangulate base, abmptly apiculate,
the larger ones (2.2-)2.8-^.5 x 0.9-1.6 m m , 2.1-3
times as long as wide, all veinless or faintly 1-2-
nerved dorsally. Peduncles very slender 6-17 m m ,
ebracteate; capitula (3-)6-10-fld, the receptacle ±1-2
m m ; bracts incurved, narrowly lanceolate 0.7-1.5
m m , 1-nerved, persistent; pedicel columnar or tur
binate 0.25-0.5 m m ; perianth (4—6-merous) not stri
ate, the corolla thinly strigulose above middle, the
rest glabrous or almost so; calyx 1.4—3 m m , the teeth
narrowly ovate acute 0.5-1.3 m m ; corolla (?dark red,
described by Clos as black) 4.3-5.5 m m , the narrowly
ovate lobes 1.2-2 m m ; androecium 28-34-merous,
11-16 m m , the stemonozone 1-1.2 m m , the tube 3.5-
5 m m , corneously thickened internally but lacking
differentiated nectary; ovary shortly stipitate, thinly
pilosulous toward apex. Pods planocompressed, when
well fertilized elliptic-oblanceolate in profile, 2.5^.5
x 1.1-1.4 cm, attenuate at base, at apex broadly
rounded and abmptly apiculate, (l-)2-4-seeded, the
sutural keels slender, in dorsal view ±1 m m wide, the
papery valves plane except where low-bullate over
seeds, lustrously fuscous-castaneous, finely trans-
verse-venulose, thinly strigulose overall; dehiscence
and ripe seeds not seen.
O n dry open slopes of quebradas in the coast
range and Andean foothills, attaining 1100(-?) m,
localized in the Chilean provinces of Atacama and
Coquimbo in lat. 27°45/S. — M a p 11. — FL
IX-X(-?).
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 101
I/D. Series P A U C I F L O R A E Barneby
Calliandra sect. Androcallis ser. Pauciflorae Bar
neby, ser. nov. Frutices xeromorphi microphylli;
phyllotaxis disticha; foliorum primariorum stipulae
parvae rigidulae vel spinescentes; fohomm formula
i/l-6(-7), foliolis 1.5̂ 4- m m usque longis; capitula
(l-)2-5-flora. — Sp. typica: C. pauciflora (A.
Richard) Grisebach.
Stiffly repeatedly branched xeromorphic shrubs;
phyllotaxy distichous; stipules of primary lvs in
durated, in one sp. spinescent; lf-formula i/l-6(-7),
the lfts ±1.5^- m m ; capitula (l-)2-5-fld. — Spp. 2,
endemic to Cuba.
The two Pauciflorae have much in common, but no
one common character that will dissever them from
all other xeromorphic microphyll species of the
genus. The Brazilian C. depauperata is in most
respects similar, but seems more closely akin to C.
brevipes. Calliandra pauciflora is uniquely armed but
has conventional leaves with 3-7 pairs of plane
leaflets, remarkable only for their small size. In C.
enervis the stipules are not truly spinose, but the
leaflets are reduced to one pair per pinna and are
uniquely carnosulous and biconvex. The characters
unique to each are such as occur widely in xeromor
phic Mimosaceae.
59. Calliandra pauciflora (A. Richard) Grisebach,
PL Wright. 180, in nota sub C. colletioidi. 1860 =
M e m . Amer. Acad. Arts II, 8: 180. 1861. Acacia
pauciflora A. Richard in Sagra, Hist. Phys. Cuba,
PL Vase. 461. 1846. — "Crescit in insula Cuba." —
Holotypus, 7?. de la Sagra s.n, P (hb. Rich.) n.v, but
examined by Bassler, 1990: 203. — Provisionally
but incorrectly referred by Bentham (1875: 634) to
genus Pithecellobium. — Anneslia pauciflora Brit
ton & Rose, N. Amer. Fl. 23: 75. 1928. Calliandra
pauciflora subsp. pauciflora Bassler, Gleditschia
18: 204. 1990.
Anneslia nipensis Britton & Rose., N. Amer. Fl. 23: 76.1928.
— ". . . between Piedra Gorda and Woodfred, Oriente,
Cuba, January 13,1910, [J. A.] Shafer 3710." — Holotypus,
NY!. — Calliandra nipensis Morton, Contr. Ocas. Mus.
Hist. Nat. Colegio "De La Salle" 10: 238. 1951. C. pauci
flora subsp. nipensis Bassler, Gleditschia 18: 205. 1990.
Calliandra pauciflora sensu Combs, Trans. Acad. Sci. St.
Louis 7: 419. 1897; Leon & Alain, 1951: 238.
C. nipensis sensu Leon & Alain, 1951: 238.
Thorny microphyll shrubs, stiffly repeatedly
branched, either broad and diffuse 3-10 d m or poten
tially arborescent to 6 m, with virgate livid long-
shoots, the primary lvs early shed and the lvs at
anthesis mostly clustered on acaulous brachyblasts,
the nodes of long-shoots armed with a pair of ascend
ing straight or outwardly curved spinescent stipules,
the young branchlets, lf-axes, and the dorsal face and
margin of some lfts finely pilosulous with white hairs
0.1-0.6 m m , the minute firm lfts plane sublustrous,
the few-fid capitula solitary on brachyblasts, either
shortly pedunculate or subsessile; phyllotaxy distic
hous. Stipules of primary lvs 1—2.5(—3.5) m m , taper
ing from confluent base, externally nerveless, those
of brachyblasts much smaller, not indurated. Lf-
formula i/(2-)3-6(-7); lf-stks 0.4—1.4 m m ; pinna-
rachises 2-6 m m , the longer interfoholar segments
0.1-0.6 m m ; lfts sessile, accrescent upward along
rachis, the furthest pair sometimes shorter and broader,
the blades mostly linear-oblong or -oblanceolate from
oblique or shallowly semicordate base, straight, ob
tuse, the penultimate pair 1.3-3.6 x 0.5-1.1 m m ,
2.6^4 times as long as wide; venation immersed, the
slightly eccentric midrib sometimes barely percep
tible on dorsal face. Peduncles 1-3.5(-4-) m m , ebract
eate; capitula (1-)2-4-fid, the receptacle <1 m m ; flo
ral bracts hnear-lanceolate or narrowly ovate 0.8-1.2
m m , 1-nerved, persistent; fls sessile, the perianth
either 4- or 5-merous, glabrous or nearly so, both
calyx and corolla strongly 4—5-nerved, 4—5-angulate;
calyx 2.1-3.4 m m , the campanulate or narrowly cam
panulate tube 0.7-1.2 m m diam, the narrowly lance
olate acute teeth 0.8-1.4 m m ; corolla (3.5-)4.1-6
m m , the erect lanceolate lobes 0.8-1.4 m m ; androe
cium (8-)12-26-merous, ±14—17(—?) m m , the tube
(2.8-)3.5-7 m m , from 1 m m shorter to 1.5 m m longer
than corolla, the stemonozone ±0.5 m m , the tassel
described as pink, red, or purple-violet; ovary at
anthesis glabrous; disc 0. Pods (scarcely known) 2-6
x 0.4—0.7 cm, dark brown, glabrous; seeds (Bassler,
1990) 3-5 x 5 m m , pleurogrammic.
In arid savanna and in carrasco on dry stony hill
sides, ±100-800 m, discontinuously scattered over
Cuba, from Pinar del Rio to Oriente (vide Bassler,
1990, Karte 3). — Fl. IV-VI, probably at other times.
Calliandra pauciflora is the one species of its
genus armed with tmly spinescent stipules, though
only those stipules that subtend primary leaves of
long-shoots are thus modified, not those of the florif-
erous brachyblasts. In general habit the species re
sembles the sympatric Sphinga prehensilis (C.
Wright) Barneby & Grimes, but has much shorter pe
rianth and androecial tube and, of course, the fruit
and dehiscence of genuine Calliandra.
102 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
60. Calliandra enervis (Britton) Urban, Symb. An-
till. 9: 437, in nota sub C. bullata. 1928. Anneslia
enervis Britton, Bull. Torrey Bot. Club 41: 18.
1914. — "Mountains of northern Oriente, Cuba;
type from Camp La Gloria, south of Sierra Moa, [J.
A.] Shafer 8274, December, 1910." — Holotypus,
NY!; isotypus, NY!. Fig. 12
C. bullata Urban, Symb. Antill. 9: 437. 1928. — "[CUBA.]
Prov. Oriente prope Minas de Iberia ad sinum Taco cr. 800
m. alt.... : [E. L. Ekman] n. 3814." — Holotypus, +B; iso
typus, NY!. — Anneslia bullata Britton & Rose, N. Amer.
Fl. 23: 194. 1928. — Equated with C. enervis by Leon &
Alain, 1951: 238; Bassler, 1990: 206, Karte 2, t. X(f, j, k), t. XI(c).
C. enervis sensu Leon & Alain, 1951: 238.
Stiffly repeatedly branched, microphyll, arborescent
shrubs flowering when l-3.5(-4) m tall, with virgate,
± zigzag, fuscous, evanescently foliate long-shoots
and very short, often crowded, densely thatched
brachyblasts, except for rudimentary caducous
pubemlence of young stems glabrous throughout,
inconspicuously armed at some nodes by pairs of
firm ascending, ±5-angulate, finally deciduous pri
mary stipules, the small fleshy lfts lustrous above, the
few-fid capitula subsessile on brachyblasts; phyl
lotaxy distichous. Stipules of primary lvs ±1-1.8 m m .
Lf-formula i/1, each If 4-foliolate; petioles and pinna-
rachises cuneate, shallowly excavated ventrally, 0.6-
1.8 m m ; lfts sessile, oblong-obovate or obovate from
shallowly semicordate base, broadly obtuse or ob
scurely apiculate, 1.6-4 x 1.2-2.2 m m , 1.3-2.3 times
as long as wide, the externally nerveless blade bi
convex along the hne of the midrib, plane toward
margin. Peduncles 1.5 m m or less; no bract seen;
capitula (l-)2-5-fld, the receptacle scarcely 1 m m ;
floral bracts subulate ±1 m m ; perianth either 4- or 5-
merous, glabrous, the calyx and corolla both 8-15-
nerved; calyx campanulate 1.7-2 x 0.8-1 m m , the
lance-subulate teeth 0.6-0.8 m m ; corolla 3.7-4 m m ,
the lanceolate lobes 1.5-2 m m ; androecium 18-24-
merous, 12-14 m m , the stemonozone ±0.9 m m , the
tube 1.6-2 m m , the tassel red; intrastaminal disc 0;
ovary subsessile, glabrous. Pods erect, in profile 28-
40 x 5-6.5 m m , 3^1-seeded, glabrous overall, the su
tural ribs in dorsal view 1.5-1.8 m m wide, the plane
leathery, dark brown valves externally nerveless;
seeds not seen, according to Bassler (1990) 4-5 x 3
m m , pleurogrammic.
In thorny thickets, on serpentine bedrock, 800-
1000 m, endemic to n.-e. Cuba in prov. Holguin and
adj. Guantanamo (cf. Bassler, 1990, Karte 2). — Fl.
XII-II(-?). FlG. 12. Calliandra enervis (Britton) Urban.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 103
Although the primary stipules of C. enervis are
somewhat indurated they are deciduous, hardly to be
termed spinescent. The tiny, unijugate leaflets, bullately
thickened at mid-blade, are peculiar to this one species.
I/E. Series AMBIVALENTES Barneby
Calliandra sect. Androcallis ser. Ambivalentes Bar
neby, ser. nov., inter ser. Androcallin et ser. Macro-
phyllas quasi intermedia, pinnis semper exacte 1-
jugis cum his congruae sed fohohs 3-7(-10)-, nee
1/2-21/2(-3)-jugis et plemmque majoribus (2-16 c m
usque longis) diversae. — Sp. typica: C. guildingii
Bentham.
Shmbs and trees, intermediate in lf-formula or in
lft-size between ser. Androcallis and ser. Macrophyl
lae, artificially separated from both; lf-formula i/
(2-)3-7 (in C. haematocephala var. haematocephala
-9, 10), the distal or penultimate lft-blades (1.5-)
2-9(-10.5), in C. haematocephala var. boliviana to
16.5 c m long. — Spp. 9, scattered within the range of
sect. Androcallis in tropical Mexico and Central
America, and in n. and equatorial S. America, thence
sub-Andean s. to Bohvia.
It has long been evident that ser. Nitidae Bentham
(= sect. Androcallis ex parte) and ser. Macrophyllae
Bentham cannot be separated by any one leaf-character,
but preservation of these relatively populous groups
remains conceptually useful. Segregation, as a ser.
Ambivalentes, of those intermediate species that differ
from ser. Macrophyllae by more numerous leaflet-pairs
and from sect. Androcallis by larger leaflet-blades, pre
serves, however artificially, Bentham's system.
61. Calliandra medellinensis Britton & Rose ex
Britton & Killip, Ann. N e w York Acad. Sci. 35: 135.
1936. — "[Club Campestre,] vicinity of Medelhn,
Antioquia, Columbia, March 23, 1927, [Rafael A.]
T o w 109r — Holotypus, NY!.
Closely resembling C. riparia except for fewer,
sometimes proportionately wider lfts, and for inflo
rescence sometimes pseudoracemose as in C. falcata,
known mostly (perhaps only) from plants in cultiva
tion, possibly of hybrid origin (C. riparia x falcata!).
Stipules of C. riparia. Lf-formula i/(3-)4—7; petioles
6-20 m m ; pinna-rachises 1.8-4 cm, the longer inter
foholar segments 5-7 m m ; lft-pulvinules 0.4—0.6 x
0.7-0.9 m m , deeply wrinkled; lfts scarcely accrescent
distally, the blades narrowly oblong-elliptic from
broadly semicordate base, at apex either deltately
acute or obtuse apiculate, the penultimate pair 20-27
x 5-9 m m , 3^-4 times as long as wide; venation of C.
riparia. Peduncles when axillary to primary lvs ±1.5-
3.5 cm, when terminally racemose (as in C. falcata)
much shorter; capitula ±11—22-fld, the sessile gla
brous fls homomorphic as to perianth but the androe
cial tube of one or more (sub)terminal fls longer and
more dilated at orifice, the calyx striate, the corolla
scarcely so; calyx 2-2.5 x 1-1.5 m m , the teeth 0.3-0.4
m m ; corolla 7.2-8.2 m m , the lobes 1.4-2 m m ;
androecium of peripheral fls 20-24-merous, 3.2-3.6
cm, the pallid tube either included or far exserted
6.5-17 m m , the tassel pink-carmine; ovary glabrous;
no nectarial disc seen. Pod and seed unknown.
In parks and gardens, and on roadsides, 950-1500
m, known only from inter-Andean valleys of n.-centr.
Colombia: in urban cultivation in Antioquia and
Tolima and from the old Guaduas-Honda road in Cun
dinamarca. — Fl. IX-III(-?). — Carbonero de pluma.
Calliandra medellinensis is an ambiguous taxon,
not certainly known outside cultivation. In general
aspect and in most macromorphic details the plants
resemble C. riparia, but have fewer leaflets that tend
to be proportionately wider. Of the four known col
lections, three (including the type) have capitula axil
lary to primary leaves, but one (Soejarto 3379, N Y )
has the terminal, pseudoracemose inflorescence of C.
falcata, which differs, however, in much wider leaf
lets. Both C. riparia and C. falcata are commonly
planted far beyond their natural range of dispersal,
and both are recorded from Colombian urban habi
tats. The character-combinations observed in C. med
ellinensis are plausible attributes of hybrid C. riparia
x C. falcata, but experimental evidence is required to
confirm or refute this hypothesis.
62. Calliandra mollissima (Willdenow) Bentham,
London J. Bot. 3: 97. 1844. Inga mollissima Hum
boldt & Bonpland ex Willdenow, Sp. PL 4(2): 1023.
1806. — "Habitat in America meridionali," the lo
cality specified by Kunth, Mimoses 61, t. XIX. 1819:
"Crescit prope Querocotillo in regione calidissima
Provinciae Jaen de Bracamoros [Cajamarca, Pem]
inter flumina Amazonum et Rio de Chota, alt. 230
hexapodamm [±825 m]." — Holotypus, Humboldt
& Bonpland 3663, B - WILLD 19050, seen in Micro
form, 36: box 31! = F Neg. 1250; isotypi, P-HBK
n.v, P (hb. Bonpland.)\, F (fragm ex P)!. — Mimosa
mollissima Poiret, Encycl. Suppl.l: 46. 1810.
Feuilleea mollissima O. Kuntze, Revis. Gen. PL 1:
188. 1891.
C. chotanoana Harms, Repert. Spec. Nov. Regni Veg. 17:
442. 1921. — "Peru: Dep. Cajamarca, Prov. Cutervo, Tal
des Rio Chotano, unter Querocotillo 1000-1100 m
104 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
(Weberbauer no. 7123. Mai 1915 . . . )." — Holotypus, fB; clastotypus (fragm.) F + F Neg. 7235!; isotypi, F!, GH!, K!.
Inga mollissima sensu Humboldt, Bonpland & Kunth, 1824:
300; de Candolle, 1825: 439. Calliandra mollissima sensu
Bentham, 1875: 541; Harms, 1921: 88; Macbride, 1943:72.
Macrophyllidious shrubs and treelets 2-A m, with
terete, commonly glabrate branches, the new branch-
lets, lf-axes, and peduncles densely pilosulous with
incurved or straighter spreading, sordid gray hairs to
0.6(-0.8) m m , the bicolored, chartaceous, reticulately
venulose lfts either glabrous facially except for pilo
sulous midrib or pilosulous with fine pallid erect-
ascending hairs, the capitula of white-stamened fls
arising singly and geminate from axils of mature
coeval lvs or from recently defohate nodes of homoti
nous branchlets; phyllotaxy distichous. Stipules erect,
firm, narrowly triangular to lanceolate or deltate
±1.5-5.5 m m , weakly 5-7-nerved dorsally, persistent
or tardily deciduous. Lf-formula i/4—7; petioles 5-20
m m , at middle 0.6-1.4 m m diam; rachis of longer pin
nae (1.5—)3—8.5(—9.5) cm, the longer interfoholar seg
ments 8-18(-20) m m ; lft-pulvinules 0.2-0.5 x 0.5-
0.8 m m , finely cross-wrinkled; lfts gently or abmptly
accrescent from base of rachis upward, in outline
asymmetrically elliptic-obovate or subrhombic-
obovate from shallowly semicordate base, broadly
obtuse, the terminal pair in larger lvs 17—42 x 10-
18(-20) m m , 1.6-2.2 times as long as wide; primary
venation of 3-4 nerves from pulvinule, the midrib
moderately or scarcely displaced, the strong inner
most posterior primary nerve incurved-ascending
through 3A of blade or more, the outer ones much
shorter, the midrib pinnately branched distally, a close
reticulum of sinuous venules elevated on both faces,
more sharply so above. Peduncles (6-) 11-27 m m ,
randomly bracteate below middle or ebracteate; capit
ula 11-27-fld, the receptacle 1-2 m m diam; bracts
subulate 0.6-1.2 m m , persistent; fls (sub)sessile, het
eromorphic, of subequal length but the androecium
of one or more central ones modified; perianth thin-
textured, either 4- or 5-merous, either glabrous except
for minute granules at tip of corolla or pilosulous-
strigulose externally, both calyx and tube of corolla
sharply striate-nerved, the corolla-lobes externally
nerveless; P E R I P H E R A L FLS: calyx campanulate
1.4-2 x 0.8-1.4 m m , the depressed ovate-deltate teeth
0.15-0.4 m m ; corolla whitish, 4.3-5.4 m m , the lobes
0.6-1.6 m m ; androecium 9-12-merous, 26-33 m m ,
the stemonozone 0.3-0.45 m m , the tube 4-5.2 m m ;
ovary glabrous; disc 0; C E N T R A L FLS: perianth
almost of peripheral fls but the androecium trumpet-
shaped, its tube 8.5-13.5 m m , the dilate orifice 2.5-
4.5 m m diam; intrastaminal disc ±0.4 m m ; ovary 0.
Pods (few seen) ±7-10.5 x 1 cm, the valves either gla
brous, or pubemlent, or densely pallid-pilosulous;
seeds unknown.
In deciduous brush-woodland at 400-1200 m, on
steep slopes of valleys tributory to rio Maranon in
deptos. Amazonas (prov. Bongara and Bagua) and
Cajamarca (prov. Cutervo and Chota) within lat. 5°-
6°30'S in n. Pem. — M a p 29. — FL X, XII, V, the full
cycle unknown.
As defined by m y description, C. mollissima is var
iable in pubescence, C. chotanoana corresponding to
the glabrescent variant. Leaf-formula (i/4—7) and rel
atively ample leaflets (to ±2-4 x 1-2 cm) together
(but not separately) exclude C. mollissima from either
ser. Nitidae or ser. Macrophyllae as newly circum
scribed and provide an exemplar of an ideal member
of ser. Ambivalentes.
63. Calliandra guildingii Bentham, London J. Bot.
3: 96. 1844. — "St. Vincent's [Lesser Antilles],
Guilding." — Holotypus, K!; clastotypus (fragm. +
photo), NY!. — Feuilleea guildingii O. Kuntze,
Revis. Gen. PL 1: 188. 1891. Anneslia guildingii
Britton & Rose, N. Amer. Fl. 23: 56. 1928.
C. decrescens Killip & Macbride, Publ. Field Mus. Nat.
Hist., Bot. Ser. 13 (Fl. Peru) 3: 71. 1943. — "[PERU.]
Loreto: Mishuyacu, Klug 152." — Holotypus, US!; iso
typus, NY!. C. guildingii sensu Grisebach, 1864: 225 (but not introduced
in Trinidad); R. O. Williams, Fl. Trin. Tob. 1(4): 298. 1931.
Macrophyllidious trees (2-)3-12(once reported 20)
m tall with gray trunk and plagiotropic long-shoots,
often appearing and sometimes tmly glabrous, but the
lf-axes and either the margin or the principal veins of
lfts pilosulous or micropuberulent with sordid hairs
0.1-O.4(-0.5) m m , the lvs moderately bicolored, lus
trous and reticulate on both faces, the capitula bome
singly or less often geminate at stipulate nodes of very
short, efoliate or 1-foliate brachyblasts axillary to
coeval primary lvs, scattered along the leafy growth of
current year; phyllotaxy distichous. Stipules broadly
or narrowly triangular, rarely lanceolate, ±l-8(-l 1) x
0.6-1.3(-2) m m , finely 3-8-nerved dorsally when
young, becoming smooth, deciduous. Lf-formula i/2-
5(-6), the lfts conspicuously accrescent, the small first
pair seldom exactly opposite; lf-stks (2-)4-17(-19) x
0.7-1.5 m m , openly shallowly grooved ventrally, bi-
cupulate at apex; rachis of longer pinnae (1.6-)2-
6.5(-7) cm, the furthest interfoholar segment (7-)9-18
m m ; lft-pulvinules 0.4-1.9 x 0.5-1.4 m m , wrinkled;
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 105
• CALLIANDRA GUILDrNGII
¥ C MOLLISSIMA
M A P 29. Distribution of Calliandra guildingii Bentham
and C. mollissima (Willdenow) Bentham in northwestern
South America.
proximal pair of lfts (often of unequal size) asymmet
rically rhombic-ovate, obscurely if at all acuminate,
5-22(-32) x 3-12(-20) m m , the distal pair inequilat
erally or subdimidiately ovate-elliptic-acuminate from
semicordate (broad-cuneate) base, at very apex either
acutely triangular or obtuse apiculate, the largest
(2-)2.5-8.2 x (0.8-)l^t cm, ±2-2.8 times as long as
wide; venation palmate-pinnate, the midrib either gen
tly incurved or straight diagonal, the inner posterior
primary nerve incurved-ascending well beyond mid-
blade, the outer posterior ones much shorter, all these
and the tertiary and reticular venules sharply promin
ulous on both faces. Peduncles 7-25(-34) m m , usu
ally but not always bracteate near middle, thickened
and arrect in fmit; capitula 10-24(-28)-fld, the recep
tacle 1.5-3 m m diam; bracts subulate 0.7-1.2 m m ,
persistent; fls commonly heteromorphic, one or more
(sub)terminal ones scarcely different as to perianth
but with dilated and exserted, whitish staminal tube,
these fls pistillate and nectariferous but sometimes ill-
developed (or perhaps absent), the rest staminate, with
slender, either included or rarely exserted staminal
tube; perianth of all fls thin-textured, glabrous or re
motely either strigulose or pilosulous, either 4- or 5-
merous, the calyx striate but the corolla not; PERIPH
E R A L FLS: pedicel often cryptic 0.15-0.5 x 0.4-0.8
m m ; calyx campanulate 1.3-3 x 1-2.5 m m , the de
pressed or vestigial teeth 0.1-0.4 m m (one or more
sinuses often more deeply split in late anthesis);
corolla 5.8-9(-10) m m , the ovate (often unequal)
lobes 1.2-2.4 m m ; androecium 10-22(-24)-merous,
attaining 42 m m , the tube 4.5-12 m m , the whole deep
crimson or sometimes the tube palhd and only the tas
sel red; ovary at anthesis glabrous. Pods acroscopic, in
broad profile 6.5-10.5 x 0.75-1.1 cm, thinly pubem
lent overall or glabrescent, the thickened sutural ribs
4-4.5 m m wide in dorsal view, sulcate lengthwise, the
hgnescent valves prominently, either obliquely or sub-
vertically venulose; seeds (few seen) ±5-6 per pod, in
broad view 9.5-10 x 5-6 m m , the papery testa pale
brown or castaneous, often crumpled, pleurogram 0.
In moist lowland forest (Trinidad; state of Miranda,
Venezuela; Amazonia) and ascending (n.-w.
Venezuela; n. Colombia) into submontane forest and
cloud-forest to 900-1850 m, discontinuously dis
persed from n.Trinidad w. through the Venezuelan
coastal cordillera to state of Yaracuy, thence s. and s.-
w. to the Casiquiare in Venezuelan Amazonas, the
eastern Andes in state of Tachira, and the lower Cauca
valley in Antioquia, Colombia, s.-e. Ecuador (Napo),
and n.-e. P e m (Amazonas, Loreto). — M a p 29. — Fl.
intermittently throughout the year. — Niaure (Trini
dad); kunchai (Pem). — Described from St. Vincent
Island, where probably cultivated in the botanical gar
den by Rev. Lansdown Guilding, and cultivated on
Martinique in 1814 (BM!), but not since seen in the
Lesser Antilles and not mentioned by R. A. Howard in
volume 4 of Flora of Lesser Antilles (1988).
Calliandra guildingii resembles C. falcata, which
has larger, more persistent stipules, only incipiently
acuminate leaflets, and flowering brachyblasts
arranged in leafless or only randomly leafy pseudo-
racemes. Whereas C. guildingii is found in moist low
land and upland cloud-forest, C. falcata is adapted to
xeromorphic, semideciduous communities. A parallel
instance of related but ecologically differentiated
species is known in Senna robiniifolia and S. viciifo-
lia, as described by Irwin and Barneby (1982: 520,
sequ.). The species further is suggestive of an ex
treme form of C. surinamensis in which the leaflets
are reduced in number to three or five, exceptionally
six pairs per pinna and reciprocally dilated. Its hg
nescent, vertically venulose pod recalls that of C.
magdalenae, but the papery seed-testa lacking pleu
rogram is that of the C. surinamensis complex.
64. Calliandra falcata Bentham, London J. Bot. 3:
97. 1844. — "Raised in Mr. Knight's nursery from
Mexican seeds." — N o typus found at K in 1994,
but collections by Fendler, Funk, and Birschell from
Caracas (all K!) cited as C. falcata by Bentham,
106 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
1875: 541, are authentic. — Feuilleea falcata O.
Kuntze, Revis. Gen. PL 1: 187. 1891.
C.fulgens Hooker fil., Bot. Mag. 124: t. 7626. 1898. — De
scribed from plants, thought to be of Mexican origin, that
flowered in Regent's Park London, in 1888, and in the
Palm House at Kew in 1897. — Specimens from both
sources, K!; lvs of Kew plant sent to Britton by Sir Arthur
Hill in 1927, NY!. —Anneslia fulgens Britton & Rose, N.
Amer. Fl. 23: 56. 1928.
C. serjanioides Urban, Symb. Antill. 2: 262. 1900. — "Cole-
batur olim in horto botanico Martinique: Duss." — Holo
typus, perhaps *B; no isotypus found. — Said to differ
from C. guildingii by 4 (not 2-3)-jugate leaflets undulately
crenate (not entire), characters no longer significant. Cal
liandra falcata was cultivated on Martinique as early as
1841 (Perrotets.n.,G).
(?) C. amblyphylla Harms, Repert. Spec. Nov. Regni Veg.
17: 88. 1921. — "[AFRICA.] Kamerun: Kult. im Bot.
Garten Victoria (no. 70)." — Holotypus, *B; no isotypus or
photo found. — Described as related to C. guildingii but different in blunter leaflets, the protologue in close agree
ment with plants grown by Broadway on Trinidad in 1921
and thence at Buitenzorg in 1941.
C. falcata sensu Pittier, 1927: 44.
Macrophyll, drought-resistent trees attaining (2-)
3-6(-8) m with one or several trunks and stiff terete
pallescent, densely foliate long-shoots, the young
stems and lf-axes variably pubemlent, subglabrous,
or pilosulous with fine soft erect-incurved hairs to
0.2-0.5 m m , the plane, stiffly papery, scarcely bi
colored lfts glabrous or glabrous ciholate to pilosu
lous along principal nerves or pilosulous overall, the
dense capitula arising, singly or fasciculate by 2-3,
from a crowded series of efoliate or only depauper-
ately foliate nodes either terminating or lateral to the
main leafy axis of the season, forming a pseudo
raceme shortly exserted from coeval foliage; phyl
lotaxy distichous. Stipules deltate or lanceolate 2-10
x 1.5-2.8 m m , when young (5—)7—11 -nerved dor
sally, becoming thick and smooth, either glabrous or
pilosulous dorsally, persistent or tardily deciduous.
Lf-formula i/2—3(^4), some lvs of any plant at least 9-
foliolate and most larger lvs 12- or 16-foliolate, the
proximal lfts much smaller than the distal ones, and
often not exactly opposite; lf-stks (3-)4-25(-28) m m ,
at middle 0.8-1.3(-2) m m diam, shallowly grooved
ventrally; rachis of longer pinnae 1.7-4.5(-5) cm, the
longest (furthest) interfoholar segment (7-)8-24 m m ;
lft-pulvinules 0.4-2 x 0.6-1.4 m m , wrinkled; lfts
heteromorphic, the small lowest pair rhombic-ovate,
the distal pair falcately elliptic or semi-ovate from
shallowly semicordate base, shortly or obscurely
acuminate and at very apex obtuse or obtuse mu-
cronulate, the larger ones (2.5-)3-9(-10.5) x
1.2-3.6(-4) cm, 2-2.6(-2.7) times as long as wide;
venation palmate-pinnate, the primary nerves 3(^1),
the midrib gently incurved or almost straight, dis
placed to divide blade 1:2-2.5, the inner posterior pri
mary nerve incurved-ascending well beyond mid-
blade, the outer one(s) much shorter, all these
together with tertiary and reticular venules sharply
prominulous on both faces. Peduncles \2-A2> x
0.9-1.5 m m , bracteate near or below middle; capitula
10-26-fld, the receptacle 1.5-5 x 2-4.5 m m , the fls (?
always) heteromorphic, one or more central ones with
pallid dilated, well-exserted staminal tube; bracts
ovate 0.4—0.9 m m , persistent; perianth either 4- or 5-
merous, minutely stigulose-pilosulous except for
sometimes glabrescent calyx, weakly or impercepti
bly nerved; P E R I P H E R A L FLS: pedicel (sometimes
not externally differentiated) 0.1-0.5 x 0.4-0.9 m m ;
calyx campanulate 1.3-2.8 x 1.2-2.2 m m , the de
pressed obtuse teeth 0.2-0.4 m m ; corolla 7-11 m m ,
the deltate-ovate obtuse lobes (1.3—)2.1^4.2 m m ;
androecium (14—)16-26-merous, 32-45 m m , the tube
(5.7-)6-10 m m , the stemonozone 0.8-1.2 m m , the
tassel dark red or crimson; intrastaminal nectary 0;
ovary (often rudimentary or lacking) pilosulous;
C E N T R A L FL(S): seldom well observed, the peri
anth broader but little longer than that of peripheral
fls, the whitish androecial tube exserted 5-7 m m from
corolla, at orifice 3-4 m m diam. Pods stiffly ascend
ing, in broad view (6-)7-10 x 1.3-1.8 cm, ±5-6-
seeded, the dark brown, sharply obliquely venulose
valves thinly brown- or grayish-strigulose; seeds in
broad view elliptic ±9.5 x 5-7 m m , the testa brown,
darker-speckled, smooth, the pleutogram deeply U-
shaped.
In drought-deciduous woodland, thickets, and es-
pinares, along the Caribbean coast at 70-650 m but
ascending to 1000-1500 m in the interior, locally
plentiful in n. Venezuela, from Distrito Federal and
Aragua w. to Carabobo, Yaracuy, and Falcon; culti
vated in West Indies and elsewhere. — M a p 30. — Fl.
all months from VIII to III, also in V, most copiously
following rains. — Clavellina, clavellino amarillo,
cimbra-potro.
The presumed relationship of C. falcata to C.
guildingii is based on resemblance in many charac
ters; differences in stipules, stressed in Key III, are dif
ferences in size rather than kind. The different habitats
are surely significant. The inflorescence architecture
of C. guildingii is that expected in sect. Androcallis,
but that of C. falcata is at least sometimes that estab
lished in sect. Calliandra, a terminal efoliate pseudo
raceme, here interpreted as independently derived by
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 107
M A P 30. Distribution of Calliandra falcata Bentham and C. haematocephala Hasskarl var. haematocephala and var. bo-
liviana (Britton) Barneby in South America.
108 M E M O I R S OF T H E N E W Y O R K BOTANICAL G A R D E N [VOL. 74(3)
suppression or partial suppression of many distal pri
mary leaves. The loss of distal leaves is perhaps
related to the ecology of the species.
The nomenclature of C. falcata is not well docu
mented. N o type of the species was found at K e w in
1994/6, but its identity is vouched for by material
from near Caracas identified by Bentham and agree
ing with the original description. Calliandra falcata
was cultivated on Martinique in the mid-nineteenth
century and seems to have been dispersed from the
Botanical Garden there to similar institutions in the
Old World. K e w possesses authentic material of C.
fulgens, which in context of C. falcata is somewhat
unusual in its narrow leaflets, but which cannot be
accommodated elsewhere in the genus as presently
known. Its reputed origin in Mexico is no doubt a
mistake. It has been equated (in herb. K ) with the
related C. haematocephala, but the leaflets are fewer
than in any wild Bolivian matieral of that species and
the flower lacks the characteristic reflexed fringe of
sterile filaments in the mouth of the filament tube
which is its signature. N o type of either C. serjan-
ioides, proceeding from Marintinique in 1900, nor of
C. amblyphylla Harms, dating to about 1921 in equa
torial Africa, survived the disaster at Berlin, and as no
duplicate material of either has been located, inter
pretation depends entirely on the protologues. Typical
C. falcata was at the Trinidad Botanical Garden at
this time, and contemporaneously at Buitenzorg.
65. Calliandra haematocephala Hasskarl, Retzia 1:
216. 1855. — Holotypus infra sub var. haemato
cephala indicatur.
Macrophyllidious shrubs and treelets 1-5 m tall, di
verse in indumentum and in number and size of lfts,
glabrous throughout or almost so to densely softly
pilosulous (more precisely described under each vari
ety), the hemispherical, prior to anthesis globose
capitula arising singly from either imbricately
thatched or from variably elongating efoliate, axillary
brachyblasts, these commonly immersed in foliage,
less often pseudoracemose at tip of currently elongat
ing stems, the capitula notable for densely crowded
flowers and usually bright red filaments. Stipules firm,
ovate-triangular or lanceolate 3-11 (-13) x 1.4-4.5
(-5) m m , striately venulose when young, becoming
thickened, pallid or blanched and externally smooth in
age, persistent. Lf-formula i/3—9(—10), the size and
number of lfts mutually adjusted; petiole of larger lvs
0.9_4.5(-5) cm, at middle 0.7-2 m m diam, at apex
dilated into a pair of homy cupules; rachis of well-
developed pinnae 5-11 cm, the longer (distal) interfo
holar segments 0.8^1.5 cm; lft-pulvinules (0.7-)
1-3 x 0.8-1.5 m m , coarsely cross-wrinkled; lfts vari
ably accrescent distally, the blades inequilaterally
elliptic, (ob)ovate-, or lance-elliptic from shallowly
semicordate base, obtuse or shortly or obscurely
acuminate (the acumen itself obtuse apiculate), the
larger ones 2.2-14(-16.5) x 0.7-54(-6) cm, 1.9-3.6
times as long as wide; primary nerves from pulvinule
2—3(-^4), the gently incurved midrib displaced to
divide blade 1:1.4—2.4, ±6-12-branched on its anterior
side, the inner posterior nerve nearly as strong,
incurved-ascending well beyond mid-blade, the outer
1-2 much shorter, the primary nerves prominulous on
both faces, the secondary and reticular nervules also
sharply raised but progressively finer. Peduncles
l-4(-6) cm, either bracteate or not; capitula densely
20-84-fld, the fls contiguous, the clavate receptacle
2-7(!) m m ; floral bracts ovate, subulate, or flabellate
0.8-2.4 m m , incurved, ±5-8-nerved, persistent;
pedicels either evident or cryptic, (0.3-)0.5-1.2 x
0.6-1.5 m m ; perianth glabrous, either 4-merous, or 5-
merous, or one cycle 4- and the other 5-merous, the
calyx and the corolla-tube striate, the corolla lobes im
perceptibly venulose; calyx campanulate (1-)1.4—
3.7(^4) x 1.3-2.9 m m , the orifice often undulately
tmncate, but sometimes cleft into depressed-ovate or
deltate teeth 0.5 m m or less; corolla red or reddish
(5-)6.5-l 1.3(—12) m m , the ovate lobes 1-3.3 m m ; an
droecium (20-)22-^tOH14)-merous, (22-)30-44 m m ,
the stemonozone 0.7-1.5 m m , the tube (5.5-)
6-11(—13.5) m m , dilated upward into a tmmpet ±3-^-
m m diam either a little shorter or to 4(-4.5) m m longer
than the corolla, the crimson-scarlet or in cultivation
rarely white or pink filaments in one cycle, a few sep
arating from stamen-tube well below orifice, the mem
branes between filaments produced into curved erose
appendages that form a fringe or corona within the ori
fice; nectary around base of glabrous ovary 0.4—0.9 x
0.5-1 m m . Pods to 5 per peduncle, in broad view 7-13
x 0.8-1.3(-l.65) cm, the sutural ribs in dorsal view
2.5-4 m m wide, the plane recessed, hgnescent valves
fuscous nigrescent glabrous, transversely venulose;
seeds (few seen) in broad view ±8.5-10 x 5.5-7.5 m m ,
the smooth crustaceous testa brown, minutely speck
led or mottled, the pleurogram incomplete.
In seasonally dry subtropical woodland and
woodland-savanna transition, 240-800 m., native and
locally abundant in the foothills of the Bolivian
Andes, between 14°S and 18°S latitude, the dispersal
given in greater detail under the varieties; widely dis
persed in cultivation. — Powder-puff tree; pompon;
bellota; flor de la cruz.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 109
As broadly defined by the foregoing description, C.
haematocephala is distinguished among unijugate
calhandras not by any one character nor by any com
bination of characters derived from foliage or indu
mentum, both of which are inordinately variable, but
by the following features of the inflorescence: flowers
not less than 25 per capitulum, in bud crowded into a
blackberry-like ball; perianth glabrous, the calyx and
the corolla-tube striate; filaments monocyclic, several
of them separating from the tube below its orifice,
where the interfilamental membranes are produced
into a corona of inwardly reflexed, pallid erose scales
that contrast in color with the blood-red tassel; and a
nectarial disc, in bisexual flowers, that embraces the
glabrous ovary.
Parts of the C. haematocephala complex have been
revised by Cowan (1963) and by Nevhng and Elias
(1971). Cowan recognized C. haematocephala sens.
str., which is widespread in cultivation but not exactly
matched by any wild plant, as specifically distinct
from C. inaequilatera Rusby, which is known from
native populations as well as from cultivars. Nevling
and Elias, after analysis of more material, found no
substantial differences between these two, and sug
gested further that C. boliviana Britton, which has
precisely the flowers of C. haematocephala but sim
plified leaves and larger leaflets and is, moreover,
sympatric with C. inaequilatera, could well be in
cluded, at some infraspecific level, in an expanded C.
haematocephala. The new material now available for
comparative study confirms this view of C. boliviana.
I here include, further, two shadowy taxa: C. fulgens
Hooker f, supposedly of Mexican origin but known
only from cultivated stock; and C. novaesii Hoehne,
described from eastern Sao Paulo, Brazil, where it has
not since been encounted in the wild. M y conclusion
that C. haematocephala consists of one polymorphic
species composed of relatively few native populations
and several more or less ambiguous cultivars, all of
which are conveniently disposed in two varieties, is
expressed in the following key.
Key to the varieties of C. haematocephala
1. Lfts commonly (where sympatric with var.
boliviana always) 5-10 pairs per pinna, but
variable in amplitude; exceptionally, in
cultivation only, 3 ^ pairs per pinna, but the
largest then only 1-1.5 cm wide; sympatric with the following on the headwaters of Beni,
not yet encountered in the Mamore watershed;
extensively cultivated, in several variants, in
the Old and New World tropics and elsewhere
under glass 65a. var. haematocephala
1. Lfts 3-4 pairs per pinna and the larger (distal)
ones 4.5-16.5 x 2-6 cm; headwaters of rios
Beni and Mamore in s. Beni, La Paz, and Sta.
Cruz, Bolivia 65b. var. boliviana
65a. Calliandra haematocephala Hasskarl var.
haematocephala. C haematocephala Hasskarl,
1855, I.e., sens. str. — "ex horto bot. Calcuttensi sub
nomine Ingae heteroxyli [ad hortum botanicum
bogoriensem] fuit missa." — Lectotypus (Cowan.
1963: 95), L 901280 (hb. Hasskarl.) n.v, but frag
ments from Hasskarl's herbarium at L sent to E. D.
Merrill by R. C. Bakhuisen van den Brink in 1950
(A!) are presumed authentic and are supplemented
by the photo of L 908,107-641, N Y ! and by speci
mens from plants grown as C. haematocephala at
BO: Merrill s.n. in 1903, de Wit s.n. in 1941, both
NY!. — Feuilleea haematocephala O. Kuntze,
Revis. Gen. PL 1: 188. 1891. Anneslia haemato
cephala Britton in Britton & Wilson, Bot. Porto Rico
6: 348. 1926.
C. novaesii Hoehne, RevistaMus. Paul. Univ. Sao Paulo 10:
654, t. II. 1918. — "Museu Paulista: n. 5603, Dr. Campos
Novaes, Municipio de Campinas. Ex Herv. Commissao
Geog. e Geol. de Sao Paulo." — Holotypus, presumably SP (n.v.), but the cited photograph and description decisive, as suggested by Harms, Repert. Sp. Nov. Regni Veg.
17: 89. 1921. C. inaequilatera Rusby, Mem. Torrey Bot. Club 6: 28. 1896. — "[Bolivia. La Paz:] Vic[inity of] Guanai, 1892 (]M.
Bang] 1586." — Holotypus, NY!; isotypi, GH!, K!, NY!. — Equated with C. haematocephala by Nevling & Elias,
1971:81. C. haematocephala sensu Hooker, Bot. Mag. 86: t. 5181.
1860; Koorders, Exkursionsfl. Java 4, Abt. 7: 876. 1926
Cowan, 1963: 95; Isely, 1973: 81; Bailey, Hortus Third 2
201 (but pinnae exactly 2 per leaf, not 2 pairs!). 1976
Everett, Encycl. Hort. 2: 559 + photo. 1981.
Calistemon [sic] linearis sensu L. Lowis, Familiar Indian
Flowers t. 7. 1878.
Calliandra inaequilatera sensu Everett, Gard. Chron. Ill,
85: 184. 1929 & 87: 203. 930; Cowan, 1963: 64, fig. 10.
Lfts in plants outside cultivation 5-9-jug. and the
larger ones 2.2-7.5 x 0.7-2.8 cm; lfts all either
facially glabrous ciholate or variably pilosulous,
either along nerves of hypophyllum or on both faces;
rarely in cultivation lfts only 3^-jug, but then not
over 1.5 c m wide.
Native in premontane forest at 240-600 m on the
upper forks of rio Beni in lat. 14°-16°S, Bohvia (Ixi-
amas, Rurrenabaque, Guanai, Isapuri, Covendo). —
Fl. IV-VII. — M a p 30. — Specimens from parks,
gardens, perhaps occasionally naturalized, have been
seen from United States, Mexico, Greater and Lesser
Antilles, Trinidad, Curacao, Pem, s.-e. Brazil (S.
Paulo), India, s. China, Malaya, Micronesia, Hawaii.
110 M E M O I R S OF THE N E W Y O R K BOTANICAL G A R D E N [VOL. 74(3)
Strictly typical C. haematocephala has relatively
small (to about 2-4.5 x 0.7-1.2 cm), facially glabrous
leaflets; it is commonly cultivated in Asia and is most
closely approached, but not precisely matched, by R. S.
Williams s.n. from the rio Kaka near Isapuri (NY) and
(ex char.) the holotypus of C. novaesii. Calliandra in
aequilatera, the form most frequently grown in North
America, differs to random degrees in facially pubes
cent leaflets mostly reaching 5-7.5 x 1-2.8 cm. In oc
casional leaves of this form one of a pair of pinnae may
have 5-7 pairs of leaflets whereas the other has only
four pairs. In C. fulgens, which may have arisen in cul
tivation, the leaflets are 3^1 pairs in all pinnae. A form
cultivated in the Caroline Islands (Salsedo 119, N Y )
has the leaf-formula of C. fulgens but broader leaflets.
The ultimate provenance of the plants grown at Cal
cutta and in Java from which Hasskarl described C.
haematocephala is unknown. Nevhng and Ehas, hav
ing established the taxonomic equivalence of C.
haematocephala and C. inaequilatera, assumed that
the former must have been introduced from the region
of Guanai in Bolivia, either by Hasskarl himself or by
H. A. Weddell, both of w h o m are known to have trav
eled in the region between 1842 and 1853, in search of
Cinchona. According to one hypothesis, plants were
collected by Hasskarl himself, perhaps described by
him on board ship between P e m and Java, and attrib
uted to the Calcutta garden in order to conceal his role
in smuggling living Cinchona out of Boh via. Alterna
tively, it was Weddell who obtained seeds of the Cal
liandra, which passed to Calcutta either through Paris
or London. There is, however, only circumstantial
evidence in favor of either hypothesis and none to
favor one over the other. It is worth noting that Has
skarl described in the same paper (Retzia 1: 214.
1855) a Calliandra sancti-pauli, hkewise grown at
Bogor, this said to have originated in Sao Paulo (see
synonymy of C. foliolosa, p. 000) and to have come to
Java by way of Utrecht. Several channels of commu
nication between Bogor and Europe were open, and in
absence of positive evidence the route taken by Cal
liandra haematocephala is unknowable.
65b. Calliandra haematocephala Hasskarl var. boli
viana (Britton) Barneby, stat. nov. C. boliviana
Britton, Bull. Torrey Bot. Club 16: 327. 1889. —
"[Bolivia. La Paz:] Guanai, 2,000 ft ([H. H. Rusby]
1314; Mapiri, 2,500 ft. ([H. H. Rusby] 1315)." —
Syntypi, NY!. — Feuilleea boliviana O. Kuntze,
Revis. Gen. PL 3(2): 63. 1898.
C. boliviana sensu Killeen & al., Guia Arb. Bolivia 432, fig.
69. 1994.
Lfts 3-4 pairs per pinna, the first pair often not
quite opposite, the larger (distal) lfts 5—14(—16.5) x
1.9-3.2 cm, all either glabrous overall, or glabrous
ciholate, or densely softly pilosulous overall; inflo
rescence and fmits of var. haematocephala.
In premontane scrub-woodland, shrub-savanna,
and in sunny places along rivers in premontane forest,
290-950 m, of bicentric range in the foothills of the
Bohvian Andes in deptos. La Paz and Sta. Cruz: on
tributaries of no Beni in Nor and Sur Yungas; and on
headwaters of rio Mamor, in distritos Sara, A. Ibafiez,
and Ichilo; in cultivation in Paraguay (Boqueron). —
M a p 30. — Fl. III-VIII.
In La Paz the stems and leaflets are consistently
glabrous, differing in this feature as well as in leaf-
formula from most sympatric var. haematocephala. In
Santa C m z glabrous and softly pubescent forms are
found close together, although not, so far, in mixed
populations.
66. Calliandra erythrocephala H. Hernandez &
Sousa, Syst. Bot. 13: 519, figs. 1,3. 1988. — " M e x
ico, Guerrero, Mpio. Atoyac de Alvarez, Las Golon-
drinas, a 22 k m al N E de El Paraiso camino a Filo
de Caballo, alt. 1000 m, 7 Sep 1983, E. Martinez S.
& J. L. Villasenor 4237." — Holotypus, M E X U
n.v; isotypus, NY!.
Macrophyllidious trees 3-5 m, appearing glabrous
when adult but the lf-axes and principal lf-venation
minutely pubemlent in vernation, the ample thin-tex
tured lfts lustrous olivaceous subconcolorous, the ca
pitula of vivid red fls arising singly from efoliate axils
of thatched brachyblasts axillary to homotinous or
lately shed primary lvs of long-shoots; phyllotaxy
distichous. Stipules subtending primary lvs narrowly
triangular or lanceolate ±2.5-5 x 1-1.5 m m , 6-8-
nerved when young, becoming dry brittle, those of
brachyblasts similar but somewhat shorter, persistent.
Lf-formula i/3^4(—5); lf-stks 5-16 m m , at middle 0.8-
1.2 m m diam, shallowly grooved ventrally; rachis of
longer pinnae ±3.5-8.5 cm, the longer (distal) inter
foholar segments 1-3 cm; lft-pulvinules 0.8-1.8 x
1-1.3 m m , coarsely wrinkled; lfts accrescent distally,
the blades subsymmetrically lance-elliptic-acuminate
from semicordate base, straight or a little incurved
beyond middle, at very apex either acute or obtuse-
apiculate, the furthest pair 3-10.5 x 0.8-3.5 cm,
2.3-3.7 times as long as wide; venation pinnate or es
sentially so, the weak primary nerve posterior to the
centric or moderately displaced midrib no stronger
and hardly longer than the first secondary nerve on
posterior side of blade, the major secondary nerves
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 111
±7-10 on each side, all these together with tertiary and
reticular venules prominulous on both faces. Peduncles
(2-)2.5-6 cm, ebracteate; capitula 8-16-fld, the recep
tacle ±2.5 m m diam; bracts obtusely deltate, <1 mm,
persistent; fls subhomomorphic, the calyx of the fur
thest a httle broader but hardly longer than that of pe
ripheral fls; pedicels scarcely differentiated externally,
in section 0.5-0.7 x 0.9-1.1 mm; perianth sub-
membranous, 4—5-merous, dark red, glabrous except
for micropuberulent orifice of calyx, the calyx ±8-10-
nerved, the corolla externally nerveless; calyx campan
ulate subtumid ±2.2-2.7 x 2 mm, the broad depressed
teeth ±0.3 mm, but 1-2 sinuses sometimes deeper;
corolla (in protologue "6-")l 1-13 mm, the lance-ovate
lobes 1-2.8 m m ; androecium 20-22-merous, ±5 cm,
the tube as long as corolla or exserted to 2.5 mm; disc
of peripheral fls 0.6-1.1 mm; ovary not seen. Pods de
scribed and figured as drooping, 15 x 1.5 cm, glabrous,
the valves thickly membranous; seeds in broad view "9
x 6 mm," the testa speckled, pleurogrammic.
In subdeciduous and moist montane forest, in
Guerrero at 1830-2230 m, in Oaxaca at 1000-1130
m, locahzed in s. Mexico and Honduras: Oaxaca,
mun. Pochutla; Guerrero, mun. Quechultenango, and
Atoyac de Alvarez; near 15°N, 86°W in Honduras
(Gualaco). — Fl. VI-XI.
Calliandra erythocephala fits neatly into the arbi
trarily defined ser. Ambivalentes having the geminate
pinnae and ample leaflets of ser. Macrophyllae but
the leaflets 4-5 pairs per pinna, hke those of ser. Ni-
tidae in number but much larger. Its pinnately veined
leaflets resemble those of C. laevis but are 3-4 pairs,
not exactly 1 pair, per pinna.
67. Calliandra rhodocephala J. D. Smith, Bot. Gaz.
(Crawfordsville) 49: 454. 1910. — "Prope viae fer-
reae pontem [or. versus] Puerto Barrios, Depto. Yz-
abal, Guatemala, Maj. [26], 1909, Charles C. Deam
n. 6015." — Holotypus, US!; isotypus, NY!. — An
neslia rhodocephala Britton & Rose, N. Amer. FL
23: 56. 1928. FlG. 13
C. rhodocephala sensu Standley & Steyermark, 1946: 26.
Amply macrophylhdious, arborescent shrubs 2.5-
5.5 m with terete pallid branchlets, except sometimes
for microscopically pubemlent upper face of princi
pal venation glabrous throughout, the chartaceous lfts
bicolored, lustrous dark green or brown-purplish-
olivaceous above, paler duller beneath, the capitula of
vivid red fls borne in axils of coarsely thatched
brachyblasts. Stipules stiffly chartaceous, lance-ovate
from shallowly cordate or dilated base, obtuse,
4.5-14 x 2.2-5.5 m m , early becoming pallid, pluris-
triate, persistent. Lf-formula i/21A-31/2, that of some
brachyblast lvs (not further described) HWi-l; peti
oles stout 2-8 x 1.3-3.5 mm, openly grooved ven
trally, homy-dilated at apex; rachis of longer pinnae
4—11.5 cm, the longer interfoholar segments 2-4.5
cm; lft-pulvinules 1.8-3.3 x 0.9-1.6 mm, coarsely
wrinkled; lfts distally accrescent, inequilaterally el
liptic-acuminate from shallowly semicordate base, at
very apex deltately acute, the furthest pair 9-16 x
4—6.5 cm, 2.3-2.9 times as long as wide; primary ve
nation palmate, the gently incurved midrib displaced
to divide blade 1:1.5-2, the inner posterior primary
nerve divergent from midrib shortly above pulvinule,
incurved-ascending well beyond mid-blade, the outer
primary nerve much shorter, all these together with
secondary and reticular venules obtusely prominu
lous on upper face, more sharply so beneath. Pedun
cles stout, ascending 23-28 mm, l(-2)-bracteate near
or below middle; capitula densely 28-46-fld, globose
in bud, the fls homomorphic, the receptacle ±2 m m
diam; bracts ovate ±1.5 m m , striate, persistent;
pedicels (scarcely or obscurely differentiated exter
nally) 0.3-0.7 x 0.5-0.7 mm; perianth 4-merous, the
calyx finely sharply striate, the membranous red
corolla almost nerveless externally; calyx campanu
late or turbinate-campanulate 2-4 x 1.6-2.5 m m , the
broad depressed teeth ±0.2-0.4 mm; corolla 6.5-10.3
mm, the ovate-oblong lobes 2.8-4 m m ; androecium
14-16-merous, 26-31 mm, the tube 5.5-7.5 m m , in
cluded or shortly exserted, the stemonozone 0.8-1.3
mm, the tassel red; intrastaminal nectary 0.5 x 0.4—
0.6 mm; ovary linear-ellipsoid glabrous. Pods (seen
only after dehiscence) 9-12 x 1-1.2 cm, the coria
ceous valves glabrous; seeds unknown.
In wet forest of coastal plain and bordering foot
hills, l-350(-550) m, locally plentiful around the
Gulf of Honduras in e. Guatemala and n. Honduras;
s.-e. Nicaragua; and reported (Standley & Steyer
mark, 1946) from Belize. — Fl. V, VIII-I, the cycle
incompletely known.
Distinctive characters of C. rhodocephala are its
large stipules, its leaf-formula, and its relatively few,
all-red filaments. It resembles the distantly allopatric
C. haematocephala var. boliviana in leaf-formula, but
the androecium is quite different.
D/F. Series MACROPHYLLAE
B e n t h a m
Calliandra sect. Androcallis ser. Macrophyllae
Bentham, London J. Bot. 3: 94. 1844. — Sp. lecto-
typica: C. trinervia Bentham.
112 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
FIG. 13. Calliandra rhodocephala J. D. Smith.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 113
Calliandra ser. Macrophyllae sensu Bentham, 1875: 537-541, exclus. spp. # 1, 3-5, 21.
Anneslia Tergeminae, Laeves Britton & Rose, 1928: 49-50, in clave, nom. nud.
Macrophyll trees and shmbs; phyllotaxy distic
hous; pinnae exactly 1-jug. and lfts 1-, Wi-, 2-, excep
tionally (C. brenesii) Vz-jug., the distal pair mostly 4-
19 cm, but in few, more xeromorphic species only
(1.5-)2^ cm. — Spp. 11, mostly of moist, lowland
or montane forest, but some xerophytic, in tropical
deciduous woodland; widespread from tropical Mex
ico, Central America, and the Lesser Antilles s. to s.-
e. Brazil, Paraguay, and sub-Andean Bolivia.
68. Calliandra trinervia Bentham, London J. Bot. 3:
94. 1844. — Typus infra sub var. trinervia indicatur.
Macrophyll arborescent shrubs and trees com
monly 10 m or less but in far western Amazonia
attaining 30(-35) m x 2 dm dbh, with gray-pallescent
trunks and annotinous branches, commonly glabrous
throughout except for thinly minutely pubemlent
corolla but occasionally the young stems and lf-axes
thinly pilosulous or (var. pilosifolia) densely setose-
pilose, the ample, thinly chartaceous lfts bicolored,
lustrous dark green (when dry brunnescent) above,
paler beneath, the epiphyllum sometimes pubemlent
along primary nerves, rarely pilose along nerves
beneath and cihate, the capitula borne either a) singly
(paired) at 1-2, stipulate but efoliate nodes of small
thatched brachyblasts axillary to a contemporary or
lately fallen If, or b) in var. paniculans at nodes of
extended axillary and terminal pseudoracemes; phyl
lotaxy distichous. Stipules most often bluntly deltate
or broadly triangular 0.6-4 x 1-3.5 mm, less often
lanceolate and 3-6 m m or (var. pilosifolia) attaining
10(-22) x 1-2 m m , striate at least when young but
sometimes thickened in age, becoming palhd and
externally nerveless, persistent. Lf-formula i/1, Wi, 2,
the proximal posterior lfts, when present, Vi or <Vi as
big as the further pair and inserted well below mid-
rachis; lf-stk of larger primary lvs (0.3-)0.5-4(-4.5)
cm, at middle 0.6-1.7 m m diam, distended and
bicupular at apex, the ventral groove wide and shal
low; pinna-rachises of larger lvs (1-) 1.4-4.5 cm;
lft-pulvinules 1.5-3(-3.4) x 0.8-1.6 mm, coarsely
cross-wrinkled; blade of lfts inequilaterally or sub-
dimidiately elliptic or ovate from cuneate or broadly
cuneate base, either shortly bluntly or attenuately
acuminate, the distal pair in larger lvs 5.5-16(-17) x
(2-)2.5-6(-7) cm, ±1.9-3.1 (-3.7) times as long as
wide; venation palmate-pinnate, of 3 ^ primary
nerves from pulvinule, the gently incurved midrib
displaced to divide blade 1:1.8-2.5, pinnately ±7-10-
branched on both sides, the strong inner posterior
primary nerve incurved-ascending through at least 2A
length of blade and often further, the outer l(-2)
much shorter, all these usually prominulous on both
faces, especially in young lvs, the tertiary and reticu
lar venulation either sharply defined on both faces of
blade or immersed on upper one. Peduncles (0.35-)
10-40 mm, minutely bracteate either above or below
middle (exceptionally ebracteate or the bract decidu
ous); capitula 9-27-fld, the floral receptacle sub-
spherical or claviform 1-3 mm; bracts ±0.5-1 mm,
incurved, persistent; fls (of most capitula, perhaps po
tentially of all) heteromorphic, many peripheral ones
staminate, few bisexual, the perianth of 1-3 central
fls scarcely longer than that of peripheral ones but
often stouter and its androecial tube always much
longer, distended distally into a pallid trumpet; peri
anth of all fls either 4- or 5-merous, thin-textured, the
striate calyx usually glabrous, the corolla-tube com
monly pubemlent or appressed-strigulose but some
times glabrous, rarely (var. peruicola) papillate:
PERIPHERAL FLS: pedicel often not or scarcely dif
ferentiated externally except by discoloration, visible
in cross-section, or distinct externally and to 0.7 m m ;
calyx either campanulate, or deeply campanulate, or
cylindric to cylindric infundibuliform, prevalently
0.8-3.7 x 0.6-1.4(-l.8), in var. carbonaria to 2.7
mm, the broad obtuse teeth 0.15-0.4 m m (one sinus
may be more deeply split), in rare var. stenocylix
(q.v.) longer and longer-toothed; corolla (5—)5.5—12.5
(in vars. pilosifolia and stenocylix potentially to 14)
mm, the ovate, sometimes unequal lobes (1-) 1.2-2.8
mm; androecium 12-28(-32)-merous, (21-)25-52
mm, the stemonozone ±0.8-1.2 m m , the pallid tube
(5-)7-18(-22) mm, as long as corolla or far exserted,
the tassel pink or carmine, exceptionally white; ovary
at anthesis glabrous; C E N T R A L FL(S): like the
peripheral ones but the androecial tube 15-25 mm,
expanded to 2-4.5 m m diam at orifice; a tub-shaped
intrastaminal nectary present but no ovary. Pods pur
plish-brown or fuscous, glabrous or exceptionally cil
iate, in profile (7-)8-25 x 0.9-2 cm, the sutural ribs
(2.5-)3.5-7 m m wide in dorsal view, the plane hg
nescent valves usually weakly venulose, the
obliquely transverse nerves immersed or only slightly
obtusely elevated; seeds compressed-lentiform, in
broad view oblong or oblong-elhptic 12-22 x (6-)
7-12 mm, the smooth thin testa becoming papery,
castaneous or light brown, lustrous, pleurogram in S.
America 0, in N. American var. arborea present.
114 M E M O I R S OF T H E N E W Y O R K BOTANICAL G A R D E N [VOL. 74(3)
Calliandra trinervia, as defined in the foregoing
description, is the one common Amazonian species of
the genus characterized by amply 4-, 6- or 8-foholate
leaves, palmate-pinnate venation of leaflets, bicolored
filaments, and weakly venulose pod. Sympatric or
closely vicariant relatives are: C. bombycina, ecologi
cally modified in seasonally dry woodland of the
Huallaga Valley in P e m and morphologically differen
tiated by striate corolla and more numerous stamens,
red throughout; C jariensis, local in Para, and C. har-
risii, of the Andean foothills in Bolivia and of extra-
Amazonian Brazil, together characterized by pinnate
venation of the leaflets; C. glyphoxylon of the west
slope of the Andes in Ecuador, which see for differen
tial characters; and C. hymenaeodes, of Atlantic low
lands in the Guianas, deceptively similar except for
simply pinnate, not bipinnate, leaves. Outside Amazo
nia the species is represented further by scarcely dif
ferentiated varieties localized in northwestern Pem,
inter-Andean southern Colombia, and tropical North
America, as described in the following pages.
Key to the varieties of C. trinervia
1. Distributed in South America; seed-coat without
pleurogram. 2. Calyx of peripheral fls 0.8-3.3(-4) mm,
at most XA as long as the corolla, this in
Brazil <10 mm, to 12.5 m m only in Amazonian Peru and adj. Colombia and
Ecuador. 3. Peduncles arising either from densely
thatched axillary brachyblasts <1 cm or appearing directly axillary to coeval lvs.
4. Young stems and lf-axes either glabrous or pilosulous with hairs
<0.5 mm; widespread. 5. Corolla not papillate; valves
of pod weakly obliquely and openly venulose or evenulose;
Amazon basin and inter-Andean
Colombia. 6. Androecium palhd proximally,
pink-carmine distally; lowland and premontane
Amazonia, 80-1100
(-1300) m 68a. var. trinervia
6. Androecium dark red;
upper Cauca Valley in
Colombia, 1300-2100 m 68b. var. carbonaria
5. Corolla papillate; valves of
pod densely horizontally
venulose; isolated in n.-w.
Peru (Tumbes) 68c. var. peruicola
4. Young stems and lf-axes pilose with
straight, vertically erect hairs to
0.8-2 mm; s.-w. Venezuela and
adj. Brazil 68d. var. pilosifolia
3. Peduncles arising from efoliate nodes
of extended, either lateral or terminal
branchlets, separated one from the next
by clearly evident internodes, forming
pseudoracemes of capitula 1-5 cm;
sub-Andean, from s.-w. Colombia to
n.-e. Bolivia 68e. var. paniculans
2. Calyx of peripheral fls 6.5-8 mm, nearly
l/2 as long as corolla, this 13-14 mm; local
on upper rio Solimoes in Amazonas,
Brazil 68f. var. stenocylix
1. Distributed in s. Mexico (s.-e. from Guerrero)
and Central America (s.-e. to centr. Costa
Rica); seed-coat pleurogrammic. . . . 68g. var. arborea
68a. Calliandra trinervia Bentham var. trinervia.
C. trinervia Bentham, 1844, I.e., sens. str. — "On
the Rio Negro, North Brazil, [Riedel, commun.]
Langsdorff." — Holotypus, dated and numbered
"1838 [year of acquistion from LE, not of collec
tion], No. 20," K (hb. Benth.)! = N Y Neg. 7949!;
presumed isotypus, Riedel s.n., collected at Barra
[do Rio Negro = Manaus], Oct 1828 (fl, fr), NY!. —
Feuilleea trinervia O. Kuntze, Revis. Gen. PL 1:
189. 1891.
C. trinervis var. parvifolia Huber, Bol. Mus. Paraense Hist.
Nat. 5: 379. 1909. "[Para:] Rio Mapuera, acima do Castan-hal [±0°10,S, 58°10'W], 7 XII. 07 ([A Ducke] 9064)." —
Holotypus, M G = F 602347, photo + clastotypus (If, capit
ulum)!. C. rotundifolia Killip & Macbride in Macbride, Field Mus. Nat. Hist., Bot. Ser. 13 (Fl. Peru), 3(1): 73. 1943. —
"[Peru.] Loreto: Mishuyacu, Klug 421." — Holotypus,
US!; isotypus, NY!.
Seldom more than 10 m tall, commonly glabrous
except for pubemlent corolla-tube, the lf-axes rarely
pubemlent with incurved hairs <0.4 m m . Stipules
usually deltate 1.5-4 x 1-3.5 m m , locally (in s.-e.
Colombia and adj. Venezuela) lanceolate 3-6 x 1-2
m m . Petioles (0.3-)0.5-4(-4.7) cm; lfts 2-3(-4) per
pinna, the distal pair mostly (5.5—)7—16 x 3-7 cm;
P E R I P H E R A L FLS: calyx glabrous or exceptionally
pubemlent, campanulate to cylindric, 0.8-3.7(-4) x
0.6-14(-l.8) m m , 1.5-3.5(-4) times as long as diam,
the teeth not more than 0.4 m m ; corolla (5—)6—11
(-12.5) m m ; androecium 12-28(-32)-merous, 26-52
m m , the tube (5—)6—14(—18) m m . Pod of the species.
In lowland rain forest, often riparian, sometimes
entering the understory, and in igapo woodland, 80-
1100(—1300) m, intermptedly widespread over and
almost confined to the n.-w. half of the Amazon basin,
from the upper Putumayo, Caqueta, and Apoporis
Valleys in Colombia e. to w. Para in Brazil, s. along
the Andean foothills to the Bopi Valley in Bolivia;
one record from French Guiana (Bas-Camopi), and
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 115
one from the headwaters of the Orinoco in T.F Ama
zonas, Venezuela. — Map 31. — Flowering inter
mittently throughout the year, the rhythm of anthesis
of particular populations not known.
Variation in length and in length proportionate to
diameter of the calyx, from the short-campanulate
extreme in the typus of C. trinervia to the tubular
extreme exemplified by Schultes & Cabrera 12787
(NY), is continuous, and the variety could be further
divided only arbitrarily by application of this crite
rion. I have found no reliable correlation between fea
tures of the calyx and indumentum, though the popu
lations in central Amazonia are more frequently
glabrous and have on the average shorter calyces than
those further west. Over most of the variety's range
the stipules are obtusely deltate and not more than 3
m m long, but occasional collections from Colombia,
not otherwise different, have the lanceolate stipules
characteristic of var. pilosifolia. Variation in length of
peripheral corollas and androecia appears random.
From available samples the pod seems to be narrower
in central Amazonia than in Amazonian Ecuador and
Pem. Most collections from western Amazonia here
referred to var. trinervia or var. paniculans have been
misidentified as C. carbonaria. Calliandra rotundi-
folia Killip & Macbride, which I confidently reduce
to var. trinervia, was perceived by its authors as dis
tinct from Peruvian var. paniculans, but was not
directly compared with genuine C. trinervia, a species
inexactly characterized in Flora of Peru.
68b. Calliandra trinervia Bentham var. carbonaria
(Bentham) Barneby, stat. nov. C. carbonaria Ben
tham, London J. Bot. 3: 95. 1844. — "[COLOM
BIA.] Province of Popayan, near the river Palace
and the Rio Blanco, where the inhabitants call it
'Carbonero,' Hartweg, n. 964." — Holotypus, K
(herb. Bentham)!; isotypi, K (herb. Hooker.)! = N Y
Neg. 1991, +B = F Neg. 12331, GH!, NY! = N Y
Neg. 9323. — Feuilleea carbonaria O. Kuntze,
Revis. Gen. PL 1: 187. 1891.
C. carbonaria sensu Bentham, 1876: 538 (exclus. pi. oer
sted.); Britton & Killip, 1936: 134 (exclus. Klug 1716).
Trees attaining 7.5(-?) m, except for sometimes
residually pubemlent peduncles glabrous throughout.
Stipules ovate-deltate 1.5-2.8 m m . Petioles 9-35 x
0.8-1.3 m m ; lfts 2 or 3 per pinna, the pulvinules
±2-2.5 m m ; distal pair of lfts inequilaterally lance- or
elliptic-acuminate or -caudate from asymmetrically
cuneate base, the larger ones 8-12 x 2.2-4 cm, 2.6-
3.7 times as long as wide. Peduncles ±2-3.3 cm;
capitula 18-27-fld; bracts 1 m m or less; fls sub
sessile, so far as known homomorphic, the perianth 4-
merous, red, weakly striate except for externally
nerveless corolla-lobes; calyx campanulate or hemi
spherical 2-3 x 1.1-2.2 m m , the teeth 0.2-0.7 m m ;
corolla 8.5-10 m m ; androecium 25-32-merous,
27-32 m, the tube 9-12 m m , a few filaments separat
ing from tube below its distended orifice, this ran
domly fringed with rudimentary filaments. Pods to 22
x 1.6 cm, glabrous, the plane coriaceous valves
obliquely venulose and reticulate; seeds unknown.
Along streams in montane woodland, 1300-2100
m, localized in the upper Cauca Valley in Cauca,
Colombia. — M a p 31. — Fl. I, IV, VIII-IX, the full
range not exactly known. — Carbonero.
Bentham's descriptions of C. trinervia and C. car
bonaria, published simultaneously and each based on
only one collection, call for no contrast between them
other than the shghtiy narrower (falcate-oblong, not
falcate-ovate) distal leaflets of C. carbonaria com
bined with shghtiy longer (2, not 1, m m ) calyx and
shorter (2, not 3V2, lines) corolla. Specimens of C.
trinervia collected in Amazonia in recent decades
show that both calyx and corolla vary greatly, and
independently, in length, and no alternative criteria
have been found. Calliandra carbonaria is conse
quently interpreted as a montane derivative of the
trinervia complex feebly distinguished by leaflets on
the average a little narrower, uniformly red filaments,
and dispersal.
68c. Calliandra trinervia Bentham var. peruicola
Barneby, var. nov., omnino fere cum var. arborea
(mexicano-centramericana) congrua, sed ab ea flo
rum periphericorum perianthio 4-, nee 5-mero,
leguminis suturis setulosis (nee glaberrimis) val-
vulisque confertim transverse (nee aperte oblique)
venulosis, necnon patria peruviana pacifica abstans.
— PERU. Depto. Tumbes, prov. Jaramillo: Bosque
Nacional de Tumbes, Quebrada Trapazola near
Campo Verde, 21.XII. 1967 (fl, fr), D. R. Simpson
(with J. Schunke V.) 435. — Holotypus, NY.
Arborescent to 4—5 m, with weakly geotropic
branches, in aspect and inflorescence closely resem
bling var. arborea; distal pair of lfts elliptic-acumi
nate 5.5-7 x 2-3 cm. Calyx of peripheral fls ±2.4 x
1.1 cm, striate, glabrous; corolla ±6.6 m m , glabrous,
thinly papillate; androecium 14-merous, the tube ±7
m m , the filaments uniformly red. Pods 7-9.5 x 1-1.3
cm, the dilated sutures setulose, the valves densely
subhorizontally venulose.
116 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
M A P 31. South American distribution of Calliandra trinervia Bentham vars. trinervia, paniculans Barneby, stenocylix
Barneby, pilosifolia (Cowan) Barneby, and carbonaria (Bentham) Barneby.
In moist quebrada bottoms, 600-800 m, known
only from the type-locality near 3°40'S, 80°20'W, in
the Peruvian department of Tumbes. — M a p 31. —
FL XI-XII(-?).
The one known locality of var. peruicola is remote
from other varieties of C trinervia, but the plant has
no substantial differential characters other than the
papillate corolla (which occurs in some populations
of Central American var. arborea), the pilosulous su
tures of the pod, and the dense horizontal venulation
of its valves. The dark red corolla and androecium
recall Colombian var. carbonaria.
68d. Calliandra trinervia Bentham var. pilosifolia
(Cowan) Barneby, stat. nov. C. pilosifolia Cowan,
M e m . N e w York Bot. Gard. 10: 142. 1958. — "B.
Maguire & L. Politi 28629 ... along trail from Base
Camp, 125 m, Cerro Sipapo, Terr. [Federal] A m a
zonas, Venezuela, Jan 25, 1949." — Holotypus,
NY!; isotypus, K!; paratypi, Maguire & Politi
28721, Maguire & al. 37510, both NY!.
Trees ±4-5 m, with habit, foliage, perianth and an
droecium of the species, but the young stems and lf-
axes densely pilose with straight, vertically erect
hairs to 0.8-2 m m , the lfts ciliate on margin and on
principal nerves of hypophyllum. Stipules of primary
lvs lanceolate 4-11 (-22) x 1-2 m m , striate, glabrous
dorsally. Petioles 3-20 m m ; lfts 2, 3, or 4 per pinna,
the proximal pair opposite, the distal pair 7.5-13 x
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 117
3-5.5 cm. Calyx glabrous; corolla tube either glabrous
or thinly strigulose; PERIPHERAL FLS: pedicel
0.4-0.7 m m ; calyx 2.2-4 x 1-1.7 mm; corolla 9.5-14
mm; androecium 16-26-merous, 30-52 mm, the tube
12-22 m m , the tassel either crimson or (rarely) white;
C E N T R A L FL(S): androecial tube 28-34 mm, ex
panded at orifice to 3.5^.5 m m diam; disc 0.4-0.8 x
0.5-0.6 mm. Pod not seen.
In wet forest or at forest margins, in Venezuela on
terra firme at 125-600 m, on the upper rio Negro in
Brazil in flooded riparian forest, of local dispersal on
and near the Orinoco-Negro divide in s.w. Amazonas,
Venezuela (C. Sipapo and Casiquiare region) and n.-
w. Amazonas, Brazil (within lat. 3°N and 0°30'S). —
Map 31. — Fl. in Venezuela I—II, in Brazil VII, the
whole season not estabhshed.
The pilose indumentum of C. pilosifolia is its single
feature consistently foreign to C. trinervia sens. lat.
and falls short of a specific character. Lanceolate stip
ules, much exaggerated in Maguire 28421 (NY), are a
noteworthy feature of the variety, but occur also in
some forms of glabrate var. trinervia. The flowers of
the Venezuelan type and paratypes of C. pilosifolia are
relatively large, but not larger than those of all C.
trinervia. A smaller-flowered variant has been distin
guished in manuscript as C. julianii Brunner & Forero.
68e. Calliandra trinervia Bentham var. paniculans
Barneby, var. nov, ab aliis speciei varietatibus inflo-
rescentiae tantum forma diversa, capitulis axi
florigero efoliato e foliomm primariorum axillo et
nonnumquam terminali producto pseudoracemoso-
paniculatis (nee e brachyblastis brevissimis imbrica-
tim stipulatis orientibus) praestans. — Pem. Depto.
Amazonas, prov. Bagua: rain forest along rio Santi
ago 3-5 km above mouth, 250-300 m, 8-13 Oct
1962 (fl), /. /. Wurdack 2168. — Holotypus, NY.
Trees 6-28(-35) m, sometimes precociously flow
ering at 4—5 m, the young stems and the If- and inflo
rescence-axes strigulose with incurved or subap
pressed hairs <0.45 mm, the lfts glabrous. Stipules
obtusely deltate or ovate-triangular 1.5-3.5 x 1.3-3
mm, striate. Petioles 1-3 cm; lfts 2, 3,4 per pinna, the
small proximal pair, when both present, not exactly
opposite, the distal pair 8-16(-17.5) x (2.5-)3.2-6.5
cm, 2.1-3.2 times as long as wide. Axis of florigerous
brachyblasts 1-5 cm; PERIPHERAL FLS: calyx
2.3-3.3 x 1.6-2.2 m m , the teeth 0.3-0.6(-0.8) mm;
corolla 5.8-8.2 mm; androecium (10-)12-16(-20)-
merous, 34-^4(-50) mm, the palhd tube (5.5-)7-ll
m m , as long as corolla and to 3.5 m m longer, the tas
sel pink-carmine. Pods 10-18 x 1.2-2 cm.
In rain forest, often along river banks, surviving
disturbance, above (200-)250 m and ascending in
Ecuador to 1400(-1500) m, intermptedly dispersed
along e. foothills and intermontane valleys of the
Andes between 1°30'N in Colombia and 16°S in
Bolivia, most common in Ecuador and n. Pem. —
Map 31. — FL III, V, VIII-XII. — Machete vaina;
urcu shimbillo (Pem).
At full anthesis var. paniculans is readily distin
guished from vicariant var. trinervia by its pseudo
racemose inflorescence, but less so when immature or
in fmit. It has often passed in herbaria as allopatric
(Colombian) C. carbonaria, but differs not only in
inflorescence architecture but also in bicolored, only
distally carmine, not uniformly red androecium.
68f. Calliandra trinervia Bentham var. stenocylix
Barneby, var. nov, habitu, pube necnon foliolorum
inflorescentiaeque forma cum var. trinervia arete
consimilis, sed ab ea calyce subcylindrico elongato
6.5-8 (nee 0.8^) m m longo longidentato insigniter
diversa. — BRAZIL. Amazonas, mun. Amatura:
Estacao Ecologica Jutaf-Solimoes, margem direita
do rio Sohmoes, 8 May 1986 (fl mat), C. A. Cid
Ferreira (& al.) 7354. — Holotypus, NY.
Trees attaining 7 m, like var. trinervia in habit, fo
liage, and inflorescence, glabrous except for strigu
lose perianth. Stipules (few seen) lanceolate ±4—6 x
1.5-2 mm. Petioles ±1 cm; lfts 3 per pinna, the distal
pair ±13-14 x 5-7 cm. Peripheral fls staminate, the
calyx 6.8-8 x 2.4 mm, its teeth ±2.2 mm, the corolla
13.5-14 mm, its lobes to 2.2 mm; androecium ±4.7
cm, 28-merous, the tube 17 mm, the tassel pink; mod
ified tmmpet fls present but not closely examined.
Pod unknown.
In forest on terra firme, near 150-200 m, known
only from the type-locality on rio Solimoes near 68°W
in state of Amazonas, Brazil. — Map 31. — Fl. IV-V.
The var. stenocylix differs materially from wide
spread var. trinervia only in the modified calyx,
which is about half as long as the corolla, not one-
third or less as long. Its narrow stipules are shared
with var. pilosifolia and with some Brazilian and
Colombian forms of var. trinervia , and its relatively
numerous (±28) filaments with some sub-Andean
forms of the same.
68g. Calliandra trinervia Bentham var. arborea
(Standley) Barneby, comb. nov. C. arborea Stand-
ley in Yuncker, Field Mus. Nat. Hist., Bot. Ser.
17(4): 365, pi. XII (phototypus). 1938. — "HON
D U R A S . . . near El Rincon, about 10 miles west of
118 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Siguatepeque, Dept. Comayagua . . . July 24, 1936,
T. G. Yuncker, R. F. Dawson & H. E. Youse 6047." —
Holotypus, F!; isotypi, K!, NY!; paratypi, Yuncker
&al. 5972, F!, NY!.
Anneslia centralis Britton & Rose, N. Amer. Fl. 23: 52.
1928. — "Type from near San Pedro, Sula [San Pedro
Sula], Honduras, April, 1889, C. Thieme 5214." — Holoty
pus, NY!. — Calliandra centralis Standley, J. Arnold Arb.
11: 30. 1930. — Equated with C. emarginata by Standley & Steyermark, 1946: 22.
A. rekoi Britton & Rose, N. Amer. Fl. 23: 53. 1928. —
"[MEXICO.] Tepinapa, Oaxaca, altitude 100 meters,
March 23, 1919. B. P. Reko 4130." — Holotypus, US!;
clastotypus + photo, NY!. — Calliandra rekoi Standley,
Publ. Field Columbian Mus., Bot. Ser. 4(8): 308. 1929.
A. splendens Britton & Rose, N. Amer. Fl. 23: 53. 1928. —
"[MEXICO.] Road between Tumlata [= Tumbal ] and El
Salto [de Agua], Chiapas [±17°25'N, 92°20,W], 1895, E.
W. Nelson 3396." — Holotypus, US!; clastotypus (lvs, fls),
NY!. — Calliandra splendens Standley, Publ. Field Columbian Mus., Bot. Ser. 4(8): 308. 1929.
Calliandra rivalis Lundell, Bull. Torrey Bot. Club 64: 549. 1937. — " . . C. L. Lundell 6610, collected in the rocky
bed of Rio Frio at San Agustin, Mountain Pine Ridge, El
Cayo District, British Honduras, July 26, 1936." — Holotypus, MICH!; isotypus, NY!. — Mistakenly equated by
Standley & Steyermark 1946: 22, with C. emarginata.
Arborescent shrubs, flowering at 1.5-3 m but attain
ing 3-9 m with trunk to 1.5 m dbh, glabrous except for
micro-puberulent lf-axes and often thinly granular-
papillate inflorescence, the capitula arising singly or
2(-3) together from few-stipulate, mostly efohate
brachyblasts axillary to coeval or new-fallen lvs. Stip
ules 0.8-3 x 0.4-0.9 m m . Lf-formula exactly i/2; peti
oles (5-) 14—40 m m ; pinna-rachises (10-) 12-30 m m ;
lft-pulvinules (1-) 1.5-3.4 m m ; lft-blades inequilater
ally lance-elliptic-acuminate or subdimidiately ovate-
acuminate from inequilaterally cuneate or shallowly
semicordate base, the distal pair (4.5-)6.5-l 1 x 1.4-5
cm, 2.1-3 times as long as wide. Peduncles 14—36
m m ; capitula 12-22-fld; bracts 0.3-0.7, rarely 2-3
m m ; P E R I P H E R A L FLS: calyx 1.2-3.4 x 0.9-1.6
m m , weakly 5-10-nerved; corolla either greenish-
white or reddish, often punctate, 6.4-7.5(-l 3) m m ;
androecium 10-18(-24)-merous, 19-33 m m , the tube
commonly 4.5-7.5 m m , included or shortly exserted,
exceptionally attaining 15.5 m m , long-exserted. Pods
in profile 8.5-18 x 1.1-1.7 cm, the valves glabrous,
either almost smooth or obhquely, weakly or (in Costa
Rica) sharply venulose; seed-testa brown, often speck
led, pleurogrammic.
In moist lowland forest, wooded ravines, and as
cending to premontane and montane forest, 100-
1650 m, discontinuously widespread in s. Mexico
(Guerrero to n. Oaxaca and adj. Veracruz, Chiapas)
and Central America (Guatemala, s. Belize, Hon
duras, centr. Costa Rica). — M a p 32. — FL V-VIII.
— Quita (Costa Rica).
As defined here, var. arborea is morphologically
intermediate between C. trinervia var. trinervia, of
which it has the general aspect and the relatively large,
more or less acuminate leaflets, and C. tergemina var.
emarginata, which it resembles in pleurogrammic
seed-coat. Without fmit it can be reliably distin
guished from var. trinervia only by its distantly dis
junct dispersal in the Northern Hemisphere (see maps
31,32).
The segregates reduced above to synonymy of var.
arborea each have some slight peculiarity: Anneslia
splendens an anomalously long androecial tube; A.
rekoi relatively short leaflet-pulvinules; and A. cen
tralis the pulvinules of the last with (apparently) dark
red tassel. The last-mentioned has been equated in the
literature with C. emarginata, and could be a variant
of it with acuminate leaflets.
69. Calliandra bombycina Spmce ex Bentham,
Trans. Linn. Soc. London 30: 538. 1875. — "East-
e m Pem on the Huallaga . . . Spruce, n. 4235." —
Holotypus, labeled: "In fluvii Huallaga ripis secus
fl. Mayo ostia [near 6°35'S, 76°20,W, in San
Martin]," K (herb. Benth.)! = N Y Neg. 7990!; iso
typi, K (hb. Hook.)!, NY!. — Feuilleea bombycina
O. Kuntze, Revis. Gen. PL 1: 187. 1891.
C. bombycina sensu Bentham, 1876: 410, in nota sub
C. trinervia: Macbride, 1943: 68, ex parte, exclus.
syn. C. boliviana.
Slender macrophyllidious, arborescent or some
times sarmentose shrubs (2-)3-7 m, glabrous except
for rudimentary pubemlence of some new branchlets
and for the ventral face of some lf-stks, the ample
papery lfts sublustrous olivaceous above, paler dull
beneath, the large capitula of red fls arising singly
from either lateral or subterminal, densely or loosely
thatched, mostly efoliate brachyblasts, the peduncles
subtended by relatively large, thick-textured stipules,
most capitula therefore elevated above coeval foliage.
Stipules obtuse deltate-triangular from subcordate
base, 2.5-5.5 x 2^.5 m m , dorsally 11-19-nerved,
persistent. Lf-formula i/(l1/2-)2, the smaller lower
pair of lfts often unequal in size and not quite oppo
site, the anterior 1ft sometimes wanting; lf-stks (1-)
1.5-5 cm, at middle 0.8-1.9 m m diam, shallowly sul
cate ventrally; rachis of pinnae 2-7 cm, one member
of some pairs shorter than its fellow, all dilated at
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 119
M A P 32. Distribution of Calliandra trinervia Bentham var. arborea (Standley) Barneby in Mexico and Central America.
apex into narrowly winged sockets accommodating
the pulvinules; lft-pulvinules 2-4.5 x 1-2 m m ,
coarsely wrinkled; lft-blades obliquely elliptic, (ob)
ovate- or broadly lance-elliptic from inequilaterally
broad-cuneate base, nearly always shortly acuminate
and at very apex obtuse mucronulate (shallowly
emarginate), the distal pair (7-)8-16 x (2.8-)3.8-7.5
cm, (1.7-)2-2.8 times as long as wide; venation
palmate-pinnate, the forwardly incurved midrib dis
placed to divide blade ±1:2, the inner posterior
primary nerve incurved-ascending well beyond mid-
blade, the outer 1-2 primary nerves shorter, the sec
ondary and reticular venules sharply elevated on both
faces. Peduncles 12-23 m m , bracteate near middle;
capitula 20-28-fld, the clavate receptacle 2-4 m m ,
the fls homomorphic, subsessile or obscurely stoutly
pedicellate; bracts ovate or oblanceolate 0.8-1.8 m m ,
striately nerved, persistent; pedicels 0.4-1 x 0.6-1.2
m m ; perianth 5-merous, the calyx sharply striate,
glabrous or almost so, the red corolla strigulose-
pilosulous, striate but more finely so; calyx campan
ulate 2-3.5 x 2.2-3.2 m m , the broad obtuse teeth 0.25-
0.9 m m , but one or more sinuses often more deeply
split in age; corolla broadly tubular-campanulate
8.5-13 x ±4-5 m m , the ovate lobes 2-3.2 m m ;
androecium 44—54-merous, 45-62 m m , the stemono
zone 1.8-2.2 m m , the tube 8-12 m m , either a little
longer or a little shorter than corolla, the tassel vivid
red or red-violet; intrastaminal disc drum-shaped
0.7-1 m m ; ovary glabrous. Pods (few seen) probably
erect, in profile ±9.5-13 x 0.8-1.2 cm, ±8-seeded,
the valves and sutural ribs smooth glabrous; seeds
unknown.
In seasonally dry forest, 200-650 m, localized in
the lower valley of rio Huallaga and its tributaries, be
tween 6°S and 8°S in depto. and prov. San Martin,
Pem. — M a p 33. — FL XII-V, IX, the full cycle not
documented by specimens.
Calliandra bombycina resembles C. trinervia in
many respects, but differs in mostly 8-foliolate
leaves, finely striate corolla, and more numerous (44—
54, not 12-32) filaments of uniformly dark red color.
Spmce recorded that the flower-heads were worn as
hair-omaments by the native women.
70. Calliandra glyphoxylon Spmce ex Bentham,
Trans. Linn. Soc. London 30: 539. 1875 (var. ex
clus.). — "Pallatanga in Ecuador . . . Spruce, n.
5571." — Holotypus, K (hb. Benth.)! = N Y Neg.
7992; isotypi, C = F Neg. 218041, K (hb. Hook.)!,
NY!. — Feuilleea glyphoxylon O. Kuntze, Revis.
Gen. PL 1: 188. 1891.
C. pallatangensis Harms, Repert. Sp. Nov. Regni Veg. 17: 87. 1921. — "Ecuador: Pallatanga, 2000 m (A. D. Sodiro
no. 393.—Aug. 1891)." —Holotypus, *B = FNeg. 7252!.
Macrophyll arborescent shrubs 1.5-4 m with
densely foliate, pallid branches, the new stems and
lf-axes and the dorsal face and margin of lfts pilosu
lous with fine straight erect hairs to 0.4-0.8 m m , the
chartaceous low-convex lfts lustrous olivaceous
above, glabrous and a httle paler beneath, the dense
hemispherical capitula arising singly from the first
node of efohate brachyblasts axillary to coeval lvs
of homotinous branches, nestled in foliage. Stipules
deltate-ovate to lance-acuminate 3-10 x 1-2.4 m m ,
striate-nerved, tardily deciduous. Lf-formula i/2, the
smaller proximal pair alternate; lf-stks 1.5-8 x 0.8-
1.5 m m , distended at apex into 2 cupules, the ventral
groove continuous; pinna-rachises of larger lvs
1—2.5(—3) cm, the proximal pair of lfts inserted far
below middle; lft-pulvinules in dorsal view 0.7-1.2
m m , the lfts in ventral view appearing sessile; lfts
inequilaterally or subdimidiately elhptic or ovate from
shallowly semicordate base, shortly obtusely acumi
nate, the larger ones 5-9.5 x (1.8-)2-4.5 cm, 1.45-
2.7 times as long as wide; venation of 3-4 nerves from
pulvinule, the straight or more often gently incurved
midrib forwardly displaced to divide blade ±1:1.5-2,
pinnately few-branched on each side, the strong inner
posterior primary nerve incurved-ascending through
± % of blade, the outer posterior ones much shorter, all
these together with tertiary venules and sinuous retic
ulum prominulous on both faces of blade. Peduncles
8-25 m m , at least sometimes bracteate above middle,
120 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
MAP 33. Distribution of Calliandra glyphoxylon Spruce ex Bentham, C. bombycina Spruce ex Bentham, and C. jariensis Barneby in South America.
the bract sometimes caducous or perhaps obsolete, ca
pitula ±30-45-fld, the floral receptacle 2 m m or less,
the fl-buds prior to anthesis compressed into a ball;
floral bracts subulate ±1 mm, tardily deciduous; fls
homomorphic as to perianth, the stamen-tube of upper
ones sometimes longer; perianth either 4- or 5-mer
ous, submembranous, glabrous except for microciho-
late calyx-teeth and occasionally minutely pubemlent
corolla-lobes, the calyx weakly striate-nerved, the co
rolla not so, both fuscous-nigrescent when dry;
pedicels scarcely differentiated externally, 0.3-0.5 x
0.5-0.6 mm; calyx campanulate 2-2.3 x 1.3-1.6 mm,
the teeth 0.3-0.6 mm; corolla 8-9.5 mm, the lobes
2-2.6 mm; androecium 16-18-merous, 23-34 mm,
the stemonozone ±0.8 mm, the tube 8-13.5 mm, as
long as corolla or well-exserted, the tassel at first
white or pale pink, becoming crimson in age; intrast
aminal nectary of outer, mostly staminate fls ±0.6
mm. Pods (few seen) 9.5-11 x 1.2-1.4 cm, either
glabrous or minutely remotely pubemlent, the stout
sutural keels in dorsal view 3-4 m m wide, the re
cessed valves densely transverse-venulose.
In thickets on steep slopes of quebradas between
600 and 2100 m, known only from the basins of rios
Chanchan and Chimbo on the slope of the Ecuado
ran Andes, near lo±50'-2°20'S in prov. Chimborazo.
— Map 33. — FL V-IX. — Palo de las siete vueltas,
"the branches traversed under the bark by seven or
more, slightly spiral striae."
Calliandra glyphoxylon so closely resembles the
widespread polymorphic C. trinervia (sens, lat.) that
its status as an independent species is precarious. In
practice, however, its copious loose indumentum,
relatively large coarse stipules, relatively short peti
oles, and densely many-flowered capitula (all de
scribed in Key IV) in the context of a narrow discrete
dispersal on the Pacific slope of the Andes distin
guish it handily.
71. Calliandra jariensis Barneby, sp. nov, omnibus
notulis cum affini C. trinervia congrua nisi folio-
lorum venatione quasi pinnata, nervo primario pos
teriori debili quam nervos secundarios e costa ortos
nee longiori nee fortiori praestans. — BRAZIL.
Para: Jari, estrada entre Monte Dourado e Planalto
A, 19 Mar 1969 (fl), N. T. Silva 1807 . — Holo
typus, NY. — A wood sample (n.v.) was collected.
Fig. 14
Macrophyll trees 3-10 m with trunk attaining 1-
1.5 dm dbh, glabrous except for minutely puberulent
peduncles and fl-buds or for sometimes remotely
minutely ciholate lfts, the ample chartaceous lfts dull
olivaceous above, paler brownish-olivaceous beneath,
the small hemispherical capitula arising singly from
stipulate but efoliate nodes of brachyblasts axillary to
coeval or new-fallen primary lvs, immersed in hor-
notinous foliage. Stipules triangular-subulate ±1-1.5
mm, firm, externally nerveless or faintly striate when
young, those of primary lvs deciduous, those of
brachyblasts persistent. Lf-formula \l\Vi, the odd prox
imal 1ft of each pinna inserted far below mid-rachis
SILK TREE, GUANACASTE, MONKEY'S EARRING 121
FiG. 14. Calliandra jariensis Barneby.
122 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
and <Vi as long as distal pair; lf-stks including fuscous
pulvinus 1.5^.3 cm, at middle 0.6-1.1 m m diam,
dilated and bicupulate at apex, the ventral groove shal
low and narrow; rachis of pinnae 11-25 m m ; lft-
pulvinules (1-) 1.5-2.5 m m , lustrous, cross-wrinkled;
distal lfts inequilaterally broad- or ovate-elliptic
from semicordate base, very shortly obtusely (some
times obscurely) acuminate, the distal pair 6-13 x
2.7-7 cm, 1.8-2.2 times as long as wide; venation
essentially pinnate, the gently incurved midrib dis
placed to divide blade ±1:1.35-1.5, giving rise on
anterior side to ±7-8 incurved secondary nerves
brochidodrome well within the loosely revolute mar
gin, the inner of 2 posterior primary nerves incurved
to anastomosis well short of mid-blade, the outer one
very short, these all, together with transverse tertiary
and openly reticular venules finely prominulous on
both faces. Peduncles slender 8-18 m m , ebracteate;
capitula 14—18-fld, the fls outwardly homomorphic
(though some functionally staminate), the receptacle
not over 2 m m ; bracts subulate, scarcely 0.5 m m ,
persistent; perianth membranous glabrous 4-merous,
the calyx and corolla weakly nerved or the calyx
sometimes weakly striate; calyx in external view
either campanulate or turbinate-campanulate 1-1.2 x
1 m m , but the lower half of calyx solid (= a cryptic
pedicel) and the tme calyx reduced to a shallowly
campanulate subtruncate limb 0.5-0.6 m m ; corolla
5.2-5.6 m m , the lobes 1.4—2 m m ; androecium
(14—) 16-22-merous, 16-27 m m , the stemonozone
<1 m m , the tube ±4.5-5.5 m m (a few filaments often
separating near mid-tube), the tassel reportedly
either white and distally rubescent, or red; ovary of
bisexual fls arising at level of top of stemonozone,
obscurely stipitate, at anthesis glabrous. Pods (few
seen) almost linear beyond the backwardly attenuate
base, ±17-21 x 1 cm, minutely appressed-puberulent
overall, the hgnescent sutural keels in dorsal view ±4
m m wide, the plane recessed valves coarsely venose
lengthwise with subparallel, randomly braided fibers;
seeds unknown.
In virgin and disturbed (capoeirao) rainforest on
terra firme below 200 m, known only from the lower
Jarf basin in mun. Almeirim, state of Para, Brazil. —
M a p 33. — Fl. III-VI. — Mororo.
Calliandra jariensis differs from other Macrophyl
lae, and in particular from C. trinervia var. trinervia
for which it has been mistaken, in the venation of the
leaflets, which lack the strong inner posterior nerve
otherwise universal in the series. Its small flowers with
minute tme calyx and its vertically venulose pods are
further distinctive features.
72. Calliandra coriacea (Willdenow) Bentham, Lon
don J. Bot. 3: 95 (with query, the plant itself un
known). 1844. Inga coriacea Humboldt & Bonpland
ex Willdenow, Sp. PL 4(2): 1010. 1806. — "Habitat
in America meridionali." — Holotypus, Humboldt
s.n. in B-WILLD 79077, seen in Microform!. —
Mistakenly equated by Bentham, 1875: 539, with
Calliandra emarginata.
C. glyphoxylon var. glaberrima Bentham, Trans. Linn. Soc.
London 30: 539. 1875. — "[Colombia.] Valley of the Mag
dalena, Triana." — Holotypus, Triana s.n. from "Vallee du
Magdalena, province de Mariquita [= n. depto. Tolima]," K
(hb. Hook.)! = N Y Neg. 2002\; presumed isotypi, Triana
6837, collected at Espinal, prov. Mariquita, 500 m, Jan
1853, COL!, Triana s.n., G!. — Calliandra glaberrima
Britton & Kilhp, Ann. New York, Acad. Sci. 35: 134. 1936.
C. rivalis Lundell, Bull. Torrey Bot. Club 64: 549. 1937. —
"C. L. Lundell 6610, collected in the rocky bed of Rio Frio at San Agustin, Mountain Pine Ridge, El Cayo District,
British Honduras, July 26, 1936." — Holotypus, MICH!;
isotypus, NY!. C. anthoniae Grimes, Brittonia 45: 25, fig. 1. 1993. — "SURINAM, below Hendrik Creek, Coppenam River
Headquarters, 30 Jul 1944 (fl fr), B. Maguire 25068." —
Holotypus, NY!; isotypus, U!. Inga coriacea sensu Kunth in Humboldt, Bonpland & Kunth, 1842: 297, who had seen no specimen.
Anneslia tergemina Kleinhoonte, 1940: 322; non Mimosa
tergemina Linnaeus. Calliandra glaberrima sensu Woodson & Schery, 1950: 257.
C. coriacea var. aquae-nigrae Barneby, in adnot., nom. nud.
Macrophyllidious arborescent shrubs fertile when
(1.5-)3-10(-12) m tall, with trunk attaining 1.5 d m
dbh and terete gray virgate long-shoots, glabrous
throughout except for sometimes minutely pubemlent
or barbellate peduncles and ventral face of lf-axes,
the chartaceous lfts at maturity lustrous dove-gray
or brown-ohvaceous above, paler dull beneath, the
capitula arising singly from lowest 1-3 elaminate
nodes of brachyblasts axillary to homotinous lvs, or
occasionally also from random primary axils; phyl
lotaxy distichous. Stipules mostly ovate or depressed-
deltate, rarely broad-lanceolate, 1-3.5 m m , weakly
striate when young, becoming indurate and externally
nerveless, persistent. Lf-formula i/lVz, each pinna 3-
foliolate, composed of a terminal pair of lfts and a
smaller posterior one near base of pinna-rachis; lf-stk
of primary lvs, including nigrescent cross-wrinkled
pulvinus, 6-39 m m , at middle 0.5-1.2 m m diam, the
ventral groove shallow; rachis of pinnae 5-24 m m ,
either longer or shorter than petiole; lft-pulvinules
(dry) either pallid or nigrescent, cross-wrinkled,
in dorsal view (0.8-)l.l-1.8(-2) x 1-2.4 m m ; lfts
inequilaterally narrow- or broad-elliptic from anti-
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 123
cally cuneate, postically decurrent or incipiently
semi-cordate base, mostly short-acuminate and at
very apex obtuse apiculate (exceptionally shallowly
emarginate, or attenuate and acute), the two distal lfts
(2-)4-10 x l-3.2(-3.5) cm, (1.8-)2.5-5 times as long
as wide; primary venation of (3-)4(-5) nerves from
pulvinule, the gently incurved, pinnately branched
midrib forwardly displaced to divide blade ±1:1.5-2,
the inner posterior nerve almost as strong, incurved-
ascending well beyond mid-blade, the outer posterior
one(s) much shorter, all these together with tertiary
and reticular venules prominulous on both faces. Pe
duncles (6-)l 1—44 mm, almost always bracteate near
or above middle; capitula 11-28-fld, the subglobose
receptacle 1.5-2.5 m m diam; bracts mostly <1 mm,
persistent; fls of each capitulum homomorphic as to
perianth but the androecia sometimes dimorphic, that
of some terminal fls a httle longer and dilated into a
trumpet; pedicels cryptic, <0.6 mm; perianth green
ish-white, glabrous (micropuberulent), either 4- or 5-
merous, the calyx finely striate, the corolla not so;
calyx campanulate or deeply campanulate 1.4—3
(-3.3) x 1.1-2.8 m m , the teeth 0.1-0.35 mm; corolla
8-13 m m , the lobes 1.6-2.7 mm; androecium of pe
ripheral fls 20-58-merous, 3-5 cm, the tube 1-3.5
cm, always at least shortly exserted, the stemonozone
0.6-1.4 m m , the tassel either pink or red-purple;
intrastaminal nectary 0. Pods standing erect from pla
giotropic branches, in broad view 9-13 x 0.8-1.3 cm,
glabrous overall, the dark-castaneous sutural ribs in
dorsal view 3.5-4 m m wide, the plane recessed hg
nescent valves faintly or weakly cross-venulose;
seeds (few seen) broad-elliptic-oblong in broad view,
±7.5-11.5 x 5.5-8 m m , the testa pale brown, papery,
loosely enveloping the embryo, becoming fragile and
sometimes narrowly winged around the periphery;
pleurogram 0.
On rocky river banks and gravelly shores, some
times forming riparian thickets of great extent,
20-1100 m, of discontinuous range between 1° and
18°N in Central America and n. South America: scat
tered between centr. Panama (prov. Code Colon, and
Panama) and extra-Amazonian Colombia (prov.
Choco to middle Magdalena Valley); on black water
streams at 100-230 m on the headwaters of the Orin
oco in Venezuela (Amazonas); in interior Guianas
(headwaters of Essequibo in Guyana, on the Nickerie
Lucie, upper Coppename, Gran, Tapanahoni and Go-
nini rivers in Surinam) and adj. Para, Brazil (middle
and lower rios Trombetas and Paru do Oeste); and
remotely disjunct in Belize (Cayo). — Map 34. — Fl.
in Belize, Panama and w. Colombia II-V, VII-IX, in
Venezuela XII-IV, VII, in the Guianas and Para V-IX,
XI, the full cycle not documented.
The comprehensive concept of C. coriacea ex
pressed in the foregoing synonymy and description
has been built up by accretion of geographic elements
that were mistaken at first for closely related but dis
tinct taxa. The historic nucleus of C. coriacea, based
on a unicate specimen collected at an unrecorded
locality, very likely in Colombia, by Humboldt, was
unknown to Bentham, who redescribed it as a var.
glaberrima of the upland Ecuadorean C. glyphoxylon,
obviously different in copious loose pubescence,
larger stipules, shorter leaf-stalks, 8 (not 6)-foholate
leaves, capitula of not less than thirty flowers, and
few (16-18) stamens per flower. The typical, Colom
bian C. coriacea has all flowers of the capitulum
homomorphic or almost so, with slenderly tubular
androecium 19-35 m m long. In southwest Venezuela
the otherwise similar riparian calliandra has at least
incipiently heteromorphic flowers, with slightly
shorter corolla mostly 8-10 (not 10-13) m m long and
androecial tubes mostly 10-17 m m long. In prelimi
nary studies these plants were segregated as a var.
aquae-nigrae Bameby, a manuscript name that has no
nomenclatural status. The plants of the Guianas and
adjoining Brazil were described as C. anthoniae in
order to correct their previous misidentification, by
Anthonia Kleinhoonte, as C. tergemina (Linnaeus)
Bentham, and to provide a name, no longer needed,
for an account of Calliandra for Flora of the
Guianas, then in preparation. As monographic studies
progressed, nothing of substance could be found to
distinguish C. anthoniae from Colombian C. cori
acea or, for that matter, from the localized Belizean
populations described by Lundell as C. rivalis.
Calliandra coriacea differs ideally from superfi
cially similar C. trinervia (sens, lat.) in nearly elliptic
leaflets with subequally convex anterior and posterior
margins, in fmits stiffly ascending from the branches
like those of C. angustifolia, and in smaller seeds. In
Venezuela, however, the leaflets are sometimes a little
wider on the anterior side of the costa, and the atti
tude of the fmits needs confirmation in the field. The
species as a whole is more easily separable from
related C. angustifolia by its 3(not 4)-foliolate pinnae
and by much larger distal leaflets. Differential char
acters of the also related C. antioquiae are brought out
in the protologue of that species, next following.
73. Calliandra antioquiae Bameby, sp. nov., habitu
foliisque C. coriaceae praesimilis sed ab ea stipuhs
majusculis (4-8 m m longis), foliolis adaxialiter vix
124 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
M A P 34. Distribution of Calliandra coriacea (Willdenow) Bentham in South America.
venulosis (nee reticulatis), corolla striata (nee laevi),
legumine parvulo (5 c m usque longo) necnon semi-
nibus areolaris graviter discrepans. — C O L O M B I A .
Antioquia: vicinity of Medellfn, anno 1927 (fl., fr.)
Rafael A. Tow 109A . — Holotypus, NY. C. glaberrima sensu Britton & Killip, 1936: 134, minore ex
parte, quoad pi. toroanam.
Macrophyllidious treelets 2 m and probably taller,
with virgate long-shoots and densely thatched brachy
blasts, glabrous except for sometimes pubemlent
ventral face of lf-axes or minutely ciholate lfts, the sub-
coriaceous brown-olivaceous lfts scarcely bicolored,
the capitula of reportedly red fls arising singly from
elaminate nodes of brachyblasts axillary to coeval or
lately fallen lvs. Stipules firmly papery, triangular-
lanceolate 4—8 x 1.5-3 m m , lustrous, striately nerved,
persistent. Lf-formula \l\Vi, each pinna consisting of a
perfect distal pair of lfts and one smaller posterior one
inserted 1-2 m m above base of pinna-rachis; petioles
15-26(^0) m m , at middle 0.7-1 m m diam; rachis of
pinnae 7-12 m m ; lft-pulvinules 0.6-1.2 x 1.3-2 m m ,
coarsely cross-wrinkled; lft-blades narrowly or
broadly elhptic from inequilateral base, obtuse apicu
late, the distal pair 3.5-5.6 x 0.7-2 cm, ±2.8-3.7 times
as long as wide; midrib displaced to divide blade
±1:2-2.5, almost straight, the inner posterior primary
nerve almost as strong, produced well beyond mid-
blade, the outer primary nerves progressively much
shorter, the secondary and weak reticular venules
prominulous dorsally, immersed on upper face. Pe
duncles stout, 2-3.5 cm, bracteate below middle; ca
pitula ±20-fld, the subglobose receptacle ±2.5 m m
diam; bracts ovate 1-1.5 m m , striate, persistent; fls
homomorphic; pedicels cryptic 0.3-0.4 m m ; perianth
5-merous glabrous, both calyx and corolla promi
nently striate; calyx campanulate 1.8-3.4 x 2.2^4 m m ,
the depressed-deltate or semicircular teeth 0.25-0.8
m m ; corolla broadly tubular, slightly dilated upward,
8-9.5 m m , the lobes 1.5-2 m m ; androecium 38-58-
merous, 28-40 m m , the stemonozone 0.6-1.6 m m , the
tube 6-8.5 m m , 1-3 m m shorter than corolla; ovary at
anthesis glabrous; intrastaminal nectary 0. Pods 3-5 x
0.6-0.65 cm, glabrous, the marginal ribs in dorsal
view ±2 m m wide, the recessed valves hgnescent, um-
bonate over 1-2 seeds, sharply cross-venulose; seeds
(few seen) compressed-ellipsoid, in broad view ±6.5 x
3.5 m m , the close-fitting testa smooth but not highly
lustrous, brown, the pleurogram narrowly U-shaped 5
x 2 m m .
On rocky riverbanks, forming riparian thickets,
150(-?) m, known only from the lower Cauca basin in
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 125
Antioquia, Colombia (Medellin; rio Nechi). — Fl.
VII(-?).
The typus of C. antioquiae was mistaken by Britton
for the habitally similar C. glaberrima, from which it
differs substantially in relatively large striate stipules,
weakly veined (not sharply reticulate) upper face of
leaflets, prominent striate venation of the corolla,
small narrow pod, and especially in the areolate seeds.
The color of the flower-parts remains to be described
in detail.
74. Calliandra angustifolia Spmce ex Bentham,
Trans. Linn. Soc. London 30: 539. 1875. — ". . .
Eastern Pem, very abundant on the banks of the
Huallaga and the Mayo rivers, Spruce n. 4466." —
Holotypus, K (hb. Benth.)! = N Y Neg. 1993; iso
typi, +B = F Neg. 7227!, G!, K (hb. Hook.)!, OXF!.
— Feuilleea angustifolia O. Kuntze, Revis. Gen. PL
1: 187. 1891.
C. subnervosa Bentham, Trans. Linn. Soc. London 30: 540.
1875 — "[Ecuador.] Guayaquil, Tafalla." — Holotypus, G-BOISS ex hb. Pavon. (in 3 folders, annotated by Ben
tham)!; isotypi, BM!, M A = G photo 294091, fragm. ex Hb.
MA, F!. — Feuilleea subnervosa O. Kuntze, Revis. Gen.
PL 1: 189. 1891. — Note that the "stipels" described in the
protologue are the dilated striate terminal appendages of lf-
stks.
C. sodiroi Harms, Repert. Sp. Nov. Regni Veg. 17: 87. 1921.
— "Ecuador: Prov. Chimborazo, Paramo Navag (A. Sodiro
no. 392 — Aug. 1891)." — Holotypus,^ = F Neg. 1260V, clastotypus (fragm.), F!.
C. stricta Rusby, Mem. New York Bot. Gard. 7: 255. 1927.
— "[Bolivia.] San Buena Ventura, Beni, 1000 feet, M. Car
denas, December 1, 1921 (no. 1739)." — Holotypus, NY!.
— San Buena Ventura = present Rurrenabaque.
Macrophylhdious arborescent shrubs with geo-
tropic or horizontal branches and notably dense tough
wood, sometimes fertile when 1 m tall or less but po
tentially attaining 4(-6) m with trunk to 2 d m dbh, the
year's growth consisting of simple virgate barren
branches and many, often crowded floriferous short-
shoots, the young stems and most lf-axes and pedun
cles sordidly pilosulous with fine wavy or incurved
hairs to 0.2-0.7 m m , often glabrous in age, the lfts
firm bicolored, ± lustrous, facially glabrous, often
microciliolate, the incipiently umbelliform capitula
arising singly from nodes of either fohate or elaminate
brachyblasts axillary to homotinous lvs; phyllotaxy
distichous. Stipules lanceolate, lance-attenuate, or
deltate-acuminate, from ± dilated, subcordate base,
those of primary lvs 3-11 x 1.4-2.4 m m , all striate,
usually glabrous dorsally but occasionally pilosulous,
tardily deciduous. Lf-formula i/2, the distal pair
longer, the anterior 1ft of proximal pair often smallest;
lf-stks 4-12(17, but few >10) m m , at middle
0.35-0.95 m m diam; rachis of pinnae 3—8(—13) m m ,
the proximal pair of lfts inserted well below middle;
lft-pulvinules transversely elhptic 0.5-0.9 x 0.8-1.4
m m ; lft-blades elliptic from shallowly semicordate
base, obtuse-mucronulate or -apiculate, the distal pair
(16-)19-42 x (4-)5-12 m m , (2.9-)3-4.1(-4.5) times
as long as wide, the proximal pair half as long or less;
midrib nearly straight, forwardly displaced to divide
blade ±1-1.5-2, the inner posterior primary nerve
incurved-ascending well beyond mid-blade, the 2-3
outer posterior primary nerves progressively much
shorter, all these together with secondary and reticu
lar venulation prominulous on both faces, a little
more sharply so beneath. Peduncles 7-26 m m , bract
eate near middle, or above middle, or close under the
capitulum, the bract (perhaps absent in some plants)
deciduous; capitula (14—)16-24-fld, the receptacle
1.5-4 m m diam; bracts narrowly lanceolate 0.7-2
m m , persistent; fls heteromorphic, but scarcely dif
ferent in length or number of filaments, the 1-3 distal
ones a little broader but not longer, their androecium
scarcely longer but greatly dilated distally, and fur
nished with an intrastaminal nectary; perianth of all
fls 4(5)-merous, glabrous overall, the calyx delicately
striate, the corolla submembranous, not externally
venulose; P E R I P H E R A L FLS: pedicel often cryptic,
in vertical section 0.35-0.6 m m ; calyx campanulate
1.4-2.6 x 0.9-1.5 m m , the obtuse teeth 0.1-0.4 m m ;
corolla 6-7.5 m m , the lobes 1.3-2.7 m m ; androecium
(6-)9-10-merous, 22-32 m m , the whitish tube 11-16
m m , the stemonozone 0.6-1.5 m m , the tassel pink;
intrastaminal nectary 0; (sub)terminal fl(s) a little
broader but scarcely longer than the rest, the androe
cium hardly longer but greatly dilated distally, the
orifice of the tube 4.5-7 m m diam, commonly fur
nished with an ascending membranous lobe between
each pair of filaments; intrastaminal nectary to 0.7
m m tall. Pods glabrous or thinly puberulent-pilosu-
lous, in profile 6-9 x (0.6-)0.7-l.l cm, the dilated
sutural ribs in dorsal view 2.5-4 m m wide, the plane
recessed valves almost nerveless externally; seeds ob
tusely rhombic in broad view, piano-compressed,
6.5-9 m m in long diam, the papery, dull brown testa
wrinkled on the seed faces, peripherally narrowly
winged; pleurogram 0.
O n stream banks and on islands in river rapids,
mostly at 10-400 m, often forming extensive colonies
and leaning over the water, scattered along the Andean
foothills of w. Amazonia from s.-e. Colombia s.
through Ecuador and P e m to n.-e. Bolivia (Beni, Santa
C m z ) , in Pem occasional as a river bank colonist
126 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
e.-ward to the Brazilian border, in Ecuador crossing
the cordillera and reappearing on the Pacific lowlands;
in Colombia frequently cultivated, to 1600 m or more,
in the Cauca and middle Magdalena valleys and in
Choco; cultivated on Puerto Rico. — Map 35. — Fl.
nearly yearlong. — Chipara (w. Ecuador).
All but rare random leaves of C. angustifolia are ex
actly 8-foholate, and the larger distal pair of leaflets
are relatively small in context of ser. Macrophyllae,
seldom attaining 4 cm in length. The androecium is
reduced to about eight to ten stamens, and the pods
stand erect on hgnescent peduncles. This set of mor
phological characters effectively separates C. angusti
folia from all related species.
75. Calliandra harrisii (Lindley) Bentham, London
J. Bot. 3: 95. 1844. Inga harrisii Lindley, Bot. Reg.
25: t. 41. 1839. — Described from plants cultivated
in England, "imported from Mexico by Thomas
Harris, Esq. of Kingsbury," where it flowered in
Feb. 1835. — Feuillea harrisii O. Kuntze, Revis.
Gen. PL 1: 188. 1891. FiG. 15
C. cylindrocarpa Bentham, London J. Bot. [1: 169. 1842, nom. nud.] 3: 96. 1844. — "Tropical Brazil, near Rio Janeiro, Sello, Pohl, Tweedie, n. 1218, Gardner n. 22 and 23." — Lectotypus, Gardner 23 , K! = NY Neg. 7957 (right); isotypi, GH!, NY!; paratypi omnes, K!. — Equated with C. harrisii by Bentham, 1875: 540. Clelia ornata Casaretto, Nov. Stirp. Bras, [decas 10]: 84. 1845. — "Reperi in insulis sinus Fluminensis (bahia do Rio de Janeiro)." — Holotypus to be sought at TO. — Equated with C. harrisii by Bentham, 1875: 540. — The monotypic genus Clelia was dedicated to Clelia Durazzo-Grimaldi of Genoa, a patroness of Botany. Calliandra harrisii sensu Lindley, Bot. Mag. 72: t. 4238. 1846; Bentham, 1876: 410; Glaziou, 1905: 187.
Macrophyllidious, weakly arborescent or subsar-
mentose shrubs attaining 3-4 m, with smooth gray
annotinous long-shoots, the young growth commonly
villosulous with fine whitish hairs to 0.3-0.6 m m but
sometimes early glabrate, or glabrous from the first,
the plane, thin-textured lfts bicolored, dark green but
opaque above, paler beneath, the incipiently umbelli
form capitula arising singly at the first 1-2 nodes of
newly activated brachyblasts either axillary to an old
primary If or at a recently defoliate primary node, the
homotinous foliage mostly hyster- or synanthous.
Stipules deltate or ovate-triangular 1-2.5 mm, some
times faintly striate when young but early thickened
and then veinless externally, often gibbous dorsally,
blanched in age, persistent. Lf-formula i/Wr, lf-stks at
maturity ±1.5-3.5 cm, at middle 0.4-0.6 m m diam;
rachis of pinnae 7-11 mm, the odd exterior 1ft
inserted well below middle, smaller than the distal
pair but over half as long; lft-pulvinules ±1-1.2 m m ;
distal lfts obliquely obovate-elliptic or oblanceolate
from semicordate base, either obtuse, or deltate-
apiculate, or retuse-apiculate, the larger ones (1.5—)2—
4.8 x (0.9-) 1-2.2 cm, 1.7-2.4 times as long as wide;
venation palmate-pinnate, the straight or gently in
curved midrib nearly centric at mid-blade, giving rise
on each side to 5-8 major secondary nerves brochi-
dodrome well within the margin, the 1-2 posterior
primary nerves very short, the tertiary venulation
either lax irregular or sharply closely reticulate, be
coming pallid and weakly prominulous on each face.
Peduncles 8-38 mm, usually ebracteate; capitula
7-12-fld, the receptacle <2 mm; floral bracts linear-
spatulate or subulate, 1 m m or less, persistent; fls
heteromorphic, the peripheral ones either subsessile
or distinctly pedicellate, the terminal one tmly ses
sile, scarcely longer than the rest but much broader, in
random capitula abortive; PERIPHERAL FLS: pedi
cel 0.4-1.5 mm; perianth either glabrous except for
minutely ciholate corolla-lobes, or the calyx thinly
pilosulous, the latter coarsely ±15-20-veined, the
corolla nearly veinless externally, reddish distally;
calyx in profile cuneiform (clove-shaped) 2.3-4.5
mm, at orifice abmptly dilated to 1.6-2.4 m m diam,
the incurved depressed-ovate, broadly obtuse teeth
0.2-0.4 m m , dorsally gibbous, purple-carmine;
corolla subcylindric 7.7-11.5 m m , the erect, oblong
obtuse, often unequal lobes 0.4—2.1 mm; androecium
20-28-merous, when fully expanded 2.8-6 cm, pallid
below middle, carmine distally, the tube 4—6.5 mm,
the stemonozone 0.6-1.4 m m ; ovary sessile,
glabrous, in some fls rudimentary; intrastaminal nec
tary 0; C E N T R A L FL: calyx broadly campanulate 2-
3 m m diam; stamens nearly twice as many as in outer
fls; nectarial disc 0.55-0.8 m m tall. Pods in profile
linear straight (5-)6-10 cm, obtusely 4-angular and
±5-6 m m diam, the sutural ribs dilated and in both
dorsal and lateral views as wide as the fmit itself, in
lateral view concealing the woody valves but sepa
rated by a narrow groove, the whole in cross-section
obtusely 8-shaped, the ribs coarsely nerved length
wise, micropuberulent glabrescent.
On exposed rocky slopes, on stream banks, and in
thin bmsh-woodland, mostly below 400 m, best
known from Cabo Frio and shores and islands of
Guanabara Bay in Rio de Janeiro, Brazil, where first
collected in 1815 by Prince Maximilian (BR); once
recorded from near 13°50/S in interior Bahia (Barra
da Estiva to Larguinha) and once from central Minas
Gerais (Glaziou, P); disjunct on Cerro Leon in Parque
Nacional Defensores del Chaco, Paraguay, and at 800
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 127
• CALLIANDRA ANGUSTIFOLIA
• C. HARRISII
Map 35. Distribution of Calliandra angustifolia Bentham and C. harrisii (Lindley) Bentham in South America.
m near Camiri, in the Andean foothills of Santa Cmz,
Bolivia, near 19°45'S; cultivated at S. Paulo, Brazil,
and formerly under glass in Europe. — Map 35. —
Fl. VIII-X(-?), the flowers precocious or accompa
nying new leaves.
Within its interrupted range south of the Hylaea C.
harrisii is the only member of ser. Macrophyllae with
primarily pinnate venation; in other respects it resem
bles C. tergemina sens. lat. The plumply obtusely an-
gulate pod is unique in the genus; its sutural ribs are
so far enlarged at expense of the narrow valves as al
most or quite to conceal them from external view, and
the cross-section has become bluntly 8-shaped. The
dispersal of C. harrisii is difficult to account for, the
known populations being so far apart and apparently
adapted to diverse associations and microclimates.
76. Calliandra tergemina (Linnaeus) Bentham,
London J. Bot. 3: 96. 1844, the epithet transferred
from Inga tergemina Willdenow, which is based on
Mimosa tergemina Linnaeus. — Typus infra sub
var. tergemina indicator.
Macrophyllidious shrubs and slender trees flower
ing when (0.6-)1.5-6(-10) m tall, with smooth palhd
annotinous and older branchlets, the epidermis split
ting lengthwise and exfoliating in papery strips, the
new growth often glabrous except for ventrally puber
ulent lf-axes and pubemlent peduncles but the whole
plant at times variously pubemlent or pilosulous, the
membranous or (in sunny sites) subcoriaceous lfts bi
colored, dull green above, paler beneath, the slenderly
pedunculate capitula borne mostly solitary on short
128 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
FiG. 15. Calliandra harrisii (Lindley) Bentham. Reproduced from Bentham in Martius, Fl. bras. 15(2): t. 104.
1876.
thatched brachyblasts axillary to coeval lvs of pliant
long-shoots, or some directly from primary lf-axils;
phyllotaxy distichous. Stipules subtending primary lvs
narrowly triangular or lanceolate 1^4.5(—5.5) m m ,
striately 4—9(-13)-nerved, deciduous, those of short-
shoots commonly smaller and more persistent, when
persistent becoming blanched and nerveless exter
nally. Lf-formula i/l(-2, but only in random, excep
tionally in nearly all lvs); petioles (2-)5-28(-34) x
0.3—0.85(—1.1) m m , the terminal appendage either
setiform or lanceolate; rachis of longer pinnae (2.5-)
3.5_24(-34) m m , the proximal lft(s) inserted 1-4.5
(-6) m m from base; lft-pulvinules 0.3-1.1 x 0.3-1.1
m m , cross-wrinkled, sometimes obscurely so; lfts
obliquely obovate, semiobovate, or asymmetrically
elliptic-(ob)lanceolate from shallowly semicordate
base, either obtuse-mucronulate, or deltately subacute,
or shallowly emarginate-mucronulate at the rounded
or very shortly bluntly acuminate apex, the distal pair
(1.3-)1.6-7.5(-8) x (0.5-)0.6-3 cm, exceptionally
8-12 x 3-6 m m , (1.4-)1.8-3.4(-3.8) times as long as
wide; venation usually palmate-pinnate, the inner of
2-3 posterior primary nerves incurved-ascending at
least to and usually well beyond mid-blade, the sec
ondary and reticular venulation variably prominulous
or subimmersed, in microphyll forms rarely 1-nerved.
Peduncles (5-)l 1^10 x 0.3-0.7 m m , either ebracteate
or bracteate above (rarely below) middle; capitula
(8-)12-21(-26)-fld, the receptacle 1-3 m m ; bracts
commonly triangular-subulate or linear-lanceolate
0.7-2 m m , exceptionally setiform to 3 m m or deltate
0.4 m m , 1-3-nerved, persistent; fls potentially, in fact
almost always, dimorphic, one or more (sub)terminal
ones fertile, with broader calyx, a dilated, further
exserted staminal tube, and an intrastaminal nectary,
the perianth of all fls either greenish, or whitish, or
pink to vivid carmine, either 4- or 5-merous, usually
glabrous but randomly stigulose or pilosulous, the
calyx-tube alone, or the corolla-tube also (but not the
lobes) prominently striate; P E R I P H E R A L FLS: pedi
cel 0-0.8 m m ; calyx campanulate 0.6-3.6(^1) x
0.4-1.4(-l.5) m m , the teeth minute obtuse to subulate
acute, 0.1-0.9(-l) m m ; corolla (4-)4.3-9(exception-
ally to 10, 12) m m , the lobes 0.7-2 m m ; androecium
(8- 10-)12-28-merous (in Lara, Venezuela, to 42-
merous), the stemonozone 1 m m or less, the tube
4—7(exceptionally -10) m m , the tassel either red-
carmine throughout, or pallid proximally and pink-
tipped, or white-ochroleucous throughout, the fila
ments sometimes united beyond the tube into fascicles
of varying number; ovary (mostly sterile) glabrous at
anthesis; disc 0. Pods plagio- or geotropic (not stiffly
erect), in broad profile narrowly oblanceolate tapering
downward into a stipehke base (4.5-)5.5-15 x 0.6-1.4
cm, glabrous or rarely finely pilosulous, when well
fertilized 5-9-seeded; seeds plumply compressed-
ellipsoid, in broad view 7-11 x 3.5-5.5 m m , the testa
either smooth or cmmpled, castaneous often dark-
speckled, either pleurogrammic or not, the pleuro
gram when present narrowly U-shaped.
As defined by the foregoing description, C.
tergemina is a variable species of wide dispersal,
polymorphic in indumentum, amplitude of leaflets,
depth of calyx, color of the stamens, and in occur
rence of a pleurogram on the seed coat. At the time
that Bentham (1875: 536, sequ) published his sum
mary account of Calliandra, comparatively few col
lections of C. tergemina and its relatives were acces
sible. From this biased evidence a glabrous Antilhan
C. tergemina characterized by a tiny calyx (±1 m m or
less), and continental larger-leaved C. emarginata
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 129
(Guerrero), C. seemanni (Panama) and C. canescens
(Veracruz), the last three collectively different in
longer calyces and C. canescens yet further in pubes
cent foliage, seemed plausibly distinct taxa. It is now
apparent that the formulae known to Bentham
amounted to a random sample of character-states that
exist in Mexico and Central America, for many
another syndrome of pubescence, leaf-size, leaf-
texture (and prominence of venulation), leaflet-outline,
length (absolute and proportional) of corolla and sta
mens has been discovered since. Those known to
Britton and Rose in 1928 were either maintained or
described as species of Anneslia (ser. Tergeminae,#4-
18, inch). This classification was rejected by Standley
and Steyermark (1946: 22, sub C. emarginata) as un
realistic. Standley and Steyermark recognized in the
flora of Guatemala only two species in this complex,
the glabrous C. emarginata, and C. mexicana, differ
ent solely in leaflets at least dorsally pilose, each
equally variable in other features and representing
probably only one species. McVaugh (1987: 155) de
scribed for Flora Novo-Galiciana a glabrate form of
C. emarginata growing with an otherwise uniformly
pubescent population. Development of indumentum,
which sometimes extends to the calyx and perianth, is
visually arresting, but has no practical taxonomic ap
plication. Equally striking is the range of variation in
size and outline of leaflets, which tend to be propor
tionately broader and more obtuse as they increase in
length. But the variation is continuous; moreover an
occasional vigorous shoot with ample leaflets arises
from a small-leaved parent branch. A like pattern of
continuous variation devalues color of the perianth
(white, greenish, pink, carmine) and androecium,
which varies from white through particolored (white/
carmine) to carmine throughout, as a useful taxo
nomic character. Texture of leaflets is modified by
age, season, and microhabitat. The small calyx char
acteristic of C. tergemina as represented on the Lesser
Antilles and coastal northeastern Venezuela coincides
with a striate corolla tube; but in west-central Carib
bean Venezuela the calyx becomes insensibly deeper
and the venulation of the corolla by degrees less dis
tinct, thereby passing into forms outwardly indistin
guishable from Central American examples of C.
emarginata. On the other hand, Pittier 8822 (NY)
from near Valencia, Carabobo has the longer calyx of
emarginata combined with the striate tube of typical
tergemina. In fact C. tergemina, by historic accident
the first to be named, is nothing more than a periph
eral expression of a specific type widely dispersed
around the western Caribbean and northward. More
difficult to compress into one species is variation in
the seed-coat, which has no pleurogram in the An
tilles, but is without exception pleurogrammic in
Mexico. However, the Panamanian type of C. see-
mannii has the relatively deep calyx of C. emarginata
combined with the plain testa of typical C. tergemina.
The alternative to a conservative taxonomy, already
adopted in part by Standley and McVaugh, is an intri
cate Hasslerian proliferation of names based on par
ticular specimens.
Provisionally included in my concept of var. emar
ginata is a series of relatively microphyll races or
forms in which the larger distal leaflets, even of pri
mary leaves, are less than 2 cm or even less than 1.5
cm long. Their biological status requires further
study, but is is likely that they are not a monophyletic
group but parallel derivatives of var. emarginata
s.lat, different one from the next in details of pubes
cence, venation, and color of perianth and androe
cium. The following list of specimens indicates the
scattered occurrence of the microphyll populations
within the range of typical emarginata: MEXICO.
Sinaloa, Mazatlan, Rose 14155 (NY); Guerrero: San
Marcos, Macqueen 407 (K); Veracruz: mun. Actopan,
7?. Acosta P. 534 (NY); Oaxaca: Niltepec, R. M. King
1805 (NY). EL SALVADOR. Ahuachapan, C. E.
Hughes 1235 (NY). BELIZE. El Cayo distr., T. B.
Croat 23531 (NY, one leaf on incipient branchlet to
30 x 13 mm).
Key to the varieties of C. tergemina
1. Calyx of peripheral fls of the capitulum (1.2-)
1.4-3.6(-4) mm, prominently 10-18-nerved;
corolla-tube not or only faintly striate-nerved;
seeds nearly always pleurogrammic; tropical
Mexico, Central America, and remotely scattered
in n.-w. Venezuela and Colombia. Distal lfts
commonly over 3 cm, reaching 7.5 cm or even
8-12 x 3-6 cm, but randomly as small as in
the next 76a. var. emarginata
1. Calyx of peripheral fls of the capitulum 0.6-
1.3 mm; calyx and corolla-tube alike striate-
nerved; seeds lacking pleurogram; Lesser
Antilles and n.-e. Venezuela. Distal lfts
mostly 1.6-2.8 cm 76b. var. tergemina
76a. Calliandra tergemina (Linnaeus) Bentham var.
emarginata (Willdenow) Bameby, comb. nov. Inga
emarginata Humboldt & Bonpland ex Willdenow,
Sp. PL 4(2): 1009. 1806. — "Habitat in America
meridionali"; but the tme origin stated by Kunth,
Mimoses 54, pi. 17. 1820: "in litore occidentali
Regni Mexicani, prope Acapulco [Guerrero]." —
Holotypus, Humboldt 3859 in B-WILLD, seen in
Microform! = F Neg. 12381. — Mimosa emarginata
130 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Poiret, Encycl. suppl. 1: 39. 1810. Calliandra emar
ginata Bentham, London J. Bot. 3: 95. 1844. Feuil
leea emarginata O. Kuntze, Revis. Gen. PL 1: 187.
1891. Anneslia emarginata Britton & Rose, N.
Amer. FL 23: 54. 1928, exclus. syn. Inga coriacea.
Inga canescens Chamisso & Schlechtendal, Linnaea 5: 592.
1830. — "[Schiede & Deppe] 678. [MEXICO. Veracruz:]
Inter Marantial et Puente del Rey [= Puente Nacional, n.-
w. of Cd Veracruz]." — Holotypus, n.v. — Calliandra
canescens Bentham, London J. Bot. 3: 96. 1844. Feuilleea
canescens O. Kuntze, Revis. Gen. PL 1: 187. 1891. Annes
lia canescens Britton & Rose, N. Amer. Fl. 23: 55. 1928.
Calliandra tetraphylla G. Don, Gen. Hist. 2: 392. 1832. —
"Native of Mexico. Mimosa tetraphylla, Sesse & Mo? in
herb. Lamb." — Holotypus, OXF! = photo, NY!. — Cal
liandra tetraphylla Bentham, Trans. Linn. Soc. London 30:
544. 1875. — Feuilleea tetraphylla O. Kuntze, Revis. Gen. PL 1: 189. 1891.
Inga semicordata Bertoloni, Novi Comment. Acad. Sci.
Inst. Bononiensis [Fl. Guatim.] 441. 1840. — "Habitat in
Guatimala." — Holotypus, / Velasquez, n.v. — Referred, with reservation, to C. seemanni by Bentham, 1875: 540,
and to C. emarginata by Standley & Steyermark, 1946: 22.
Calliandra seemanni Bentham in Seemann, Narr. Voy. Her
ald 1: 116, pi. XXII. 1853. — "[PANAMA.] On banks of
several rivers in Veraguas." — Holotypus (quoted by Hem-
sley, Biol, centr.-amer. 1: 358): Seemann 1193, from Tole,
Veraguas [8°15'N, 81°40'W], K (hb. Hook.)! = N Y Neg.
7994; isotypus, BM!. — Feuilleea seemanni O. Kuntze,
Revis. Gen. PL 1: 189. 1891. Anneslia seemanni Britton &
Rose, N. Amer. Fl. 23: 54. 1928.
C. rupestris T. S. Brandegee, Zoe 5: 199. 1905. — "[MEXICO. Sinaloa:] ... in a canyon near Cofradia [Sep-Oct, 1904, T. S. Brandegee]." — Holotypus, UC!. —Anneslia
rupestris Britton & Rose, N. Amer. FL 23: 54. 1928 C. purpusii T. S. Brandegee, Univ. Calif. Publ. Bot. 6: 180.
1915. — "[MEXICO.] . on Cerro de Picacho, Oaxaca,
[C. A. Purpus] No. 7199." — Holotypus, U C 174965\. — Anneslia purpusii Britton & Rose, N. Amer. Fl. 23: 55.
1928. C. langlassei Harms, Repert. Sp. Nov. Regni Veg. 17: 87.
1921. — "Mexico: Chichihualco [±25 km n.-w. of
Chilpancingo, Guerrero; cf. McVaugh, Candollea 13: 195.
1951], 1200 m (Langlasse, no. 1042. - V 1899)." — Holo
typus, ̂ B; isotypus, K! = photo s.n. by J. N. Rose in 1927,
NY!. — Anneslia langlassei Britton & Rose, N. Amer. Fl.
23: 53. 1928. C. mexicana T. S. Brandegee, Univ. Calif. Publ. Bot. 10:
183. 1922. — "[MEXICO.] .. . near Remulatero [= Remu-
dadero], Vera Cruz. [C. A. Purpus] No. 8726." — Holotypus, U C 2132961. —Anneslia mexicana Britton & Rose,
N. Amer. Fl. 23: 55. 1928. — Provisionally interpreted as
"no more than a pubescent variety" of C. emarginata by
Standley & Steyermark, 1946: 24. Anneslia yucatanensis Britton & Rose, N. Amer. Fl. 23: 53.
1928. — "Yucatan, 1917-1921, C. F. Gaumer 24240." —
Holotypus, NY!; isotypi, K!, NY!. — Calliandra yucata
nensis Standley, Publ. Field Columbian Mus., Bot. Ser.
4(8): 309. 1929. — Equated by Standley & Steyermark,
1946: 22, with C. emarginata. A. sinaloana Britton & Rose, N. Amer, Fl. 23: 54. 1928. —
"[MEXICO.] Mazatlan, Sinaloa, January 17-19, 1897, [J.
N f Rose 1380." — Holotypus, NY!. — Calliandra
sinaloana Standley, Trop. Woods 34: 40. 1933. A. cruziana Britton & Rose, N. Amer. Fl. 23: 54. 1928. —
"[MEXICO.] Veracruz. Type from Cameron [= Camaron],
Nov. 15, 1926, [C. A.] Purpus 11000." — Holotypus, N Y ! = N Y Neg. 9314; isotypi, K (2 sheets)!. — Calliandra
cruziana Standley, Publ. Field Columbian Mus., Bot. Ser.
4(8): 212. 1929. — Equated by Standley & Steyermark,
1946: 22, with Calliandra emarginata. A. juchitana Britton & Rose, N. Amer. FL 23: 55. 1928. —
"[MEXICO.]... Juchitan, Oaxaca, January 31. 1896, Seler
J988." — Holotypus, E. & C. Seler 1988, NY! = N Y Neg.
9317; isotypus, K!. — Equated with C. emarginata by
Standley & Steyermark 1946: 22.
A. leucotricha Britton & Rose, N. Amer. Fl. 23: 55. 1928. —
"[MEXICO.] Acasonica [= Acosonica, ±19°20'N,
96°35,W], Veracruz, 1919, [C. A.] Purpus 8391." — Holo
typus, NY! = N Y Neg. 9318. — Non Calliandra leucathrix
Standley, 1938. A. deamii Britton & Rose, N. Amer. Fl. 23: 56. 1928. —
"Guatemala, along path to Motagua River from Gualan
[depto. Zacapa], June 14, 1909, [C. C ] Deam 6258." — Holotypus, NY!. — Calliandra deamii Standley, Publ. Field
Columbian Mus., Bot. Ser. 4(8): 309.1929. — Equated with
Calliandra mexicana T. S. Brandegee by Standley & Stey
ermark, 1946: 24. Calliandra tolimensis L. Uribe, Caldasia 1(4): 9. 1942. —
"Tipo en el Herb, de la Univ. Javeriana, No. 644; encon-
trada en marzo de 1941 entre Alpujarra (Tolima) y el rio
Cabrera, a 1200-1300 m. lat. . . . [3°20'N, 75°W]" —
Holotypus, C O L 776709!. — C. uribei Killip & Dugand,
Caldasia 3(11): 35. 1944, a legitimate substitute, non C.
tolimensis Taubert, 1895. Inga emarginata sensu Humboldt, Bonpland & Kunth,
1824: 296; de Candolle, 1825: 438. Calliandra emarginata sensu Bentham, 1875: 539, ex parte, exclus. Inga coriacea;
Standley, 1922: 384; McVaugh, 1987: 154.
Calliandra langlassei sensu Standley, Contr. U.S. Natl. Herb. 23(5): 1: 658. 1926.
C. seemani sensu Woodson & Schery, 1950: 258.
Shrubs and trees flowering when (0.6-)l-6(-10) m
tall, quite glabrous to finely pilosulous; petioles (4—)
5-28 x 0.4—0.85 m m ; lfts membranous to charta
ceous, either glabrous or glabrous ciholate, or pilosu
lous on one or both faces, the distal pair (1.3—)1.6—
7.5(-8) x (1.4-)1.8-3.4(-3.8) cm; peduncles 1 6 ^ 0 x
0.4-0.7 m m ; perianth either glabrous, or weakly
strigulose, or pilosulous, the calyx 1.2-3.6(^1) x 0.7-
1.4(1.5) m m , the teeth 0.1-0.9(1) m m ; corolla (4-)
4.6-9(exceptionally 11,12) m m ; filaments commonly
white pink-tipped, exceptionally white, randomly
deep red-carmine throughout; pods 6-15 x 0.6-1.4
cm, usually glabrous, sometimes finely pilosulous;
seeds in broad view 8.5-11 x 3.5-5.5 m m , almost
always pleurogrammic.
In bmsh-woodland, short-tree forest, glades in oak-
woods, and riparian or on drier ridge habitats in sub
montane forest, flourishing in disturbed woodland and
on roadsides, 1-1050 m and recorded in El Salvador
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 131
at 1100, in Colombia at 1300, and in Chiapas at 1500
m, discontinuously widespread from n.-w. subtropical
Mexico to Yucatan Peninsula, s.-e. through Central
America to inter-Andean and n.-w. Amazonian
Colombia and n.-w. Venezuela: in w. Mexico s.-ward
from 27°N (Sa. de Alamos) and on the Gulf lowlands
from the Tropic line; in Colombia on the Magdalena
valley s. to Tohma, e. very locally to Serrania de la
Macarena at ±74°W in Meta and to arenitic hills at
70°30'W in Vaupes; in Venezuela very local in Lara;
formerly cultivated in Cuba, in Florida, and under
glass n.-ward. — Map 36. — Fl. nearly throughout the
year, most abundantly XI-V. — Clavellino, a generic
term for Calliandra.
Note: Sousa 5208 from Oaxaca, Mexico, is de
scribed on label (NY) as having red flowers with
golden yellow stamens, "hermosa, facilmente orna
mental," a color scheme unknown elsewhere in the
genus.
76b. Calliandra tergemina (Linnaeus) Bentham var.
tergemina. C. tergemina (Linnaeus) Bentham,
1944, I.e., sens. str. Mimosa tergemina Linnaeus,
Sp. PL 517. 1753. — "Habitat in America merid
ionalis — Holotypus (Howard, 1988: 352): Acacia
frutescens non aculeata, flore purpurascente
Plumier ed. Burmann, PL amer. 5, t. X, fig. 1 (top
left + misplaced caption at bottom right). 1755!. —
Inga tergemina Willdenow, Sp. PL 4(2): 1008.
1806. — Feuilleea tergemina O. Kuntze, Revis.
Gen. PL 1: 189. 1891. Anneslia tergemina Britton &
Rose, N. Amer. FL 23: 53. 1928. Calliandra tergem
ina Standley, Publ. Field Columbian Mus., Bot. Ser.
4(8): 309. 1929, comb, superflua.
Inga caripensis Willdenow, Sp. PL 4(2): 1009. 1806. — "Habitat in Nova Andalusia prope Caripe [estado Sucre,
Venezuela]." — Holotypus, Humboldt 578 in B-WILLD
19015, seen in Microform!. — Mimosa caripensis Poiret,
Encycl. suppl. 1: 39. 1810. — Equated with 7. tergemina
by Humboldt, Bonpland & Kunth, Nov. Gen. Sp. 6(qu):
298. 1824. Mimosa tergemina sensu Jacquin, Select. Stirp. Amer. Hist.
265, t. 177, fig. 81 (one fl). 1763. Inga tergemina sensu de
Candolle, Prodr. 2: 437. 1825. Calliandra tergemina sensu
Grisebach, Fl. Brit. W. I. 225. 1864; Duss, Fl. Phan. AntilL
franc. 253. 1897; Stehle & Quentin, FL Guadeloupe
Depend. Martinique 695. 1978; Howard, 1988: 352.
Shrubs and treelets mostly <4 m; petioles very slen
der 0.3-0.5 m m diam; lfts thin-textured, seldom over
3 cm, obhquely obovate to inequilaterally elliptic-
(ob)lanceolate, either obtuse or acute; peduncles (5-)
11-28 x 0.3-0.4 m m ; pedicels at most 0.25 m m ,
commonly 0; perianth glabrous; calyx 0.6-1.2 x
0.4-0.75(-l), the teeth 0.1-0.4 m m ; corolla 4.3-7
m m ; filaments bicolored, pink distally; pods 5.5-12 x
0.65-0.8 cm; seeds in broad view 7-9 x 3.5^.5 m m ,
pleurogram 0.
In drought-deciduous brush-woodland and savanna
margins, sometimes invading cleared pasture, below
300 m, locally plentiful in the Lesser Antilles (Do
minica to Grenada) and along the Caribbean coast of
Venezuela from Paria peninsula in Sucre w. through
Distrito Federal and Carabobo to e. Lara; an immature
specimen from state of Bolivar at 7°22'N, 61°49'W (E.
Sanoja 1185, N Y ) is provisionally referred here, but its
varietal status is uncertain. — M a p 36. — Fl. VHI—II,
perhaps randomly through the year. — Bois-patate,
madam-di-poule (Antilles); clavellino (Venezuela).
77. Calliandra macqueenii Bameby, sp. nov., C.
tergeminae var. emarginatae (Willdenow) Bameby
proxima et quoad habitum et foha similis, sed ab ea
peduncuhs abbreviatis 2.5-5 (nee 10-40) m m longis,
capitulis confertissime 20-40 (nee plerisque 12-20)
-floris, receptaculo ±5-7 (nee 1-3) m m longo,
androecii tubo longe exserto 15-16 (nee 4—7, raris-
sime 10) m m longo, necnon leguminibus ut videtur
acroscopicis (nee plagiotropicis penduhsve) diversa.
— M E X I C O . Guerrero: 15 k m N on road toward
Mexico from outskirts of Acapulco, 130 m, 8 Feb
1992 (fl, fr jun), D. J. Macqueen (with Contreras,
Lewis & Nileshwar) 414. — Holotypus, K (2 sheets).
Macrophyll bushy shrubs 1-1.5 m with smooth
gray-brown branches, the young shoots and both faces
of lfts ± pilosulous with spreading and narrowly
ascending hairs to ±0.5 m m , the dull olivaceous sub
concolorous lfts densely papillate, the dense capitula
of pale-pink ochroleucous-stamened fls shortly pedun
culate from axillary brachyblasts; phyllotaxy distic
hous. Stipules firm-papery, narrowly lanceolate or
ovate-lanceolate 3-9 x 0.8-1.3 m m , 5-8-nerved,
becoming dry and brittle. Lf-formula i/2; lf-stks 9-21
m m , at middle ±0.5-0.7 m m diam, shallowly openly
sulcate ventrally; rachis of pinnae 10-21 m m , the
smaller proximal pair of lfts inserted well below mid
dle; lft-pulvinules 0.7-1 x 0.55-0.7 m m , finely wrin
kled; lfts inequilaterally elhptic or obovate-elhptic
from semicordate base, deltate- or triangular-apiculate,
the blade of distal pair 2.2-5 x 0.9-2.2 cm; venation
palmate-pinnate, the gently incurved midrib displaced
to divide blade ±1:2, 5-8-branched on each side,
the strong inner posterior primary nerve produced
beyond mid-blade. Peduncles stout 2.5-5 m m , some
times bracteate; capitula ±20-40-fld, the receptacle
132 M E M O I R S OF T H E N E W Y O R K BOTANICAL G A R D E N [VOL. 74(3)
calliandra tergemina
• var. TERGEMINA
Y var. EMARGINATA
M a p 36. Distribution of Calliandra tergemina (Linnaeus) Bentham var. tergemina and var. emarginata (Willdenow)
Barneby.
5-7 m m ; bracts lanceolate or ovate, <1 m m , persis
tent; perianth 4- or 5-merous, the calyx sharply finely
striate, the membranous corolla faintly striate below
middle, thereafter swollen, externally veinless, both
calyx and corolla thinly pilosulous, especially dis
tally; calyx campanulate 1.9-2.3 x 1.2-1.6 m m , the
teeth 0.3-0.5 m m ; corolla ±9-10 m m , the obtuse
lobes ovate ±2 m m ; androecium ochroleucous 18-20-
merous, ±4 cm, the stemonozone 0.7 m m , the tube
15-16 m m , well exserted, no intrastaminal nectary
seen; ovary at anthesis glabrous. Pods apparently
erect from branchlets, subfalcately linear-oblanceo-
late in profile, (6-)8-14 x 0.65-0.8 cm, the sutural
ribs ±1.5 m m wide in dorsal view, the thinly hgnes
cent valves finely thinly pilosulous overall, faintly
venulose; seeds (few seen) in profile oblong ±10 x 5
m m , the hard smooth testa brown darker-speckled,
pleurogrammic.
In brush-woodland near 130 m, known only through
the type-collection, from hill country on the Mexico
City highway ±15 k m n. of Acapulco (H^lTM,
99°47,W), Guerrero, Mexico. — Fl. I—III.
Calliandra macqueenii is presumably a recent de
rivative of C. tergemina var. emarginata, with which it
has much in common, but differs substantially in too
many details to escape formal description. As listed in
the Latin diagnosis, the principal differential characters
include the following: shortly pedunculate capitula of
some 20 to 40 flowers crowded along a distinctly elon
gate receptacle; an extremely long, far-exserted stami
nal tube; and pods stiffly erect on hgnescent peduncles.
The species is dedicated to D. J. Macqueen (OXF),
who has recently amassed an extensive herbarium of
Mexican and Central American Calliandra of all types
but in particular of species akin to C. houstoniana, an
important set of which has been presented to NY.
78. Calliandra brenesii Standley, Publ. Field Mus.
Nat. Hist., Bot. Ser. 18 (FL Costa Rica), 2: 491.
1937. — "[Costa Rica. Alajuela: Cataratas de San
Ramon, May, 1931, [Alberto M ] Brenes 13180." —
Holotypus, Brenes 13680 (not 13180), F!; isotypus,
NY!; paratypi, Brenes 4424,13503, F!, NY!.
C. brenesii sensu J. G. Laurito, Brenesia 25-56, cover. 1986 (photo in color).
Slender sarmentose macrophyllidious shrubs 2-3
m, with smooth palhd terete branchlets, glabrous
throughout, the papery lfts lustrous dark brown-oliva
ceous above, paler beneath, the capitula of dark red fls
arising singly on subfiliform pliant, mostly geotropic
peduncles from axils of short, mostly efoliate, loosely
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 133
thatched brachyblasts; phyllotaxy distichous. Stipules
erect, linear-lanceolate (4-)6-13 x 0.6-1.6 m m ,
weakly striate toward base, pallid in age, persistent. Lf-
formula i/Vz, the lvs bifoholate; petioles 2-17 x 1.1-
1.6 m m , openly shallowly sulcate ventrally, homy-
dilated at apex; rachis of each pinna at most 3 m m ,
often reduced or almost so to pulvinus and homy api
cal dilation; lft-pulvinules 0.8-1.5 x 1-2 m m , coarsely
wrinkled; lft-blades obhquely or subdimidiately ovate-
or lance-acuminate from postically shallow-cordate,
antically cuneate base, at very tip either deltately acute
or obtuse mucronulate, the larger ones 10-19 x
3.3-6.5 cm, 2.3-3.3 times as long as wide; primary
venation palmate, the strong, gently incurved midrib
displaced to divide blade ±1:2, the inner posterior pri
mary nerve incurved-ascending at least 2A length of
blade, the outer posterior one much shorter, these all
prominulous on both faces of blade, the numerous sec
ondary and sinuous tertiary venules much weaker. Pe
duncles 2.8-2.5 cm, at middle 0.25-0.6 m m diam,
bracteate either above or below middle; capitula
(9-)16-24-fld, the plumply claviform receptacle 1.5-
2 m m ; bracts lanceolate or narrowly ovate 0.8-2 m m ,
persistent; pedicels (often not well differentiated ex
ternally) 0.3-0.9 x 0.4-0.6 m m ; fls homomorphic, the
perianth either 3- or 4-merous, often asymmetrical,
glabrous throughout; calyx campanulate 2.8-3 x
1.3-2.5 m m , the teeth 0.3-0.9 m m , one sinus often
more deeply split in age; corolla 9-10.5 m m , the
lance-ovate lobes 2-3.2 m m ; androecium 24—30-mer-
ous, 2.9-3.6 cm, the stemonozone 0.6-1.4 m m , the
tube 8.5-12 m m , either included or shortly exserted,
the intrastaminal nectary 0.6-1.4 m m tall; ovary gla
brous. Pods (seen only after dehiscence) ±10-11 x
0.85-1.2 cm, the coriaceous valves glabrous, obhquely
venulose; seeds unknown.
In understory of wet premontane forest, (?-)500-
650(-?) m, apparently localized in upland Alajuela,
Costa Rica. — M a p 37. — Fl. V-VI, IX, the full
cycle unknown.
Notable characters of the morphologically special
ized C. brenesii are: loss of indumentum; bauhinia-
like, bifoholate leaves with short petiole and subob-
solete pinna-rachis; an often trimerous perianth; and
scarlet androecium.
79. Calliandra laevis Rose, Contr. U.S. Natl. Herb. 5:
194. 1899. — "Collected by J. N. Rose near Colomas
[= Colomos, in mun. Rosario, 23°05'N, 105°30'W],
State of Sinaloa [Mexico], July 18, 1897 (No. 1753)
and at Dolores, Territorio de Tepic [= Nayarit],
August 6, 1897 (No. 3365)." — Lectoholotypus
(Britton & Rose, 1926: 57, by indirection), Rose
1753, US!; isolectotypi, K!, NY!; paratypus, Rose
3365, US!. — Anneslia laevis Britton & Rose, N.
Amer. Fl. 23: 57. 1928. FiG. 16
C. laevis sensu Standley, 1922: 384; McVaugh, 1987: 165.
Macrophylhdious trees 5-12 m with simple trunk
attaining 5 d m diam and terete pliant branchlets,
except for sometimes minutely granular peduncles
glabrous throughout, the epidermis early sphtting and
flaking in strips, the annotinous and older branchlets
gray, the stiffly chartaceous lfts olivaceous, scarcely
paler beneath, the hemispherical capitula arising soli
tary or geminate either a) directly from coeval lf-
axils, or b) from efoliate nodes of axillary brachy
blasts, or c) in default of distal lvs forming a small
terminal pseudoraceme; phyllotaxy distichous. Stip
ules ovate-deltate 0.4—1.6 m m , venulose if at all only
when young, early caducous. Lf-formula i/1, the lvs
all exactly 4-foliolate; petioles 5-19 m m , at middle
0.6-1 m m diam, either shallowly sulcate ventrally or
subterete; rachis of pinnae 5—14(—17) m m ; lft-pulvin
ules 1.3-2 x 0.7-1.1 m m , shallowly wrinkled; blade
of lfts elliptic, elhptic-oblanceolate or lanceolate,
from inequilaterally cuneate base, sometimes shortly
acuminate, at very apex either deltately acute or
obtuse and mucronate, the larger ones 5-11 (-12) x
1.3-3.6 cm, (2.8-)3^t.6 times as long as wide; vena
tion pinnate, the subcentric midrib straight or almost
so, the major incurved-ascending secondary nerves
9-14 on each side, these together with tertiary and
sinuous reticular venules either bluntly prominulous
or almost immersed on either or both faces. Pe
duncles 14—25 m m , ebracteate; capitula hemispheri
cal, 6-14-fld, the subglobose receptacle 1-1.5 m m
diam; bracts ovate or subulate 0.2-0.6 m m , persis
tent; fls commonly dimorphic, one or more central
ones slightly larger or broader, with greatly elongate,
tmmpet-like androecium, but sometimes all fls of the
trumpet type or the fls at mid-capitulum intermediate
in form between the outer peripheral and innermost
ones; perianth (4—)5-merous, glabrous, the calyx pal
lid, the petals whitish or tinged with red, the androe
cial tube white or pallid, the tassel pink or carmine
distally; P E R I P H E R A L FLS: pedicel none or to 0.6 x
0.4 m m ; calyx campanulate 1.3-2.2 x 0.9-1.2 m m ,
faintly 5-nerved, the teeth 0.1-0.2 m m , wider than
long, obtuse; corolla 6-7 m m , the ovate lobes 1.3-2.1
m m ; androecium 12-15-merous, 23-32 m m , the ste
monozone 0.4—0.5 m m , the tube 6-9 m m ; intrasta
minal nectary 0; ovary subsessile, linear-ellipsoid,
glabrous; DISTAL FL(S): calyx sometimes to 3 m m ;
134 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
• C A L L I A N D R A LAEVIS
• C. BRENESH
M a p 37. Distribution of Calliandra laevis Rose and C. brenesii Standley in Mexico and Central America.
corolla almost of peripheral fls but sometimes wider,
sometimes attaining 7.5 m m , the dilated androecial
tube 9-16 m m , 2.5^4 m m wide at orifice. Pods geot
ropic, linear-oblanceolate 8-20 x 1.2-1.7 cm, 5-9-
seeded, the rigid, light brown valves glabrous, errati
cally venulose; seeds oblong or rhombic-oblong in
broad view, plumply biconvex, 10-12 x 6-8.5 m m ,
the smooth testa lustrous castaneous, fuscous-speck
led, the pleurogram narrowly U-shaped.
In rocky ravines and on stream banks, in subdecid-
uous mixed woodland and in pine-oak forest, 750-
1500 m, locally plentiful on the seaward mountains of
w. Nueva Galicia, Mexico, from extreme s.-e. Sinaloa
s. through Nayarit to w. Jalisco, in lat. 18°30'-23°
30'N. — M a p 37. — Fl. VII-X.
Calliandra laevis resembles C. tergemina var.
emarginata in form of the flower and seed and in the
hanging pods, but is distinct in treelike stature, in the
exactly 4-foliolate leaves, and especially in the pin
nately nerved leaflets.
I/G. Series H Y M E N A E O D E A E
Barneby
Calliandra sect. Androcallis ser. Hymenaeodeae
Bameby, ser. nov. monotypica, adspectu fohohsque
paucis amplis ser. Macrophyllis similis, sed foliis
simpliciter, nee bis pinnatis singularis. — Sp. unica:
C. hymenaeodes (Persoon) Bentham.
Like ser. Macrophyllae in few and ample lfts, but
lvs simply, not twice pinnate. — Sp. 1, of lowland
Guianas.
Prior to development of the characteristic fmit,
Calliandra hymenaeodes is readily mistaken for an
Inga, but differs from all compatriot ingas in lack of
leafstalk nectaries.
80. Calliandra hymenaeodes (Persoon) Bentham,
Trans. Linn. Soc. London 30: 537 ("hyme-
neaeoides"). 1875. Mimosa hymenaeodes Persoon,
Syn. PL 2: 262. 1806. — "Richard. Hab. in Cajenna."
— Holotypus, P!. — Inga hymenaeodes Desvaux, J.
Bot. 3: 70. 1814. Feuilleea hymenaeodes O. Kuntze,
Revis. Gen. PL 1: 188. 1891. Fig. 17
C. patrisii Sagot, Ann. Sci. Nat. Bot. VI, 13: 324. 1882. —
"Vidi in Herb. Prodromi a Patris in Guyana Gallica lecta."
— Holotypus, G-DC n.v. (but description unmistakable).
Inga hymenaeoides sensu de Candolle, Prodr. 2: 436. 1825.
Calliandra hymenaeoides sensu Jansen-Jacobs, 1976: 649. Calliandra patrisii sensu Lemee, 1952: 55.
Slender macrophyll trees ±3-9 m with virgate
branches, glabrous except for sometimes ciholate
stipules or pubemlent lf-stks and primary nerves of
hypophyllum, notable in the genus for simply pinnate
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 135
FiG. 16. Calliandra laevis Rose.
136 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
FiG. 17. Calliandra hymenaeodes (Persoon) Bentham.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 137
2-4-foliolate lvs, the papery lfts lustrous dark-oliva
ceous above, paler beneath, the capitula arising singly
from lower efoliate nodes of condensed brachyblasts
axillary to coeval lvs; phyllotaxy distichous. Stipules
firmly chartaceous, lance-triangular 2.5^- x 0.8-1.8
m m , striately nerved, becoming dry pallid, persistent.
Lvs paripinnate, the lfts 1- or 1-2-jug.; lf-stks 6-27
m m , the one interpinnal segment, when present,
longer than the petiole, the ventral groove open, shal
low, obscurely bridged at insertion of lower lft-pair;
lft-pulvinules 1.2-2 m m , strongly wrinkled; lfts
inequilaterally broad-elhptic or obovate-elliptic from
postically rounded, antically cuneate base, obscurely
obtusely deltate-acuminate, when two pairs the distal
pair a httle larger, these 4—10 x 2.5^4.5 cm; venation
of 3 primary nerves, all incurved-ascending from pul-
vinule, the midrib forwardly displaced to divide blade
1:2-3, the inner posterior primary nerve produced
well beyond mid-blade, the outer much shorter, the
secondary nerves and reticulum of venules prominu
lous on both faces, the margin narrowly revolute. Pe
duncles ±2-2.5 cm, 1-bracteate near middle; capitula
12-20-fld, the receptacle 1-2 m m diam; fls sessile,
glabrous [probably heteromorphic, but incompletely
known]; P E R I P H E R A L FLS: calyx deeply campanu
late 2.4—2.7 x 1.1-1.3 m m , the tube striately nerved,
the teeth ±0.3 m m ; corolla 7-9 m m , the lobes 1.5-2.1
m m ; androecium crimson distally, not seen complete.
Pod unknown.
In lowland forest, apparently endemic to the coastal
plain of the three Guianas, from the lower Essequibo
in Guyana to Cayenne in French Guiana. — M a p 38.
— Fl. XI, III-IV, the full cycle not established.
I/H. Series LONGIPEDES Barneby
Calliandra sect. Androcallis ser. Longipedes Bar
neby, ser. nov. monotypica, fohomm formula foh
ohsque amplis cum ser. Macrophyllis congrua, sed
habitu toto dispar: caules abbreviati herbacei e
rhizomate subterraneo nascentes; folia longissime
petiolata; pedunculi axillares sed quasi scapiformes,
praecoces. — Sp. unica: C. longipes Bentham.
Lvs resembling those of ser. Macrophyllae in
ample size and in number of lfts, but stems function
ally herbaceous from subterranean rhizome, shorter
than the elongate petioles and scapiform peduncles,
these developing precociously from nodes anterior to
lvs. — sp. 1, pyrophytic, of the Brazilian Planalto,
Paraguay, and adj. n.-e. Argentina.
Except for exaggerated petioles, the foliage of
monotypic ser. Longipedes is essentially that of some
shmbby or treelike Macrophyllae such as C. trinervia,
from which C. longipes is probably derived. Examples
of functionally herbaceous, fire-pmned species evolv
ing in harmony with campo limpo of Planaltine Brazil
are known in the genera Andira (Fabaceae) and
Chamaecrista (Caesalpiniaceae). Granted that C.
longipes has been selected by habitat, the subter
ranean spurs that give rise to its precocious flowers
and shortly after to the season's leaves are homo
logous to the rapidly elongating brachyblasts of the
woody Macrophyllae.
81. Calliandra longipes Bentham, Trans. Linn. Soc.
London 30: 538. 1875; & in Martius, Fl. bras. 15(2):
410, t. ciii. 1876. — "... in provincia Goyaz inter
Capao S. Joao et Ribeira dos Muencos [misreading
of Macacos]: Burchell n. 6538, 7188, 7499; prope
Paracatu prov. Minas Geraes: Lund." — Lectotypus,
Burchell 7499, K (hb. Hook.)! = N Y Neg. 1950; iso
typus, GH!. — Feuilleea longipes O. Kuntze, Revis.
Gen. PL 1: 188. 1891. FiG. 18
C. longipes fma. nana Chodat & Hassler, Bull. Herb.
Boissier II, 4: 484. 1904. — "[PARAGUAY] ... ad ripam
fluminis Tapiraguay [s.-e. of San Estanislao], Aug., [Has
sler] n. 4312." — Holotypus, G n.v.
C. longipes var. valenzuelensis Chodat & Hassler, Bull.
Herb. Boissier II, 4: 484. 1904. — "[PARAGUAY] ... in campis siccis glareosis pr. Valenzuela, Jan., [Hassler.] n.
7005." — Holotypus, G n.v.; isotypus, NY!. C. pyrophila Fernandez Casas & Schinini, Fontqueria 4: 29,
fig. 1, 2. 1983. — "PARAGUAY, Caaguazu, entre Yhu y
San Bias . . Fernandez Casas 3855 & Molero, 23-IX-1980." — Holotypus, M A n.v.; isotypi, NY!, and (not seen)
CTES, G, MO. C. longipes sensu Glaziou, 1905: 187; Burkart, 1952: 110;
Hoc, 1992:204, Map 1.
Functionally herbaceous, macrophyllidious sub
shrubs of singular habit, with obhquely or horizon
tally creeping, shallowly buried rhizomes said to be
come 1 m long, the precocious scapiform peduncles
arising at or shortly below soil-level, from 1-2 lowest,
efoliate nodes of the erect fohate stems, these hyster-
anthous, attaining 1-3 dm, ±3-5-lvd, hke the lf-axes
and lfts either glabrous or thinly evanescently pilosu
lous, the mature lfts chartaceous venulose; phyllotaxy
distichous. Stipules triangular-ovate or lanceolate 1.5-
3 m m , weakly several-nerved, persistent. Lf-formula
\I\V2, the lfts exactly 6 per If; petioles 4.5-7.5 cm, ±1
m m diam, at apex dilated into a shallow cupule; rachis
of pinnae ±2.5-3 cm, the odd (posterior) 1ft inserted
below middle, as long as or scarcely longer than ter
minal pair; lft-pulvinules discolored ±1 m m ; lft-
blades obhquely obovate or elliptic from shallowly
cordate base, obtuse or sometimes subemarginate, the
138 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Map 38. Distribution of Calliandra longipes Bentham and C. hymenaeodes (Persoon) Bentham in South America.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 139
FiG. 18. Calliandra longipes Bentham. Reproduced from Bentham in Martius, Fl. bras. 15(2): t. 103. 1876.
larger ones 5.5-10 x 2-3.5 cm, 1.8-3.5 times as long
as wide; venation palmate-pinnate, the gently
incurved midrib a little forwardly displaced from
mid-blade, giving rise on each side to 4—8 major
incurved-ascending (and some weaker intercalary)
secondary nerves and an open tertiary reticulum of
veinlets, the inner posterior primary nerve produced
to anastomosis almost to or shortly beyond mid-
blade, the whole venation becoming pallid and
prominulous on both faces. Peduncles at anthesis
6-18 cm, bracteate near or below middle; capitula
12-25-fld, the homomorphic fls subsessile, the
depressed-convex or claviform receptacle 2.5-4.5
m m diam; bracts subulate, linear-oblanceolate, or
-spatulate 0.7-2 x 0.4—0.5 mm, persistent; perianth
glabrous except for brownish-puberulent calyx-teeth
and minutely ciholate corolla-lobes, the pallid calyx
finely ±25-nerved, the distally pink corolla almost
nerveless externally; pedicels 0.3-0.8 x 0.6-0.8 mm;
calyx 2.4—2.7 x 2-2.5 m m , the subtmncate teeth
0.45-1.2 x 0.8-1 mm, at apex incurved and gibbous
dorsally; corolla 8.5-13.3 m m , the ovate lobes
2.4—2.8 mm; androecium 66-82-merous, 21-33 mm,
the stemonozone 1.8-3 mm, the whitish tube 9-18
mm, the tassel red; no intrastaminal nectary. Pods
erect, narrowly oblanceolate 6.5-12.5 x 0.75-0.85
cm, the sutural ribs 3-4- m m wide in dorsal view, the
cmstaceous glabrous valves recessed when young,
becoming low-convex at maturity, sinuously venulose
lengthwise; seeds obovoid-elhpsoid, variably rhombic-
angular, the testa smooth, castaneous or dull gray, the
U-shaped pleurogram pallid but not incised.
In campo subject to fire, ±200-750 m, apparently
local in scattered stations on the Brazihan Planalto, in
n.-e. Argentina, and Paraguay: in Brazil known from
s. Goias, adj. s.-e. Mato Grosso, and w. and centr.
Minas Gerais (Paracatu; Sa. do Inficionado); in
Paraguay from Sa. de Amambay s. to centr. Caaguazu
and Cordillera; in Argentina local in s. Misiones. —
Map 38. — Fl. IX-I.
Calliandra longipes varies in lengths of corolla and
stamen-tube, but flower-size is not significantly cor
related with distribution. The type of C. longipes,
from Goias, and that of var. valenzuelensis, from
Paraguay, have equally long perianth and stamen-
tubes, whereas those of fma. minor and C. pyrophila,
both from province Caaguazu in Paraguay, have
shorter perianth and included stamen-tubes.
II. Sectio ACISTEGIA Barneby
Calliandra sect. Acistegia Bameby, sect. nov. Fru-
tices saepe ramosissimi, microphylhdii, xerophili;
foliorum primariorum stipulae basi spicula patula
(nunc in calcar parvum reducta) armatae; foliorum
formula i/2-18, foholis majoribus 2-9 m m usque
longis; peduncuh e brachyblastis axillaribus nas
centes; praeter aciculos stipulatos a speciebus uni-
jugis microphyllidiis seriei Androcallidis vix diver-
sae. — Sp. typica: C. haematomma (de Candolle)
Bentham.
Calliandra ser. Pedicellatae Bentham, London J. Bot. 3: 100. 1844, sens, str., quoad speciem typicam, reliquis ex-pulsis. — Sp. lectotypica: C. pedicellata Bentham. Anneslia Haematostomae Britton & Rose, 1928: 52, max. ex parte, nom. nud.
Microphyll shmbs and shmblets; stipules associ
ated with primary lvs armed dorsally at base with a
spicule (sometimes reduced to a conical spur) simu
lating and often mistaken for a spinescent stipule; lf-
formula i/2-18, the longer lfts 2-9 mm; peduncles
arising from short axillary brachyblasts. — Spp. 2,
140 M E M O I R S OF THE N E W Y O R K BOTANICAL G A R D E N [VOL. 74(3)
one pluriracial, of the Bahamas, Greater Antilles, and
Virgin Is. (St. Croix).
The peculiar stiff spicule arising from the back of
the primary stipules in sect. Acistegia was correctly
described by Bentham (1844: 102, 103) and by Urban
(1928: 28), but was carelessly misinterpreted by Brit
ton and Rose (1928: 40) as a spinescent stipule, with
which it is by no means homologous although appar
ently similar in defensive function. It is a feature
unique in the genus.
82. Calliandra haematomma (de Candolle) Ben
tham, London J. Bot. 3: 103. 1844. — Typus infra
sub var. haematommate indicatur.
Erect or exceptionally diffuse-depressed, intricately
stiffly branched, microphyll shrubs (0.2-)0.8-2.5(-4)
m tall with gray or blanched older stems, virgate long-
shoots, and acaulous brachyblasts, armed at all or at
most nodes with a pair of straight, ascending or de
clined, slenderly tapering or more stoutly conical,
spinescent spicules arising from the exterior base of
stipules associated with (often caducous) primary lvs,
the young growth variably pilosulous or pubemlent
and commonly glabrescent, the firm plane lfts most
often glabrous but sometimes ciliate, or glabrous ven
trally and thinly strigulose dorsally, or finely pilosu
lous overall, the peduncles and corollas often more
densely pubescent but sometimes glabrous, the capit
ula arising singly (exceptionally geminate) from efoli
ate axils of tightly thatched brachyblasts; phyllotaxy
distichous. Stipules of primary lvs lanceolate or nar
rowly to broadly triangular 0.4—3(-3.5) x 0.4-1.2 m m ,
(3-)5-9-nerved, becoming dry, disintegrating but not
disjointing in age, those of brachyblasts smaller, per
sistent. Lf-formula i/2—3, 3-12, 14—18, but the small
anterior first 1ft sometimes suppressed and the lfts
then of uneven number; lf-stks 0.6-12 x 0.2-0.4 m m ;
rachis of pinnae (0.7-)l-33 m m , the longer interfoh
olar segments 0.4-4- m m ; lft-pulvinules obsolete or to
0.25 m m diam; lfts distally accrescent, the blades obo
vate, oblance-obovate, or narrow-oblong, from shal
lowly semicordate base, broadly obtuse and some
times incipiently apiculate, the penultimate pair 2-9 x
0.8-4 m m , 1.4-4-.1 times as long as wide, the terminal
pair often proportionately wider than the rest; vena
tion prominulous dorsally or sometimes on both faces
of blade, rarely subimmersed, the straight midrib dis
placed to divide blade ±1:1.5-2, 2^-branched on the
broader side, 1 weak posterior primary nerve incurved-
ascending well short of mid-blade. Peduncles 3-25
m m , either bracteate or not, the bract either below
or above middle, <1 m m ; capitula 7-21-fid, the recep
tacle ±1-1.5 m m diam; floral bracts ovate or lanceo
late 0.3-1.6 m m , persistent; fls sessile, homomorphic
as to perianth but some lacking pistil, most bisexual,
the perianth 5(-6)-merous, the calyx dry, brown, stri
ately 15-20-nerved, commonly glabrous externally
(except for pubemlent orifice) but occasionally pu
bemlent beyond the tube or silky-pilosulous overall,
the corolla usually densely subappressed-pilosulous
with white, ± lustrous hairs, in two varieties charac
teristically or randomly glabrous overall; calyx cam
panulate (1—)1.3—3.3 x 1-1.8 m m , the narrowly or
broadly ovate or depressed-ovate teeth 0.2—1.3(—1.8)
m m , one or two sinuses sometimes more deeply split;
corolla 3-5.1 m m , the erect, narrowly ovate or lance
olate lobes (0.5-)0.9-1.6 m m ; androecium (8—)9—17
(-20)-merous, (12.5-)14-25 m m , the tube 2-5.5 m m ,
the stemonozone 0.4—1.2 m m , the tassel most often
bright red, seldom pink, in var. glabrata whitish; no
intrastaminal disc; ovary at anthesis villosulous, 2-5
(-6)-ovulate. Pods 1-6 per capitulum, erect, in broad
profile 3-7 x 0.4-0.9 cm, the sutural keels in dorsal
view ±1.4—2.2 m m wide, the coriaceous, recessed
valves not or only faintly cross-venulose, finely pilose
or pubemlent overall, exceptionally glabrate; seeds in
broad view orbicular-obovate or elliptic-oblong
±5.5-6.5 x 2.5-4.8 m m , the smooth hard testa light
brown, speckled or mottled.
Except for the closely related C. pedicellata, sym
patric in Hispaniola but instantly recognized by its
umbelliform capitula, C. haematomma is the only
calliandra with dorsally spiculate primary stipules. It
is heteromorphic in pubescence of leaves and flowers
and in number and size of leaflets; moreover, the
spicules vary greatly in development, even along one
branch. As material has accumulated, discontinuities
between the segregate species and varieties have be
come increasingly elusive, and the taxonomic level of
several has become precarious. In recognition of this
state of affairs, a comprehensive interpretation of C.
haematomma seems appropriate. The key that follows
separates seven varietal taxa, but these are uniform
neither in area of dispersal nor in degree of internal
variability. The var. glabrata, localized in southeast
ern Jamaica, has one unique character, a white tassel,
which coincides with glabrous corolla, of uncommon
occurrence elsewhere. The vars. locoensis and tor-
tuensis are known each from a single population, and
are essentially monomorphic, so far as known. The
Cuban var. colletioides has on the average fewer
leaflets and simultaneously longer spicules than the
rest, and var. rivularis on the average longer leaf-stalk
and bigger leaflets. Over most of Hispaniola and in
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 141
scattered stations eastward var. haematomma has most
commonly glabrous leaflets and silky-pubescent co
rollas, but forms, otherwise identical, that combine
glabrous leaflets with glabrous (glabrescent) corollas
or pubescent leaflets with either glabrous or silky
corollas, occur at random in the same range, some
times even in one population. It came as a surprise to
find that the Bahaman populations of C. haematomma
have appreciably ampler leaflets than those of His
paniola, such as to deserve equivalent taxonomic
status. There seems to be no partem to the variation
observed in length of calyx and corolla or the propor
tion between them.
The epithet haematomma (Gr. haimat-, blood +
omma, eye or hght, in reference to the brilhantly col
ored filaments) appeared first in the corrupt form
haematoma, and was shortly afterward republished by
Kurt Sprengel as haimatostoma (haimat- + stoma,
mouth, without obvious significance). Both epithets
were attributed to Bertero and it is not known which
was actually written by him. In the protologue of C.
haematomma that appeared in de Candolle's Memoires
sur la famille des Legumineuses, anterior in composi
tion although posterior in pubhcation to that in the Pro-
dromus, the epithet is written haematomma, and it was
this form that Bentham transferred to Calliandra and
that has become standardized in m o d e m literature of
the Antillean flora. Herein C. haematostoma Sprengel
is treated as an independent, taxonomically super
fluous synonym of C. haematomma de Candolle. The
two spelhngs, as they occur in subsequent literature,
are interpreted as orthographic variants.
Key to the varieties of C. haematomma
1. Endemic to s.-e. Jamaica; corolla glabrous and longer pinna-rachises 6-15 mm; filament-
tassel whitish 82f. var. glabrata
1. Allopatric, not Jamaican; corolla mostly
white-silky, but if (sub)glabrous the longer
pinna-rachises 2.5-5 mm; filament-tassel red
or rarely pink. 2. Lf-stk of primary lvs 4-12 mm; longer
lfts 6-8.5 mm; local in n.-e. Dominican Republic and adj. Haiti.. . ... 82d. var. rivularis
2. Lf-stk of primary lvs commonly 0.6-4
m m , to 6 m m only on the Bahamas. 3. Lfts of longer pinnae mostly 2-8 pairs,
but 9-12 pairs in var. tortuensis of
n.-w. Haiti. 4. Lfts 2-3 pairs on rachis 0.7-2.5
m m ; e. Cuba and perhaps extreme
s.-w. Hispaniola 82a. var. colletioides
4. Lfts (3-)4—12 pairs on rachis 3-8
m m ; Bahamas, Hispaniola, s.-w.
Puerto Rico, St. Croix.
5. Lfts of longer pinnae 9-12
pairs, He Tortue, n.-w. Haiti 82e. var. tortuensis
5. Lfts of longer pinnae (3-)4-8
pairs; range as given in choice
4(2). 6. Bahama Is.; rachis of longer
pinnae 7-21 m m and ampler lfts 4-10 m m 82c. var. correllii
6. Allopatric; Hispaniola e.-
ward; rachis of longer
pinnae 3-8 m m and ampler
lfts 2.2-6 mm.82b. var. haematomma
3. Lfts of longer pinnae 11-18 pairs;
local on serpentine in s.-w. Puerto
Rico. . . . . 82g. var. locoensis
82a. Calliandra haematomma (de Candolle) Ben
tham var. colletioides (Grisebach) Bameby, comb.
nov, based on autonym generated by C. colletioides
var. gonavensis Urban, 1929 (vide infra). C. col
letioides Grisebach, PL Wright. 1: 180. 1860 =
M e m . Amer. Acad. Arts II, 8: 180. 1861. —
"[CUBA. Oriente:] in praeruptis montium prope
Nouvelle Sophie, etc. . . . [C. Wright] 153." —
Holotypus, G O E T n.v; isotype, A!. — Feuilleea
colletioides O. Kuntze, Revis. Gen. PL 1: 187.
1891. Anneslia colletioides Britton, Bull. Torrey
Bot. Club 41: 18. 1914.
C. colletioides sensu Bentham, 1875: 575; in Hooker, Icon.
12: pi. 1167. 1876; Leon & Alain, 1951: 238; Bassler,
1990: 201, fig. 5(a), Karte 2, t. X(a). Anneslia colletioides
sensu Britton & Rose, 1928: 75. C. colletioides var. gonavensis sensu Liogier, 1985: 28.
Commonly erect and 1-2 m; lvs either glabrous, or
glabrous ciholate, or thinly pubemlent dorsally; stip-
ular spicules (1—)2—11 (—13) m m ; petiole of primary
lvs 0.6-1.6 m m and longer rachises (0.7-) 1-2.5 m m ,
the interfoholar segments 0.4—1 m m ; lfts 2-3 pairs, the
penultimate 3-6 x 2-3.5 m m ; calyx glabrous except
sometimes for pubemlent teeth; corolla white-silky.
In xeromorphic shmb communities and on savanna,
mostly on limestone but occasionally on serpentine,
locally plentiful below 200 m along and near the s.
coast of Oriente, Cuba, from Cabo Maasi w. to Santi
ago, and in scattered stations, inland and on the n.
coast of the province, to Holguin and Nicaro; one
record from Camaguey. — M a p 39. — Fl. XII-IV,
VII-VHI.
82b. Calliandra haematomma (de Candolle) Ben
tham var. haematomma. C. haematomma Bentham,
1844, I.e., sens. str. Acacia haematomma de Can
dolle, Prodr. 2: 456 ("haematoma"). 1825; & M e m .
Legum. 8: 448, t. LXVIII. 1827. — ". . . decouverte
142 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
... Saint-Domingue par M. Bertero." — Holotypus,
G-DC!. — Feuilleea haematomma O. Kuntze, Revis.
Gen. PL 1: 188. 1891. Calliandra haematomma
("haematostoma") var. genuina Urban, Symb.
Antill. 2: 263. 1900.
Acacia haematostoma Bertero ex Sprengel, Syst. 3: 137. 1826. — "Hispaniola." No typus known to survive, but doubtless based on part of the same collection described by
de Candolle, as indicated by Steudel, Nomencl. Bot. 5:
1840, who illegitimately adopted the posterior
haematostoma as correct. Anneslia haematostoma Britton,
Mem. Brooklyn Bot. Gard. 1: 50. 1918.
Calliandra haematostoma var. pubescens Urban, Symb. An
till. 2: 263. 1900. — "Hab. in Sto. Domingo: Bertero, Rob.
Schomburgk, n. 104." — Syntypi, ̂ B; no duplicates found.
— Calliandra haematomma var. pubescens Macbride,
Contr. Gray Herb., n. ser. 59: 6. 1919. — Equated with An
neslia haematostoma by Britton & Rose, 1928: 74.
C. haematostoma var. minutifolia Urban, Symb. Antill. 2:
263. 1900. — "Hab. in Haiti prope Cadets, 1200 m. alt.:
Picarda n. 869." — Holotypus, *B; no isotypus found. —
C. haematomma var. minutifolia Macbride, Contr. Gray
Herb., n. ser. 59: 6. 1919. Anneslia minutifolia Britton &
Rose, N. Amer. Fl. 23: 75. 1928. Calliandra minutifolia Urban, Ark. Bot. 24A(4): 6. 1931; non C. minutifolia Pit-
tier, 1927. C. picardae Alain, Brittonia 20: 156. 1968.
(?) C. colletioides var. gonavensis Urban & Ekman ex Urban, Ark. Bot. 22A(8): 28. 1929. — "Haiti, insula Gonave in
Trous-Louis . . . m. Aug . . . [Ekman] 8814." — Holotypus,
^B (sterile, with half a pod); isotypus, K!. — Calliandra
colletioides subsp. gonavensis Bassler, Gleditschia 18: 202.
1990. Mimosa spartioides Vahl ex de Candolle, Mem. Legum. 8:
448, pro syn. 1827.
Calliandra haematomma sensu Bentham, 1875: 545.
Commonly l-3(-4) m tall, but in shallow soils, on
rocks, or in windswept places depressed or diffuse; lvs
glabrous to finely pilosulous, sometimes in one popu
lation; stipular spicules 0.5-7 m m , sometimes lacking
at some nodes, exceptionally 0; petiole of primary lvs
l-3(-4) m m and longer rachises 3-8(-10) m m , the
longer interfoholar segments 0.3-1.4(-2) m m ; lfts
3_8(-10) pairs, the penultimate 2.2-6 x 0.9-
3(—3.5) m m ; calyces usually glabrous to sometimes
ciholate orifice, rarely silky-pilosulous overall; corolla
commonly white-silky, sometimes minutely pubem
lent, sometimes glabrous.
In xeromorphic shrub communities between sea
level and 1300 m, commonly on limestone, locally
plentiful, widespread over much of Hispaniola and
known from single localities in s.-w. Puerto Rico
(Guayanilla) and on St. Thomas, Virgin Is. — M a p
39. — Flowering almost yearlong, but most prolifi
cally VII-XII. — Clavelina; tabacuela.
82c. Calliandra haematomma (de Candolle) Ben
tham var. correllii Bameby, var. nov, cum var.
haematommate, in Hispaniola endemica, in insulis
bahamensibus vicarianti et ab hac vix nisi foliolis
amplioribus, ut infra descriptis, diversa. — BA
H A M A IS. Great Abaco: ±1.5 mi. N W of Marsh
Harbour, 13 Mar 1975 (fl, fr jun), D. S. Correll
(with F G. Meyer) 44596. — Holotypus, NY.
C. haematomma sensu Bentham, 1844: 103, ex parte, quoad
pi. baham. swainsonianam; Britton & Millspaugh, Bahama
Fl. 159. 1920; Correll & Correll, Fl. Bahama Archipel.
610. 1982.
Erect, potentially arborescent, \-A m, with pliant
long-shoots; lvs glabrous or almost so; stipular
spicules 0.3-3.5 m m ; petiole of primary lvs l-6(-7)
m m and longer rachises (6-)7-21 m m , the longer in
terfoholar segments (1.7—)2-4 m m ; lfts 5-8 pairs, the
penultimate (3.5-)4-10 x 1.5-4 m m ; calyx glabrous;
corolla white-silky.
In coppice below 50 m, locally abundant on the
Bahama archipelago from Grand Bahama and Abaco
s.-e. to Inagua and South Caicos (not recorded from
N e w Providence, nor from Andros and associated
islets). — M a p 39. — Fl. nearly throughout the year,
most prolifically XI-III.
Correll (1982: 612) called Bahaman C. haema
tomma "apparently endemic," though this is paradox
ical for a species described from Hispaniola. It is
agreed, nevertheless, that the Bahaman populations
of C. haematomma indeed represent an infraspecific
taxon endemic to the Archipelago.
82d. Calliandra haematomma (de Candolle) Ben
tham var. rivularis (Urban & Ekman) Bameby, stat.
nov. C. rivularis Urban & Ekman, Ark. Bot. 24(4):
5. 1932. — "Haiti, Massif du Nord prope Valliere ad
Grande Savane . . . cr. 250 m alt., m. Majo [1928]
flor.: IE. L. Ekman] n. H 9948." — Holotypus, fB;
isotypi, K!, NY!.
Shmbby, 1-2 m; lfts glabrous, sometimes pallid;
stipular spicules 1-6 m m ; petiole of primary lvs 4—12
m m and longer rachises 12-25 m m , the longer inter
foholar segments 2-3 m m ; lfts 7-9 pairs, the penulti
mate 6-8.5 x 1.8-3 m m ; calyx glabrous; corolla thinly
silky; pods (little known) glabrous or glabrescent.
In riparian thickets, sometimes in pine forest, 120-
700 m, locally plentiful in the foothills of Cord. Cen
tral in n.-e. Dominican Republic (Libertador, Santiago
Rodriguez, n.-w. Santiago) and adj. Haiti (Nord).
Notable for relatively long petioles and ample leaf
lets, features perhaps reflecting the riparian habitat,
but not essentially different in flower or fmit from C.
haematomma sens. lat.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 143
CALLIANDRA H A E M A T O M M A
var. CORRELLH
var. COLLETIOIDES
var. GLABRATA
var. H A E M A T O M M A
MAP 39. Distribution of Calliandra haematomma (de Candolle) Bentham var. haematomma, var. correllii Bearneby, var. colletioides (Grisebach) Barneby, and var. glabrata Grisebach in the West Indies. Not mapped are vars. rivularis (Urban & Ekman) Barneby, tortuensis (Alain) Bameby, locoensis (Garcia & Kolterman) Bameby, and gonavensis.
82e. Calliandra haematomma (de Candolle) Ben
tham var. tortuensis (Alain) Bameby, stat. nov. C.
tortuensis Alain, Brittonia 20: 156. 1986. — "HIS
PANIOLA. Haiti: ... W of La Vallee, Tortue Island,
28 Dec 1928, E. & G. Leonard 11565." — Holo
typus, NY!; isotypus, US!.
Slender shrub to 1 m tall; lvs thinly minutely pilo
sulous overall; stipular spicules 0.5-1.5 mm; petiole
of primary lvs 1.5-3.5 m m and longer rachises 9-12
mm; lfts 9-12 pairs, the penultimate ±7.55 x 2 mm;
calyx glabrous; corolla white-silky.
On rocky bank of lagoon near La Vallee, Tortue I.,
Haiti (dep. Nord Ouest), known only from the type-
collection. — Fl. XII-I(-?).
The typus of C. tortuensis was annotated by Brit
ton as a new species of Anneslia, but this name was
left unpublished. The specimens have nothing that
could exclude them from C. haematomma sens, lat.,
but are peculiar in the combination of number and pu
bescence of leaflets.
82f. Calliandra haematomma (de Candolle) Ben
tham var. glabrata Grisebach, Fl. Brit. W. I. 224.
1860, based in part on Acacia pilosa Bertero ex de
Candolle, Prodr. 2: 455. 1825. — ". . . in Jamaica
(Bertero)." — Holotypus, G-DC!, verified by
Urban, I.e. infra. —Anneslia pilosa Britton & Rose,
N. Amer. Fl. 23: 75. 1928. Calliandra pilosa Urban,
Ark. Bot. 24A, 4: 6. 1931. Fig. 19
C. haematomma var. glabrata sensu Urban, Symb. antill. 2: 263. 1900. C. haematomma sensu Fawcett & Rendle, Fl. Jamaica 4: 143. 1920. C. pilosa sensu Adams, Fl. PL Jamaica 332. 1972.
Arborescent shmbs 2-4 m with virgate branches,
differing from all other varieties of C. haematomma
in whitish, not red or pink filaments; lfts mostly
glabrous, but sometimes ciholate or thinly strigulose
dorsally; stipular spicules to 4.5 mm; petiole of pri
mary lvs 1-2.5 m m and longer rachises 6-15 m m , the
longer interfoholar segments 1-2.1 m m ; lfts 6-8(-9)
pairs, the penultimate 3.7-7 x 1-1.7 mm; calyx
glabrous; corolla glabrous, or its tube thinly strigu
lose and its lobes glabrous.
On dry bmshy hillsides and in coppice along the
shore, 1-180 m, locally plentiful but apparently con
fined to the coast and foothills of s.-e. Jamaica, from
Kingston Bay in Saint Catherine and Saint Andrew
parishes w. into Clarendon Parish. — Map 39. — Fl.
IX-II.
De Candolle described the leaflets of Acacia pilosa
as up to 13 pairs per pinna. His material was sterile,
144 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
FiG. 19. Calliandra haematomma (de Candolle) Bentham var. glabrata Grisebach.
but his description of the stipules and their dorsal
spicule is definitive for C. haematomma in the con
text of the Jamaican flora. The first anterior leaflet of
each rachis is sometimes wanting and the leaflets
therefore of odd number; the 13 pairs may be a slip
for 13 leaflets, seen in some modem gatherings.
82g. Calliandra haematomma (de Candolle) Ben
tham var. locoensis (Garcia & Kolterman) Bameby,
stat. nov. C. locoensis Garcia & Kolterman,
Caribbean J. Sci. 28: 57, figs. 1, 2. 1992. — "Puerto
Rico, Mun. Yauco, Bo. Sosua Alta, Bosque Estatal
de Sosua, en las orillas de una quebrada que desem-
boca en el Rio Loco ... 17 feb. 1991, Garcia &
Caminero 3425." — Holotypus, n.v; isotypi, NY!,
and (not seen) JBSD, K, M O , RAC, UPR, US.
Shmbs to 3 m tall; lfts facially glabrous, ciholate;
stipular spicules 3-13 mm; petiole of primary lvs 2-4
m m and longer rachises 25-35 m m , the longer inter
foholar segments 1.4—1.8 mm; lfts of larger lvs
14-18(-19) pairs, the penultimate ±5-9 x 1—1.7(—3)
mm; calyx glabrous except for ciholate teeth; corolla
white-silky; pod thinly pilosulous glabrescent.
On creek banks, on serpentine bedrock, 170-200
m, locally plentiful but known only from the type-
locality in the basin of rio Loco, near 18°05'N, 66°
54'W, in s.-w. Puerto Rico. — FL I—III(—?).
The var. locoensis is visually arresting in the context
of C. haematomma because of the relatively long leaf-
rachises and numerous leaflets, but has everything else
in common with var. haematomma. The var. tortuensis
approaches it in leaflet-number, but is probably an in
dependent modification in the same direction. The one
known Puerto Rican population of C. haematomma, at
Guayanilla, is strictly typical var. haematomma.
83. Calliandra pedicellata Bentham, London J. Bot.
3: 102. 1844. — "Haiti, [C. A.] Ehrenberg [385]."
— Holotypus, collected Jun, 1831, "hB. — Feuilleea
pedicellata O. Kuntze, Revis. Gen. PL 1: 188. 1891.
Anneslia pedicellata Britton & Rose, N. Amer. Fl.
23: 76. 1928. Fig. 20
C. pedicellata sensu Bentham, 1875: 545; Urban, Symb. antill. 8: 254. 1920; Alain, 1985: 30.
Erect, few-stemmed, xeromorphic microphyll
shmbs 0.6-3 m with virgate long-shoots densely
clothed in short recurved lvs, armed at most nodes with
a pair of either ascending or declined post-stipular
spicules or conical spurs (0.5-)l-^-.5(-5.5) m m , the
young stems, lf-axes and peduncles thinly or remotely
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 145
FiG. 20. Calliandra pedicellata Bentham.
pilosulous with weak fine hairs 0.3-1 mm, the plane
coriaceous lfts facially glabrous, sometimes remotely
ciholate or microscopically granular-ciholate, slightly
discolorous, the long-pedunculate umbelliform racemes
of long-pedicellate fls arising singly from 1-2 efohate
nodes of very short brachyblasts axillary to distal pri
mary lvs, these units of inflorescence together forming
a compact or loosely elongate terminal psudoraceme.
Stipules of primary lvs ovate or lanceolate 1-3.5 x
0.5-1.1 (-1.4) mm, the papery blades appressed, stri
ate, persistent, those of brachyblasts similar but
smaller, lacking basal spurs. Lf-formula i/(4-)
5-9(-10); lf-stk of primary lvs, including pulvinus,
0.6-1.6(—2.1) mm, dorsiventrally compressed, shal
lowly grooved ventrally; pinna-rachises of primary lvs
5—12(—15) mm, the longer interfoholar segments
0.8-1.6 mm; lft-pulvinules 0.1-0.2 x 0.15-0.25 mm;
lfts usually a httle accrescent upward except for the
often shorter and broader terminal pair, the blade of
those near and above mid-rachis oblong or narrow-
oblong from semicordate base, some a trifle widened
toward apex, all obtuse or apiculate, the longer ones
(3-)3.2-5.5(-5.7) x l-1.8(-2) mm, 2.3-3.6 times as
long as wide; venation palmate-pinnate, immersed or
nearly so ventrally, bluntly prominulous dorsally, the
midrib forwardly displaced to divide blade 1:1.5-2,
2-3(-4)-branched on one or both sides, a weak inner
(or only) posterior primary nerve incurved-ascending
to anastomosis well short of mid-blade. Peduncles
(1.5—)2—3.5 cm, bracteate above middle; capitula um
belliform (10-)12-24(-26)-fld, the floral axis 1.5-3.5
mm; bracts papery, narrowly ovate or linear-subulate
0.5-1.3 mm, incurved, persistent; pedicels all equilong
or some distal ones a httle shorter, the longest (2-)
2.5-8 mm; fls homomorphic (except some staminate,
some bisexual), the glabrous perianth (4—)5(-6)-mer-
ous; calyx campanulate or hemispherical 1.8-2.3 x
2-2.8 mm, the tube ±15-nerved, the triangular or de
pressed-ovate teeth 0.4-1.1 mm; corolla 4.8-6 mm, the
ovate lobes (0.9-) 1.2-2.4 mm; androecium 16-20-
merous, 14—21 mm, the stemonozone 1.3-2.1 mm,
thickened internally, the tube 3.6-5 mm, the tassel dull
white or ochroleucous; disc 0; ovary usually villosu
lous, sometimes glabrous at anthesis. Pods in profile
4—7.5 x 0.6-0.8 mm, subappressed-silky-pilose over
all, the sutural ribs in dorsal view 2-2.5 m m wide, the
stiffly leathery valves low-convex over each of 3-6
seeds, these (few seen) ± 5 x 4 mm, hght brown, finely
pleurogrammic.
In cactus-thornscrub, deciduous thom-forest, and
xeromorphic thickets, from near sea level to 380 m,
discontinuously widespread in parts of w. Hispaniola:
n.-w. and centr. Haiti, in departements Nord-Ouest,
Artibonite and Ouest; s.-w. Dominican Republic
in provinces Independencia, Barahona, Azua, and
146 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Peravia. — Fl. intermittently, most prolifically fol
lowing rains.
Calliandra pedicellata has the characteristic, dor
sally appendaged stipules of sect. Acistegia and in
vegetative characters closely resembles the polymor
phic and sympatric C. haematomma. Its umbelliform
capitula and whitish tassel of filaments are, however,
substantially different.
III. Sectio ACROSCIAS Bameby
Calliandra sect. Acroscias Bameby, sect. nov. mono
typica, inflorescentia determinata ab omnibus con-
generibus diversa; suffmtices, caulibus herbaceis e
rhizomate subterraneo ortis in capitulum umbelli-
forme terminale exeuntibus, ahter sect. Androcallidis
species plurijugas microphylhdias simulantes. —
Sp. unica: C. brevicaulis M. Micheh.
Functionally herbaceous subshrubs arising anew
each year from subterranean caudex or rhizome, each
stem (and rare lateral branches) terminating in an um
belliform capitulum, in other respects resembling
plurijugate microphyllidious spp. of sect. Androcal
lis. — Sp. 1 (2 vars.) of campo habitats in cisparagua-
ian Paraguay, adj. Brazil (Mato Grosso do Sul), and
n.-e. Argentina (Misiones, Corrientes). — Potential
pyrophytes.
The determinate inflorescence of C. brevicaulis
was first described by Spegazzini (1926) and has not
been observed elsewhere in the genus. Spegazzini
seems to have thought that C. longipes also has de
terminate peduncles, but in fact they are axillary,
though elongate and subscapose.
84. Calliandra brevicaulis M. Micheh, Mem. Soc.
Phys. Geneve 28(7): 82. 1883. — Typus infra sub
var. brevicauli indicatur. FiG. 21
Functionally herbaceous or subherbaceous, fire-
resistent subshrubs of campo habitats, the one or few
stems erect or incurved-ascending from obhque hg
nescent rootstock, commonly simple 1-4 dm and
monocephalous, less often 2-5-branched distally and
each efoliate or 1-lvd branch monocephalous, excep
tionally attaining 7 dm, the units of inflorescence all
terminal to their axis of origin, the stems, lvs, and fls
glabrous to variably pilose, pilosulous, or silky-
strig(ul)ose, the small imbricate lfts almost always
facially glabrous, either cili(ol)ate or not, bicolored,
lustrous dark olivaceous above, paler beneath, the pro
portionately large umbelliform capitula shortly
emersed from foliage, white to pink or crimson. Stip
ules narrowly ovate, narrowly lance-elliptic, or linear-
lanceolate 2-5.5(-7) x 0.5-2.2(-2.5) mm, weakly
3-5(-7)-nerved, tardily deciduous, expanded at base
into a shallow cupule nidulating the pulvinus of each
lf-stk. Lf-formula (ii-)iii-viii/20-42; lf-stk of longer
lvs 3-7.5 cm, the petiole (4—)6—13 cm, the rachis
cupulately distended at insertion of pinnae, the longer
interpinnal segments 5—13(—18) mm, the ventral
groove bridged between pinnae; pinnae scarcely (but
sometimes randomly) graduated, the rachis of longer
ones 2.5-7.5(-8) cm, the longer interfoholar segments
0.7-1.9 mm; lft-pulvinules 0.1-0.3 mm; lfts subequi
long except at very ends of rachis, in outline narrowly
lance-oblong from obtusangulate base, acute or mu-
cronulate, those near mid-rachis (3.3-)3.5-7.5 x 0.75-
1.9 mm, 3-5 times as long as wide; venation palmate-
pinnate, the straight or distally subporrect midrib for
wardly displaced to divide blade l:2-2.7(-3), giving
rise on posterior side to 2-4(-5) widely diverging sec
ondary veinlets brochidodrome well within the plane
margin, the 1-2 posterior primary nerves very short,
the whole venation pallid and finely prominulous dor
sally, immersed or almost so above. Peduncles (mea
sured from furthest If) 2.5-10.5 cm, bracteolate or not;
capitula cymose-umbelhform 12-26-fld, the clavate
receptacle including short terminal pedestal 2-5 mm;
floral bracts often 0, those subtending random outer
most fls hnear or linear-oblanceolate 2-4.5 mm, de
ciduous; fls of most capitula dimorphic, the peripheral
ones long-pedicellate and the central one either short-
pedicellate or subsessile, stouter than the rest but
hardly longer and not otherwise modified, in some
capitula wanting or degenerate; PERIPHERAL FLS:
pedicel to 5-26 x 0.4—1.1 mm, sulcate lengthwise,
dilated under the calyx proper into a solid turbinate
head 0.8-1.7 m m tall; perianth (5-7-merous) thin-tex
tured, whitish, brownish or pink-tinged, either
glabrous or variably pubescent, weakly few-nerved;
calyx narrowly or openly campanulate 3-5.2(-6) x
2.3-3.5(-4) mm, the ovate or depressed-ovate teeth
(0.4-)0.8-1.4 mm; corolla vase-shaped (8.2-)9-14
mm, the ascending ovate lobes often unequal, the
longest ±2.5-4.5 mm; androecium 34-44-merous,
4.5-6(-7) cm, the stemonozone (1-) 1.2-2.4 mm, the
tube 2.5-5(-6) mm; no intrastaminal nectary; ovary
subsessile, at anthesis smooth or micropapillate. Pods
\-i per capitulum, erect, in profile 5-9 x 0.6-1.2 cm,
the sutural ribs 2.5 m m wide or more, the recessed
valves not perceptibly venulose, densely gray- and
sordid-pilose overall.
Calliandra brevicaulis is instantly recognized by
its truly terminal peduncles, and is further notable in
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 147
FiG. 21. Calliandra brevicaulis M. Micheli.
its area of dispersal for short stems arising annually
from an often strongly hgnescent, sometimes creeping
rootstock, and for the proportionately great size of the
umbelhform capitula. The plants are diverse in length,
orientation, and distribution of hairs on stem, leaves,
and flowers, several states of which were described by
Hassler as varieties or forms. Much of the variation
seems random, even though relatively stabihzed within
a population. There is, however, a strong correspon
dence between dispersal, leaflet-number, and pubes
cence of the flowers which calls for recognition of two
infraspecific taxa.
148 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Key to the varieties of C. brevicaulis
1. Perianth puberulent, silky-strigulose, or sometimes
pilosulous externally; lfts of longer pinnae 30-
42 pairs; centr. and n. cisparaguaian Paraguay;
and in immediately adj. Mato Grosso do Sul,
Brazil 84a. var. brevicaulis
1. Perianth glabrous externally; lfts of longer pinnae
20-33 pairs; w.-centr and s. cisparaguaian
Paraguay s. into s.-w. Misiones and Corrientes
in Argentina 84b. var. glabra
84a. Calliandra brevicaulis M. Micheli var. brevi
caulis. C. brevicaulis M . Micheli, 1883, I.e., sens.
str. — "[PARAGUAY.] Caaguazu in campis; Mart.,
[Balansa] n. 1436." — Holotypus, G n.v; isotypus,
K!. — C. brevicaulis var. genuina Chodat & Has
sler, Bull. Herb. Boissier II, 4: 484. 1904. C. brevi
caulis var. genuina Hassler, Feddes Repert. Sp.
Nov. Regni Veg. 8: 556. 1910 (presented as a new
combination for revised circumscription).
C. brevicaulis var. genuina fma robusta Chodat & Hassler,
Bull. Herb. Boissier II, 4: 484. 1904. — "[PARAGUAY] in
campis Ipe hu Sierra de Maracayu . . . [Hassler] n. 4980."
— Holotypus, G n.v.; isotypi, BM!, GH!, K! [the plant so
numbered at A = var. glabra]. — C. brevicaulis var. pubes
cens fma. robusta Hassler, Feddes Repert. Sp. Nov. Regni
Veg. 8: 556. 1910. C. brevicaulis var. puberula Hassler, Repert. Spec. Nov.
Regni Veg. 8: 555. 1910. — "Paraguay: Villa Sana . . . Fiebrig no. 4862 in Herb. Hassler." — Holotypus, G n.v.;
isotypi, BM!, GH!, K! [calyx ciliate but otherwise
glabrous, intermediate to var. glabra]. C. brevicaulis var. puberula fma. hirsutula Hassler, Feddes
Repert. Sp. Nov. Regni Veg. 8: 555. 1910. — "Paraguay: in
campis siccis in regione cursus superioris fluminis Apa, Has
sler no. 7765." — Holotypus, G n.v.; isotypi, A!, GH!, K!. C. brevicaulis var. pubescens Hassler, Feddes Repert. Sp.
Nov. Regni Veg. 8: 556. 1910. — "Paraguay: in campis arenosis pr. Estrella . . . Hassler no. 10135 leg. Rojas." —
Holotypus, G n.v.; isotypi, BM!, NY!. C. brevicaulis var. pubescens fma. intermedia Hassler, Fed
des Repert. Sp. Nov. Regni Veg. 8: 556. 1910. — "Paraguay: in campo pr. flumen Carimbatay . . . Hassler
4560." — Holotypus, G n.v.; isotypus, BM!.
C. brevicaulis var. tomentosa Hassler, Feddes Repert. Sp.
Nov. Regni Veg. 8: 556. 1910. — "Paraguay: . . . prope
Cerro Cora . . . Hassler no. 9736." — Holotypus, G n.v.
As described for the species except as modified by
key to varieties.
In campo, often on sandy soils, ±250-550 m, locally
plentiful in cisparaguaian Paraguay, in deptos. A m a m
bay, Alto Parana, Canindeyu, and Caaguazu, and in im
mediately adj. mun. Amambai in Mato Grosso do Sul,
Brazil. — M a p 40. — Fl. IX-III. — Nino sote.
84b. Calliandra brevicaulis M. Micheh var. glabra
Chodat & Hassler, BuU. Herb. Boissier II, 4: 484.
1904. — "[PARAGUAY.] in campis pr. Itacumbi...
[Hassler] n. 1628; . . . pr Caraguatay . . . n. 3292;
Cordillera de Altos ... n. 2974 ... pr. San Estanislao
. . . n. 4201." — Lectoholotypus, Hassler 2974, G!;
isotypus, K!. — Calliandra glabra Hassler, Feddes
Repert. Sp. Nov. Regni Veg. 8: 555. 1910.
C. brevicaulis var. glabra fma. roseiflora Chodat & Hassler,
Bull. Herb. Boissier II, 4: 484. 1904. — "[PARAGUAY] in
campis pr. Valenzuela . . . [Hassler] n. 6991." — Holo
typus, G n.v.
C. breviflora var. glabra fma. grandiflora Hassler, Feddes
Repert Sp. Nov. Regni Veg. 8: 555. 1910. — "Paraguay: in
campis pr. San Estanislao; Hassler n. 4201 . . . pr. Valen
zuela . . . Hassler no. 6991." — Syntypi, G n.v.
C. brevicaulis var. glabra fma. parviflora Hassler, Feddes
Repert. Sp. Nov. Regni Veg. 8: 484. 1910. — "Paraguay:
Cordillera de Altos, Hassler no. 1628, 2974, 3292." —
Syntypi, G n.v. C. brevicaulis sensu Spegazzini, 1925: 190; Burkart, 1952:
111, fig. 12; Hoc, 1992: 209, fig. 3 + map 1.
C. parvifolia sensu Britton, Ann. New York Acad. Sci. 7:
101. 1892 (Morong 412, NY!).
As described for the species except as modified by
key to varieties.
In lowland campo, openings in matorral, in palm-
savanna, and along rights of way, 150-300 m, scat
tered in w.-centr. and s. cisparaguaian Paraguay, in
deptos. San Pedro, w. Caaguazu, Cordillera, Paraguari,
and Itapua, thence s. in Argentina to s.-w. Misiones
(Posadas) and Corrientes (Paso de los Libres). — M a p
40. — Fl. XII—III, VIII. — Plumerito; anchiguito.
IV Sectio CALLIANDRA
Calliandra Bentham sect. Calliandra. Calliandra
Bentham, 1840, I.e., sens. str. — Sp. typica (conser-
vanda): C. houstoni (L'Heritier) Bentham = C. hous
toniana (Miller) Standley. Calliandra sect. Eucal-
liandra Grisebach, F. Brit. W . Ind. 224. 1864, nom.
superfl.
Anneslia Salisbury, Parad. lond. 64. 1807, nom. rejic. — Sp.
unica: A. falcifolia Salisbury, nom. illegit. = Calliandra
houstoni (L'Heritier) Bentham = C. houstoniana (Miller) Standley.
Inga sect. Annesleya G. Don, Gen. hist. 2: 396. 1832. — Sp.
lectotypica: Inga houstoni (L'H6ritier) G. Don = Calliandra
houstoniana (Miller) Standley. — Because Don did not cite
the anterior generic name Anneslia, his sect. Annesleya must be treated as an independent proposition.
Inflorescence terminal to stems or branches of cur
rent year, pseudoracemose, efohate either throughout
or at least at distal nodes, either simple or rarely (ser.
Comosae) paniculately branched; no brachyblasts;
peduncles arising directly from nodes of primary in
florescence-axis, only exceptionally bracteate beyond
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 149
MAP 40. Distribution of Calliandra brevicaulis M. Micheli var. brevicaulis and var. glabra Chodat and Hassler in South America.
base. — Series 4, in dispersal almost coextensive with
the genus, but not extending into temperate latitudes.
TV/A. Series CALLIANDRA
Calliandra Bentham sect. Calliandra ser. Callian
dra. Calliandra ser. Racemosae Bentham, London
J. Bot. 3: 111. 1844, nom. superfl. — Sp. typica:
eadem ac typus sectionis Calliandrae.
Anneslia Houstonianae, Palmeranae, Wendlandianae, Brit
ton & Rose, 1928: 52, nom. nud.
Primary axis of pseudoraceme simple, terminal to
homotinous stem or branches; phyllotaxy distichous;
primary nerves of lfts conventionally palmate, or re
duced to the costa, or immersed, not parallel length
wise along blade; androecial tube included or scarcely
exserted. — Spp. 39, forming 2 geographical assem
blages: 6 spp. native in Mexico and Central America,
some widely cultivated elsewhere; 33 spp. elaborately
differentiated in interior e. Brazil, many of them mor
phologically specialized and highly locahzed. The
two groups appear monophyletic but are not separable
by any one technical character.
85. Calliandra lintea Bameby, sp. nov, ut videtur C.
bahianae et C. nebulosae Bameby affinis, ab ilia
floribus magis numerosis (utriusque capituli
plerumque 7-11, nee 3-6) et perianthio parvulo,
haud rubro-granuloso, ab hac foliorum formula
ix-xiii/36-50 (nee iv-vi/24-35), fohohsque 3-5.5
(nee 5-8) m m usque longis distans. — BAHIA.
Mun. Lencois: O n trail to Barro Branco ±5 k m n. of
Lencois, 13 Jun 1981 (fl, fallen fr), S. A. Mori (with
B. M. Boom) 14400. — Holotypus, C E P E C ; isotypi,
K, NY.
Microphyllidious shmbs 8^10 dm with stout, sim
ple or distally few-branched, virgate long-shoots but
no axillary brachyblasts, the young growth densely
livid-granular and sometimes in addition minutely
thinly strigulose or pilosulous with white hairs <0.3
m m , the leaves scarcely bicolored, dorsally either
granular or resin-spotted, ventrally sublustrous, gla
brous or micropuberulent, ciholate, the capitula borne
solitary or 2-3 together at bracteate (efoliate) nodes
near apex of main stem and branches in shortly
exserted pseudoracemes; phyllotaxy distichous. Stip
ules narrowly triangular or broad-subulate ±1-1.5 x
0.7-1 m m , not externally venulose, caducous. Lf-
formula ix-xiii/36-50; lf-stk of longer lvs (4.5-)
5.5-10 cm, the dilated petiole 3^1.5 x 1.2-2 m m , the
longer interpinnal segments 4—8(-9.5) m m , the ven
tral groove bridged at insertion of pinnae; pinnae
subequilong or decrescent near apex of lf-stk, the
rachis of longer ones (2.5-)3.5-5.5 cm, the longer
interfoholar segments 0.5-0.9 m m ; lft-pulvinules
±0.2 x 0.3-0.45 m m ; lfts equilong except at very ends
of rachis, the blades linear or linear-lanceolate from
auriculate base, obtuse, straight or a trifle incurved at
apex, the longer ones 3-5.5 x 0.7-1.2 m m , (3.7-)
4.2-6.1 times as long as wide; venation prominulous
only abaxially, palmate, the simple or distally 1-2-
branched midrib displaced to divide blade 1:3.2-5,
the weak inner posterior primary nerve produced
beyond mid-blade, the outer 1-2 much shorter. Pedun
cles 2-5.5 cm, ebracteate, the hemispherical recepta
cle 2-3.5 m m diam; capitula (5—)7—1 l(-12)-fld; bract
of peripheral fls deltate-ovate <1 m m , deciduous, that
of inner fls wanting; pedicels 0.5-0.6 x 1-1.2 m m ,
perceptible only in longitudinal section; perianth 4-
merous, moderately carnosulous, appearing nearly
glabrous, sometimes thinly minutely strigulose or
remotely minutely resin-spotted (not red-granular);
calyx shallowly campanulate or broadly turbinate
1.7-2 m m , faintly 4(-12)-nerved, the depressed-
deltate teeth 0.3-0.4 m m ; corolla 5-5.5 m m , the
broadly ovate obtuse lobes 2-2.2 m m , not nerved
externally; androecium 42-72-merous, 22-23 m m ,
the stemonozone 1-1.5 m m , the homy tube 4.5-5 m m ,
intrastaminal nectary 0, the tassel white rubescent.
Pods 1-3 per capitulum, in profile 6.5-10 x 0.7-0.9
cm, the sutural ribs and the plane recessed valves
densely minutely granular and pallid-puberulent;
150 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
seeds (few seen) ±8 x 4.5 m m , the lustrous brown
testa mottled, pleurogrammic.
In rocky campo, 420-1200 m, locally plentiful but
known only from the n.-e. slopes and foothills of Sa.
do Sincora near Lencois and Andaraf, within lat.
12°30'-13°S in interior Bahia, Brazil. — FL VI,
IX-XL
Calliandra lintea resembles C. bahiana in habit and
leaf-formula but differs in relatively small and more
numerous flowers, the perianth not red-granular. Cal
liandra nebulosa, disjunct on the southwestern foot
hills of Sa. do Sincora, has lower leaf-formula and
somewhat larger leaflets. For measurements see the
Latin diangosis. The epithet linteus (Lat., of sheets)
alludes to the town of Lencois, the type-locality.
86. Calliandra nebulosa Bameby, sp. nov, habitu
toto, inflorescentiae forma, necnon perianthii fere
glabri (nulla parte rubro-granulosi) calyce corollae
fere 3-4-plo breviori C. linteae Bameby proxima,
ab hac autem notulis sequentibus, haud gravibus sed
adhuc consistentibus diversa: foliorum formula
iv-vi/24-32(-37) nee ix-xiii/36-50; foliola majora
5-7.6(-8) x 1.2-1.7(-2.4) nee 3-5.5 x 0.7-1.2 m m ;
capitula 3-6(-7)- nee (5-)7-12-flora; corolla
6.2-7.5 nee 5-5.5 m m longa; androecium 104— 112-
nec 42-72-merum. — BRAZIL. Bahia: 2.5-5 k m s.
of Vila do Rio de Contas on side road to w. of road
to Livramento., ±13°36'S, 41°50'W, 28 Mar 1977,
R. M. Harley & al 20092. — Holotypus, CEPEC;
isotypi, K, NY.
C. squarrosa var. crassifolia Bentham in Martius, Fl. bras. 15(2): 419. 1876. — "[Bahia.] . habitat in planitie alta
Chapada das Minas Novas inter Gamelleira et Caitete:
Martius'' — Holotypus, Martius Obs. 1882, with com
ment: "Planta gossypiumfovens," M!.
C. squarrosa sensu Bentham, 1876: 418, ex parte (Martius
s.n.), typo excluso.
Microphyllidious, sometimes arborescent shrubs
1.5—3(^4) m, branched from base, with virgate, simple
or distally few-forked long-shoots and no axillary
brachyblasts, the young stems and lf-axes pilosulous
with shining white hairs to 0.25-0.75 m m and at least
the dorsal face of lf-stks and the peduncles thinly
red-granular (but perianth not so), the thick-textured,
contiguous or imbricate lfts bicolored, lustrous oliva
ceous above, paler and at least thinly granular be
neath, minutely ciholate, the capitula of few obese,
nearly glabrous fls borne singly and geminate in a
few distal lf-axils, nestled in foliage; phyllotaxy dis
tichous. Stipules ovate or lance-subulate 1-3 m m ,
thin-texmred, not striate, caducous. Lf-formula iv-vi/
21-32(-37); lf-stk of longer lvs 2-4.2 cm, the dilated
petiole 1.5-4.5 x 1.4-2 m m , the longer interpinnal
segments 5-9.5 m m , the ventral groove bridged at
insertion of pinnae; pinnae subequilong or distally a
little shorter, the rachis of longer ones ±3-5(-5.4) cm,
the longer interfoholar segments (0.9-) 1-2.2 m m ; lft-
pulvinules 0.15-0.25 x 0.45-0.7 m m ; lft-blades lin
ear or narrowly linear-oblong from obtusely deltate-
auriculate base, straight or incipiently sigmoid,
obtuse, the longer ones 5-7.6(-8) x 1.2-1.7(-2.4)
m m , (3.1-)3.7-5.4 times as long as wide; venation
prominulous only dorsally, the midrib displaced to
divide blade l:2.5-3.5(-7), simple or distally l(-2)-
branched on posterior side, a weak inner posterior
primary nerve sometimes produced beyond mid-
blade. Peduncles 2-5.5 cm, ebracteate, dilated at apex
into a depressed-convex receptacle scarcely 1 x
2.5-3.5 m m , the capitula 3-6(-7)-fld; floral bracts
ovate, <1 m m , caducous from below young fl-buds;
pedicels scarcely or not differentiated externally,
in section 0.4—1 x 0.7-3 m m ; perianth 4-merous,
obese, nearly glabrous, the calyx-rim and tip of co
rolla sometimes thinly microciliolate, the greenish-
ochroleucous corolla sometimes faintly resin-spotted;
calyx shallowly turbinate-campanulate 1.2-2.1 x 2-
4.5 m m , the low-deltate teeth 0.25-0.4 m m ; corolla
6.2-7.5 m m , the dorsally convex lobes 1.8-4 m m , in
section 0.55-0.8 m m thick; androecium 104-112-
merous, 2.5-3.3 cm, the homy stemonozone 1.1-2.5
m m , the tube 6-7 m m , the tassel white rubescent; no
intrastaminal disc; ovary at anthesis glabrous. Pods
(not seen fully ripe) ±7 c m x 7(-?) m m , the margin
and the recessed valves alike densely brown-granular
and sparsely minutely gray-pilosulous overall; seeds
unknown.
About outcrops in cerrado near 950-1100 m,
localized (but locally plentiful) on and w. of the crest
of Chapada Diamantina in lat. 13°30'-14°S (Vila do
Rio de Contas, Livramento do Brumado, Ituacu, Iga-
porao, Caitete), interior Bahia. — Fl. II-VI, VIII.
Calliandra nebulosa resembles C. lineata in habit,
in almost glabrous, nowhere granular perianth, and in
calyx three to four times shorter than the corolla, but
differs from it in lower leaf-formula, somewhat am
pler leaflets, fewer flowers in the capitulum, longer
corolla, and more numerous stamens; for measure
ments see the Latin diagnosis.
This species was first collected by Martius be
tween Caitete and Rio das Contas (M! = F Neg.
6167), but the specimen was strangely misidentified
by Bentham as C. squarrosa, with which it has little
in common. A second specimen from Martius (M = K
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 151
Neg. 19445) is mislabeled "Vao do Parana, Minas
Gerais."
The epithet nebulosa is translated from Portuguese
brumado and alludes to the habitat on Rio B m m a d o .
87. Calliandra bahiana Renvoize, Kew Bull. 36: 69.
1981. — Typus sub var. bahiana infra designata.
Microphyllidious shrubs (1-) 1.6-3 m with erect
and incurved, defoliate annotinous and older branches,
densely leafy distally, the young stems and lf-axes
pilosulous with whitish hairs to 0.2-0.9 m m inter
mixed with reddish-brown granular trichomes, the pe
duncles and perianth of fls either densely exclusively
red-granular or both red-granular and gray-pilosulous,
the lvs bicolored, the firm plane lfts lustrous dark-
olivaceous and either glabrous or microscopically
pubemlent above, paler dull and both pubemlent and
red-brown-granular beneath, the stout peduncles
borne, either 2-3 together or solitary, in the axils of
several distal primary lvs and sometimes, beyond
these, shortly pseudoracemose; no axillary brachy
blasts; phyllotaxy distichous. Stipules ovate or bluntly
deltate or triangular-lanceolate 0.8^1 m m , not exter
nally venulose, caducous. Lf-formula xi(-xiii)/20-33;
lf-stk of longer lvs 3-6.5 cm, the petiole (consisting
largely of pulvinus) 2-5 m m , at middle 1.5-2 m m
diam, the longer interpinnal segments 4-8 m m , the
ventral groove bridged at insertion of pinnae; pinnae
either subequal or randomly shorter distally, the rachis
of longer ones 3-5.5 cm, the longer interfoholar seg
ments 0.9-1.9 m m ; lft-pulvinules 0.15-0.3 x 0.35-0.7
m m ; lfts equilong or almost so, contiguous or imbri
cate, the blades narrowly oblong or linear-oblong from
shortly obtusely auriculate base, obtuse, the longer
ones 4.5-8 x 1.2-2.5 m m , (2.5-)3-4.1 times as long as
wide; venation palmate, the straight midrib forwardly
displaced to divide blade 1:2.5-3, weakly 1-2-
branched beyond middle on posterior side, the inner
posterior primary nerve incurved to or beyond mid-
blade, the 1-2 outer ones much shorter, all bluntly pro
minulous dorsally, faintly so or immersed above. Pe
duncles stout (1.5-)2.5-6 cm, ebracteate; capitula 3-5
(-6)-fld, the low-convex or subtmncate receptacle
wider than long; floral bracts obtusely deltate or broad-
lanceolate 2-4(-6) m m , ventrally concave, deciduous;
perianth obese, variably granular or granular and pilo
sulous (see key to varieties), mostly 4-merous but the
corolla rarely 5-merous; pedicels often poorly differ
entiated externally but in longitudinal section 0.6-1 x
2.5-3.7 m m ; calyx shallowly campanulate 3-8 x
5.5-7.5(-9) m m , the teeth mostly depressed-deltate
obtuse 0.6-1.2 m m , rarely triangular 2 m m ; corolla
9—13(—14) m m , the broadly ovate lobes 4.5-5.5(-7)
m m , in section 0.8-1.3 m m thick; androecium
(2.8-)3.2-4.4 m m , (98-)114-204(-218)-merous, the
homy stemonozone 1.4—3.5 m m , the tube 7-11 m m ,
the tassel white mbescent; intrastaminal disc 0; ovary
at anthesis either glabrous or distally pubemlent. Pods
6-9 x 1.1-1.2 cm, densely softly sordid-velvety-pilo-
sulous overall; seeds ± 9 x 4 m m , the testa brown, mot
tled, pleurogrammic.
Calliandra bahiana is notable in the context of ser.
Calliandra for relatively high leaf-formula, capitula of
few, very large multistaminate flowers, and thick-
textured perianth abundantly red-granular overall. T w o
geographic variants are here distinguished.
Key to the varieties of C. bahiana
1. Perianth white-pilose as well as red-granular, the white trichomes predominant, ± masking
the red; Sas. Curral Feio and do Tombador
(lat. 10o20'-Ho30'S); one record from Catoles near 13°18'S, 41°53,W. 87a. var. bahiana
1. Perianth densely and almost exclusively red-granular, white trichomes either 0 or very
few and short, the red predominant; vicinity
of Vila do Rio de Contas (lat. 13°32-36'S). .... 87b. var. erythematosa
87a. Calliandra bahiana Renvoize var. bahiana. C.
bahiana Renvoize, 1981, I.e., fig. 1(9), 5B, sens. str.
— "Bahia... Serra do Curral Feio ... Harley & al.
16985." — Holotypus, collected 15 k m n.-w. of
Lagoinha, 8 Mar 1974, CEPEC!; isotypi, K!, NY!.
C. bahiana sensu Lewis in Stannard, 1995: 378.
As described for the species, except as modified by
key to varieties.
About sandstone outcrops in cerrado or cerrado—
campo mpestre transition, 950-1200 m, locahzed on
and near the crest of Chapada Diamantina between
10°20'S and 11°30'S (Sas. Curral Feio and do Tomba
dor) and disjunct at 13°18'S (Catoles), in n.-centr.
Bahia. — Fl. XII-IIL
87b. Calliandra bahiana Renvoize var. erythe
matosa Bameby, var. nov., a var. bahiana vix nisi
perianthio toto rubido-granuloso, haud vel parcis-
sime brevissime griseo-puberulo (nee ubique pilo-
sulo) necnon dispersione allopatrica diversa. —
BAHIA: 5 k m s. of Rio de Contas on road to Livra
mento do B m m a d o , 16 April 1991 (fl, fr), G. P.
Lewis (with S. M. M. de Andrade) 1986. — Holo
typus, CEPEC; isotypi, K, M O , NY.
152 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
In disturbed cerrado, campos gerais, and campo ru-
pestre with arenitic outcrops, sometimes on river
banks, 975-1600 m, locally common but known only
from mun. Rio de Contas in interior Bahia, Brazil. —
Fl. III-VII.
88. Calliandra lanata Bentham, Trans. Linn. Soc.
London 30: 553. 1875; & in Martius, Fl. bras. 15
(2): 422. 1876. — "Habitat in campis inter fluvium
Fermoso [far. n. Minas Gerais, near the Goias line]
et Rio de Contas [near 13°40'S, 41°45,W in centr.
Bahia], Minas Geraes: Martius." — Holotypus,
Martius Obs. 1897, collected probably at Rio de
Contas, hence not in Minas Gerais, M! = F Neg.
67(55 — Feuilleea lanata O. Kuntze, Revis. Gen. PL
1:188.1891. FiG. 22
C. sericea Renvoize, Kew Bull. 36: 77, fig. 3(27) 6C. 1981.
— "BAHIA ... Rio de Contas to Pico das Almas, [R. M.]
Harley et al. 19822r — Holotypus, CEPEC!; isotypi, K!,
NY!.
C. lanata sensu Bentham, 1875. 553 (loco erroneo 'Minas
Gerais'); 1876: 422; Stannard, 1995: 380, fig. 23E-G.
C. sericea sensu Lewis, 1987: 176, fig. 11C,D, pi. 10E,F.
Stout erect, amply multifoholate shrubs with few
simple, distally foliate stems 1.5-3.5 m, notable
for dilated stipules and for efoliate, densely white-
pilose-tomentose, compactly pseudoracemose inflo
rescence, the firm plane subconcolorous lfts densely
finely cihate and thinly pilose facially when young,
glabrescent and moderately lustrous in age. Stipules
ovate-suborbicular from shallowly cordate or broadly
rounded base, 13-30 x 10-25 m m , either obtuse or
abmptly deltate-apiculate, the chartaceous blade dor
sally pilose but glabrescent, especially toward the
margin, weakly nerved, internally glabrous and fla-
bellately multistriate, tardily deciduous. Lf-formula
v-viii/34-50; lf-stks of fully expanded lvs (4-)6-l 1
cm, the stout petiole 12-20 m m , at middle 2-4 m m
diam, the longer interpinnal segments 6-13 m m ;
pinnae subequilong, the rachis of longer ones 8-13
cm, the longer interfoholar segments 1.8-2.6 m m ; lf-
pulvinules 0.45-0.7 x 0.7-1 m m , cross-wrinkled; lfts
proximally decrescent, thence subequilong, the blade
linear-oblong or linear-lanceolate from postically
obtusangulate base, obtuse, the larger ones 10-14.5 x
2.5-3.5 m m , ±4 times as long as wide; venation
externally indistinct, the straight simple or faintly
branched midrib a little eccentric, the inner posterior
primary nerve expiring without anastomosis ± at mid-
blade, the 1-2 outer primary nerves progressively
shorter. Peduncles 2-4 per node of inflorescence, the
longer ones 2-5 c m x 3.5-4.5 m m , none bracteolate;
capitula 3-6-fld, the receptacle hemispherical, wider
than long; bracts ovate 4-5.5 m m , caducous; fls
homomorphic, sessile or almost so, the pedicel (seen
in vertical section) ±0.5 x 2 m m ; perianth densely
white-pilose-tomentose externally, glabrous within,
of subcoriaceous texture; calyx (without vesture)
broadly turbinate-campanulate 7.5-10 m m , the un
equal sinuses between teeth 1.4—3 m m deep; corolla
(also without vesture) 9-14 m m , the ovate lobes 1.5-
5.5 m m ; androecium 136-156-merous (3 counts),
31-36 m m , the greatly thickened, almost homy
stemonozone 2.7-3.4 m m , the tube 7-10 m m , the tas
sel white, distally lilac in age; no independently
developed nectary around base of ovary, but the ste
monozone nectariferous internally in elevated vertical
lines; ovary pilosulous, at least above middle. Pods
erect, ±4.5-14 x 1-1.3 cm, densely pilose-tomentose
overall, the valves recessed below massive sutural
keels; seeds not seen.
In campo mpestre, ±1000-1400 m. known only
from upper Rio de Contas at points within 10-12 k m
to the n. and n.-w of the town of Rio de Contas and in
adj. mun. Abaira, at foot of Pico de Almas, near 13°
15'S on the slope of Chapada Diamantina in centr.
Bahia.- Fl. X-III (VI).
Calliandra lanata is instantly recognized by its
ample, internally flabellate-striate stipules, dense
white indumentum of the inflorescence, and large
flowers. By mistake Bentham attributed the species to
Minas Gerais, and for this reason the name was not
used in Renvoize's revision of the Bahian species
(1981), where the plant was redescribed as C. sericea.
89. Calliandra fuscipila Harms, Repert. Sp. Nov.
Regni Veg. 17: 91. 1921. — "Brasilien: Bahia (Ph.
v. Liitzelburg in Herb. Munchen, no. 245, 1914.)."
— Holotypus, labeled "Minas de Contas, 1000 m,
Aug. 1913," M ! = K Neg. 19441 = IPA Neg. 1327;
isotypus, *B = F Neg. 1242\.
C. fuscipila sensu Renvoize, 1981: 69, fig. 1(8); Lewis, 1987: 173; Stannard, 1995: 379.
Either erect or diffuse, microphyll shrubs 0.5-2 m,
with fuscous defohate glabrate older stems, the closely
leafy homotinous branches, all lf-axes, and all units of
inflorescence densely beset with red-brown, granular
or coralloid trichomes mixed with scattered minute
white hairs to 0.2-0.3 m m , the lvs subconcolorous, the
small crowded, thick-textured, minutely ciholate lfts
lustrous but either glabrous or microscopically papil-
late-puberulent on upper face, both reddish-granular
and inconspicuously pilosulous dorsally, the few-fld
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 153
FiG. 22. Calliandra lanata Bentham
154 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
capitula shortly stoutly pedunculate, commonly 2-3
together, in (l-)2-5 furthest lf-axils, nestled in foliage.
Stipules obtusely deltate ±1 m m , caducous, not venu
lose externally. Lf-formula vii-x/15-21 ["25," proto-
logue]; lvs subsessile, the lf-stk of longer ones 2-4.2
cm, the petiole, consisting largely of pulvinus, 1.5-2
m m , the longer interpinnal segments 2.5-5 m m , the
ventral groove bridged at insertion of pinnae; pinnae
abmptly shorter at apex of lf-stk, otherwise subequi
long, the rachis of longer ones 14—22 m m , the longer
interfoholar segments 0.6-1 m m ; lft-pulvinules
0.1-0.2 m m , a little wider than long; lfts abmptly
decrescent at each end of rachis, otherwise subequi
form, the blades narrowly oblong or oblong-elliptic
from obtusely short-auriculate base, rounded at apex,
the longer ones 2.4-3.4 x 0.9-1.3 m m , (2-)2.4-
3.2 times as long as wide; venation of 2(-3) primary
nerves from pulvinule bluntly prominulous on dorsal
face, the straight simple midrib forwardly displaced to
divide blade ±1:3, the weak posterior nerve expiring
near or shortly beyond mid-blade. Peduncles 3-10
x 1.5-1.9 m m , ebracteate, the receptacle depressed-
hemispherical scarcely 2 m m diam or subtmncate;
bracts deltate ±1 m m , caducous; fls homomorphic,
essentially sessile, the discoid pedicel not differenti
ated externally, ±0.3 m m in section; perianth densely
reddish-granular overall, the corolla firm, thick-
textured, both calyx and corolla evenulose externally;
calyx shallowly campanulate 1.8-2.1 m m , the deltate
teeth 0.4-0.7 m m ; corolla 5.2-6 m m , the ovate, ven
trally concave lobes 1-3.2 m m ; androecium 50-56-
merous, ±3 cm, the stemonozone 0.6-1 m m , the tube
3.3-4 m m , the tassel bright red; ovary at anthesis
glabrous. Pods (Ganev 472, H U E F S ) erect, in profile
5.5-6 x 1 cm, the stout sutural ribs ±4 m m wide in dor
sal view, the woody valves not externally venulose, the
whole fmit densely fuscous-granular.
About rock outcrops in campo mpestre, ±1350-
1650 m, apparently localized on the e. and n.-e.
slopes of Pico das Almas within a radius of ±25 k m
of Vila do Rio das Contas, interior Bahia, Brazil. —
FL II-III, VII, the full cycle not estabhshed.
Calliandra fuscipila is notable for many tiny
crowded leaflets, red-granular indumentum of pedun
cles and capitula, a hemispherical calyx, and red fila
ment-tassel. It somewhat resembles C. mucugeana
but is taller and has more numerous and smaller
crowded leaflets.
90. Calliandra feioana Renvoize, Kew Bull. 36: 75
["feioanum"], fig. 2(24), 6B. 1981. — "Brazil,
Bahia . . . Serra do Curral Feio, Lagoinha to Minas
do Mimoso, Harley et al. 16972." — Holotypus,
CEPEC!; isotypi, K!, NY!.
C. feioanum sensu Lewis, 1987: 173.
Erect microphyll shmbs ±1 m with simple or few-
branched stems, lacking brachyblasts, the new stems,
all lf-axes, and peduncles pilose with fine straight sub-
vertically erect, palhd hairs to 1-1.5 m m mixed (espe
cially the peduncles) with small reddish trichomes, the
lvs moderately discolorous, the plane, facially gla
brous lfts a little darker and more lustrous above than
beneath, ciliate, the peduncles arising solitary or 2(3)
together in distal lf-axils, as long as or scarcely longer
than the subtending If; phyllotaxy distichous. Stipules
herbaceous lance-ovate 3.5-5 x 1-1.5 m m , ±5-nerved,
pilose dorsally, deciduous. Lf-formula iv—viii/16—21
(-25); lf-stks 1.5-2.3 cm, the petiole of lower ones
attaining 4—7 x 0.7-1 m m , that of further ones almost
0, the longer interpinnal segments 3-7 m m , the ventral
groove bridged; pinnae subequilong (randomly
shorter), the rachis of longer ones 1.6-2.7 cm, the
longer interfoholar segments 0.7-1 m m ; lft-pulvinules
0.15-0.2 x 0.3-0.4 m m ; lft-blades narrowly oblong-
elhptic from bluntly auriculate base, deltate or obtuse
at apex, the longer ones 3.6-4.5 x 1-1.5 m m , 3-3.8
times as long as wide; venation obtusely prominulous
on both faces, the straight midrib forwardly displaced
to divide blade 1:2-3, shortly 2-branched beyond mid
dle, the inner posterior primary nerve incurved-
ascending a little beyond mid-blade, the 1-2 outer
posterior nerves much shorter. Peduncles ±2-4 cm,
ebracteate; capitula 4-8(-9)-fld, the clavate receptacle
1.5-2(-2.5) m m ; floral bracts linear-lanceolate ±3.5-
4.5 x 0.3-0.5 m m , early dry caducous; fls homo
morphic, 5-merous, the calyx (dry) ±15-nerved, the
corolla more faintly so, the calyx-tube glabrous but the
teeth thinly pilosulous near apex, the corolla remotely
granular-papillate, its lobes thinly pubemlent or thinly
pilose near apex; pedicels broadly turbinate ±0.8 x 1.2
m m ; calyx ±2-2.5 m m , the deltate teeth 0.5 m m ;
corolla 7 m m , the moderately carnosulous lobes ±2
m m , at apex 0.4 m m thick in section; androecium 52-
merous, ±31 m m , the stemonozone 1.8 m m , the tube
6.2 m m , the tassel white rubescent; ovary at anthesis
thinly pilosulous around top. Pod unknown.
About sandstone outcrops in campo cerrado, 950-
1000 m, known only from the type collection, from
near 10°22'S, 41°20'W, on Sa. Curral Feio in interior
Bahia. — Fl. II-III.
The affinities of C. feioana are not manifest in the
one known collection, which in technical characters
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 155
is suggestive of either C. hirsuticaulis or C. fuscipila.
It differs from both of these in prevailingly longer
petioles and peduncles and in androecial tassel white
rubescent, not red at full anthesis; and further from C.
hirsuticaulis in short (±0.5, not 1.2-3.6 m m ) calyx-
teeth and from C. fuscipila in longer (3.5-4.5, not ±1
m m ) floral bracts.
91. Calliandra asplenioides (Nees) Renvoize, Kew
Bull. 38: 79, fig. 4(33). 1981. Acacia asplenioides
Nees, Flora 4: 303. 1821. — "Urn Valos im Campo
Geral [leg. Pr. Maximilian]." — Holotypus, BR!. —
Bokermann (1957: 236) identified Quartel del Valo
with contemporary Valo Fundo, situated at ±15°
10'S, 41°40'W on the Bahia-Minas boundary, vis
ited by Maximilian in February 1817. — Inga
mertensioides Nees & Martius, [Beitr. fl. bras, in]
Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol.
Nat. Cur. 12: 35, t. 5. 1824, nom. substit. illegit.
Calliandra asplenioides Bentham, London J. Bot.
1: 527. 18429 comb, provis. C. mertensioides Ben
tham, London J. Bot. 3: 106. 1844, comb, illegit.
Feuilleea asplenioides O. Kuntze, Revis. Gen. PL 1:
184. 1891.
C. dendroides Renvoize, Kew Bull. 36: 71, fig. 1(11), 5(c, d). 1981. — "Brazil, Bahia, Harley & al. 15568 . . . Serra do Sincora, Barra da Estiva." — Holotypus, CEPEC!; isotypus, K!. C. mertensioides sensu Bentham, 1875: 550; 1876: 419, var. exclus.; Glaziou, 1905: 188 (n. 14655, P!).
C. asplenioides sensu Lewis, 1987: 171.
Shrubs resembling C. fasciculata in habit, stature,
and inflorescence, fertile when 0.5-1.5, exceptionally
2-3 m, or even arborescent to 5 m, the young stems
and lf-axes varying from pilose with patent fine
straight hairs ±0.4—1 (-1.4) m m to microscopically
pubemlent or minutely granular-resinous, the firm,
plane or ventrally low-convex lfts bicolored lustrous
dark above, lighter brown and often densely resinous-
punctate beneath, facially glabrous but sometimes
remotely ciholate; phyllotaxy distichous. Stipules
ovate-triangular, lanceolate or linear, mostly obtuse,
0.8̂ 4- m m , weakly 1-5-nerved dorsally, deciduous.
Lf-formula ii-iv(-vi)/(12-)14-25(-30); lf-stks 1-6
(-7) cm, the petiole including fuscous pulvinus 1.5-
7(-12, seldom >4) m m , the one or the longest inter
pinnal segment 5-15 mm, the ventral groove bridged
at insertion of pinnae; pinnae subequilong or a little
shorter distally, the rachis of longer ones
(3.5-)4-7(-8) cm, the longer interfoholar segments
1.8-4 m m ; lft-pulvinules 0.1-0.45 mm, at least twice
broader than long; lfts usually ± decrescent at each
end of rachis, the blades sessile against rachis, in out
line narrowly oblong from shallowly cordate or
obliquely subtmncate base, broadly obtuse or obtuse
and obscurely depressed-deltate at apex, the larger
ones 6-12(-14) x (2-)2.3-4.5 mm, 2.4 x 3.7(-4)
times as long as wide; venation palmate, the nearly
straight midrib forwardly displaced to divide blade
1:(1.9—)2.2—3.6, weakly 1-3-branched near or above
mid-blade, the inner posterior primary nerve incurved-
ascending beyond mid-blade, the outer posterior ones
1-3, short and weak, the whole venation either weakly
prominulous or immersed above (but epidermis often
wrinkled when dry), sharply prominulous beneath.
Peduncles 1—5(—6) per node, (1-) 1.5-6 cm, ebract
eate; capitula densely 4—12(-?)-fld, the clavate or
depressed-hemispherical receptacle 2-4 m m diam;
bracts of lower fls deltate-ovate or subulate 0.4—1.2
mm, caducous, of inner fls often 0; fls homomorphic,
either sessile or contracted at base into a turbinate or
dmm-shaped, discolored pedicel 0.4—0.8 x 0.7-2.1
mm; perianth glabrous or thinly pubemlent distally,
faintly or imperceptibly nerved; calyx (disregarding
pedicel) 1.3-3.3 x l-2.5(-4) mm, the depressed-
deltate or shortly subulate teeth 0.3-1 mm; corolla
5-8(-9) mm, the ovate lobes 2-3.5 m m (some sinuses
shallower); androecium (20-)24-54-merous, (18-)
22-34 mm, the stemonozone 1.4—2.2 mm, internally
thickened but not disciferous, the tube 5-8 mm, the
tassel either white, or white and distally pink, or pink
to red overall; ovary at anthesis glabrous. Pods 1-3
per capitulum, erect, linear-oblanceolate 4.5-8 x 0.7-
1 cm, hgnescent, resinously granular overall and
sometimes in addition finely pilosulous, the sutural
ribs in dorsal view ±2.5-3 m m wide, the plane re
cessed valves evenulose or almost so; seeds (few
seen) oblong-elliptic in broad view, ±6.5-9 x 4.5 m m ,
the smooth testa light brown, the narrowly U-shaped
pleurogram 4.5-7 x 2.5-3 mm.
In campo mpestre, in pockets of quartzite outcrops,
and on rocky stream banks and cliff-ledges, 700-
300 m, scattered along and near the crest of Chapada
Diamantina and Sa. do Espinhaco from Sa. Agua da
Rega and Rui Barbosa near 12°20'S in interior Bahia
s. to Sa. da Caraca near 20°05/S in centr. Minas
Gerais. — Fl. XII-IV, the fmit long persistent.
Among Bahian calhandras of ser. Calliandra with
deltate calyx-teeth C. asplenioides is recognized by
pinnae 3-4 pairs in almost all larger leaves, relatively
broad leaflets, and filaments about 24-54 per flower.
In Minas Gerais the sympatric C. fasciculata, similar
in leaf-formula, has linear or linear-lanceolate, not nar
rowly oblong, leaflets about 4-8, not 2.4-4.5, times as
156 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
long as wide. The resinous spots or glandular tri
chomes that are a feature of C. asplenioides in Minas
are rare or absent in C. fasciculata, which has usually
longer floral bracts 2.5-15 (not 0.6-1.2) m m long.
92. Calliandra viscidula Bentham, London J. Bot. 3:
109. 1844. — "Brazil, Serra Jacobina [Bahia],
Blanchet, n. 2620." — Holotypus, K (hb. Hook.,
commun. Moricand)! = N Y Neg. 7975!; isotypus,
BM!. — Feuilleea viscidula O. Kuntze, Revis. Gen.
PL 1: 189. 1891.
C. viscidula sensu Bentham, 1875: 553, var. exclus.; 1876:
423; Renvoize, 1871: 74, fig. 2(10); Harley & Simmons,
1987: 115; Lewis, 1987: 177.
Robust, densely and relatively coarsely foliate
shrubs 1-3 m, except for glabrous or exceptionally
thinly pubescent faces of lfts softly densely pilose
throughout with lustrous white, erect hairs ±0.8-2
m m , the foliage subconcolorous, the firm lfts moder
ately lustrous above, dull and sometimes obscurely
resin-spotted beneath, ciliate, the stoutly pedunculate,
well-furnished, hemispherical capitula solitary and
geminate either in a few furthest lf-axils, or shortly
pseudoracemose, or both. Stipules erect submembra-
nous, ovate short-acuminate 5-12 x 2.5-7 m m , stri
ately many-nerved from point of attachment, glabrous
or thinly pilose dorsally, deciduous. Lf-formula
iii-v/14-20; lf-stks (1—)1.5—7 cm, the petiole together
with (or consisting largely of) dusky pulvinus 2-10 x
1.5-2.4 m m , the longer interpinnal segments 8-16
m m , the ventral groove bridged at insertion of pinnae;
pinnae either subequilong or a httle graduated in
either direction, the rachis often recurved, that of
longer pinnae 4-8.5(-10) cm, the longer interfoholar
segments 2.5-5.5 m m ; lft-pulvinules 0.4-1 x 0.6-1.2
m m ; lfts a httle decrescent at each end of rachis, oth
erwise subequilong, the blade oblong or narrowly
lance-oblong from broadly obtusely auriculate base,
abmptly obliquely short-apiculate, the longer ones
(10-)11.5-20 x 4.2-6.3 m m , 2.4-3.5 times as long as
wide; primary venation of 4-6 nerves from pulvinule,
the nearly straight midrib forwardly displaced to divide
blade ±1:2-3, 1-3-branched at or beyond mid-blade,
the innermost posterior primary nerve incurved-
ascending well beyond mid-blade, the outer ones pro
gressively shorter, the sinuous reticulum of secondary
venules either prominulous on both faces or almost
immersed beneath. Peduncles stout (1.5-)2-4(-6) cm,
charged near middle and beyond with 1 or to 5 papery,
ovate-acuminate deciduous bracts, the further ones
sometimes forming an involucre under the capitulum;
capitula 8-12-fld, the subsessile fls homomorphic;
bracts papery, narrowly or broadly ovate-acuminate
±5-8.5 x 2.3-6 m m ; pedicels stout, a httle wider than
long, 0.9-2 x 1.4-2.5 m m ; perianth either distally or
overall pilose with divergent lustrous white hairs and
micro-granular, the 4—5-merous calyx obtusely striate-
venulose, the 4(-5)-merous corolla scarcely so; calyx
turbinate-campanulate 5.6-10(-l2) m m , the lance-
ovate teeth 2-5.5 x 0.8-3.7 m m ; corolla 8-14.5 m m ,
1.2-4.5 m m longer than calyx, the lanceolate or ovate
lobes 2.8-5.3 x 1.5-3.3 m m ; androecium 58-92-
merous, commonly 5-7 c m but sometimes scarcely
half as long, the thickened stemonozone 2.2-4 m m ,
the tube 7-12 m m , the tassel opening white, some
times mbescent; intrastaminal nectary 0; ovary at
anthesis glabrous. Pods 1-3 per capitulum, in profile
oblanceolate 5.5-9.5 x 1-1.5 cm, the stout sutural
keels ±2.5 m m wide in dorsal view, the recessed
valves coarsely ascending-venulose, the whole densely
softly pallid-pilose or the valves only thinly so; seeds
in broad view ± 8 x 5 mm., the testa pale brown dark-
speckled, the U-shaped pleurogram ± 6 x 2 m m .
In campo mpestre on sandstone bedrock, 850-
1000 m, known by numerous collections from lower
slopes of Sa. do Sincora near Palmeiras, Lencois, and
Mucuge in the upper Paraguacu basin, and by the
type and five m o d e m (to 1993) collections from
campo-caatinga transition at 520 m upward on the
sources of rio Itapicum near Jacobina, within 11-
13°S in interior Bahia. — FL II-VI, X-XI, perhaps
intermittently through the year.
A m o n g upland Bahian species of sect. Calliandra
the robust C. viscidula may be recognized by a com
bination of characters: ovate striate stipules 5-12 m m
long; pinnae 3-5 pairs per leaf; ample leaflets to
11-20 m m long; peduncular bracts usually more than
one; capitula 8-12-flowered, and lance-ovate calyx-
teeth 2-5.5 m m long. Contrary to the epithet the
plants are usually not viscid, though the leaflets are
sometimes thinly resin-spotted dorsally.
93. Calliandra fasciculata Bentham, J. Bot.
(Hooker) 2: 140. 1840. — Typus infra sub var. fas
ciculata indicatur.
Microphyllidious shrubs (0.4-)l-2(-?) m with fus
cous defoliate annotinous and older stems and
densely foliate new ones, the erect-ascending stems
paniculately few-branched distally, often glabrous
except for pilosulous ventral face of lf-axes and the
inflorescence often ± resinously papillate, but the new
stems sometimes and the peduncles often pilose with
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 157
fine white spreading hairs to 0.8-1.8 m m , the lvs bi
colored, the firm narrow plane crowded lfts glossy
and glabrous (exceptionally thinly strigulose) on upper
face, paler dull and often minutely glandular-papillate
dorsally, rarely ciholate, the stout peduncles mostly
fasciculate in upper lf-axils or by suppression of dis
tal lvs forming a shortly exserted pseudoraceme of
hemispherical few-fld capitula, the bracts and fls
varying from glabrous to thinly pilose. Stipules
herbaceous, when well developed resembling lfts in
shape and texture, narrowly lance-oblong to linear-
lanceolate and commonly 3-14 x 0.6-2.3 m m , less
often subulate and only 1.5-3 m m , exceptionally
dilated to ovate 6 x 3.5 m m , in any case deciduous or
early caducous. Lf-formula ii—iv(—v)/( 17—) 19—35; lf-
stks (0.9-)1.3-4(-5) cm, the petiole including livid
pulvinus (3—)4—12 m m , the one or the longer interfo
holar segment 6-14 m m , the narrow ventral groove
bridged at insertion of pinnae; rachis of longer pinnae
4—7(-8) cm, the longer interfoholar segments 1.4—2.8
(-3) m m , lft-pulvinules 0.2-0.45 m m , broader than
long, transversely wrinkled, the lft-blades sessile
against pinna-rachis; lfts subequilong except at very
ends of rachis, in outline linear or linear-lanceolate
from deeply auriculate base, obtuse or deltately sub
acute, either straight or incipiently falcate, those near
mid-rachis (6-)7-12.5 x (l-)1.2-2.3 m m , (4.8-)5-8
times as long as wide; midrib simple or faintly 1-2-
branched beyond middle on posterior side, displaced
to divide blade ±1:2-3, the inner posterior primary
nerve ascending, almost parallel to midrib, well be
yond mid-blade, the 1-2 posterior ones very short or
obscure, all venation not or only faintly raised on
ventral face of blade, a little more prominulous be
neath. Peduncles 1-5 per node, (2-)2.5-7 cm, angu-
late-sulcate lengthwise, ebracteate, expanded at apex
into a ± hemispherical receptacle broader than long;
capitula 5-8-fld, the fls homomorphic, appearing ses
sile or nearly so; bracts thinly herbaceous, ovate,
lanceolate, or lance-acuminate (2—)2.5—15(—19) x
(1.2-) 1.8^4 m m , but sometimes reduced to scales <1
m m , deciduous; pedicels either turbinate, to 1 m m , or
distinguished from calyx externally only by discol
oration; perianth either 4- or 5-merous, either
glabrous overall or thinly white-pilose (-pubemlent)
distally; calyx (1.2-)1.5-6.5(-12) m m , the tube
(0.9-) 1.5-3 x 2-3 m m , the teeth varying in length
from 0.4, when depressed-deltate, to 4(-9) m m , then
lanceolate and sometimes of unequal lengths; corolla
6-9 m m , the erect ovate lobes 2.3-4 m m (the sinuses
often unequal), the whole varying from 3 times
longer than calyx to a trifle shorter; androecium
(23-)26-42 m m , 38-56(-68)-merous, the tube ±6-8
m m , the stemonozone 1.4—2.6 m m , thickened inter
nally, the tassel opening white, rubescent; intrasta
minal nectary 0. Pods 1-2 per capitulum, in profile
linear-oblanceolate (4.5—)5—11 x 0.7-0.95 cm, the
stout sutural ribs 2.2^ m m wide in dorsal view, the
stiffly coriaceous or hgnescent, recessed valves either
smooth or weakly ascending-venulose from either su
ture, reddish-brown or fuscous at maturity, either
glabrous overall or thinly pilose with erect hairs,
sometimes in addition microscopically resin-spotted;
seeds 5-8, much subject to predation, not seen whole.
In campo cerrado, especially about outcrops, on
rocky stream banks, and in rock-pockets of campo
mpestre, 820-1350 m, locally plentiful in two some
times sympatric and nearly coextensive varieties
along and near the crest of Sa. do Espinhaco between
latitudes 16°S and 20°S in Minas Gerais (the range of
each variety more precisely indicated below). — Fl.
VIII-V.
Calliandra fasciculata, endemic to upland northern
and central Minas Gerais, closely resembles sympatric
C. asplenioides (=C. mertensioides Bentham) in habit,
and typical var. fasciculata is hardly different in inflo
rescence and individual flower. Bentham (1876: 420)
speculated that when better known in the field the two
might prove to be conspecific. The proportionately
broader, scarcely auriculate and broadly obtuse
leaflets of C. asplenioides that, unlike those of C. fas
ciculata, are, at least in the common range, resinous-
punctate or -papillate dorsally, provide a rehable crite
rion; the only specimens of ambiguous identity that I
have seen are either immature or fragmentary.
The taxonomic status of C. bracteosa, likewise
questioned by Bentham (1876: 419), is more difficult
to interpret. Its variably modified calyx, accompanied
by enlarged floral bracts and often by similarly exag
gerated stipules, are the only features that distinguish
it from typical C. fasciculata, though the perianth usu
ally bears a few, exceptionally many, long straight tri
chomes. The lengths of calyx-teeth and bracts are,
however, poorly correlated and the perianth is some
times almost glabrous. This random inconstancy is
outbalanced by multiple resemblances in foliage and
corolla, and in consequence C. bracteosa is here re
duced to varietal status. The dispersal of the two vari
eties of C. fasciculata is peculiar. The var. fasciculata
is best known from the environs of Diamantina and
Serro. It is represented south of this region by only
one collection each from Sas. do Cipo and Caraca, and
to the northwest by one collection from Sa. do Cabral.
Around Diamantina var. bracteosa is as c o m m o n as
158 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
var. fasciculata, and on Sa. do Cip6 much more fre
quent; northwest from Diamantina it is known only
from Grao Mogol and Montes Claros. On the crest of
Sa. do Cip6 C. fasciculata is sympatric with the en
demic C. linearis, which has essentially the flowers of
var. fasciculata but differs in dwarf stature and rhi
zomatous habit of growth.
Key to the varieties of C. fasciculata
1. Calyx (disregarding cryptic pedicel) 1.2-2.4 x 2-3 m m , the deltate teeth 0.4-0.8(-1.3) mm; floral bracts 0.8-1.8 m m ; perianth glabrous; stipules commonly lanceolate or linear-lanceolate 1.5-4 m m , rarely like those of var. bracteosa linear-lanceolate to 12 m m or (Sa. do Cabral) dilated to ovate 93a. var. fasciculata
1. Calyx (as above) 3-5.5 x 1.8-2.5 m m , the lanceolate or narrowly subulate teeth (1.7-) 2-3.5 m m , distinctly longer than the tube; floral bracts 4—15(—19) m m ; perianth thinly pilose or minutely puberulous distally; stipules lanceolate or linear-lanceolate (3-) 4—13(—19) m m 93b. var. bracteosa
93a. Calliandra fasciculata Bentham var. fascicu
lata. C. fasciculata Bentham, 1840, I.e., sens. str. —
"Brazil, Pohl." — Holotypus, Pohl 1456, K!; iso
typus, Pohl 3480/1456, NY!. — Feuilleea bentha-
miana O. Kuntze, Revis. Gen. PL 1: 184. 1891.
C. fasciculata sensu Bentham, 1844: 108; 1875: 550; 1876: 420. C. mertensioides sensu Glaziou, 1905: 188; non Bentham.
Characters of the species, modified by key to
varieties.
Habitat of the species, characteristically developed
around Diamantina, Mendanha, Gouveia, and Serro in
lat. ±18°-18°40'S, collected once on Sa. do Cipo
(where var. bracteosa is dominant) and once near
Caraca in lat. ±20°10/S; once recorded, in a form with
dilated stipules, from Sa. do Cabral in lat. 17°40'S.
93b. Calliandra fasciculata Bentham var. bracteosa
(Bentham) Bameby, stat. nov. C. bracteosa Ben
tham, London J. Bot. 5: 104. 1846. — "Diamond
district, prov. Minas Geraes, Gardner, n. 4523 and
4524." — Lectotypus, Gardner 4524, K (hb.
Hook.)! = N Y Neg. 1965 ; paratypus, Gardner 4523
from Formigas [= Montes Claros], K (hb. Hook.)!.
— Feuilleea amplebracteata O Kuntze, Revis. Gen.
PL 1: 184. 1891.
C. bracteosa sensu Bentham, 1875: 550; 1876: 419.
Characters of the species, modified by key to
varieties.
Habitat of the species, best known from Diaman
tina and vicinity, extending n. to Montes Claros and
Grao Mogol, and s. through Serro to Sa. do Cipo,
only at its n.-most stations occurring independently
of var. fasciculata, the whole range contained in lat.
16°30'-19°30'S.
94. Calliandra linearis Bentham, London J. Bot. 5:
103. 1846. — "Serra da Lapa, Brasil, Riedel." —
Holotypus not found at K in 1994, presumably at
LE but not verified; isotypus, *B = F Neg. 7247!. —
Feuilleea linearis O. Kuntze, Revis. Gen. PL 1: 188.
1891. FIG. 23
C. linearis sensu Bentham, 1875: 549; 1876: 418.
Dwarf glabrous microphyllidious subshrubs from
creeping rhizomes, the erect and ascending homo
tinous stems 5-15 cm, herbaceous and sometimes
fire-pmned, together forming mats or diminutive
thickets several dm diam, the firm plane lfts scarcely
bicolored but lustrous above and dull beneath, the rel
atively large capitula solitary (geminate in furthest lf-
axils and one terminal), or by suppression of distal lvs
shortly pseudoracemose. Stipules herbaceous, linear-
subulate 1-3.5 mm, 1-nerved, persistent or tardily
deciduous. Lf-formula i—iii(—iv)/l 3—18, the pinnae of
some further lvs always 2 pairs or more; lf-stks 1-3
cm, the petiole 4—15 x 0.5-0.9 mm, the ventral groove
(in 2-jugate lvs) bridged between pinnae, the one or
longest interpinnal segment 5-12 m m ; rachis of
longer pinnae ±2.5-6 cm, the longer interfoholar seg
ments 1.5-2.8 mm; lfts subequilong except at very
ends of rachis, subsessile, the pulvinules <0.2 mm,
the blades linear from shortly obtusely auriculate
base, straight or obscurely falcate, the longer ones
6-10 x 0.85-2 mm, (4.5-)5-10.4 times as long as
wide; venation simple, the almost centric, unbranched
midrib prominulous only beneath, in broader lfts
accompanied by a weak posterior primary nerve pro
duced to or beyond mid-blade. Peduncles stout
(1.5-)2-6 cm, sulcate lengthwise, usually ebracteo-
late (bracteole when present 2-2.5 m m ) ; capitula 5-
11-fld, the fls sessile homomorphic, the receptacle ±3
m m diam; floral bracts lanceolate 1-2.5 m m , decidu
ous; perianth glabrous, the calyx externally nerveless,
the corolla faintly openly veined, the androecium at
first white, pink in age; calyx including the solid,
broadly turbinate base 2.4-4 mm, the teeth 0.6-1.2
mm; corolla 8-11 mm, the lobes mostly 1.4-4 m m
(some randomly fused); androecium 52-92-merous,
(19-)21-34 mm, the tube 4.5-7 m m , the stemono
zone 1-2.2 mm, a little thickened internally at base
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 159
FiG. 23. Calliandra linearis Bentham.
but lacking a defined nectarial disc. Pods 1-2 per ca
pitulum, erect, in profile 5.5-10.5 x ±1 cm, the red-
brown or dull brown hgnescent valves glabrous, the
stout sutural ribs smoothly rounded, the recessed
valves sinuously venulose; seeds (few seen) in broad
view obovate 7-8 x 5-5.5 mm, the testa light brown
smooth, the elliptic incomplete pleurogram 4.5-5 mm.
In white sand of campo mpestre and about quartzite
outcrops, 1080-1300 m, locally plentiful but confined
to the crest of Sa. do Espinhaco (Sa. da Lapa; Sa. do
Cipo) in centr. Minas Gerais, Brazil. — Fl. XI-II.
In montane central Minas Gerais, where section
Calliandra is represented by few species, C. linearis
is instantly recognized by dwarf stature, rhizomatous
caudex, and two or three, rarely four, pairs of pinnae
per leaf Except for its distinctive habit it closely re
sembles some forms of sympatric C. fasciculata, of
which it may be an ecologically induced derivative.
95. Calliandra elegans Renvoize, Kew Bull. 36: 77,
fig. 3(31), 7B. 1981. — "Brazil, Bahia . . . Serra do
Rio de Contas, Mato Crosso, Harley et al. 19928."
— Holotypus, CEPEC!; isotypi, K!, NY!.
C. elegans sensu Lewis, 1987: 172.
Coarsely multifoliolate shrubs to 2-3.5 m tall with
stout virgate stems distally few-branched and efoli
ate, the young stems and the If- and inflorescence-
axes densely but shortly pilosulous-tomentulose with
grayish-brownish hairs <0.6 m m (very few short, col
ored trichomes on calyx or corolla), the firm bicol
ored plane lfts facially glabrous ciholate, on upper
face lustrous dark olivaceous-brown, paler duller and
faintly resin-spotted beneath, the few-fld capitula
arising (singly) 2^4 together from efoliate nodes
of small, shortly exserted pseudoraceme; phyllotaxy
distichous. Stipules (caducous) thinly herbaceous,
broadly ovate-flabellate ±3-5 x 2.5-4 m m , not exter
nally venulose. Lf-formula iv-v/30-35; lf-stks 4—5
cm, the petiole 10-17 mm, at middle 1.6-2 m m diam,
the longer interpinnal segments 6-17 m m , the ventral
groove bridged; pinnae of random lengths, the longer
ones ±10-14.5 cm, the longer interfoholar segments
2.5-3.5 mm; lft-pulvinules 0.6-1.1 x 0.7-0.9 m m ;
lfts decrescent at each end of rachis, otherwise sub
equilong, the blades narrowly oblong from shallowly
semicordate base, obtuse, nearly straight, the longer
ones 10-14 x 3-4 mm, 3-3.7 times as long as wide;
midrib displaced to divide blade ±1:2, weakly 2-
branched on posterior side above mid-blade, the inner
posterior primary nerve incurved-ascending well be
yond mid-blade, the 2(-3) outer posterior ones much
160 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
shorter, the venation bluntly prominulous on both
faces. Peduncles 12—18(—20) mm, ebracteate; capit
ula 4-6-fld, the hemispherical receptacle 1-1.5 x 2-
2.5 m m ; floral bracts minute ephemeral; cryptope-
dicels broadly turbinate 0.8 x 1.6 mm; perianth 4-
merous, greenish red-tinged, thinly white-pilosulous
around top of each cycle, faintly resin-spotted, not
papillate; calyx shallowly campanulate 2 x 2 mm, the
depressed-deltate teeth 0.5-0.8 mm; corolla 6.5-7
m m , the unequally cleft lobes 2.1-3.5 mm; androe
cium 42-54-merous, 2.3-3 cm, the stemonozone
1.2-1.5 mm, the tube 4.5-6 mm, the tassel white;
ovary at anthesis glabrous. Pod unknown.
In rocky campo, 1050-1200 m, known only from
the headwaters of rio de Contas at 13°18,-29,S,
41°47'-49/W, in municipios Rio de Contas and
Abaira, in interior Bahia. — Fl. Ill—IV.
In overall habit and in number of pinna-pairs C.
elegans resembles C. viscidula, but it differs in well-
developed petioles, nearly twice as many leaflets (to
30-35, not 14—20, pairs), and much shorter, not stri
ate calyx.
At the type-locality of C. elegans, between Mato
Grosso and Rio de Contas, Harley collected also the
set of C. lanata that became type of C. sericea Ren
voize, and a unicate (Harley 19928A, CEPEC, in bud),
which has the foliage of C. elegans but dense pallid
cauline indumentum that could be derived from sym
patric C. lanata. Another ambiguous unicate (Harley
19822A, CEPEC, in young flower) taken at a point 12
km north-northwest of Rio de Contas is suggestive of
introgression between C. lanata and C. bahiana, but
has not been certainly identified.
96. Calliandra mucugeana Renvoize, Kew Bull. 36:
83,fig.4(38), 8E. 1981. —"Brazil,Bahia.. .Mucuge
Harley etal. 16095." — Holotypus, CEPEC!; isotypi,
K!, NY!. Fig. 24
C. mucugeana sensu Harley & Simmons, 1986: 115; Lewis, 1987: 175.
Small shrubs forming diminutive thickets 2-5(-6)
m tall, the simple and when older widely branched
stems fuscous defoliate glabrate, the homotinous
branches densely leafy and together with lf-axes mi
nutely pilosulous with erect whitish hairs ±0.1-0.2 m m
and in addition microscopically reddish-granular, the
lvs conspicuously bicolored, dark, highly lustrous and
low-convex above, paler dull and minutely resin-dotted
beneath, the few-fld capitula borne solitary or in pairs,
either all in uppermost lf-axils or some distal ones
shortly pseudoracemose, the glabrous red-stamened fls
nestled in foliage or barely exserted; phyllotaxy erratic,
spiral or subdistichous. Stipules thin-textured, early
shed, linear or narrowly elhptic 1-3 x 0.4—1.2 m m ,
weakly 1-nerved, the nerve simple or rarely branched.
Lf-formula (ii-)iii-iv/13-18(-20); lf-stks (4.5-)7-34
mm, the petiole including discolored obese pulvinus
1.5-4 x 1-1.7 mm, the one or the longest of 2-3 inter
pinnal segments 4—11 mm, the ventral groove bridged
at insertion of pinnae; pinnae subequilong or the fur
thest pair shorter, the rachis of longer ones
(2-)2.5^K-6.5) cm, the longer interfoholar segments
1-3.6 mm; lft-pulvinules 0.1-0.25 x 0.5-1 mm; lfts de
crescent at each end of rachis, otherwise subequilong,
the blades narrowly oblong from obhquely tmncate
base, straight or almost so, broadly obtuse, those near
mid-rachis 5-9 x 1.7-3 mm, 2.5-3.5 times as long as
wide; venation of 3-4(-5) nerves from pulvinule, the
straight midrib forwardly displaced to divide blade
±1:2, usually 2-branched beyond mid-blade, the inner
posterior primary nerve produced to anastomosis well
beyond mid-blade, the outer one(s) much shorter, all
veins bluntly prominulous on lower face, less so or
immersed on upper. Peduncles 7-22 mm, ebracteate;
capitula 5-9-fld, the tmncate or low-convex receptacle
1-1.5 mm; bracts submembranous ovate 0.5-1.1 mm,
deciduous; fls appearing sessile, homomorphic; peri
anth usually glabrous except for microscopically ciho
late calyx-teeth, the corolla exceptionally strigulose
with few random and remote appressed hairs, the calyx
faintly 5-nerved, the corolla externally nerveless;
pedicels discoid 0.3-0.6 x 1.1-1.6 mm, scarcely dif
ferentiated externally from calyx; calyx submembra
nous, shallowly campanulate 1.7-2.4 x 2-2.5 mm, the
low-deltate teeth 0.3-0.5 mm; corolla 5.6-6.1 mm, the
ovate lobes 1.8-3 mm; androecium 30-36-merous, 24—
27 mm, the stemonozone 0.9-1.7 mm, thickened inter
nally, the palhd tube 4.4-5.5 mm, the tassel bright red;
ovary subsessile, glabrous at anthesis, pubemlent after
fertilization. Pods ±4.5-6.5 x 0.7-0.8 cm, densely
velutinous overall with fine erect brownish hairs, 4—5-
seeded, the sutural keels in dorsal view ±3 m m wide,
the recessed valves nearly plane hgnescent, not visibly
venulose; seeds unknown.
In campo mpestre and on rocky river banks,
600-1100 m, locally plentiful on the e. slope of Cha
pada Diamantina in lat. 12°30 -13°S, interior upland
Bahia (Palmeiras, Lencois, Brejao, Andarai, Mu
cuge). — Fl. II, VI, XI, perhaps at intervals through
out the year.
The decorative C. mucugeana is readily recognized,
in context of Bahian ser. Calliandra, by low stature,
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 161
FiG. 24. Calliandra mucugeana Renvoize.
thicket-forming or "carpet-like" habit of growth, short-
toothed calyx, and brilliant carmine tassel.
97. Calliandra calycina Bentham, Trans. Linn. Soc.
London 30: 549. 1875; & in Martius, FL bras.
15(2): 419. 1876. — ". . . in Serra da Jacobina
provinciae Bahiensis: Blanchet, n. 3683." — No
holotypus found at K in 1994; isotypi, BM!, BR! =
K Neg. 19426, G = K Neg. 19435, 6, 7!, G H
(fragm.)!. — Feuilleea calycina O. Kuntze, Revis.
Gen. PL 1: 187. 1891.
C. jacobiana Renvoize, Kew Bull. 36: 82, fig. 4(39), 8A.
1981. — "Bahia. Serra do Jacobina, Estiva, Harley & al
16540." — Holotypus, CEPEC!; isotypi, K!, NY!.
C. angusta Renvoize, Kew Bull. 36: 82, fig. 3(36), 6B, C.
1981. — "Bahia. Serra do Sincora, Mucuge, Rio Cumbuca,
Harley & al. 15937." — Holotypus, CEPEC!; isotypus, K!.
C. robusta Renvoize, Kew Bull. 36: 82, fig. 4(37), 8D. 1981. — "Bahia . . . Serra do Curral Feio, Lagoinha to Minas do
Mimoso, Harley et al. 16971." — Holotypus, CEPEC!; isotypus, K!.
C. calycina sensu Renvoize, 1981: 63, fig. 4(39); Lewis 1987: 171.
C. jacobiana sensu Lewis, 1987: 174. C. angusta sensu Harley & Simmons, 1986: 114; Lewis,
1987: 170. C. robusta sensu Lewis, 1987: 176.
Stiffly erect shrubs 0.6-2(-3) m with fuscous defo
hate annotinous and older branches, the homotinous
ones clad in crowded, narrowly ascending, upwardly
imbricate lvs, the new stems and at least the ventral
face of lf-axes pubemlent or pilosulous with fine, sub-
appressed-ascending or widely spreading white hairs
to 0.2-0.8 m m , the lvs bicolored, the glabrous or
glabrous microscopically ciholate lfts lustrous oliva
ceous above, paler and minutely resin-spotted beneath,
the few-fld capitula solitary or paired either in a few
furthest lf-axils or by reduction of furthest lvs forming
a shortly exserted pseudoraceme. Stipules erect, thin-
textured, linear or hnear-subulate (1.5-)2-6.5 x
0.2-1.6 m m , 1-3-nerved, caducous. Lf-formula i-ii/
15—30(—36), the pinnae of some lvs almost always 2-
jugate; lf-stks, including discolored pulvinus, either
1.5-3 m m , or when pinnae are 2-jugate to 8(-9) m m ,
at middle 0.9-1.3 m m diam; rachis of longer pinnae
2.5-5 cm, the longer interfoholar segments 0.8-1.5
m m ; lft-pulvinules 0.15-0.3 x 0.45-0.65 m m ; lfts
decrescent at each end of rachis, otherwise subequi
long, the blades hnear from auriculate base, deltately
acute-apiculate, straight or incipiently sigmoid, the
longer ones 5-9.5 x 1.1-1.8 m m , 3.9-6.6 times as long
as wide; midrib forwardly displaced to divide blade
1:2-4, weakly few-branched, the inner posterior
162 M E M O I R S OF T H E N E W Y O R K BOTANICAL G A R D E N [VOL. 74(3)
primary nerve produced nearly to blade's apex but
sometimes barely perceptible, the outer posterior one
very short or obscure, a weak random secondary venu
lation sometimes expressed on upper face. Peduncles
stout 12-33 m m , ebracteate; capitula 3-7-fld, the
receptacle ±1-1.5 m m diam; floral bracts minute fuga
cious, absent from some fls; fls appearing sessile,
either 4- or 5-merous, homomorphic, the perianth
commonly glabrous but the calyx-teeth sometimes
ciholate and the corolla sometimes thinly appressed-
pilosulous above middle; pedicels (differentiated exter
nally by darker color) discoid or broadly turbinate
0.4-1.2 x 0.&-2.5(-4) m m ; calyx 2.2-5 x (2.2-)3-5
m m , faintly 5-nerved, the depressed-deltate or semicir
cular teeth 0.5-1.8(-2.2) m m ; corolla 6.5-8(-9) m m ,
the ovate lobes 1.5-4.5 x 2.4-4 m m ; androecium
62-92-merous, 24-42 m m , the stemonozone 1.6-2.2
m m , comeously thickened internally, the tube 7-9 m m ,
the tassel white rubescent; ovary subsessile, at anthesis
pilosulous, especially above middle. Pods erect,
4.5-7.5 x 1-1.3 cm, 5-6-seeded, densely pilose-
tomentulose overall with erect palhd hairs, the coarse
sutural keels 5-6 m m wide in dorsal view, the deeply
recessed valves hgnescent; seeds 6.5-10 x 4—5 m m ,
the smooth testa mottled, the pleurogram ± half as
broad as the seed-face.
In campo mpestre sometimes transitional to campo
cerrado or caatinga, (500-)800-l 100 m, locally plen
tiful along and e. of the crest of Chapada Diamantina,
from Sas. do Curral Feio and da Jacobina s. through
Sa. do Tombador to Lencois and Mucuge in lat.
10°20'-13°S. — F1.XI-II.
Calliandra calycina has nearly the leaf-formula of
C. asplenioides except that the pinnae are not more
than two pairs; they have similarly short petioles and
the individual flowers are similary proportioned. The
leaves of C. calycina are usually more crowded along
the stem and the leaflets are much narrower (1.1-1.8,
not 2-4.5, m m wide) and more crowded along the
pinna-rachis.
I have not found reliable diagnostic features to con
firm the segregate C. jacobiana, C. angusta, and C.
robusta, which are treated herein as minor variants
without taxonomic status. Renvoize (1981) in his key
to Bahian Calliandra distinguished C. jacobina from
C. calycina (sens, str.), both of which were described
from Sa. da Jacobina, by color of the filaments: known
to be white mbescent in C. jacobiana, and supposed to
be red in C. calycina. Neither Blanchet nor Bentham
described the filaments of the latter as red, and
Blanchet's specimens, now more than a century old,
tell nothing certain of flower-color. Calliandra
angusta was separated from C. jacobiana by greater
stature (1.75-2, not 7.5, m ) and smaller flowers
(corolla 6, not 10, m m long); but specimens subse
quently identified by Renvoize himself (Lewis 853,
N Y ) are described in field-notes as 6-10 d m tall, while
the corolla in the type-region of C. angusta varies from
about 6 m m to 8.5 m m in length. The leaflets of the
type-collection of C. calycina are shghtiy more nu
merous than those of C. jacobina, but the plants are
otherwise extremely similar. Calliandra robusta has
calyces proportionately longer as compared to the
corolla, but is otherwise compatible with C. calycina.
A unicate specimen (Harley 16971 A, K!) collected
with the typus of C. robusta (Harley 16971) and in the
same locahty as that of C. bahiana (Harley 16985)
may provide evidence of introgression between these
sympatric species. The obese perianth and filament-
number (±124) are reminiscent of C. bahiana,
whereas the smooth perianth and upwardly imbricate
leaves suggest C. calycina. The leaf-formula (iii/30) is
intermediate, but the leaflets are more like those of C.
calycina in other respects.
98. Calliandra x cumbucana Renvoize, K e w Bull.
36: 75, fig. 3(25)p 7A. 1981. — "Brasil, Bahia . . .
Serra do Sincora, Mucuge, Rio Cumbuca, Harley et
al. 15394." — Holotypus, CEPEC!; isotypi, K!,
NY!. — Stirps forsan hybrida, inter C. calycinam et
C. viscidulam quasi intermedia, priori propior.
Shrubs attaining 3 m, with habit of C. calycina, the
young stems, lf-axes, and peduncles pilose with erect
pallid hairs to 0.75-1.2 m m mixed with few minute
reddish trichomes, the plane firm lustrous, facially
glabrous lfts remotely ciholate, the capitula solitary
and 1-2 together at most elaminate but conspicuously
stipulate, distal axils, forming a short simple pseudo
raceme or few-branched panicle scarcely exserted
from upper lvs. Stipules herbaceous, lanceolate or nar
rowly ovate-acuminate 5-9.5 x 1.2-3.3 m m , striately
7-11-nerved, deciduous. Lf-formula ii-iii/17-24; lf-
stks 1.2-2.5 cm, the petiole 3.5-5 m m , in dorsal view
1.1-1.8 m m diam, the one interpinnal segment or the
longer of two 7-12 m m ; pinnae scarcely or not dis
tally accrescent, the rachis of longer ones ±3.5^4-.5
cm, the longer interfoholar segments 1.7-2.3 m m ; lft-
pulvinules 0.15-0.25 x 0.45-0.7 m m ; lft-blades linear
from deeply auriculate base, deltately subacute,
straight or incipiently sigmoid, the longer ones 6.5-11
x 1.7-2.2 m m , 4.2-5 times as long as wide; midrib
displaced to divide blade 1:2-3, the inner posterior
nerve produced well beyond mid-blade, the whole ve
nation bluntly prominulous on both faces. Peduncles
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 163
Contrasted characters of C. calycina, C. viscidula, and suspected hybrid C. x cumbucana.
C. calycina C. viscidula C. x cumbucana
Stipules (mm)
Pinnae (-jug.)
Longer lfts (mm)
Floral bracts (mm)
Calyx
Corolla (mm)
Indumentum of perianth
2-6.5x0.2-1.6 i-ii
±5-9.4x1.1-1.8 Minute
Smooth, 3-5 mm; teeth
0.5-2 m m
6.5-8(-9)
Almost 0
5-9.5 x 1.2-3.3
ii-iii
±6.5-11x1.7-2.2
2.5-4.5x1.3-2 Venulose, 3 mm; teeth
1.3 m m
±8.5
Scattered white pili with few minute reddish
trichomes
5-12x1.5-7
iii-v
11.5-20x4-6
5-8.5 x 2.3-6
Venulose, 5.5-9.5 mm;
teeth 2-5.5 m m
8-14.5 Pilose and microglandular
±2.5 cm, ebracteate; capitula 6-9-fld, the clavate or
hemispherical receptacle 1-2.5 x 2-3 mm; bract of
outer fls papery brown, ovate 2.5-4.5 x 1.3-2 mm,
several-nerved, deciduous, that of inner fls 0; pedicels
drum-shaped ± 1 x 2 mm; perianth 4-merous, thinly
white-pilosulous distally, the calyx prominulously
±20-nerved, the corolla moderately camosulous,
externally nerveless, minutely obscurely resinous-
papillate; calyx shallowly campanulate ±3 mm, the
triangular teeth 1.3 mm; corolla ±8.5 mm, the lobes
3.5 mm; androecium 90-merous, ±42 mm, the ste
monozone 1.5 mm, the homy tube 7.5 mm, the tassel
white mbescent; ovary not seen. Pod unknown.
In damp sandy soil of riverside, among sandstone
rocks, 850 m, known from one station along rio Cum-
buca ±3 km s. of Mucuge at e. foot of Sa. do Sincora,
near 13°01'S, 41°21,W in interior Bahia. — Fl. II-?.
Mucuge, the source of the one known collection of
C. x cumbucana, is home also to C. calycina and C.
viscidula, each of which has been observed several
times, in typical form, in the immediate environs of
the town. The typus of C. x cumbucana has almost
the foliage of C. calycina, but has features, tabulated
above, that are atypical and could be derived by
introgression from sympatric C. viscidula. Pending
further observation, C. x cumbucana is provisionally
described and listed here as a suspected hybrid.
99. Calliandra hirsuticaulis Harms, Bot. Jahrb.
Syst. 42: 203. 1908. — "Bahia: Serra do Sincora,
1400 m (ULE n. 7312 — Nov. 1906)." — Holoty
pus, ̂ B = F Neg. 1244 [lapsu: hirticaulis]\; isotypi,
HBG!,K!.
C. hirsuticaulis sensu Renvoize, 1981: 75, fig. 2(33); Lewis,
1987: 174; Stannard 1995: 379.
Microphylhdious shrubs or subshrubs (2-)5-12(-20)
dm, the erect or diffuse, simple or distally few-
branched stems of the year together with lf-axes and
peduncles pilose-pilosulous with fine erect straight
whitish hairs to 0.6-1.7 m m mixed, at least on pedun
cles but often elsewhere, with reddish granular tri
chomes, the firm plane imbricate lfts either glabrous
or minutely pubemlent facially, ± lustrous on both
faces, rarely papillate, randomly ciholate, the capitula
of red-stamened fls shortly pedunculate in upper lf-
axils, often crowded into short-leafy pseudoracemes;
phyllotaxy distichous. Stipules herbaceous, ovate or
broadly lanceolate (2—)3—10.5 x 1.2-2.5 m m , mostly
6-11-nerved, commonly glabrous but sometimes pilo
sulous dorsally, ciholate, deciduous. Lf-formula
(ii-)iii-viii(-ix)/17-37(-42); lf-stks 7^10 m m , the
petiole including (or reduced to) pulvinus 1-3 m m , in
dorsal view 0.9-1.6 m m diam, the (one or) longest in
terpinnal segment 3-8 m m , the ventral sulcus bridged
at insertion of pinnae; pinnae subequilong or a little
accrescent distally, the rachis of longer ones 2.4—5(-6)
cm, the longer interfoholar segments 0.7-1.5 m m ; lft-
pulvinules 0.1-0.3 x 0.3-0.6 m m ; lfts decrescent at
each end of rachis, otherwise subequilong, the blades
hnear, narrowly (lance-)oblong, or linear-lanceolate
from shortly obtusely auriculate base, straight or gen
tly porrect beyond middle, deltately acute, the longer
ones (3.7-)4-8 x 0.9-2 m m , (2.7-)3^4.5(-5.1) times
as long as wide; venation palmate, prominulous dor
sally but only faintly so or immersed ventrally, the
midrib forwardly displaced to divide blade 1:(2.5—)
3.3—4, beyond middle 1-2-branched on posterior side,
the inner posterior primary nerve incurved-ascending
to or beyond mid-blade, the outer 1 (-2) much shorter.
Peduncles mostly <2 c m (but some lower ones attain
ing 3 cm), all ebracteate; capitula 5-9(-10)-fld, the
floral receptacle hemispherical or subtmncate ±1-2 x
1.2-2.5 m m ; floral bracts resembhng stipules in tex
ture and venulation, lanceolate or narrowly ovate
(2-)3-7 x 1-1.8 m m , deciduous; perianth either 5- or
4-merous, yellowish red-tinged, firm but scarcely
camosulous, either silvery-pilose overall or only the
164 M E M O I R S OF T H E N E W Y O R K BOTANICAL G A R D E N [VOL. 74(3)
teeth and lobes hirsute, the calyx- and corolla-tube
glabrous, the lobes sometimes (indistinctly) granular-
papillate; pedicels turbinate 0.4-1 x 1-1.6 m m ; calyx
campanulate or turbinate-campanulate (1.5—)3—5.5.
m m , the tube faintly 5-15-nerved, the lanceolate, tri
angular-lanceolate or exceptionally subdeltate teeth
(1.2-)1.4-3.6 m m ; corolla (5.2-)5.5-7.5(-9) m m , the
obovate lobes 2-3.2 m m ; androecium 30-72-merous,
(18—)21—31 m m , the cartilaginous stemonozone 1-1.8
m m , the pallid tube 3.5-6.5(-7) m m , the tassel blood-
red; ovary at anthesis glabrous. Pods (few seen) 4—6 x
0.7-0.8 cm, the sutural keels in dorsal view ±3^1 m m
wide, the recessed valves at once densely silky-pilo
sulous overall with erect sordid-white hairs <1 m m
and reddish-granular; seeds unknown.
About arenitic outcrops in campo mpestre, 1000-
1525 m, scattered across the e. slope of Chapada Dia
mantina in lat. 12°15V13°20'S (Seabra, Palmeiras,
Lencois and Mucuge on upper rio Paraguacu, and on
upper rio de Contas near Piata and Vila do Rio de
Contas), in upland interior Bahia. Fl. IX-XII.
The morphological characters, all of which or most
of which together characterize C. hirsuticaulis, are
the following: indumentum of fine erect hairs and at
least some granular trichomes; dilated stipules and
floral bracts; moderate leaf-formula of about iii-viii
/17-40; peduncles seldom over 2 cm; calyx-teeth
mostly longer but not much longer than the tube; and
blood-red androecium. The indumentum of the perianth
is denser in the upper Contas valley than northward.
Renvoize (1981: 66, in key) contrasted C. hirsuticaulis
with C. viscidula, but no close affinity between the pair
is probable.
100. Calliandra crassipes Bentham, Trans. Linn.
Soc. London 30: 555. 1875; & in Martius, Fl. Bras.
15(2): 426. 1876. — "... in subalpinis ad scaturig-
ines inter Caraboto et Caitete provinciae Bahiensis:
Martius." — Holotypus, Martius s.n., M ! = F Neg.
6163 = IPA Neg. 1324 = K Neg. 19446. — Feuilleea
crassipes O. Kuntze, Revis. Gen. PL 1: 187. 1891.
C. crassipes sensu Renvoize, 1981: 67, fig. 1(6, poor);
Lewis, 1987: 172.
Microphyllidious shrubs 0.5-2.5 m with stout vir
gate long-shoots but lacking axillary brachyblasts, the
young stems, lf-axes, and peduncles pilosulous with
pallid and either few or many red-brown granular tri
chomes mixed in varying proportions, the lvs bicol
ored, the lfts above lustrous dark brown-ohvaceous
and either glabrous, or papillate, or minutely pilosu
lous, beneath paler and beset with at least a few, often
many and crowded, red-brown trichomes, perhaps
resinous when fresh, the stoutly pedunculate capitula
of few obese fls arising singly and 2-3 together from
distal lf-axils; phyllotaxy distichous. Lf-formula v-
viii/17—23; lf-stks 2.5-^.5 cm, tapering from laterally
dilated base, dorsally low-convex, the petiole (includ
ing inert pulvinus) 3-5.5 x ±2.5 m m , the longer
interpinnal segments 3-7 m m , the ventral groove
bridged at insertion of pinnae; pinnae subequilong or
erratically graduated, the rachis of longer ones 2-
4 cm, the longer interfoholar segments 1-1.8 m m ;
lft-pulvinules 0.35-0.5 x 0.5-0.6 m m ; lfts linear-
lanceolate or linear, rarely narrow-oblong, from ob
tusely auriculate base, obtuse or deltately subacute,
the longer ones 5-7.5 x 1.3-2 m m , (2.8-)3.5^L2
times as long as wide; venation palmate, prominulous
only on dorsal face of 1ft, the straight midrib for
wardly displaced to divide blade 1:3-5, on posterior
side weakly 1-2-branched beyond middle, the inner
posterior primary nerve produced to or beyond mid-
blade, further venulation obscure. Peduncles (1.5-)
2.5-7 cm, at apex dilated into a hemispherical recep
tacle 3^- m m diam, capitula 4-7-fld; floral bracts
either linear to 6 m m (Harley 15657, N Y ) or wanting
(Harley 15521, N Y ) ; perianth 4-merous, obese,
inversely pyriform in bud, charged with scattered
short white hairs and crowded red-brown granular
trichomes; pedicels cryptic, externally hardly differ
entiated, except by discoloration, from calyx proper,
broadly turbinate 1-1.5 x 2-3 m m ; calyx 5-9.5 x
3-8 m m , the broadly lance-ovate teeth 3-7 m m , or
one sinus split to base and the calyx then quasi-
spathiform, the teeth becoming weakly few-nerved
when dry; corolla 6.5-9 m m , the broad-obovate
obtuse lobes 3.5-5 m m , usually surpassing the calyx
by 1.5-2.5 m m but sometimes a trifle shorter;
androecium 82-98-merous, 2.4-3.4 cm, the thick-
walled stemonozone 1.3-1.7 m m , the tube 6-8 m m ,
the tassel white rubescent; intrastaminal nectary 0;
ovary sessile, at anthesis glabrous. Pod unknown.
Along streams in rocky grassland at 1000-1300 m,
endemic to s. spurs of Sa. do Sincora near 13°30'S in
interior Bahia, Brazil. — FL XI—III.
Calliandra crassipes closely resembles C. bahiana
in indumentum, leaf-formula, and relatively large
obese perianth, but differs substantially in the calyx.
In C. bahiana the calyx-teeth are deltate and only
0.6-1.2 m m long; in C. crassipes lance-ovate and 3-7
m m long, not more than 2.5 m m shorter than the
corolla. In view of variation in the calyx of some re
lated species, such as C. coccinea and C. fasciculata,
the contrast between and C. crassipes and C. bahiana
may be overemphasized.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 165
101. Calliandra hirtiflora Bentham, Trans. Linn.
Soc. London 30: 550. 1875. — Typus infra sub var.
hirtiflora indicatur.
Amply microphyllidious shmbs 1-2 m with stout,
simply virgate or distally few-branched, densely fo
liate stems, commonly glabrous except for barbellate
ventral face of lf-axes but the young stems in addi
tion sparsely pilose or barbate with erect, extremely
fine, shining white hairs to ±1.4—1.8 m m , the pedun
cles and perianth of fls sometimes minutely granular,
the foliage drying brown, the lfts darker and lustrous
above, paler dull and faintly resin-spotted beneath,
either glabrous overall or randomly cihate, the stout
peduncles arising singly or 2(-3) together from dis
tal lf-axils or shortly pseudoracemose; phyllotaxy
either distichous or incipiently spiral. Stipules herba
ceous, ovate or lanceolate 5-11 x (1.8-)2-4.5(-5)
m m , several-nerved, glabrous or rarely ciliate, decid
uous. Lf-formula (iii-)iv-vii/(22-)24-38(-40); lf-stk
of longer lvs 3.5-6.5(-10) cm, the petiole including
nigrescent wrinkled pulvinus 10-22 cm, at middle
1.1-1.7 m m diam, the longer interpinnal segments
8-15 m m , the ventral groove bridged at insertion of
pinnae; pinnae either scarcely or randomly graduated
in length, the rachis of longer ones (4—)4.5-7.5 cm,
the longer interfoholar segments l-2.2(-3) m m ; lft-
pulvinules 0.25-0.5 x 0.5-0.7 m m ; lfts subequilong
or gradually decrescent toward end of rachis, the firm
plane blades linear from obtusangulate or shortly
obtusely auriculate base, deltately obtuse, the longer
ones (6-)7-12 x 1.5—4- m m , 4.2-6 times as long as
wide; midrib simple or almost so, 1-2 weak posterior
primary nerves sometimes faintly raised, the tender
young lfts becoming vertically wrinkled when dried.
Peduncles 3.2-8.5 cm, usually ebracteate; capitula
(4—)4—8-fld, the receptacle 1.5-4.5 m m diam; bracts
resembling stipules in texture and venation, narrowly
ovate or lanceolate 3-6 x 1-2 m m , caducous; fls
homomorphic; pedicels 0.8-1.7 x 1.2-1.9 m m ; peri
anth moderately thickened, ochroleucous, either a)
proximally smooth and distally granular-papillate, or
b) papillate and remotely strigulose overall, or c)
smooth but randomly ciliate, or d) distally pilose, the
calyx (4-)5-merous, the corolla mostly 4- but by
fusion sometimes 3- or 2-merous, the calyx-teeth
sometimes weakly 1-3-nerved but the rest externally
nerveless; calyx 5.5-9.2 m m , the narrowly lanceo
late to narrowly ovate, sometimes unequal teeth (3-)
3.5-7 m m ; corolla 6.5-11 (-12) m m , either as long as
calyx or well exserted, the lobes separated to vari
able depth, sometimes to the rim of stemonozone;
androecium 44—112-merous, (2-)2.4—4.1 cm, the
stemonozone 1.7-3.5(-4) m m , the tube (5-)7-9 m m ,
the tassel white, mbescent in age; ovary at anthesis
glabrous, pubescent after fertilization. Pods in profile
7-10 x 0.75-1 cm, the sutural ribs in dorsal view ±3
m m wide, the recessed valves transversely and sub-
reticulately venulose, the whole ± densely granular
or the margins egranular and thinly pilosulous; seeds
light brown, ±6.5-7 x 4—5 m m , the pleurogram 4-4.5
x 2.2-2.5 m m .
Calliandra hirtiflora proves quite variable in de
tail, and a sharply defined image of it is not easily
attained. A combination of herbaceous stipules about
5-11 x 2-4.5 m m , petioles about 1-2 c m long, pinnae
mostly 4—7 pairs, and calyx-teeth longer than the tube
separate it from presumably related taxa of Chapada
Diamantina. The material falls into two categories
that I first described as independent species but that
now seem better assessed as varieties of one.
Key to the varieties of C. hirtiflora
1. Young stems and perianth barbate with fine straight shining white hairs to 1.5 mm; corolla and pod egranular. 101a. var. hirtiflora
1. Young stems and perianth either glabrous or only remotely pilose; corolla and pod minutely granular. 101b. var. ripicola
101a. Calliandra hirtiflora Bentham var. hirtiflora.
C. hirtiflora Bentham, 1875, I.e.; & in Martius, Fl.
Bras. 15(2): 420. 1876. — ". .. in campis deserti ad
Caitete provinciae Bahiensis: Martius." — Holoty
pus, M ! = F Neg. 6164 = IPA Neg. 1328 = K Neg.
19447. — Feuilleea hirtiflora O. Kuntze, Revis.
Gen. PL 1: 188. 1891.
C. hirtiflora sensu Renvoize, 1981: 75, fig. 4(26).
Characters of the species, except as modified by
key to varieties.
In campo, described (perhaps erroneously) from
Caietete, collected since only in mun. Ituacu in inte
rior upland Bahia, Brazil. — Fl. VI-VII(-?).
101b. Calliandra hirtiflora Bentham var. ripicola
Bameby, var. nov, a var. hirtiflora cauhbus sub-
glabris, perianthio minutim granuloso recedens. —
Brazil. Bahia, mun. Andarai: rio Paraguacu, 19 Jun
1984 (fl). G. Hatschbach (with R. K u m m r o w )
48062. — Holotypus, M B M ; isotypi, K, NY.
Characters of the species, except as modified by
key to varieties.
166 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
On rocky river banks, 400-1000 m, locally plenti
ful along upper n. fork of rio Paraguacu, between
12°10'S and 12°50'S (Dr Seabra, Palmeiras, Lencois,
Andarai) in interior Bahia. — FL I, VI, IX-X, the full
cycle not known.
102. Calliandra sincorana Harms, Bot. Jahrb. Syst.
42: 204. 1908. — "Bahia: Serra do Sincora, 1400
m. (ULE n. 7310. — Nov. 1906)." — Holotypus, +B
= F Neg. 12391; isotypus, HBG! = K Neg. 18754.
C. exsudans Harms, Bot. Jahrb. Syst. 42: 203. 1908. — "Bahia: Serra da Vendinha [900 m], Sincora (ULE n. 7133.
— Nov. 1906)." — Holotypus, *B = F Neg. 7239!; clastotypus, F!; isotypi, HBG!, K!.
C. sincorana sensu Renvoize, 1981: 77, fig. 3(29); Lewis, 1987: 176.
C. exsudans sensu Renvoize, 1981: 67, fig. 1(5); Lewis,
1987: 173 ("exudans").
Shrubs 0.5-3 m with fuscous defohate annotinous
stems and densely foliate new branches, except for
thinly barbellate ventral face of lf-axes seemingly
glabrous throughout but here and there micropuberu
lent, resinous and papillate, the plane bicolored lfts
dark brown sublustrous above, paler dull beneath, the
few-fld capitula geminate in upper coeval lvs, im
mersed in foliage or shortly exserted but not pseudo
racemose; phyllotaxy distichous. Stipules lanceolate
or narrowly ovate obtuse 2.5-4 x 0.5-1.6 m m , exter
nally nerveless or faintly 3-nerved, deciduous. Lf-
formula ii-iii/17-25; lf-stk of larger lvs 12-19 m m ,
the petiole including discolored pulvinus ±5-9 m m ,
in dorsal view 0.9-1.2 m m wide, the one or the longer
of two interpinnal segments about as long; pinnae a
little accrescent distally, the rachis of furthest ones
±3.5-4.5 cm, the longer interfoholar segments 1.5-3
m m ; lft-pulvinules 0.15-0.3 x 0.4—0.5 m m ; lfts grad
ually decrescent distally, the subcontiguous or imbri
cate blades subfalcately linear from shortly bluntly
auriculate or merely inequilateral base, obliquely
deltate at apex, the longer ones 8-10.5 x 1.8-2.4 m m ,
4.2-5 times as long as wide; venation weak on both
faces, the simple or almost simple midrib forwardly
displaced to divide blade 1:2.5-4.5, a posterior pri
mary nerve scarcely perceptible. Peduncles 2.5-5 cm,
bracteate above middle or close under the capitulum,
the bract lanceolate or ovate-conduphcate 1.5-3.5
m m ; capitula 3-6-fld, the receptacle 1.5-2 m m diam;
floral bracts narrowly lanceolate ±3 m m , caducous;
fls homomorphic, the perianth 4-merous, glabrous,
copiously resinous, papillate distally, the corolla
sometimes becoming spathaceous when the lobes
adhere in pairs; pedicels 0.6-1.5 x 0.5-1.2 m m ; calyx
3.5-7 m m , the campanulate tube 1.5-2.3 m m , the
lanceolate teeth 2-5.5 m m ; corolla 6-10 m m , the
lobes variably adherent or separated to variable
lengths, ±3 m m longer than calyx; androecium 26-
42-merous, 2.2-5.2 cm, the stemonozone 0.6-2.3
m m , the tube 5-9 m m , the tassel white rubescent;
ovary at anthesis glabrous. Pod not seen.
In unrecorded habitat but to be expected about rock
outcrops in campo mpestre, 900-1400 m, known at
present only from the type of the species and from
that of C. exsudans, both collected in Serra do Sin
cora, interior Bahia. — Fl. XI-XII.
Harms compared C. sincorana with C. fasciculata,
which is similar in leaf-formula but has a much
shorter calyx. He expressed no opinion as to the
affinities of C. exsudans, which appears in Renvoize's
(1981) key to Bahian Calliandra to differ from all
relatives in flowers "covered with silver-white needle
like hairs and white exudate." These hairs, however,
are foreign filaments adhering to the sticky peduncles
and perianths, and the exudate consists partly of
wind-blown particles of quartz and partly of palhd tri
chomes such as occur also in the typus of C. sinco
rana. The typi of C. sincorana and C. exsudans are
dissimilar insofar as the perianth of the latter is a little
longer and its androecium about twice as long. In
leaf-formula, in individual leaflets, and in bracteate
peduncles they are essentially alike, and represent
probably two populations of one species.
103. Calliandra stelligera Bameby, sp. nov, pinnis
recurvis, florum forma et magnitudine, ahisque notu-
lis C. elegantem (fere sympatricam) revocans et ei
sine dubio proxime affinis, sed statura demissa, pin
nis brevioribus 6-8.5 (nee 10-14.5) c m usque longis,
fohohs minus numerosis ±19 (nee 30-35)-jugis
usque utrinque dense minutim pubemlis, necnon
androecio 28-32 (nee ±54)-mero saturate rubro (nee
ex albo rubescenti) diversa, ulterius indumento
ex parte e trichomatibus stelliformibus constanti in
genere suo unica. — BRAZIL. Bahia: mun. de Piata:
estrada Inubia-Piata, descida da Serra do Atalho para
Iniibia, 1350 m, 13°06'S, 41°56,W, 20 Aug 1992 (fl),
Wilson Ganev 917. — Holotypus, HUEFS 11360.
Relatively amply plurifoholate subshrub ±5 d m
tall, stiffly widely branched above middle, the new
stems with all If- and inflorescence-axes densely
pubemlent overall with simple pallid hairs <0.2 m m
mixed with minute mfous granules and, especially on
peduncles but randomly elsewhere, with minute plur-
iradiate stellae, the pallidly ohvaceous lfts subconcol
orous, dull and equally pubemlent on both faces, the
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 167
solitary and geminate peduncles forming a short ter
minal efohate panicle of pseudoracemes exserted <1
d m from foliage; phyllotaxy distichous. Stipules
firmly herbaceous, narrowly ovate ±1.5-2 x 0.8-1.2
m m , externally nerveless, tardily deciduous. Lf-for
mula iii/18-19; lf-stks ±4 cm, the petiole 5-10 m m ,
at middle ±2 m m diam, the longer interpinnal seg
ments 12-15 m m , the ventral groove bridged at in
sertion of pinnae; pinnae scarcely graduated, arcu-
ately recurved, 5-8.5 cm, the longer interfoholar
segments ±3-4 m m ; lft-pulvinules ±0.5-0.6 x 1 m m ,
faintly cross-wrinkled; lfts a little decrescent at each
end of rachis, otherwise subequilong, the blades
oblong from shallowly auriculate base, broadly obtuse,
those near mid-rachis 11-16 x 2.6-3 m m , 2.6-3 times
as long as wide; venation bluntly low-prominulous
dorsally, obscurely so ventrally, the straight midrib
displaced to divide blade ±1:2-2.5, weakly 1-2-
branched distally, the inner posterior primary nerve
incurved-ascending well beyond mid-blade, the outer
2-3 posterior ones progressively much shorter. Pe
duncles ±1.5-2.5 cm, ebracteate; capitula 5-12-fld,
the clavate receptacle 1.5-2 m m ; bracts deltate-ovate
to 0.8 m m , caducous; fls homomorphic, the perianth
4- or the corolla by fusion of lobes 3-, 2-merous,
gray-silky-puberulent overall but more densely so
above middle; pedicels disciform 0.3-0.4 x 1.5-1.8
m m ; calyx campanulate 2 x 2.4—2.7 m m , the low-
deltate teeth ±0.3 m m ; corolla 6.5 m m , the ovate lobes
±2 m m ; androecium ±2.5 cm, 28-32-merous, the car
tilaginous tube 2 m m , the stemonozone 1.5-1.6 m m ,
intrastaminal nectary 0; ovary at anthesis glabrous.
Pod unknown.
In campo mpestre near 1350 m, known only from
the type-locality in Chapada Diamantina near 13°S,
42°W, in interior Bahia, Brazil. — Fl. VIII(-?).
Calliandra stelligera seems akin to C. elegans but
is, so far as known, a dwarfer shrub with shorter pin
nae, fewer, densely pubemlent leaflets, and fewer,
bright red filaments; for measurements see the Latin
diagnosis. The stellate trichomes of the peduncles
have not been seen elsewhere in this genus.
104. Calliandra coccinea Renvoize, Kew Bull. 36:
69. 1971. — Typus sub var. coccinea indicatur.
Microphyllidious subshrubs 5-10 dm with stiff
straight annotinous stems widely few-branched and
densely leafy distally, thinly pubemlent nearly
throughout with fine erect white hairs <0.2 m m , the
young stems, the lf-axes, the dorsal face of lfts and all
units of inflorescence in addition beset with red-
brown granular or coralloid trichomes, the shortly
pedunculate capitula solitary in a few distal lf-axils,
nestled in foliage; phyllotaxy distichous. Stipules
erect, narrowly ovate or lanceolate 1.5^4.5 x 0.7-1.8
m m , faintly 1-nerved dorsally, caducous. Lf-formula
(ii-)iii-iv/(22-)25-37; lvs nearly sessile, the lf-stk of
longer ones 6-18 m m , the petiole often reduced to
discolored pulvinus, 1-2.5 x 1.2-1.5 m m , the longer
interpinnal segments 2.5-9 m m , the ventral groove
bridged at insertion of pinnae; pinnae either equilong
or the furthest shorter, the rachis of longer ones
30-47 m m , the longer interfoholar segments 0.45-
1.1 m m ; lft-pulvinules 0.1-0.25 m m ; lfts equilong ex
cept at very ends of rachis, the blades oblong-elliptic
from obtusely auriculate base, obtuse or deltately
subacute, the longest 3^.5 x 1.1-1.5 m m , 2.5-3.2
times as long as wide; venation faint, the straight sim
ple midrib forwardly displaced to divide blade 1:2.5—
5, prominulous only dorsally, a short posterior pri
mary venule sometimes faintly perceptible. Pedun
cles 5-12 m m , ebracteate; capitula 3-6-fld, the fls
sessile, the receptacle subtmncate; bracts resembling
stipules in form and texture, 1.5-4-.5 m m , caducous;
perianth firm, reddish-granular overall, the calyx
4-merous, the corolla (2-)3-4-merous, further
described in key to varieties; androecium 42-51-
merous, 24—40 m m , the stemonozone 0.6-2.1 m m ,
the tube 5-7 m m , the tassel red; stamen-tube thick
ened internally, but no nectary differentiated; ovary
substipitate, glabrous at anthesis. Pods (of var. coc
cinea) ±4.5 x 0.7 cm, densely tomentulose overall
with mixed simple gray and brown coralloid tri
chomes, the recessed valves not evidently venulose;
seeds not seen.
The five known collections of C. coccinea are
nearly uniform in foliage, in reddish-brown granular
indumentum of the perianth, and in the red-tasseled
androecium, but are remarkably dissimilar in the
inner whorl of the perianth. I feel obliged to discrim
inate taxonomically between these florally differenti
ated forms.
Key to the varieties of C. coccinea
1. Fl-buds prior to anthesis pyriform, obscurely
minutely umbonulate at tip; corolla 3-merous,
±7 mm, the sinuses between the 3 broadly
ovate lobes of about equal depth. . . . 104a. var. trimera
1. Fl-buds prior to anthesis narrowly flask-
shaped, long-attenuate; corolla 4-merous,
7.5-13.5 mm, either once deeply cleft and
spathiform, entire or minutely 4-toothed at
apex, or deeply cleft into 2-3 lobes, one of
them apically 2-denticulate 104b. var. coccinea
168 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
104a. Calliandra coccinea Renvoize var. trimera
Bameby, var. nov., a var. coccinea corolla trimera, ra-
diatim symmetrica vix 7 m m longa, lobis late ovatis;
flores ante anthesin pyriformes, obscure umbonulati.
— BRAZIL, Bahia, mun. Rio de Contas: Serra das
Almas a 5 k m N W de Vila do Rio de Contas, 21 Mar
1980 (fl), S. A. Mori & F. Benton 13532. — Holo
typus, C E P E C ; isotypi, M O , NY. FiG. 25
Habit, lvs and indumentum of the species; calyx 4-
merous ±3 m m , the tube 1.3 m m , the teeth ±2.6 m m ;
corolla 3-merous, 6.5-7 m m , 3-lobed nearly to mid
dle, the ovate lobes ±3 m m .
In campo mpestre between 1000 and 1200 m,
known only from one collection from the foothills of
Pico das Almas in interior Bahia. — FL III—IV.
104b. Calliandra coccinea Renvoize var. coccinea.
C. coccinea Renvoize, 1981, sens. str. — "Bahia.
Serra do Rio de Contas: Rio de Contas . . . Mato
Grosso Harley et al. 19980." — Holotypus, C E P E C
n.v; isotypus, K!. FiG. 25
C. coccinea sensu Lewis, 1987: 172, pi. 10D.
Habit, lvs and indumentum of the species; calyx 4-
merous 4.7-5.4 m m , the tube 1.7-2.4 m m , the teeth
2.7-3.3 m m ; corolla 2^4-merous, 7.5-13.5 m m , slen
derly flask-shaped, either a) once cleft to below mid
dle and spathiform, the attenuate limb entire or
minutely 4-denticulate at apex, or b) 3-cleft to near
middle or further, 2 lobes acute, 1 lobe 2-denticulate.
Habitats of var. trimera, 1000-1600 m, known
only from upper Rio de Contas in vicinity of Abaira,
Piata, and Vila do Rio de Contas, in lat. 13°09 -33'S.
— FL XII-IV, VII.
105. Calliandra involuta Mackinder & Lewis, Kew
Bull. 45: 683, fig. 2. 1990. — "Brazil, Bahia
Serra Larga ('Serra Larguinha'), Municipio de
Lencois, near Caete-Acu, 19 Dec. 1984, Lewis et al.
C F C R 7248." — Holotypus, SPF n.v; isotypus, K
(2 sheets)!.
Shrubs attaining 1.5 m with virgate long-shoots
and without axillary thatched brachyblasts, the new
stems, lvs, and inflorescence pilose with fine erect,
glossy white or ochroleucous hairs to 1.4-2.2 m m
mixed with very small, red or livid granular tri
chomes, nowhere resinous, the lvs subconcolorous
but the lfts facially differentiated, ventrally convex,
medially sulcate and intricately rugulose-crumpled,
dorsally concave and carinate by the midrib, the mar
gin strongly revolute, the capitula arising singly from
a few furthest lf-axils; phyllotaxy distichous. Stipules
FiG. 25. Calliandra coccinea Renvoize var. trimera Bameby (left, perianth; center, habit) and var. coccinea (right, perianth).
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 169
triangular-lanceolate 3.5-6 m m , resembhng lfts in
form, texture, and pubescence, caducous. Lf-formula
(ii-)iii/10-15; lvs subsessile, the lf-stk of longer ones
±2-3 cm, the stout petiole ±2-4 x 1.5-2 m m , the
longer interpinnal segments 8-13 m m , the ventral
groove bridged at insertion of pinnae; pinnae of ran
d o m lengths but the second pair often longest, the
rachis of these 3-5.5 cm, the longer interfoholar seg
ments (3-)3.5-6 m m ; lft-pulvinules ±0.2 x 0.8-1 m m ;
lfts decrescent at each end of rachis, narrowly oblong-
elhptic from postically acutangulate base, broadly
obtuse, those near mid-rachis (discounting revolute
edges) 8.5-11 x 2.7-3.6 m m , 2.8-3.2 times as long as
wide; midrib only a trifle forwardly displaced from
mid-blade, straight, weakly 2(-3)-forked on posterior
side near and beyond middle, one posterior primary
nerve incurved-ascending to mid-blade, no secondary
venulation. Peduncles ±3.5-5.5 cm, ebracteate, the
low-convex, ±5-8-fld receptacle 2.5-3 m m diam; flo
ral bracts narrowly ovate 4—5.5 x 1.6-2 m m , faintly
striate, persistent; pedicels externally differentiated
only by discoloration, disciform ±0.5-1.4 m m ; peri
anth greenish-yellow, each cycle pilose from middle
upward, glabrescent proximally, minutely reddish-
granular, the calyx faintly ±15-nerved, the corolla
submembranous, its lobes hardly 0.2 m m thick in sec
tion; calyx 4-merous, turbinate-campanulate 3-3.5
m m , the narrowly lanceolate teeth of unequal length,
the longest ±2.5 m m , each prominulously 3-nerved;
corolla (?always) 3-merous, 5.3-5.7 m m , the obtuse
lobes ±2.2 m m ; androecium 30-32-merous, ±28 m m ,
the internally thickened stemonozone ±1 m m , the
tube 4-4.5 m m , the tassel rose-red; ovary at anthesis
glabrous. Pod not seen.
In relict cloud forest near 1400 m, known only
from near the crest of Sa. Larga w. of Lengois, at n.
end of Sa. do Sincora, in interior Bahia. — Fl. XII-I.
Calliandra involuta resembles C. asplenioides in
general habit of growth, but is distinguished at a
glance by its indumentum of glossy hairs mixed with
livid granules, its fewer, ventrally bullate, revolute
(not involute) leaflets, and persistent floral bracts.
106. Calliandra santosiana Glaziou [Mem. Bull. Soc.
Bot. France 3: 188. 1905, nom. nud.] ex Bameby, sp.
nov, inter congeneres brasihenses planaltinos conju-
gato-pinnatos stipuhs parvis (1-3 m m ) extus en-
erviis, fohohs foliorum m a j o m m valde numerosis
(70-88-jugis usque) parvis (±6.5-8.5 x 1.1-1.6 m m
usque) secus rachin elongatam (8-13.5 c m longam)
confertis, capituhs paucifloris omnino glabris, nec
non filamentis vivide rubris praestans. — BRAZIL.
Minas Gerais: Biribiry, 23 Mar 1892 (fl), A. Glaziou
19113. — Holotypus, P; isotypus, K.
Erect, fmtescent but of unknown stature, glabrous
except for rudimentary pubemlence of some lf-axes,
notable for slender, sparsely foliate, distally branched
stems and elongate unijugate microphylhdious pin
nae, the lvs subconcolorous, glossy on upper face, the
capitula of red-stamened fls borne singly and 2^4
together at nodes of efoliate branchlets axillary to
distal lvs, together forming a loose narrow panicle;
phyllotaxy distichous. Stipules narrowly lanceolate or
subulate 1-3 x 0.4—0.6 m m , externally nerveless,
deciduous. Lf-formula i/70-88; lf-stks including obese
nigrescent pulvinus 3-9 m m , at middle 0.6-1.4 m m
diam; rachis of longer pinnae 8-13.5 cm, the inter
foholar segments 0.85-1.3 m m ; lft-pulvinules 0.25-
0.4 m m ; lfts shghtiy decrescent near each end of
rachis, otherwise equilong, the blades linear or linear-
lanceolate from shortly auriculate base, straight or
scarcely falcate, deltately subacute, the larger ones
5.5-8.5 x 1.1-1.6 m m ; venation faint, the simple
midrib forwardly displaced to divide blade 1:2, the
weak inner posterior primary vein expiring near or
short of mid-blade, the upper lft-face not clearly venu
lose but somewhat rugulose when dry. Peduncles
14—24 m m , ebracteate; capitula 5-9-fld, the receptacle
±1.5-2 x 2 m m ; bracts minute caducous; fls (sub)ses-
sile; perianth glabrous; calyx campanulate or turbinate-
campanulate 1-2 x 1.5 m m , weakly 5-nerved, the
deltate apiculate teeth 0.4-0.6 m m ; corolla 5-6.5 m m ,
the ovate lobes ±3 m m ; androecium 36-40-merous,
nearly 2.5 cm, the stemonozone 1-1.3 m m , the tube
4.5-6 m m ; no intrastaminal disc; ovary at anthesis
glabrous. Pod unknown.
In white gravelly sand near 1200 m, known from
two collections from the vicinity of Diamantina
(Biribiry; rio dos Cristais) in n.-centr. Minas Gerais,
Brazil. — Fl. II-IV.
Calliandra santosiana resembles C. renvoizeana,
C. debilis, and C. longipinna, all of which are en
demic to Bahia, but differs from all in leaf-formula
and in red filaments, described by the collectors as
rouges or vermelhas insignes. The epithet santosiana
is of unknown significance.
Harms (Repert. Sp. Nov. Regni Veg. 17: 90. 1921)
misinterpreted an isotype at Berlin (no longer extant)
as a variety of C. fasciculata distinguished by small
stipules and more numerous and smaller leaflets.
107. Calliandra renvoizeana Bameby, nom. nov.
C. gracilis Renvoize, K e w Bull. 36: 73, fig. 2(19),
170 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
5(E, F). 1981. — "Brazil. Bahia [s.-w. of] Mucuge
[on road from Cascavel, ±41°25'W, 13°02'S, 6 Feb
1974, fl, fr], Harley et al. 16061." — Holotypus,
CEPEC!; isotypi, K!, NY!. — Non C. gracilis
Grisebach, 1861.
C. gracilis sensu Harley & Simmons, 1986: 114; Lewis, 1987: 174.
Virgate, thinly foliate subshrubs attaining ±12 dm,
the simple stem glabrate and vertically striped proxi
mally but pilose distally with straight spreading, lus
trous white hairs to 1.3-2 m m , the lvs glabrous
except for adaxially barbellate axes and rare random
cilia, the few-fld capitula (1—)2—3 together in upper
lf-axils and, beyond these, forming a short efoliate
pseudoraceme; phyllotaxy distichous. Stipules ovate
from shallowly cordate base, ±6-10 x 3-6 m m , thinly
herbaceous, facially glabrous, ciliate, palmately
many-nerved, deciduous long before the associated
If. Lf-formula i/40-58; lf-stks including pulvinus 5-7
x 1-1.4 m m ; rachis of longer pinnae 7-9 cm, the in
terfoholar segments 1.4-2 m m ; lft-pulvinules 0.3 x
0.6-0.9 m m ; lfts scarcely graduated, the blades nar
rowly oblong-elhptic from obtusely auriculate base, at
apex obtusely deltate, the larger ones 5-8.5 x 1.7-
2.2 m m , 3-4 times as long as wide; venation almost
fully immersed, the midrib faintly perceptible dorsally
but not prominulous. Peduncles 3.5-5 cm, ebracteate;
capitula 3-7-fld, the receptacle ±2 x 2.5 m m ; bracts
resembling stipules in shape and size, caducous; fls
subsessile homomorphic, the obese dmm-shaped ped
icel 0.9-1.2 x 1.3-2.5 m m ; perianth 5-merous,
glabrous proximally, thinly pilose distally, faintly
several-nerved; calyx turbinate-campanulate 6-9 x
5-6 m m , the ovate lobes 3-i m m ; corolla 8.5-11.5
m m , the ovate lobes 2>-^ m m ; androecium 60-65-mer-
ous, 4—5 cm, the thickened stemonozone ±2 m m , the
tube 6.5-10 m m , the tassel white mbescent. Pods in
profile ±10 x 1 cm, densely softly pilose overall, the
sutural ribs in dorsal view ±2 m m wide, the valves
externally evenulose; seeds (not seen fully ripe) ±8.5
x 4.5 m, the narrowly U-shaped pleurogram palhd.
In campo mpestre or wooded grassland, on stony
soils, 900-1000 m, known only from Sa. do Sincora
in interior Bahia, Brazil: s.-w. of Mucuge (40°25'W,
13°02'S) and between Mucuge and Andarai (41°
18'W, 12°15'S). — Fl. II-III, VI.
Calliandra renvoizeana resembles C. debilis in vir
gate habit, ovate venulose stipules, and few-flowered
capitula, but differs in more numerous and narrower
leaflets, distally pilose stems, peduncles, and flowers,
and differently proportioned, longer individual flow
ers. The type-localities of these sibling species are
separated by no more than 8' latitude and 6' longitude.
108. Calliandra longipinna Bentham, Trans. Linn.
Soc. London 30: 546. 1875; & in Martius, FL bras.
15(2): 416. 1876. — "Habitat in subalpinis ad scatu-
rigines inter Carapato [perhaps Fazenda Carabatos,
between Sincora and Maracas, visited by Martius
(cf. Urban, 1906: 39) in late October, 1818] et
Caitete [sic]: Martius." — Holotypus, Martius s.n.,
M ! (2 sheets, one = F Neg. 6166 = K Neg. 19439) +
Martius Obs. 1975, M!(same data); probable iso
typus, G!. — Feuilleea longipinna O. Kuntze, Revis.
Gen. PL 1: 188. 1891.
C. longipinna sensu Renvoize, 1981, fig. 1(10); Lewis,
1987: 184.
Virgate subshrubs with stiff, simple or few-branched,
sparsely foliate stems 3-5 dm, the short petioles and
the elongate rachises of the 1-2-jugate pinnae thinly
barbellate ventrally and the young stems, peduncles
and fls minutely reddish-pubemlent with twisted or
granular trichomes <0.2 m m , the firm, dorsally venu
lose lfts facially glabrous, subconcolorous, the few-
fld capitula solitary or geminate in some distal lf-
axils, the furthest sometimes pseudoracemose late in
season; phyllotaxy distichous. Stipules lance-ovate
1-3.5 x 0.9-1.8 m m , not or weakly venulose dorsally,
deciduous. Lf-formula i-ii/(33-)38-48; lf-stks 2-14
m m , the petiole with pulvinus 2-5(-8) m m , at middle
1-1.6 m m diam, the interpinnal segment 7-10 m m ;
rachis of longer pinnae 5.5-12 cm, the longer interfo
holar segments 1.2-2 m m ; lft-pulvinules 0.2-0.4 x
0.55-0.8 m m ; lfts subequilong except at far ends of
rachis, elliptic-oblong from obtusely semicordate
base, broadly obtuse, the largest 4.5-8.5 x 2-3.8 m m ,
2-2.5 times as long as wide; venation palmate, ob
tusely prominulous only dorsally, the straight, few-
branched midrib forwardly displaced to divide blade
±1:2.2-3.5, the inner posterior primary nerve pro
duced almost to blade's apex, the outer ones progres
sively shorter, camptodrome well within the plane
margin, the secondary venules randomly anastomos
ing. Peduncles 1.2-3 cm, ebracteate; capitula 3-6-fld,
the receptacle ±1 x 2-2.5 m m ; bracts deltate or lance-
ovate 0.8-2.5 m m , deciduous; fls homomorphic, ses
sile or nearly so, the turbinate or discoid pedicel <0.6
m m , the perianth thinly red-granular, the calyx dis
tinctly pallid-venulose but the corolla not so, the
corolla-lobes in addition minutely pallid-strigulose;
calyx turbinate-campanulate ±2.5-3 x 2.5-3.8 m m ,
the deltate teeth 0.3-1.3 m m ; corolla campanulate
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 171
±5.7-8.5 m m , the broadly ovate-deltate lobes ±1.8-
3.5 x 2.2-2.5 m m ; androecium 54-78-merous, 2.5-
3.3 cm, the thickened stemonozone ±1-3.2 m m , the
tube 4.5-6 m m , the tassel reddish-pink; ovary at
anthesis glabrous. Pod not seen.
In thinly wooded grassland near 1100 m, some
times in moist microhabitat, known with certainty
only from Sa. do Sincora (near Barra da Estiva and
Ibicoara) in upland interior Bahia, Brazil. — Fl. X-II.
This is the only Bahian species, so far as known, in
which red-granular peduncles and perianths coincide
with 1-2-jugate pinnae. One unicate specimen from
north of Barra da Estiva (Harley 15853, C E P E C ) has
a few upper leaves with three pairs of pinnae, not
allowed for in the foregoing description.
109. Calliandra debilis Renvoize, Kew Bull. 36: 73,
fig. 6A. 1981. — "Serra de Sincora [s. of Andarai
on road to Mucuge] near Xique-xique [14 Feb 1977,
fl], Harley et al. 18676." — Holotypus, C E P E C
n.v; isotypi, K!, NY!.
C. debilis sensu Harley & Simmons, 1986: 114; Lewis,
1987: 172.
Remotely foliate virgate shrubs attaining 2.5 m
with terete reddish, longitudinally striped stems,
except for white-barbellate inner margins of pin-
narachises glabrous, the firm lfts bicolored, lustrous
olivaceous above, paler and randomly resin-spotted
beneath, the few-fld capitula either solitary or gemi
nate in the furthest lf-axils and beyond these shortly
pseudoracemose. Stipules (few seen) herbaceous firm,
ovate from shallowly cordate base, ±5-6 x 3.5 m m ,
weakly palmate-nerved, deciduous. Lf-formula i-ii
/32-46; lf-stks including pulvinus 6-21 m m , the one
interpinnal segment, developed in few lvs, nearly
twice as long; rachis of longer pinnae 8-9.5 cm, the
interfoholar segments ±2-2.5 m m ; lft-pulvinules 0.5-
0.6 x 0.6-0.75 m m ; lfts subequilong, the blade
oblong-elliptic from bluntly broad-auriculate base,
deltately subacute, straight, the larger ones 7.5-9 x
2.6-3 m m , 2.8-3.2 times as long as wide; venation of
dorsal lft-face palmate, the straight, almost simple
midrib displaced to divide blade ±1:2, the inner poste
rior and one anterior primary vein produced well
beyond mid-blade, the 2-3 outer posterior ones pro
gressively much shorter, a fine secondary subvertical
venulation scarcely perceptible. Peduncles ±2.5-5 cm,
ebracteate; capitula 3-5-fld, the homomorphic fls sub
sessile, the receptacle ±1-1.5 x 1.5-2 m m ; bracts
ovate-acuminate 2-2.5 m m , weakly 5-7-nerved, ca
ducous; pedicels turbinate ±1 x 1.5-1.8 m m ; perianth
5-7-nerved, glabrous except for few random white
cilia at tip of calyx-teeth or corolla-lobes; calyx cam
panulate 2.5-2.8 x 2.7-3 m m , the deltate teeth sepa
rated by broad shallow sinuses; corolla 9 m m , the
lobes ±4 x 3.5 m m ; androecium ±56-merous, 5 cm,
the stemonozone 1 m m , internally thickened, the tube
5 m m ; no intrastaminal nectary. Pods (not seen)
described as "10 c m x 11 m m , reddish, pilose."
About sandstone outcrops in campo mpestre, 700-
1200 m, known only from the vicinity of the type-
locality in Sa. do Sincora near 12°54,S, 41°19,W in
upland interior Bahia, Brazil. — FL II—III(—?).
Calliandra debilis is in most respects hardly differ
ent from C. longipinna, but may be distinguished
technically by the larger stipules, 5-8 (not 1-3.5) m m
long and the nearly glabrous, neither strigulose nor
reddish-granular corolla.
110. Calliandra iligna Bameby, sp. nov, C. san-
tosianae Glaziou ex Bameby proxime affinis, sed ab
ea cauhbus simphcibus, fohomm multomm pinnis 2-
jugis (nee omnium 1-jugis), foliohs 43-51 (nee
7O-80)-jugis, corolla 3(nee 4)-mera androecioque
16-26(nec 36-40)-mera diversa. — BRAZIL. Minas
Gerais: 13 k m n.-w. of Congonhas do Norte, camino
a Gouveia, ±18°41'S, 43°42'W, 900 m, 13 Feb 1991
(fl), M. M. Arbo (with Menezes, Schinini & Campos)
5005. — Holotypus, NY. Fig. 26
Slender, functionally herbaceous subshrub with
erect simple, sparingly foliate stems 3-4 dm, glabrous
except for traces of pubemlence on some lf-stks and
for thinly granular peduncles, the firm plane, ventrally
lustrous lfts glabrous, minutely resin-spotted dorsally,
the peduncles fasciculate by 3-6 at \-A furthest, leaf
less nodes, together forming a depauperate, shortly
exserted panicle; phyllotaxy distichous. Stipules
green, linear-subulate or narrowly ovate 1.5-3 x
0.3-0.8 m m , deciduous. Lf-formula (i-)ii/43-51; lf-
stks 4-20 m m , the petiole 4-16 m m , at middle 0.7-0.8
m m diam, the ventral sulcus open, deep, the one inter
pinnal segment, when present, ±8-11 m m ; rachises of
further (or only) pinna-pair 5-8.5 cm, the longer
interfoholar segments ±1-1.5 m m ; lft-pulvinules
0.2-0.3 x 0.4-0.5 m m , wrinkled; lfts equilong except
at very ends of rachis, the blades linear-oblong or
lanceolate from shortly auriculate base, obtuse apicu
late or triangular-acute, those near mid-rachis
(4.5-)5-7.7 x 1.2-1.6 m m , 3.8-5.8 times as long as
wide; midrib forwardly displaced to divide blade
1:1.5-2, the one posterior primary nerve either ob
scure or produced nearly to mid-blade, both faces
172 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
FiG. 26. Calliandra iligna Bameby.
of 1ft wrinkled when dry. Peduncles ±1.5-4 cm,
ebracteate; capitula 5-9-fld, the receptacle depressed-
hemispherical ±1.3-2 m m diam; bracts submembra-
nous narrow-ovate, <1 m m , deciduous; fls subsessile,
homomorphic, the perianth glabrous, thinly resinous,
reddish; calyx campanulate 3-4-merous, 1.2-1.5 m m ,
the deltate teeth 0.3-0.5 m m ; corolla 3-merous 5 m m ,
the ovate lobes 1.4 m m ; androecium 16-26-merous,
red or reddish, ±22 m m , the tube 3.5 m m , scarcely
thickened; ovary at anthesis glabrous. Pod unknown.
About rocks, in sandy soil of campo mpestre, 900
m, known only from the type-locality near the crest of
Sa. do Espinhaco, between Congonhas do Norte and
Gouveia (18°40'S) in centr. Minas Gerais. — Fl.
II—III(—?).
Calliandra iligna (of Congonhas, or botanical Ilex)
is closely related only to the almost sympatric C. san
tosiana but is a more slender plant with simple stems,
pinnae of most leaves two pairs, fewer leaflets per
pinna, and a somewhat smaller 3-4-merous perianth
enclosing a simplified androecium. Separately the
differential characters are not strong, but when com
bined in one plant cannot be passed over as individ
ual variations. Its specific status needs, however, the
tests of field observation and more collections.
111. Calliandra paterna Bameby, sp. nov, habitu vir-
gato, stipuhs ovato-cordatis palmatim plurinerviis,
pinnis paucis (1-2-jugis) elongatis necnon perianthio
elongato 7.5-10 m m longo ultra medium piloso C.
renvoizeanae proxime affinis, sed ab ea foliorum ma-
jorum foliolis 32-39(nec fere 60)-jugis 10-13.5 (nee
5-7) m m usque longis palmatim 4—5 (nee incon-
spicue l)-nerviis diversa. — BRAZIL. Bahia: mun.
de Palmeiras, Morro do Pai Inacio, ±12°30/S,
41°27W, 19 Dec 1981 (fl), G. P. Lewis & al. 880. —
Holotypus, C E P E C 30069; isotypi, K (2 sheets),
M O , NY.
Virgately erect shrub 6-10 dm, the simple or dis
tally few-branched stems, the ventral face of lf-axes,
and the peduncles thinly hirsute with subhorizontal,
shining white hairs to ±1-1.5 m m , the loosely imbri
cate lfts bicolored, lustrous olivaceous above, paler
and minutely resin-spotted beneath, the capitula
1—2(—3) together in the axils of 2-3 distal primary lvs
and thereafter shortly pseudoracemose; phyllotaxy
distichous. Stipules erect, ovate from shallowly cor
date base, ±6-10 x 3.5-7 m m , palmately 11-21-
nerved from point of attachment, tardily deciduous.
Lf-formula (i-)ii/32-39; lf-stks l-3(-3.5) cm, the
petiole 6-22 m m , the one interpinnal segment 6-20
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 173
m m ; pinnae equilong or the distal pair a little longer,
the rachis of these ±8-11 (-13) cm, the longer inter
foholar segments 2^4.5 m m ; lft-pulvinules 0.3-0.7 x
1 m m ; lfts little graduated, narrowly oblong-elliptic
from shortly obtusely auriculate base, obtuse, the
longer ones 10-13.5 x 2.7-4.7 m m , 2.7^.7 times as
long as wide; venation palmate, the midrib displaced
to divide blade ±1:2, the inner posterior primary
nerve incurved-ascending far beyond mid-blade, the
secondary venules ascending at narrow angles, ran
domly anastomosing, the whole blade when young
appearing subvertically striate, the venation nearly
immersed in age. Peduncles 2.5-5.5 cm, ebracteate;
capitula 5-9-fld, the receptacle 2-3 x 3-4.5 m m , but
sometimes one fl displaced onto peduncle; bracts
ovate or lance-ovate 3-6.5 m m , early dry deciduous;
fls subsessile homomorphic, the broad solid pedicel
±0.7 x 1.4-2.4 m m ; perianth "yellow, shiny, ± sticky,"
the calyx 4-, the corolla 3-4-merous, both glabrous in
lower half, thinly white-pilose distally, the calyx-
lobes striately subparallel-nerved, the corolla only
faintly so; calyx 5.5-7 m m , the lanceolate teeth 3.4—5
m m ; corolla ±8.5-10 m m , the ovate lobes ±3 m m ; an
droecium (of 2 fls examined) 86-96-merous, 28-32
m m , the thickened stemonozone 1.7-2 m m , the tube
8-10 m m , the tassel white; no intrastaminal disc;
ovary at anthesis glabrous. Pods (few seen) ±10 x 1.1
cm, the dilated sutural ribs in dorsal view 2+ m m
wide, the plane recessed valves reticulately venulose,
resinously papillate, and together with the margins
densely softly pilose overall; seeds unknown.
In campo mpestre, on granite bedrock, ±1000 m,
known only from the type-locality near 12°30'S in
Sa. do Sincora, upland interior Bahia, Brazil. — Fl.
XII-I, VI; fr. V-VI(-?).
Calliandra paterna, now known from four collec
tions from Morro do Pai Inacio (whence the epithet),
is compared in the diagnosis to C. renvoizeana which
also has only one or two pairs of pinnae per leaf, but
differs in leaflets about twice as many, scarcely half
as large, and weakly one-nerved. The also related C.
debilis may be recognized by its smaller calyx (±4,
not 5.5-7 m m ) and glabrous corolla.
112. Calliandra ganevii Bameby, sp. nov, cum C.
paterna habitu, indumento e pilis patulis lucidis
constanti, stipulisque ovatis majusculis congrua et
ei manifeste affinis, sed foliorum petiolo brevissimo
vix 6 (nee 6-22) m m longo, foliolis cujusque pin
nae 11-13 (nee 32-39)-jugis, perianthii extus
subevenii (nee tenuiter multstriati) paullo longioris
calyce 7.5-10 (nee 5.5-7) m m corollaque 12-12.5
(nee 8.5-10) m m longis diversa. — BRAZIL.
Bahia, mun. de Piata: in moist campo mpestre at
1430 m, estrada Piata-Gerais da Tromba, banks of
rio de Contas, 14 M a y 1992 (fl), Wilson Ganev 294.
— Holotypus, H U E F S 10737.
Virgate subshrub ±5 dm with simple stem and sub
sessile, relatively ample but simple lvs, the stems dis
tally and the peduncles pilose with erect straight
lustrous white hairs to 1.2-1.8 m m mixed with
remotely scattered, minute granular trichomes, the
plane discolorous chartaceous lfts facially glabrous,
sometimes remotely microciliolate, the few-fld capit
ula solitary on long peduncles axillary to distal lvs.
Stipules ovate acute from shallowly cordate base, foli
aceous ±7-9 x 4.5-6 m m , apparently persistent. Lf-
formula i-ii/11-17; petiole ±3-6 x 1.5 m m , the one
interpinnal segment, when present, to 13 m m ; rachis
of pinnae 4.5-6.5 cm, the longer interfoholar seg
ments 4-6 m m ; lft-pulvinules 0.4-0.6 x 0.8-1.2 m m ,
discolored and cross-wrinkled when dry; lfts little
graduated, the blades oblong from shallowly inequi
laterally cordate base, depressed-deltate-apiculate, the
longer ones 13-16 x 3-7 m m , 2.1-4.6 times as long as
wide; venation of 4—6 primary nerves, the straight
midrib displaced to divide blade ±1:1.5, weakly 1-2-
branched beyond middle, the inner posterior primary
nerve incurved-ascending far beyond mid-blade, the
outer ones weak and short, these all finely prominu
lous on lower face of blade, immersed or almost so
above, the ventral face minutely undulate-venulose.
Peduncles 5.5-12.5 cm, ebracteate; capitula 5—8-fld,
the hemispherical receptacle ±1.5-3.5 m m ; bracts
thinly herbaceous, lance-elliptic ±4—6.5 m m , not read
ily deciduous; fls homomorphic; pedicels 0.7-1 x 1.2-
1.9 m m ; perianth 4—5-merous, the calyx and corolla
each pilose distally, glabrous proximally; calyx
7.5-10 m m , the tube 3 x 3.5—4 m m , faintly obtusely
4—5-ribbed, the hnear-lanceolate teeth unequal, the
longest 4.5-6.5 m m ; corolla 12-12.5 m m , the ovate
lobes ±3.5 m m ; androecium ±64—106-merous, 32-
43 m m , the coriaceous stemonozone 1.8-2 m m , the
tube 12-16 m m , the tassel white mbescent, several
sterile filaments separating from tube below its ori
fice; intrastaminal nectary 0; ovary at anthesis
glabrous, but densely pubemlent after fertilization.
Pod unknown.
In moist campo rupestre near 1430-1500 m,
known only from Sa. da Tromba near the source of
rio de Contas (in municipios Piata and Abaira) and
174 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
from Mucuge within 12°58'-13°16'S, 41°20'-50'W,
in interior Bahia, Brazil. — FL V-VI(-?).
Calliandra ganevii belongs to a group of micro-
species characterized by one or two pairs of pinnae per
leaf, ovate stipules, lanceolate calyx-teeth, and rela
tively long perianth (or some combination of these
attributes) that includes C. paterna, C. renvoizeana,
and C. debilis. From all of these it differs in few
(11-17) pairs of leaflets per pinna.
113. Calliandra erubescens Renvoize, Kew Bull.
36: 79, fig. 3(32), 7C. 1981. — "Brazil, Bahia . . .
Morro de Chapeau [sic], Harley et al. 19227." —
Holotypus, CEPEC n.v; isotypi, K!, NY!.
C. erubescens sensu Lewis, 1987: 173 (exclus. Harley 20073 quae = C. luetzelburgii Harms); Stannard, 1995: 378 (exclus. Harley 26102, 25746, quae = C. semisepulta).
Shrubs 1.5-3 m with fuscous defoliate annotinous
and older branches, densely foliate distally, appearing
glabrous but the new stems, lf-axes and peduncles
thinly pubemlent with minute appressed white hairs
<0.15 m m mixed with discolored granular trichomes,
the fohage conspicuously bicolored, dark brown-
olivaceous and lustrous above, paler dull and some
times remotely granular-papillate beneath, the few-
fld capitula mostly fasciculate by 2-5 in the furthest
lf-axils and beyond these forming a compact, shortly
exserted pseudoraceme. Stipules linear or narrowly
lanceolate 0.6-3.5 x 0.2-0.5 mm, deciduous. Lf-
formula ii/5-9(-l 1), a few distal lvs conjugate-pinnate;
lf-stk of larger lvs 10-21 mm, the petiole including
fuscous pulvinus 2-5 x 0.8-1.6 mm, the one interpin
nal segment ±2-3 times as long, the ventral groove
bridged at insertion of lower pinna-pair; pinnae sub
equilong, the rachis of the further pair 2.5̂ 4.5 cm, the
longer interfoholar segments 4—7 mm; lft-pulvinules
0.3-0.55 x 0.7-1.2 mm, cross-wrinkled; lfts subac-
crescent distally or subequilong, the blades oblong or
oblong-elliptic from shortly broadly auriculate base,
broadly rounded at apex, those near mid-rachis 9-
14.5 x 3.5-6 m m , ±(1.7-)2-2.7 times as long as wide;
primary venation of 4-5 nerves from pulvinule, the
midrib forwardly displaced to divide blade 1:2-
2.5, weakly 1-2-branched beyond middle, the inner
posterior primary nerve produced nearly to blade's
apex, tertiary venulation random weak, the whole
venation finely prominulous dorsally, the upper face
of blades reticulately rugulose when dry. Peduncles
1.5-5 cm, ebracteate; capitula 4-7(-8)-fld, the hemi
spherical receptacle 1-2 x 2-3 mm; fls subsessile, the
drum-shaped discolored pedicel 0.6-0.7 x 1-2 mm;
perianth pinkish-white, nigrescent, glabrous except
for sometimes microscopically ciholate orifice of
calyx or for minute pubemlence of corolla-lobes, each
calyx-tooth faintly 1-nerved, the corolla externally
nerveless; calyx shallowly campanulate 1.4—1.7 x 1.9-
2.9 mm, the depressed-deltate teeth 0.3-0.5 mm;
corolla 5-6.2 mm, the ovate lobes 2-3 mm; androe
cium 24-44-merous, 2.4-3 cm, the stemonozone 1.1-2
mm, the tube 3-6 mm, the tassel opening white,
mbescent; intrastaminal nectary 0; ovary at anthesis
glabrous. Pods (few seen) in profile (3-)4—8 cm x 6-8
mm, finely minutely pilosulous overall, the recessed
valves in addition randomly granular; seeds not seen.
In campo mpestre, on sandstone, 800-1500 m,
endemic to the crest and upper slopes of Chapada
Diamantina, in lat. n°20'-13°40'S, interior upland
Bahia, Brazil. — Fl. II-VII.
This species is manifestly close to C. luetzelburgii
and C. hygrophila, but different from the first in hav
ing two (exceptionally three) pairs, not exactly one
pair, of pinnae, and from the second in five to nine,
not two to three pairs of leaflets per pinna. Nodulated
roots were collected by H. de Lima (no. 3883, K). A
population on Pico de Almas (CFCR 6895, K) has
relatively wide leaflets (to 8-11 m m ) and "deep red"
flowers, thereby resembling sympatric C. semi
sepulta, and may be varietally distinct.
114. Calliandra semisepulta Bameby, sp. nov, C.
erubescenti (sympatrica) ut videtur proxime affinis,
praesertim caulibus homotinis nanis herbaceis vix
1.5 dm longis e rhizomate assurgentibus (nee mox
fmticosis 1.5-3 m altis) diversa, ulterius stipuhs
latioribus 0.8-2 m m latis, foliolis majoribus ±15-25
x 7-12 (nee 9-16 x 3.5-7) m m usque, pedunculis
elongates (5-)6-14 (nee 1.5-5) cm longis, corolla
majori 7-9.5 (nee 5-6.2) m m longa, filamentisque
magis numerosis 44-64 (nee 22-44) ab initio rubris
(nee albidis rubescentibus) ab ea distans. —
BRAZIL. Bahia, mun. Rio de Contas: Pico de
Almas, 8 km n.-w. of the town, 1200 m, 10 Nov
1988 (fl), R. M. Harley (et al.) 26102. — Holoty
pus, K; phototypus s.n, NY; isotypus, CEPEC.
Macrophyll, functionally herbaceous subshrubs
with slender 3^-lvd stems ±8-15 cm, diffuse and
incurved-ascending from oblique rhizomes to ±6 m m
diam, glabrous except for microciliolate lf-axes, the
ample, stiffly papery lfts moderately bicolored, a little
darker green and more lustrous ventrally, sometimes
remotely granular dorsally, the long peduncles erect
from one or more lf-axils and thence shortly pseudo
racemose. Stipules herbaceous, lanceolate or narrowly
ovate 2-4.5 m m , mostly persistent. Lf-formula
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 175
(i-)ii(-iii)/(4-)5-8; lf-stks 2-5.5 cm, the petiole 1-2.5
cm, at middle 1.4—2 m m diam, the one or the longer of
2 interpinnal segments 1-2.8 cm, the open ventral
groove strongly bridged at insertion of pinnae; pinnae
not or scarcely accrescent distally, the rachis of distal
pair (2-)3.5-6.5 cm, the longer interfoholar segments
(5-)7-17 m m ; lft-pulvinules 0.8-1.2 x 1-1.9 m m ,
deeply wrinkled; lfts subequilong or distally a little de
crescent, oblong or oblong-obovate from obhquely
subtmncate or incipiently cordate base, broadly obtuse
apiculate, the larger ones (13-) 15-25 x 7-12 m m ,
1.8-2.2 times as long as wide; venation prominulous
on both faces of lfts, the primary nerves 5-7, the mid
rib only shghtiy displaced from mid-blade, giving rise
on each side to 2-4 strong, narrowly ascending second
aries, the inner posterior primary nerve produced to or
beyond the third quarter of blade, the close tertiary
venulation sub vertically sinuous. Peduncles solitary or
fasciculate by 2-3, at anthesis (5-)6-14 cm, sometimes
1-bracteate close under the 7-12-fld capitulum; bracts
±1-2.5 m m , tardily deciduous; pedicels stout, low-
conic 0.5-1 x 0.7 x 2 m m ; fls homomorphic or almost
so, the perianth submembranous, glabrous microgran-
ular or the calyx minutely ciholate, the calyx 5-6- and
the corolla 3-5-merous, both greenish red-tinged;
calyx campanulate 2-4.2 m m , the deltate-triangular
teeth 0.6-1.9 m m ; corolla 7.2-9.5 m m , the lobes 3-5
m m ; androecium red overall, 44-64-merous, 26-29
m m , the stemonozone 1.2-1.4 m m , ± thickened inter
nally, the tube 6-6.5 m m ; intrastaminal nectary 0;
ovary at anthesis glabrous. Pod unknown.
In sandy soil about arenitic outcrops, 1160-1500
m, sympatric with C. erubescens on Pico das Almas
at points 8-10 k m n.-w. of the town of Rio de Contas
and Catoles de Cima (near 13°18'-32'S, 41°52'-
54'W) in interior Bahia, Brazil. — Fl. X-XII.
Calliandra semisepulta is narrowly sympatric with
C. erubescens and is presumably derived from it by
modification of growth-form, its short assurgent, func
tionally herbaceous stems being in marked contrast to
the frutescent framework of adult C. erubescens.
Smaller differences between these two species in stip
ules, leaflets, peduncles, and individual flowers are
noted in the Latin diagnosis. A perplexing specimen
(Lewis in C F C R 6904, K ) from the common habitat of
C. erubescens and C. semisepulta, which has the ample
fohage of the latter but stems more than 3.5 d m tall
said to be woody at base, is possibly a hybrid
derivative.
115. Calliandra germana Bameby, sp. nov, C. luet
zelburgii quam maxime affinis sed foliolis subduplo
latioribus (±9-20, nee 3-6 m m usque latis) dorso
simulac perianthio dense mfo-granulosis diversa; a
distantiori C. erubescenti imprimis cujusque folii
pinnis exacte 1 (nee 2)-jugis, foliolis dorso granulo
sis, necnon androecio 24—26-, nee 42-44-mero
recedens. — B R A Z I L . Bahia, mun. Catoles: estrada
Catoles-Barra, 1200 m, 16 Jun 1992 (fl), Wilson
Ganev 506. — Holotypus, HUEFS 10949.
Shrubs attaining 2 m with terete fuscous branches,
appearing glabrous but the lf-axes and dorsal face of
ample chartaceous lfts densely minutely reddish-
brown-granular, the bicolored lfts glossy on upper
face, paler dull beneath, the few-fld capitula arising
1-3 together from axils of diminished distal lvs,
forming a dense short panicle of pseudoracemes.
Stipules caducous (few seen), narrowly ovate ±1.5 x
0.7 m m , not striate. Lf-formula i/5-8; lf-stks 4.5-8 x
0.9-2 m m , shallowly grooved ventrally; rachis of
longer pinnae 3.5-5.5 cm, the longer interfoholar
segments 7-14 m m ; lft-pulvinules 0.6-1.1 x 0.9-1.3
m m , sharply cross-wrinkled; lfts little graduated, the
blades oblong or oblong-obovate from shallowly cor
date base, broadly obtuse or often shallowly emar-
ginate, the longer ones (in lvs below inflorescence)
14—24 x 7.5-20 m m , 1.2-1.8 times as long as wide;
venation of 5-8 nerves from pulvinule, the midrib
displaced to divide blade ±1:1.5-2, 2-3-branched on
anterior side near and above middle, the inner poste
rior primary nerve incurved-ascending nearly to
blade's apex, the outer ones progressively shorter. Pe
duncles (1.4—)2-3.5 cm, ebracteate; capitula 5-8-fld,
the receptacle ±1-1.5 m m ; bracts <1 m m , caducous;
fls nearly homomorphic, the 4-merous perianth palhd
or pink-tinged, thinly granular but otherwise glabrous
except for microciliolate orifice of calyx; pedicels
±0.2 x 0.8 m m ; calyx campanulate 2.4-2.5 x 2-2.4
m m , faintly 4-nerved, the depressed-deltate teeth
0.25-0.4 m m ; corolla 6.4-6.6 m m , the lobes 2-2.1 x
1.6 m m ; androecium 24— 26-merous, ±23 m m , the
camosulous stemonozone 1.6-1.8 m m , the tube 6-6.5
m m , the tassel at first white, pink in age; intrastami
nal nectary 0; ovary at anthesis glabrous. Pod seen
only in annotinous fragments, the ribs and valves
densely minutely pubemlent overall; seeds unknown.
In campo mpestre at ±1200 m, known only by 2
collections from the headwaters of rio de Contas near
13°15'S, 41°50'W, in upland interior Bahia, Brazil.
— FL VI-VII(-?).
Calliandra germana is manifestly close to C. luet
zelburgii but distinguished from it by notably broader,
dorsally granular leaflets, and by granular perianth.
176 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
116. Calliandra luetzelburgii Harms, Repert. Sp.
Nov. Regni Veg. 17: 89. 1921. — "Brasilien: Bahia
(Ph. v. Lutzelburg in Herb. Munchen, n. 55. 1914)."
— Holotypus, M, not found at M in 1992, at K in
1994!; presumed isotypus, +B = F Neg. 12481.
C. virgata sensu Bentham, 1876, minori ex parte, quoad plantam bahianam Martiusianam.
C. luetzelburgii sensu Renvoize, 1981: fig. 3(30); Lewis,
1987: 175; Stannard, 1995: 380, fig. 23A-D.
Diffuse or bushy shrubs 4-15 dm with fuscous de
fohate annotinous stems and densely foliate new
growth, except for minutely ciholate calyx glabrous
throughout, the lvs bicolored, lustrous dark brown-
olivaceous above, dull and paler beneath, the few-fld
capitula borne solitary or 2-4- together in a few distal
lf-axils, and beyond at efoliate nodes of a short,
scarcely exserted pseudoraceme. Stipules erect, lin
ear-lanceolate or hnear (1.5—)2—6 x 0.4-0.9 m m ,
faintly 1-nerved or externally nerveless, deciduous.
Lf-formula i/(5-)8-ll; lf-stks including pulvinus
4.5-12 x 0.8-1.3 m m ; rachis of longer pinnae 3.5-6
cm, the longer interfoholar segments 3-6 m m ; lft-
pulvinules in dorsal view 0.6-1 x 0.7-0.9 m m ; lfts
scarcely graduated, in outline linear-elliptic from
inequilateral but scarcely auriculate base, obtuse or
obtusely deltate at apex, the larger ones 12-25 x 2.8-
5.7 m m , (3.9-)4—5.3 times as long as wide; primary
venation of 4—5 nerves from pulvinule, the nearly
straight, almost simple midrib displaced to divide
blade 1:1.3-2.3, the inner posterior primary nerve
expiring (faintly anastomosing) well beyond mid-
blade, the outer posterior and one anterior one much
shorter, secondary venules few and weak, the whole
venation finely prominulous only dorsally. Peduncles
13-30 m m , ebracteate; capitula 4-6(-7)-fld, the sub
tmncate receptacle 1.5-2 m m wide; bracts ±1-1.5
m m , caducous; fls subsessile, homomorphic; pedicels
drum-shaped or almost discoid 0.6-0.9 x 1.2-1.6
m m ; calyx shallowly campanulate 1.5-1.7 x ±2 m m ,
the depressed deltate teeth 0.5-0.6 m m ; corolla 6-6.6
m m , the ovate, often unequal lobes 1-3 m m ; androe
cium 28-30-merous, 23-25 m m , the stemonozone
1.2-1.4 m m , the tube 5.5-6 m m , the tassel opening
white, rubescent. Pods (httle known) in profile ±8 c m
x 7.5 m m , the sumral ribs in dorsal view ±2.5 m m
wide, minutely pubemlent, the recessed valves
brown-papillate with short obese trichomes, subgran-
ular, obscurely venulose; seeds unknown.
In campo mpestre, 980-1600 m, locally plentiful
on the e. slope of Pico de Almas in the highlands of
interior Bahia. — Fl. III-VII.
Calliandra luetzelburgii is close to C. erubescens
and to C. hygrophila. It resembles C. erubescens in
stature, but differs in conjugate pinnae and somewhat
more numerous (8-11, not 5-8) pairs of narrower
leaflets 4-5, not 2.1-2.7, times as long as wide. Cal
liandra hygrophila differs from both in fewer (2-3)
leaflet pairs, dwarf stature, and red tassel of filaments.
117. Calliandra hygrophila Mackinder & Lewis,
K e w Bull. 45: 681, fig. 1. 1990. — "Brazil, Bahia .
. . Serra do Sincora, 3 k m S W of Mucuge ... 27
March, 1980, R. M. Harley & al. 21055." — Holo
typus, CEPEC!; isotypi, K!, M!, M O ! , NY!.
Calliandra sp. C. Mackinder & Lewis, 1986: 115.
Shrublets with procumbent, incurved-ascending,
few-branched, often abmptly flexuous stems forming
depressed thickets to 1.5-3.5 d m tall and 1 m diam,
glabrous throughout, the relatively few and ample
coriaceous lfts olivaceous sublustrous above, some
times obscurely reddish-papillate dorsally, the few-
fld capitula solitary in distal lf-axils, immersed in or
barely exserted from foliage. Stipules caducous (few
seen), narrowly lance-oblong ±1.5 x 0.5 m m , perhaps
lacking at some nodes. Lf-formula i/2—3; lf-stks in
cluding discolored pulvinus 1.5-4.5 x 1-1.4 m m ;
rachis of longer pinnae 1.5-3.5 cm, the one or the
longer of two interfoholar segments 8-16 m m ; lft-
pulvinules 0.3-0.8 x 1.1-1.8 m m , cross-wrinkled; lfts
accrescent distally, the blades obovate or elliptic-
oblanceolate from inequilateral, postically rounded,
antically cuneate base, broadly obtuse and sometimes
obscurely apiculate, the distal pair 16-29 x 6-15 m m ,
1.8-2.6 times as long as wide; venation palmate, the
scarcely excentric midrib simple or 1-2-branched on
either side, the inner posterior primary nerve
incurved-ascending nearly to blade's apex, the one
anterior and 2-3 posterior ones progressively shorter,
unbranched, all finely prominulous dorsally, only
faintly so or immersed above. Peduncles 5-13 m m ,
ebracteate; capitula 4-6-fld, the receptacle scarcely 1
m m ; bracts subulate ±0.5 m m , caducous; fls sessile
(but calyx thickened at base, as seen in section),
homomorphic; perianth reddish, glabrous except for
microscopically granular-ciholate sinuses of calyx,
the corolla thinly minutely reddish-papillate; calyx
campanulate 1.9-2.3 x 1.6-1.7 m m , the tube faintly
5-nerved beyond middle, the depressed-deltate teeth
0.3-0.7 m m ; corolla tubular 8 m m , slightly ampliate
at the limb, the often unequal, ovate teeth to 1.8-2.5
m m ; androecium 16-24-merous, 21-24 m m , the ste
monozone 1-1.9 m m , the tube 4.5 m m , the tassel
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 177
bright red. Pods narrowly oblanceolate in profile, when
well fertilized ±5-6.5 x 0.55-0.75 cm, 3-4-seeded, the
sutural ribs ±2 m m wide in dorsal view, the recessed
valves hght brown, remotely venulose, glabrous over
all; seeds reportedly "4 x 2 mm," those seen on iso
typus (NY) ±7 x 4.5 m m , the testa hght brown dark-
speckled, the U-shaped pleurogram 5 x 2 m m .
O n alluvial sands and about rocks at river's edge,
800-1280 m, very local, known only from mun. M u
cuge in the upper Paraguacu basin near 13°S in inte
rior Bahia, Brazil. — Fl. XII-IV.
Calliandra hygrophila may be visuahzed as a re
duced and specialized derivative of C. erubescens or
some c o m m o n ancestor, notable for dwarf, thicket-
forming habit of growth and leaves reduced to a for
mula of i/2—3.
118. Calliandra houstoniana (Miller) Standley,
Contr. U.S. Natl. Herb. 23: 386. 1922. — T y p u s sub
var. houstoniana infra indicatur.
Amply microphylhdious (sub)shrubs and bushy
trees flowering when 1.5—6(—8) m tall, either virgately
single-stemmed or branching upward, the trunk rarely
attaining 1(—1.4) d m diam, the terete branches and the
lf-axes either glabrous or thinly to densely pilose-
pilosulous with erect, ascending or appressed, either
white, or sordid-gray, or brown to bronze or black
hairs mostly 0.2-1 m m , the lvs bicolored, the lfts
green and commonly glabrous above, glabrous to
appressed-pilosulous beneath, appressed-ciliolate, the
inflorescence a stout and narrow or a shorter pyrami
dal, efohate or at base few-lvd pseudoraceme of either
condensed or umbelliform capitula of relatively large
fls terminal to the homotinous stem or to occasional
lateral branches; phyllotaxy distichous. Stipules thinly
herbaceous, those associated with primary lvs lan
ceolate or narrowly ovate ±3-9 x 1-2.5 m m , early
caducous, absent from many specimens, those of axil
lary buds similar but smaller, loosely imbricate. Lf-
formula (v-)vi-xxiv(-xxxi)/24-65(-69); lf-stk of
major lvs (4.5-)6-20(-21) cm, the petiole including
pulvinus 8-35(^0) m m , at middle 0.8-1.7 m m diam,
the longer interpinnal segments (4.5-)6-15(-21) m m ,
the ventral groove continuous between pinnae or
obscurely bridged; pinnae either decrescent near each
end of lf-stk or only proximally, or randomly short and
long, the rachis of longer ones (1.8-)2-10(-14) cm,
the longer interfoholar segments (0.4-)0.6-2 m m ;
lft-pulvinules 0.2-0.45 x 0.35-0.6 m m , not wrinkled;
lft-blades linear or narrowly linear-lanceolate from
either tmncate or subacutely to obtusely auriculate
base, acute or acuminate or obtuse mucronulate, either
straight or ± incurved distally, less often sigmoid-fal-
cate, the larger ones (2.5-)3-10(-ll) x
(0.4-)0.6-2.4(-2.7) m m , (3.2-)4-8(-9) times as long
as wide, all plane or low-convex on upper face; midrib
either subcentric or displaced to divide blade ±1:2,
either simple or (in broadest blades) weakly branched
on one or both sides, posterior primary nerves com
monly 0, if 1-2 then very short and weak. Axis of
pseudoracemes (0.3-)0.6-3 dm; stipuliform bracts at
each node of pseudoraceme ovate 2-A m m , early
papery caducous; peduncles fasciculate by 2-6(-7), at
anthesis 2—13(—16) x 0.7-1.7 m m , in fmit sometimes
to 2.6 m m diam, all ebracteate; floral bracts ovate or
obtusely deltate 1.2-2.4 m m , either discrete or shortly
connate at base, caducous or sometimes persistent
through anthesis; pedicels at anthesis 0.7-8 x 0.35-
2.5 m m , either terete or when short turbinate and as
broad as long, in fmit 1-3.2 m m diam; fls of each
capitulum homomorphic or nearly so, the perianth
5-merous (calyx sometimes 6-7-merous), the corolla
firm or subcoriaceous, the calyx and corolla alike
glabrous to micropuberulent, strigulose-puberulent, or
densely pilosulous, the indumentum white to sordid,
brown, bronze, black, or of mixed colors; calyx
campanulate or shallowly campanulate-patelhform
(l-)1.2-3.6(4.5) x 2.2-54(-6.5) m m , bluntly 5-
angulate and often in addition weakly several-nerved,
the teeth deltate, depressed-deltate, or narrowly ovate
to triangular, obtuse, 0.3-1.6(-3) m m ; corolla
(6.5-)7-12(-13) m m , the lobes (3-)4-7(-8) m m ,
toward tip 0.2-0.6 m m thick in section, often in age
separating down to rim of stemonozone; androecium
36-76(-100)-merous, 4-7(-8) cm, the thickened
stemonozone 2-3.5 m m , the tube ±2-4(-5) m m , the
tassel usually crimson-scarlet throughout, sometimes
pale crimson, sometimes bicolored, then pink-crimson
only beyond middle; intrastaminal nectaries not dif
ferentiated, the smooth inner wall of the stemonozone
copiously nectariferous. Pods erect-ascending, in pro
file mostly (6-)7-12(-16) x (0.9-)1.2-2(-2.3) cm, in
var. colomasensis only ±3.5-5 x 0.5-0.6 cm, the
sutural ribs in dorsal view mostly 2.5^4.5 m m wide,
the ribs and valves ahke glabrous to pubemlent or
densely velutinous to pilose with brown, mixed brown
and white, or largely white hairs to 0.3-1.2 m m , the
valves weakly cross-venulose; ovules 8-11 but seeds
seldom more than 8 per pod, the seeds in broad view
elliptic-obovate or broadly obovate ±7 x 4—7 m m , the
smooth hard testa light brown, often dark-mottled or
-speckled, pleurogrammic.
The definition of C. houstoniana embodied in the
foregoing description breaks with a long tradition of
178 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
recognizing three, four, or several specific taxa in the
group. Material available for study is now plentiful
and lately has been enriched by intensive collections
of D. J. Macqueen (OXF, generously shared with N Y )
in tropical Mexico and Central America. Since Brit
ton and Rose's summary in 1928, the close alhes of
C. houstoniana have been critically revised in their
entirety only by Macqueen and Hernandez (1997),
although McVaugh (1987) and Hernandez (1991)
have recently presented thoughtful, even though incon
clusive, analyses of variation within and between
traditionally accepted taxa. The latter can still be rec
ognized readily enough, but no longer as morpholog
ically discrete entities, and are reevaluated here as
varieties interconnected by intermediate forms. The
main divisions of C. houstoniana sens. lat. are four,
characterized as follows:
a. C. houstoniana sens, str.: Foliage relatively insen
sitive; leaflets distally incurved or falcate, highly
lustrous adaxially, and the midrib decidedly dis
placed from mid-blade; vesture of inflorescence
wholly or in part brown-bronze, fuscous, or
black; pedicels relatively short and stout, the
pseudoraceme consequently narrow and dense;
and androecium uniformly red.
b. C. grandiflora, also known as C. anomala: Foliage
more sensitive; leaflets straight or almost so,
either acute or obtuse, with subcentric midrib;
pinnae relatively short and numerous; vesture of
inflorescence white through shades of gray,
brown, and bronze to black, the color transitional
in a northwest-southeast cline between Sinaloa,
Mexico, and Honduras; and pedicels either
longer or, if no longer, more slender than in the
preceding, the pseudoraceme therefore more
open; androecium of C. houstoniana sens. str.
c. C. calothyrsus: Foliage, lfts, and inflorescence of
C. grandiflora, but perianth glabrous.
d. C. acapulcensis: Close to the preceding except for
fewer pinnae and wider leaflets, pubemlent peri
anth, and often bicolored androecium, the tassel
rose-carmine in lower half, pallid proximally.
Each of the differential characters mentioned in
this broad analysis is now known to vary indepen
dently of any other. Furthermore, each of the four ide
alized but only inexactly definable taxa varies inter
nally in density, length, and orientation of vesture, in
leaf-formula, in size and pubescence of leaflets, in
size of perianth parts, in outline of flower-buds (from
plumply to slenderly pyriform), and in depth of the
sinuses between mature corolla-lobes. The number of
potential character-syndromes seems indefinite, and
those known from specimens are much more numer
ous than the available names. Most of the segregates
described at the specific level by Britton and Rose
(1928) are minor variants. Beside C. acapulcensis,
only Anneslia colomasensis, notable for extremely
short pinnae and small narrow pod, is provisionally
preserved here as an independent taxon.
Particular notice must be taken that the North
American species numbered 118 to 122 (inch) in this
account are coextensive with the seven species
recently revised by Macqueen and H. Hernandez
(1997) under the title Calliandra ser. Racemosae.
Contemporary but independent, these two revisions of
one set of closely related taxa were undertaken and
carried out in different contexts, mine within the
framework of whole genus, the other more intensely
focused on an assemblage of species of proven or
potential value in tropical agroforestry, especially as
sources of fuel. The synonymy and the definitions of
taxa for the most part m n parallel. Such conflict as
exists is due to different emphasis on, or different
interpretation of, a morphological polymorphism
almost universal in the group, comphcated by local
hybridization between sympatric populations. The
inconvenience of two competing taxonomies pro
posed almost simultaneously for one group of species
is regrettable. In practice, that of Macqueen is strongly
recommended for the taxa of economic importance.
Insofar as m y opinions presented herein are signifi
cantly different, they should be viewed in the light of
a full generic survey. The classification of ser. Race
mose by Macqueen and Hernandez is supported by
intensive fieldwork throughout lowland and tropical
montane Mexico and Central America and is influ
enced by a study of seedling morphology. It is a plea
sure and an agreeable duty to acknowledge Dr. Mac-
queen's gift to N Y of a substantial sample of his
carefully selected and beautifully prepared specimens.
Key to the varieties of C. houstoniana
1. Lfts either obtuse or acute, but the midrib
straight or almost so, and simple, the blade ±
bicolored but not highly lustrous on upper face;
perianth either pubescent or glabrous and
pedicels either stout or slender, 1-8 x (0.35-)
0.4-1.2 m m (sometimes wider in fruit); lvs
relatively sensitive to shock, often ± contracted in dried specimens.
2. Perianth pubescent, often densely so, with
either appressed or variously spreading-
ascending hairs, these either white, gray, sordid,
brown, bronze, or black (often of mixed color).
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 179
3. Larger lfts (0.5-)0.6-1.2(-l.4) m m wide, 1 -nerved, the costa simple; pinnae
of larger lvs 11 pairs upward; filament-tassel red throughout.
4. Pods(6-)7-14x0.9-1.6(-2.3)cm; widespread 118a. var. anomala
4. Pods 3.5-5 x 0.5-0.6 cm; known only
from Sinaloa. . . . 118b. var. colomasensis 3. Larger lfts 1.5-2.4 m m wide, 1-nerved
from pulvinule but the midrib at least
faintly pinnately branched; pinnae of larger lvs 6-9(-10) pairs; filaments
bicolored, the tassel pallid in lower
half, red distally 118c. var. acapulcensis
2. Perianth glabrous (in s.-w. Mexico
sometimes micropuberulent near tip of corolla); lvs and pedicels of var.
anomala 118d. var. calothyrsus 1. Lfts sharply acute and the midrib at apex
turned upward, the blade lustrous on upper
face, paler dull beneath; perianth densely
pubescent, the color of hairs variable but
some brownish, bronze, or black, exceptionally
all or most sordid; pedicels stout, 0.7-4.5
(-5) x 0.7-2.5 mm; lvs relatively insensitive
to shock, often fully expanded in dried
specimens 118e. var. houstoniana
118a. Calliandra houstoniana (Miller) Standley var.
anomala (Kunth) Bameby, comb, nov, based im
mediately on autonym Inga anomala var. anomala
generated by /. anomala var. pedicellata de Can
dolle, Prodr. 2: 442. 1825, and ultimately on Inga
anomala Kunth, Mimoses 70, pi. 22. 1820. — ". . .
in regno Mexicano, prope urbem Pascuaro et in
declivitate montis ignivomi Jorullo [Michoacan]."
— Holotypus, P-HBK!, photo, NY!. — Calliandra
kunthii Bentham, J. Bot. (Hooker) 2: 139. 1840; &
London J. Bot. 3: 112. 1844, nom. illegit. Callian
dra anomala Macbride, Contrib. Gray Herb., n. ser.
59: 4. 1919. Anneslia anomala Britton & Rose, N.
Amer. FL 23: 73. 1928. — Equated with C. grandi
flora (L'Heritier) Bentham by Riley, Bull. Misc.
Inform. 1923: 397. 1923; Standley, Contrib. Natl.
Herb. 23: 1658. 1926.
Mimosa grandiflora L'Heritier, Sert. angl., fasc. 1. 30, "t. 42
[but this not published and the original lacking in L'Heri-
tier's papers at G-DC.]." 1789. — Described from plants
grown in England, introduced from "India orientalis"
according to Aiton, Hort. kew. 3: 441. 1789, "by Mrs
Morton . . ca 1769." — No typus found at BM, an old
specimen cultivated at Kew, from herbarium of Bishop
Goodenough, was identified by Verdcourt (annot. 1984)
with L'Heritier's plant and is recommended as neotypus,
replacing that proposed by Macqueen and H. Hernandez,
1997: 21. —Acacia grandiflora Willdenow, Sp. PL 4:1074.
1806. Calliandra grandiflora Bentham, J. Bot. (Hooker) 2:
139. 1840; & London J. Bot. 3: 111. 1844. Anneslia gran
diflora Britton & Rose, N. Amer. Fl. 23: 70. 1928.
Inga anomala var. (ft) pedicellata de Candolle, Prodr. 2:
442. 1825. — Based on crude figures of Sesse seen as
Caique des dessins 206 (NY, library!) and of Tlacoxiloxo-
chitl Hernandez, Nova PL Hist. 104, fig. 1, 2. 1651, these
here regarded as syntypi. — Referred by Britton & Rose,
1928: 71, with marks of interrogation, both to Anneslia
callistemon and to A. strigillosa; by Macqueen and H. M.
Hernandez, 1997: 21 to C. grandiflora.
Acacia callistemon Schlechtendal, Linnaea 12: 568. 1838.
— "Pr[ope] Reglam [Hidalgo, Mexico] (C. Ehrenberg),
Mexico (hb. Lehm[ann.])" — No typus seen. — Calliandra callistemon Bentham ex Jackson, Index Kew. 1: 385.
1893. C. anomala var. callistemon Macbride, Contrib.
Gray Herb., n. ser. 59: 4. 1919. — Anneslia callistemon
Britton & Rose, N. Amer. Fl. 23: 71. 1928. — Equated with C. grandiflora by Macqueen & H. M. Hernandez,
1977: 23.
Anneslia chihuahuana Britton & Rose, N. Amer. Fl. 23: 71.
1928. — "Southwestern Chihuahua, August-November 1885, [Edward] Palmer 266." — Holotypus, NY!; iso
typus, K!.
A. strigillosa Britton & Rose, N. Amer. Fl. 23: 71, syn.
dubio candolleano incluso. 1928. — "Type from Eugenio,
Sierra de la Cruz, Orizaba [Veracruz, Mexico], 1853, F.
Mueller 1594." — Holotypus, NY!. — Calliandra strig
illosa Standley ex Leavenworth, Amer. Midi. Naturalist 36:
143. 1946.
A. bella Britton & Rose, N. Amer. Fl. 23: 71.1928. — "[Mex
ico. Jalisco:] . . . between Bolanos and Guadalajara, Sep
tember 20. 1897, [J. N.] Rose 3036." — Holotypus, NY!;
isotypus, US!. — N o n Calliandra bella Bentham, 1844.
A. conzattiana Britton & Rose, N. Amer. Fl. 23: 72. 1928 —
"Type from Cerro San Antonio, Oaxaca ... June 26, 1906,
Conzatti 1424." — Holotypus, NY!. — Calliandra conzat
tiana Standley, Trop. Woods 34: 40. 1933.
A. albanensis Britton & Rose, N. Amer. Fl. 23: 72. 1928. — "Monte Alban, Oaxaca, December 29, 1895, [Caec. & Ed]
Seler 1734." — Holotypus, NY!.
A. albescens Britton & Rose, N. Amer. Fl. 23: 72. 1928-
"[Mexico.]. . extreme southern Durango, August 13, 1897, [J. N.]Rose 2254." — Holotypus, NY!; isotypus, US!.
A. pueblana Britton & Rose, N. Amer. Fl. 23: 73. 1928. —
"[Mexico.] vicinity of San Luis Tultitlanapa, Puebla, near
Oaxaca, June-August, 1908, [C. A.] Purpus 2666." — Holotypus, NY!.
A. rusbyi Britton & Rose, N. Amer. Fl. 23: 194. 1928. —
"Limon Mountain, Guerrero, [Mexico], July 2,1910, H. H.
[& Ruth] Rusby 22." — Holotypus, NY!; isotypus, NY!.
Calliandra leucothrix Standley, Field Mus. Nat. Hist., Bot.
Ser. 17: 366. 1938. — "Honduras . near El Achote, hills
above the plains of Siguatepeque, Dept. Comayagua, 1350
meters, July 26, 1936, T. C. Yuncker, R. F. Dawson & H. R.
Youse 6128." — Holotypus, F!; isotypus, NY!. — Non An
neslia leucotricha Britton & Rose, 1928.
C. anomala var. longipedicellata McVaugh, Fl. Novo-gah-
ciana5: 151. 1987. — "[Mexico.] Jal[isco] . . 11-12 mi S of Talpa, McVaugh 21323." — Holotypus, MICH!. — C.
longipedicellata Macqueen & Hernandez, Kew Bull. 52: 40, fig. 9, map 4.
Lf-formula (x-)xi-xxiv(-xxxi)/(21-)24-52(-55);
lf-stks (7-)8-20(-21) cm, the longer interpinnal seg
ments 3-12 m m ; rachis of longer pinnae (1.8-)2-4.6
180 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
(-5.5) cm; larger lfts (2.5-)3.2-7 x (0.4-)0.6-14
(-1.5) mm , (3.2-)3.7-5.5(-6) times as long as wide.
Inflorescence thinly to densely strigulose or pilosulous
with either appressed or spreading-ascending hairs,
those from Michoacan n.-w.-ward mostly palhd, from
Michoacan s.-e.-ward mostly or wholly yellowish,
brown, dusky, or black; peduncles 4—12(-16) m m ;
pedicels (l-)1.4-4.5 m m , or in W . Mexico (1.8-)2.5-8
m m , 0.35-1 m m diam at anthesis; calyx (1-) 1.2-2.8
(-3) x 2.2-4.2(-5) m m , the teeth 0.3-1.1 (-1.2) m m ;
corolla (6.5-)7-12(-14) m m , at early anthesis cleft
±1/3-J/2 its length, but the lobes later separating to rim
of stemonozone; androecium 4-7.5(-8) cm, the free
filaments scarlet or carmine throughout. Pods 7-11 x
0.9—1.5(—1.6) cm, pilose-velutinous with fuscous or
partly fuscous, rarely with white or yellowish-white
hairs.
In oak- and pine-forest, 750-2400 m, locally plen
tiful and sometimes associated with var. houstoniana,
widespread in tropical Mexico s.-ward from s. Sonora
and Hidalgo, s.-e. through upland Chiapas to
Guatemala and Honduras. — M a p 41. — Fl.
(VI-)VII-II(-III).
The taxon longipedicellata was originally separated
from C. anomala by long pedicels and winter anthe
sis, and further from C. grandiflora by Macqueen &
Hernandez (1997: 16) by details of seedlings.
Note that Macqueen and Hernandez (1997: 23)
attribute to C. grandiflora the types of Inga sericea
Martens & Galeotti and 7. elegans Martens & Galeottii
(Bull. Acad. Roy. Sci. Bruxelles 10: 318, 321. 1843),
collected by Galeotti in Oaxaca (nos. 3418,3235, both
BR, not seen by me).
118b. Calliandra houstoniana (Miller) Standley var.
colomasensis (Britton & Rose) Bameby, stat. nov
Anneslia colomasensis Britton & Rose, N. Amer. FL
23: 72. 1928. — "[Mexico.] . . . near Colomas,
Sinaloa, July 14, 1927, //. N.J Rose 1660. " —
Holotypus, NY!; isotypus, US!.
Suggesting a small-lvd variant of var. anomala, but
the fls and especially the fmits notably small, the ves
ture of inflorescence appressed, silvery-gray. Lf-for
mula ±xiii/38; lf-stks 7-8 cm, the longer interpinnal
segments 6-9 m m ; rachis of longer pinnae 1-1.7 cm;
lfts imbricate, linear-elliptic obtuse, faintly 1-nerved,
the longer ones ±2.8 x 0.5 m m . Peduncles to 8 m m ;
pedicels to 2.5 x 0.6 m m ; calyx ±1.4 x 2.2 m m , the
teeth 0.3 m m ; corolla to 7.7 m m , the lobes at maturity
to 5 m m ; androecium ±3.2 cm, of unknown color but
probably red throughout. Pods ±3.5-5 x 0.5-0.6 cm,
the valves subappressed-pilosulous with gray hairs.
In unrecorded habitat, known only from the type,
collected in young flower and almost mature fmit in
July, the "Colomas" of the protologue not precisely
identified.
In their key to Houstonianae Britton and Rose
(1928: 52) distinguished Anneslia colomasensis by
leaflets only 1.5 m m and flowers 6 m m long, but these
measurements are not confirmed by the holotype,
which has leaflets to 2.8 m m and corolla to 7.7 m m
long. The plant is nevertheless peculiar in its very short
pinnae and especially in its short narrow pod. There
seems httle of substance to separate A. colomasensis
from var. anomala, and what there is will obviously
require evaluation when further collections are se
cured. It was equated with C. grandiflora by Mac
queen and H. Hernandez (1997: 23).
118c. Calliandra houstoniana (Miller) Standley var.
acapulcensis (Britton & Rose) Bameby, stat. nov.
Anneslia acapulcensis Britton & Rose, N. Amer. Fl.
23: 72. 1928. — "[Mexico.] Acapulco, Guerrero, Oc
tober, 1894, [Edward] Palmer 59." — Holotypus (fr),
NY!. — Calliandra acapulcensis Standley, Field
Mus. Nat. Hist., Bot. Ser. 11: 159. 1936. Referred by
Macqueen and H. M . Hernandez to C. calothyrsus.
Lf-formula (v-)vi-ix(-x)/(18-)20-36(-39); lf-stk of
longer lvs 6-15 cm, the longer interpinnal segments
7-14 m m ; rachis of longer pinnae 4—8 cm; lfts hnear or
linear-lanceolate acute, straight or nearly so, the longer
ones 5.5-10.5 x 1.4-2.4(-2.7) m m , (3.2-)3.5-5.2
times as long as wide, the scarcely displaced midrib
faintly 5-8-branched on each side in young lvs, the
secondary venules scarcely visible in older ones. Inflo
rescence-axes gray-pilosulous or subglabrous, the pri
mary one mostly 7-20 cm, but out-of-season some
times shorter; peduncles 7-12 m m ; pedicels l^t m m ;
calyx 1.3-1.7 x 1.7-2.3 m m , the teeth 0.25-0.55 m m ;
corolla 6.5-9 m m ; androecium of var. calothyrsus, but
the tassel bicolored, palhd in proximal half, thence
pink-crimson. Pods in profile 6-12 x 0.9-1.4 cm,
pubemlent or pilosulous overall.
In tropical deciduous woodland, from near sea
level to 600 m, thence ascending into oak forest to
1950 m, scattered within part of the range of var.
anomala in Mexico s. of the Transverse Volcanic
Range, from w. Jalisco and Colima s.-e. to the Pacific
slope in Oaxaca. — Fl. VIII—II.
118d. Calliandra houstoniana (Miller) Standley var.
calothyrsus (Meisner) Barneby, stat. nov. C.
calothyrsus Meisner, Linnaea 21: 251. 1848. — "In
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 181
CALLIANDRA HOUSTONIANA
• var. A N O M A L A
T var. ACAPULCENSIS
M A P 41. Distribution of Calliandra houstoniana (Miller) Standley var. anomala (Kunth) Bameby and var. acapulcensis (Britton & Rose) Bameby in Mexico and northern Central America.
sylvis montosis prope flum. Mariepastonkreek, m.
Maio 1846 legit Kegel n. 1465." — Holotypus, N Y
(hb. Meisner, at present on loan to M E X U ) ; iso
typus, G O E T ace. Breteler, Acta bot. neerl. 38(1):
79, fig. 1. 1989. — Feuilleea calothyrsa [sic] O.
Kuntze, Revis. Gen. PL 1: 187. 1891. Anneslia
calothyrsus J. D. Smith, Enum. PL Guat. 1: 10. 1889.
C. confusa Sprague & Riley, Bull. Misc. Inform. 1923: 371.
1923. — "Central America. Guatemala: Alta Vera Paz; Coban, 1200 m. Tuerkheim 690." — Holotypus, K!; iso
typus, NY!; paratypi, Salvin & Godman 210, K!, Bernoulli
& Cario 1253, K!. —Anneslia confusa Britton & Rose, N.
Amer. Fl. 23: 70. 1928. C. similis Sprague & Riley, Bull. Misc. Inform. 1923: 372.
1923. — "Central America. Costa Rica: San Jose, Oersted
56." — Holotypus, K! = photo s. num., NY!; paratypus,
Oersted 54, K!. — Anneslia similis Britton & Rose, N.
Amer. Fl. 23: 71. 1928. — Equated with the preceding by
Woodson & Schery, 1950: 264, fig. 96.
C. calothyrsus sensu H. Hernandez, Anales Inst. Biol. Univ.
Nac. Mexico, ser. bot. 62: 121-131, fig. 1-5. 1991; Mac
queen & H. M. Hernandez 1997: 16, fig. 1, map 1.
C. callothyrsus [sic] Anon., Tropical Legumes: Resources
for the future 197-199, fig. 1979.
Lf-formula (vi-)ix-xviii/(30-)34-62; lf-stks of
larger lvs 8-15 cm, their longer interpinnal segments
6-10(-12) m m ; rachis of longer pinnae (4—)5-8.5 cm;
longer lfts hnear acute 4—9(-10) x 0.65-1.9 m m ,
either straight or gently incurved. Inflorescence-axes
either glabrous, or pubemlent, or pilose, but the
peduncles at most thinly so; peduncles (3—)4—12(—15)
m m ; pedicels 2-4.5 m m ; perianth commonly gla
brous, rarely micropuberulent; calyx 1.6-2.3 x 2.2-
3.2 m m , the teeth 0.25-0.6 m m ; corolla (6-)6.5-9.5
(-11.5) m m , the lobes as long as tube or separating to
rim of stemonozone; androecium 40-52-merous, usu
ally crimson throughout, occasionally pallid proxi
mally and pink distally. Pods in profile 8-11 (-12) x
1.1-1.6 cm, commonly glabrous or micropuberulent,
less often strigulose or even pilose.
In brush-woodland, along stony river banks, on
roadsides, in wasteland, and in undisturbed open for
est, (2-)50-1400(-1650) m, locally plentiful, s.-e.
Mexico (Oaxaca, Chiapas) to Belize and n.-w.
Panama, collected once in Veracmz (Jalapa, perhaps
planted) and apparently disjunct locally in upland w.
Jalisco and Cohma; cultivated and weedy in interior
Hispaniola, and widely planted in the Paleotropics;
first described from specimens collected in 1846 by
H. A. H. Kegel on the Saramacca River in Surinam,
182 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
but not since encountered in S. America. — Map 42.
— Fl. VI-III, perhaps in all months of the year.
118e. Calliandra houstoniana (Miller) Standley var.
houstoniana. Mimosa houstoniana Miller, Gard.
Diet. ed. 8: Mimosa #16. 1768, sens. str. — Lecto-
holotypus, a fmiting spm so annotated by H.
Hernandez, BM!; paratypi, B M (fr) = photo, NY!,
B M (fl) = Acacia americana non spinosa flore pur-
pureo e Vera C m z , these all annotated by P. Miller.
— Anonymos Houston ex Banks, Reliq. Houston, t.
26. 1781. Anneslia houstoniana Britton & Rose, N.
Amer. Fl. 23: 70. 1928, syn. omnibus inclusis. C.
houstoniana subsp. houstoniana Macqueen & H.
M . Hernandez, 1997: 34, fig. 5, 6, map 3.
Mimosa houstoni L'Heritier, Serf Angl. 30. 1789. — De
scribed from plants cultivated in England, probably at Kew,
in years 1786-1787. — Not found at K in 1994. —Acacia
houstoni Willdenow, Sp. PL 4: 1062. 1805. Anneslia falcifo-
lia Salisbury, Parad. Lond. 2(1); t. LXIV (titled A. grandi
flora). 1807, substit. illegit. Calliandra houstoni Bentham, J.
Bot. (Hooker) 2: 139. 1840; & London J. Bot. 3: 111. 1844.
Acacia metrosiderifolia Schlechtendal, Linnaea 12: 567.
1838. — "[Mexico. Veracruz:] in sylvis dumetisque Jala-
pae et prope Hac[ienda] de la Laguna . . . (Schiede)." —
Isotypus, OXF!. — Calliandra metrosiderifolia Bentham
ex Jackson, Index Kew. 1: 385. 1895. Anneslia alamosensis Britton & Rose, N. Amer. Fl. 23: 72.
1928. — "[Mexico.] Vicinity of Alamos, Sonora, March
14, 1910, Rose, Standley & Russell 12792." — Holotypus,
NY!; isotypus, US!. — Calliandra houstoniana subsp.
alamosensis Macqueen & H. M. Hem. 1997: 34.
A. etzatlana Britton & Rose, N. Amer. FL 23: 73. 1928. —
"[Mexico.] Near Etzatlan, Jalisco, October 2, 1903, Rose &
Painter 7525." — Holotypus, NY!; isotypus, US!. A. lucens Britton, N. Amer. Fl. 23: 194. 1928. — "Type from
the vicinity of Siguatepeque, Comayagua, Honduras . . .
February 14-27. 1928, [P. C ] Standley 56397." — Holo
typus, NY!. — Calliandra lucens Standley, J. Arnold Arb.
11:30. 1930. Calliandra houstoniana subsp. stylesii Macqueen & H. M.
Hernandez, Kew Bull. 52: 36, fig. 7 (A-C), 1997. —
"Mexico, Tabasco, 58 km west of Catazaja, Macqueen
S60V — Holotypus, M E X U n.v.; isotypus, K!.
Acacia houstoni sensu Edwards, Bot. Reg. 2: tab. 78. 1816, nom. illegit., including anterior Anneslia falcifolia Salisb.
Calliandra houstoni sensu Bentham, 1875: 556. Calliandra houstoniana sensu Standley, 1922: 386 (vernac
ular names and notes on use in medicine); McVaugh, 1987:
162.
Lf-formula (v-)vi-xii(-xiv)/40-65(-69); longer lf-
stks 6-12(-16) cm, the longer interpinnal segments
(4.5_)6_i3(_20) m m ; rachis of longer pinnae (4-)
4.5-10(-14) cm; longer lfts (4.2-)5-10(-ll) x 0.8-
1.8(-2) m m , the simple or pinnately branched midrib
curved forward toward the sharply acute apex, the
blade (4.2-)4.5-8(-9) times as long as wide. Inflores
cence strigulose-pilosulous or pilose with at least
partly, often wholly brown, bronze, or black hairs;
peduncles 2-13(-16) m m ; pedicels 0.7^.5(-5) x
0.7-2.5 m m ; calyx (1.7-)2-3.6(-4.5) x 2.4-54(-5.6)
m m , the teeth (0.3-)0.5-1.6(-3) m m ; corolla 7.5-
12(—13) m m ; androecium 36-76(-100)-merous,
(5_)55_7 cm, the tassel red beyond the often pallid
tube. Pods in profile (6-)7-14 x (l-)1.3-2(-2.3) cm,
densely pilose-velutinous with brown or black, some
times partly or almost wholly pallid hairs.
In disturbed woodland, along highways, and in pas
ture thickets, ascending from near sea level into pine-
and oak-forest, (0-)5-1800(-2000) m. in s.-w. Mexico
sometimes sympatric with var. anomala and in Chia
pas s.-ward with var. calothyrsus, locally plentiful in
tropical Mexico and Central America: in Mexico s.
ward from 27°N in s. Sonora and from 22°30'N in s.
Tamaulipas through the Transverse Volcanic Range
and interior Oaxaca and Chiapas to Guatemala,
Behze, Honduras, and El Salvador. — M a p 43. — Fl.
VII-XII(-I).
The subsp. alamosensis was distinguished by Mac
queen and Hernandez by texture of corolla and per
sistent vesture of the pod.
119. Calliandra juzepczukii Standley, Publ. Field
Columbian Mus., Bot. Ser. 8: 313. 1931. — "Mex
ico: Arriaga, Chiapas, January 7, 1926, S. Juzepczuk
1382V — Holotypus, F 6214261; isotypus, L E n.v
FIG. 27
C. juzepczukii sensu Macqueen & H. M. Hem., 1997: 37,
fig 8, map 4.
Amply, elaborately microphyllidious (sub)shrubs
with 1 to several simple or nearly simple stems 1-2.5
m, exceptionally arborescent to 4 m, the stems and lf-
axes densely pilosulous with mostly gray or with
mixed gray and sordid or brown hairs to 0.3-0.9 m m ,
the lvs bicolored, the lfts brown-olivaceous glabrous
and lustrous above, paler dull brown and minutely
strigulose beneath, ciholate, the inflorescence a simple
terminal efohate pseudoraceme of few-fld umbelli
form racemes well exserted from fohage, the individ
ual fls relatively large, coarsely thick-textured, the
corolla white-silky externally; phyllotaxy distichous.
Stipules of thin texture, early dry caducous (hence
poorly known), the blades lance-ovate ±3.5-5 m m .
Lf-formula (viii-)ix-xix/(38-)40-63, the pinnae fewer
only in random diminished distal lvs; lf-stks (5-)
6.5—13(—15) cm, the petiole 11-24 m m , at middle
(0.9-) 1.1-2.2 m m diam, the longer interpinnal
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 183
M A P 42. Distribution of Calliandra houstoniana (Miller) Standley var. calothyrsus (Meisner) Bameby in Mexico and Central America.
segments 4—8(-10) m m , the narrow ventral groove
obscurely bridged at insertion of pinnae; pinnae
shorter toward base of lf-stk, less so distally, the rachis
of longer ones (4-)5-8(-9) cm, the longer interfoholar
segments 0.8-1.4 m m ; lft-pulvinules 0.1-0.25 x 0.45-
0.6 m m , not wrinkled; lfts firm, plane or ventrally low-
convex, in outline linear-oblong from obhquely tmn
cate base, proximally straight but toward apex abmptly
bent forward and triangular-apiculate, those near mid-
rachis 4-7.5(-7.8) x 1-1.8 m m , (3.4-)3.8-^.6M.8)
times as long as wide; midrib slender, displaced to di
vide blade ±1:1.5-2.5, simple or almost so, the poste
rior primary nerves 1(2), very short and faint. Primary
inflorescence-axis (0.5-)l-3 dm; peduncles ascending
stout, solitary or fasciculate by 2-3,7-18(-24) x 1.4-3
m m , ebracteate; capitula 2-5-fld, the receptacle
turbinate tmncate 3-4 m m diam; bracts resembhng
stipules but smaller, early papery caducous; pedicels
(2-)4—7 x 1.5-2.4 m m ; perianth 5-merous, the calyx
shortly densely gray-brown-pilosulous, the fleshy-
coriaceous greenish corolla densely white-silky-
strigose externally, its lobes ±1-1.5 m m thick in sec
tion; calyx hemispherical, turbinate-campanulate or
shallowly patelliform-campanulate 3.5-6(-6.5) x
6.5-9(-10) m m , the deltate, depressed-deltate or
depressed-semicircular and acuminulate teeth 0.6-2
m m ; corolla 13.5-17 m m , the lanceolate or lance-
elliptic lobes 8-10.5 m m ; androecium 110-166-
merous, 5.5-7.5 cm, the stemonozone 3-4 m m , the
tube 5-6 m m , the tassel crimson-scarlet; intrastaminal
nectary 0; ovary linear-oblanceolate glabrous. Pods 1
per capitulum, stiffly erect, straight, 6-7-seeded, in
profile 10-12.5 x 1.3-2 cm, the sutural ribs in dorsal
view 4-7 m m wide, the woody valves densely gray-
velutinous-pilosulous overall; ripe seeds not seen.
In scmb thickets, semideciduous woodland, and
ascending into the edge of oak forest, 100-650 m and
in Chiapas to 900 m, locally plentiful on the Pacific
slope in s.-e. Mexico (Oaxaca, adj. Veracmz, and
Chiapas). — M a p 44. — Fl. VI-II. — Cabello rojo.
Calliandra juzepczukii, of tropical southeastern
Mexico, and its sibhng C. palmeri, of southwestern
Mexico, together resemble C. houstoniana sens, lat.,
but differ collectively in the very large, camosulous,
white-silky corollas. They differ from one another in
size of calyx and leaflets, characters of no great
moment in the genus, and they might better be treated
as allopatric varieties of one species.
184 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
M A P 43. Distribution of Calliandra houstoniana (Miller) Standley var. houstoniana in Mexico and Central America.
120. Calliandra palmeri S. Watson, Proc. Amer.
Acad. Arts 22: 410. 1887. — "[MEXICO. Jalisco:]
Guadalajara . . . August [1886] . . . ([E. Palmer]
279)." — Holotypus, GH!; isotypi, K!, N Y (2
sheets)!, US!. —Anneslia palmeri Britton & Rose,
N. Amer. Fl. 23: 73. 1928.
C. palmeri sensu Standley, 1922: 385; McVaugh, 1987: 166,
fig. 20; Macqueen & H. M. Hem., 1997, fig. 10, map 4.
Elaborately microphylhdious, few- but ample-lvd,
either virgately 1-stemmed or bushy and several-
stemmed shrubs 0.5-1.7 m, except for lustrous
glabrous ventral face of lfts densely pilose-pilosulous
throughout with fine gray hairs to ±0.6-1.3 m m , the
lfts paler brown and dull beneath, ciliate, the inflores
cence a long efoliate pseudoraceme of few- and large-
fid, umbelliform capitula, the perianth of individual
fls densely white-silky externally; phyllotaxy distich
ous. Stipules caducous (absent from fmiting speci
mens), thinly herbaceous, ovate or lanceolate ±5-7.5
m m wide. Lf-formula viii-xiii/44-55(-60); lf-stks
stout 12-22 cm, the petiole ±2-5 cm, at middle 2-3
m m diam, the longer interpinnal segments 1-2 cm,
the narrow ventral groove bridged at insertion of pin
nae; pinnae either distally accrescent or shorter at
both ends of lf-stk, the rachis of longer ones 8-14 cm,
the longer interfoholar segments 1.3-2.4 m m ; lft-
pulvinules 0.3-0.7 x 0.5-1 m m , not wrinkled; lfts
abmptly decrescent proximally, gradually so distally,
the firm plane blades linear-lanceolate from shortly
deltate-auriculate or semitmncate base, straight or
incipiently falcate, deltately acute, the larger ones
9-13.5 x (1.7-)2-3.5 m m , (3.5-)3.7-6.3(-6.5) times
as long as wide; venation palmate-pinnate, the midrib
displaced to divide blade ±1:2-2.5, the inner of 2 pos
terior primary nerves incurved-ascending short of
mid-blade, the midrib ±5-7-branched on posterior
side, the secondary and faint tertiary venulation either
delicately prominulous or immersed. Inflorescence-
axis simple, efohate, 2-5 dm; peduncles solitary or
geminate 4.5-18 m m , ebracteate, becoming 2.5-4
m m diam in fmit; capitula (1—)2—5-fld, the caducous
bracts ovate-suborbicular 3.5-5 m m diam, early dry
deciduous; pedicels turbinate 1-3.5 x ±2 m m ; peri
anth 5(-6)-merous, the thinly coriaceous calyx hemi
spherical, gray-brownish-strigose except for glabres
cent teeth, the fleshy, when dry coriaceous corolla
white-pilose overall, in section ±1 m m thick; calyx
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 185
FiG. 27. Calliandra juzepczukii Standley.
186 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
¥ CALLIANDRA PALMERI
• C JUZEPCZUKII
MAP 44. Distribution of Calliandra palmeri S. Watson
and C. juzepczukii Standley in Mexico.
6-9 m m , the teeth either hemispherical erose or
depressed-deltate 0.6-2 m m ; corolla 15-21 mm, the
erect lanceolate lobes 9-11.5 m m ; androecium
130-152-merous, 6.5-8 cm, the tube 5.5-7 mm, the
stemonozone 3-3.5 mm, internally thickened but
without defined discoid nectary, the tassel crimson-
scarlet. Pods stiffly erect, in broad profile 12-14 x
1.6-2.2 cm, the massive sutural ribs in dorsal view
6-8 m m wide, the woody valves low-convex over
each of 6-8 seeds, obliquely sulcate between them,
the whole pod densely gray-brown-velvety-pilose
overall; seeds in broad view 9.5-12 x 7 mm, the hard
smooth, hght brown testa finely pleurogrammic.
At edge of fields, in shrub-thickets, and openings
in pine-oak or oak forest, 1020-1650 m, locally plen
tiful in the lower valley of rio Grande de Santiago in
Jahsco and Nayarit, Mexico. — Map 44. — Fl.
(V-)VI-IX; fr. VIII-XII.
An exceptionally handsome calliandra, akin to the
preceding species, C. juzepczukii, which differs prin
cipally in smaller leaflets and shorter calyx.
121. Calliandra physocalyx H. Hernandez & Sousa,
Syst. Bot. 13: 521, figs. 2-A. 1988. — "Mexico.
Guerrero, Mpio. Tlacochistlahuaca, 33 km al N de
Omotepeque ... 26 Nov 1983, E. Martinez S. & F.
Barrie 5763V — Holotypus, M E X U n.v; isotypi,
BM!,NY!,US!.
C. physocalyx sensu Macqueen & H. M. Hem., 1997: 42,45
(fig.), map 4.
Slender shrubs 1.5-3 m with terete brown stems,
glabrous except for micropuberulent lf-stks and
perianth, the fohage olivaceous subconcolorous, the
narrow crowded lfts facially glabrous but some micro-
ciliolate, the inflorescence a terminal, efoliate or
basally few-leaved pseudoraceme of few-fld umbel
liform capitula, the tumid calyces nearly or quite
enclosing the corolla; phyllotaxy distichous. Stipules
lanceolate 3-6 m m , thinly herbaceous, caducous. Lf-
formula (iii—)iv—vi/58—70; lf-stks 4—7 cm, the petiole
7-13 m m , at middle 0.8-1.3 m m diam, the interpinnal
segments about as long, the ventral groove continuous
or weakly bridged at insertion of pinnae; pinnae a little
accrescent distally, the rachis of distal ones 7-
9.5 cm, the interfoholar segments scarcely 1 m m ; lft-
pulvinules 0.3-04 m m diam, not wrinkled; lfts
decrescent only near each end of rachis, otherwise
subequiform, the blades linear-lanceolate from shal
lowly obtusely auriculate base, deltately apiculate, the
longer ones 5.5-9 x 1-1.7 m m , (4.2-)4.5-6 times as
long as wide; midrib slender simple, subcentric at mid-
blade, further venulation immersed. Primary axis of
inflorescence ±10-16(-22) cm, its lowest 1-2 nodes
sometimes foliate, thereafter efohate; peduncles 8-17
m m , ebracteate; capitula (2-)3-5-fld, the ascending
pedicels 3-5 m m ; bracts caducous, described as ovate
acute to 9 m m ; perianth firm but not fleshy, 5-merous,
the thinly coriaceous calyx subglabrous, faintly
nerved, nearly as long as or shghtiy surpassing the
thinly gray-strigulose-puberulent corolla; calyx
broadly ventricose-campanulate ±10-13 m m diam, the
deltate-triangular teeth unequal, 2 sinuses recessed
4.5-6 m m , the rest very short, the whole calyx appear
ing 2-lipped; corolla 11-15.5 m m , the lobes 3.5-6 m m ,
recurved at apex; androecium 118-merous (1 count),
±7-7.5 cm, either red overall or proximally white to
pink and the tassel pink to red, the stemonozone ±3
m m , somewhat thickened internally, the tube ±8-11
m m ; intrastaminal disc 0; ovary sessile, densely pu
bemlent. Pods stiffly ascending, in broad view 10-12 x
0.9-1.1 m m , the sutural ribs in dorsal view scarcely 3
m m wide, the valves densely pilosulous overall and
microscopically granular; seeds in broad view ± 7 x 5
m m , the smooth brown testa finely pleurogrammic.
In disturbed and regenerating, mesophytic mon
tane forest and oak-woodland, 490-1050 m, localized
astride the Guerrero-Oaxaca state line, on the s. slope
of Sa. Madre del Sur, near 17°N, 98°W, in s. Mexico.
— Fl. IX-XI.
Calliandra physocalyx is in broad detail similar to
some elements of the C. houstoniana complex, but has
rather fewer pinnae per leaf and is unique in its blad
dery calyx that nearly or quite encloses the corolla.
122. Calliandra wendlandii Bentham, Trans. Linn.
Soc. London 30: 556 ("Wendlandr). 1875. — ". . .
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 187
Tropical America: Guatemala, Wendlandr — Holo
typus, dated 4 Jan 1857 (fl, imm fr), Hermann Wend-
land 113, GOET!. — Feuilleea wendlandii O.
Kuntze, Revis. Gen. PL 1: 189. m i . Anneslia wend
landii Britton & Rose, N. Amer. Fl. 23: 73. 1928.
C. wendlandii sensu Hemsley, 1988: 359; Standley & Steyermark, 1946: 28.
Known from only one foliate long-shoot with ter
minal efoliate pseudoraceme of few-fld umbelliform
capitula resembling those of C. houstoniana var.
anomala, either shmbby or virgate, the homotinous
stem and lf-axes densely pilosulous with sordid hairs
to ±0.6 m m , egranular, the small crowded lfts gla
brous facially, ciholate, subconcolorous, the perianth
white- and fuscous-strigose. Stipules caducous (few
seen) lanceolate ±3 x 1 mm. Lf-formula i-ii/30-
38, some lvs consisting of one proximal pair of pin
nae and one terminal pinna; lf-stks 3.5-7 mm, the one
interpinnal segment, when present, {2 mm; rachis of
longer pinnae ±2.5 cm, the longer interfoholar seg
ments ±1 m m ; lft-pulvinules ±0.2 x 0.3 mm, not
wrinkled; lfts subequilong except at very ends of
rachis, the blades linear-elliptic from shallowly auric
ulate base, obtuse, straight or gently incurved, the
longer ones 4—5.5 x 1-1.2 m m , 4-4.5 times as long as
wide, 1-nerved, the slender costa subcentric at mid-
blade, simple. Inflorescence narrowly pyramidal, the
primary axis simple ±1 dm, efoliate; peduncles stout,
2-3 per node of inflorescence, 4—7.5 mm; capitula
3-5-fld, the pedicels 1-2 x 1-1.5 mm; perianth 5-
merous, the calyx thinly brownish-pubemlent, the
corolla densely strigose, the hairs mostly white, fus
cous at tip of lobes; calyx hemispherical ±2 x 3.7
m m , the depressed-deltate, very obtuse teeth ±0.3
m m ; corolla 12 m m , the lanceolate, internally fuscous
lobes ±7 m m ; androecium 40-merous, ±4 cm, the ste
monozone 3 m m , the tube 5 m m , the tassel of un
known color (but probably red); intrastaminal disc
not seen; ovary at anthesis pilosulous. Pods not seen
fully ripe, ±6.5 x 0.7 cm, densely gray-pilosulous
overall; seeds unknown.
In unrecorded habitat, but to be sought in the pine-
oak zone, collected once in Guatemala, in 1857, be
tween Lago Izabal and Cd. Guatemala, not since
seen. — Fl. I-II(-?).
Calliandra wendlandii has the blunt leaflets with
scarcely displaced midrib, the terminal efoliate inflo
rescence, and the individual flowers of C. houstoni
ana var. anomala, of which it is clearly a near rela
tive. In the protologue Bentham twice described the
leaves as conjugately bipinnate, but the leaves of the
holotypus, annotated in Bentham's hand, have either
one pair of pinnae, or two pairs crowded together at
tip of leaf-stalk, or (in a few) one perfect pair and one
terminal pinna. Bentham's error of oversight was
copied by Britton and Rose (1928) and by Standley
and Steyermark (1946), but does not jeopardize the
status of the species, unless the typus itself represents
a freak with abnormally reduced leaf-formula.
Standley and Steyermark (1946) suggested that Cal
liandra wendlandii may not be Guatemalan. However,
if the date of collection (4 Jan 1857) is recorded cor
rectly, the typus must have been collected (Schlechten
dal, Bot. Zeit. 15: 278-280. 1858) by Wendland en
route with Skinner between Lago Yzabal and Ciudad
Guatemala. The species should be expected in the oak-
belt in southeastern Guatemala.
The typus of C. wendlandii is interpreted by Mac
queen and Hernandez (1997: 23) as decapitated C.
grandiflora.
123. Calliandra quetzal (J. D. Smith) J. D. Smith,
Enum. PL Guatemal. 8: 36. 1907. Anneslia quetzal
J. D. Smith, Bot. Gaz. (Crawfordsville) 13: 28.
1888. — "Santa Rosa, Dept. Baja Verapaz . . . July,
1887. Ex pi. Guat. cit. [H von Tuerckheim] 1324."
— Holotypus, US!; isotypi, K!, N Y (2 sheets)!.
Fig. 28
Anneslia quetzal sensu Britton & Rose, 1928: 74. Calliandra quetzal sensu Standley & Steyermark, 1946: 26.
Macrophylhdious shrubs 1-2 m, glabrous except
for some minutely ciholate lfts, with terete smooth
homotinous stems, dilated stipules, and plane charta
ceous bicolored lfts dark dull brown-olivaceous
above and paler dull cinnamon-brown beneath, the
inflorescence a terminal exserted efoliate pseudo
raceme of few-fld umbelliform capitula; phyllotaxy
distichous. Stipules resembling lfts in size and tex
ture, suborbicular-flabellate or broadly reniform
±10-24 x 12-30(-36) mm, broadly obtuse, flabel-
lately many-nerved from base, persistent. Lf-formula
ii-iii/6-9; lf-stks 4—9 cm, the petiole 2-3.5 cm, shal
lowly and narrowly sulcate ventrally, the longer inter
pinnal segment 1.5-3.7 cm, the terminal appendage
linear-lanceolate caducous; pinnae a little accrescent
distally, the rachis of furthest pair 5-9 cm, the longer
interfoholar segments 8-16 mm; lft-pulvinules 1.2-
1.6 x 0.55-0.7 mm, sharply wrinkled; lft-blades
broadly elliptic, oblong-elliptic or suborbicular from
shallowly semicordate base, the penultimate ones
±2^- x 1.3-2.2 cm, 1.2-2 times as long as wide;
venation pinnate, the midrib subcentric, straight, 4—
188 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
FiG. 28. Calliandra quetzal (J. D. Smith) J. D. Smith.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 189
7-branched on each side, the secondary nerves slen
der, brochidodrome well within the margin. Primary
axis of inflorescence ±1-2 dm, simple; peduncles
either solitary or geminate (2-)7-26 mm, ebracteate,
abmptly dilated at apex; capitula 4-5-fld, the floral
bracts subobsolete, the pedicels 3-6 x 0.5-0.65 mm;
fl-buds plumply elhpsoid, short-acuminate; perianth
firm but scarcely fleshy, glabrous, brown in dried
specimens, the calyx faintly several-nerved, the co
rolla not so; calyx campanulate 3.6-5 x 2.3̂ 4.5 mm,
the teeth either deltate or broadly ovate obtuse
1.5-1.8 m m ; corolla 9.5-15 m m , the lobes free
through 3.5-4 mm, or further in late anthesis; an
droecium 54—66-merous, 3.4-5.2 cm, the tube 5.5-7
mm, the stemonozone to 2.6 m m , the tassel white, the
intrastaminal nectary ±1.2 m m tall. Pods (not seen)
described as ±10 x 1.5 cm, glabrous.
In oak-pine forest at 1500-1700 m, apparently very
local in Baja Verapaz, Guatemala (Santa Rosa;
Salama). — Fl. IV-VI.
In context of sect. Calliandra in Central America
the rare C. quetzal is unmistakably characterized by
foliaceous stipules, bi- or trijugate pinnae, and only
6-9 pairs of ample leaflets per pinna.
IV/B. Series VIRGATAE Barneby
Calliandra sect. Calliandra ser. Virgatae Bameby,
ser. nov, monotypica ab affini ser. Calliandra his
signis conjunctis recedens: caules herbacei e cau-
dice orientes; phyllotaxis subdisticha, nonnumquam
irregularis; pinnae geminatae; androecii tubus
angustus longe exsertus. Inflorescentia pseudorace-
mosa infeme fohata, dein aphylla. — Sp. unica: C.
virgata Bentham.
Functionally herbaceous subshmb from rootstock;
phyllotaxis labile, irregular or subdistichous; lf-for
mula i/14—27; peduncles arising from furthest lf-axils
and terminally pseudoracemose, bracteate; androecial
tube slender, exserted up to twice the length of
corolla. — Sp. 1, of centr. Brazilian Planalto in Goias
and far w. Minas Gerais.
In its range of dispersal C. virgata is instantly rec
ognized by the syndrome of functionally herbaceous
stems, geminate pinnae, absence of axillary brachy
blasts, and homomorphic flowers with slender, long-
protruded androecial tube. The numerous calhandras
of upland Bahia and Minas Gerais that share its habit
of growth have either pluripinnate leaves, or ebracteate
peduncles, or fewer and thick-textured flowers, or an
included to shortly exserted staminal tube, or most of
these features combined. The twin pinnae of C. virgata
combined with its androecial structure are reminiscent
of Hylaean C. surinamensis, but this is an arborescent
shrub very different in ecology and gross aspect.
124. Calliandra virgata Bentham, J. Bot. (Hooker)
2: 140. 1840. — "Brazil, Pohl," the locahty given in
Martius, 1876: 417: "Serra dos Chrystaes [Goias]."
— Holotypus, Pohl 1474, K (hb. Benth.)! = N Y
Neg. 1963. — Feuilleea virgata O. Kuntze, Revis.
Gen. PL 1: 189. 1891. FiG. 29
C. virgata sensu Bentham, London J. Bot. 3: 105. 1844; 1875: 548; 1876: 417, t. 108.
Subshrubs from woody rootstock, the erect virgate
stems simple or almost so, 3-7 dm, dying back yearly
to the ground, the stems, lf-stks, and peduncles thinly
to densely pubemlent and sometimes in addition
finely hirsutulous, the firm plane bicolored lfts dark
brownish-green and lustrous ventrally, paler beneath,
usually glabrous or only minutely ciholate, exception
ally (Irwin 8805, NY) facially hirsutulous and cihate,
the hemispherical capitula solitary or rarely geminate
on stout peduncle in upper lf-axils, or some later ones,
by suppression of distal lvs, forming a short terminal
pseudoraceme; phyllotaxy indecisive, sometimes
subdistichous, often irregular. Stipules erect, mostly
narrow-lanceolate or subsetiform, sometimes lance-
ovate, (2-)3-9 x 0.45-1.3 mm, weakly striate, tardily
deciduous. Lf-formula i/14—27; lf-stks upward from
mid-stem 2-8 mm, some proximal ones to 10-18 mm,
the terminal appendage linear-lanceolate 1-4 mm;
rachis of longer pinnae 5.5-13 cm, the longer interfo
holar segments (2.5-)3-8 mm; lfts ascending at nar
row angles to rachis, little graduated, each sessile in an
obliquely dilated socket on rachis, the pulvinule twice
as wide as long, the blade linear-lanceolate from
inequilaterally shallow-cordate or semicordate base,
abmptly apiculate or mucronulate, straight to gently
retrofalcate, the longer ones 11-23 x (2-)2.4-4-.2 mm,
(4—)4.3-6 times as long as wide; venation of 2(3) pri
mary nerves, the midrib subcentric, the posterior
nerve produced ± to mid-blade, the short secondary
nervules widely ascending, the tertiary ones sinuous,
the whole venulation prominulous on both faces.
Peduncles (2—)2.5—8(—11) cm, bracteate above mid
dle; capitula 12-26-fld, the receptacle 2-3 m m diam,
the sessile calyces contiguous at anthesis, the outer
ones often incurved toward vertical; bracts lance-ovate
1.2-2.8 mm, persistent; fls glabrous, the calyx striate,
the pallid whitish or brownish-green corolla only
faintly nerved; calyx campanulate 2.9-5 x 1.7-3.2
mm, the ovate-deltate teeth 0.3-1 mm, often unequal;
corolla (7.5—)8.5—11.5 mm, the ovate or broad-lance-
190 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
FiG. 29. Calliandra virgata Bentham. Reproduced from Bentham in Martius, Fl. bras. 15(2): t. 108. 1876.
olate lobes 1.6-3 x 1.2-2 m m ; androecium 28-38-
merous, 29-37 m m , the stemonozone 0.6-1.6 m m , the
tube 13-25 m m , the tassel pink or carmine distally; in
trastaminal disc 0.5-1 m m tall, distinctly 5-lobed;
ovary substipitate, glabrous, the style hnear, at porose
apex ±0.2 m m diam. Pods solitary from top of recep
tacle, erect, in profile oblanceolate ±5-6 x 1 cm, 7-8-
seeded, the sutures and valves ahke ligneous, purple-
brown glabrous, the dilated sutures 3-4-ribbed
lengthwise on each side of line of dehiscence, the
recessed valves densely venulose with arborescentiy
ramulose nerves obhquely ascending from inner edge
of abaxial suture; seed-funicles deltate ±1.5 m m ; ripe
seeds unknown.
In campo cerrado of the centr. Brazilian Planalto,
650-1225 m , locally abundant in the Federal District
and adj. Goias, thence s. to and just beyond the
Paranaiba river into the Triangulo Mineiro. — M a p
45. — FL IX-III.
The record of C. virgata from Pemambuco in
Flora brasiliensis (Bentham, 1876: 418) is based on
a mislabeled and misidentified specimen of C. luet
zelburgii (Martius s.n., M ! ) .
IV/C. Series TSUGOIDEAE Barneby
Calliandra sect. Calliandra ser. Tsugoideae Bameby,
ser. nov, inflorescentia terminali saltern breviter ultra
foha exserta cum ser. Calliandra conveniens, folio-
lomm autem venatione conspicue diversa: foliolorum
costa et nervi primarii, tarn posterior quam anterior
rectae, inter se paralleli, in laminae apicem producti.
— Sp. typica: C. tsugoides Cowan.
Shrubs and subshrubs, like ser. Calliandra in ter
minal inflorescence (sometimes only shortly exserted
from foliage), but distinguished from all congeners
by parallel primary venation of lfts, the inner poste
rior nerve always and an anterior nerve often pro
duced nearly or quite to apex of lft-blade; lf-formula
(i—)ii—viii/24—80, the larger lfts to 20 m m . — Spp. 4,
of sandstone outcrops and white sand savannas, en
demic to the Guayana Highland and outlying sand
stone islands in s. Venezuela, immediately adj.
Guyana and Brazil, and Amazonian Colombia.
125. Calliandra tsugoides Cowan, Mem. New York
Bot. Gard. 10(1): 143. 1958. — ". . . B. Maguire, R.
S. Cowan, and J. J. Wurdack 30705 . . . cumbre and
slope of Cerro Yapacana, Terr. Amazonas,
Venezuela, Jan. 3, 1951." — Holotypus, NY!.
Arborescent shrubs, fertile when 0.8-10 m, with
terete, densely foliate gray- or fuscous-pilosulous or
sometimes sordidly pilose long-shoots but lacking
brachyblasts, the lvs strongly discolorous, the con
tiguous or narrowly imbricate lfts stiffly chartaceous,
on upper face dark olivaceous (brown when dry) and
highly lustrous, paler dull beneath, either glabrous
facially and thinly ciholate or on dorsal face thinly
finely strigulose or (very locally) loosely pilose, the
capitula of ± silvery-pilosulous, red-stamened fls
borne singly or in fascicles of 2-3 in the axil of dis
tal, coevally expanding lvs or, by suppression of
these, shortly pseudoracemose; phyllotaxy disti
chous. Stipules firm, linear-lanceolate or narrowly
ovate (2-)2.5-10.5 x (0.6-)0.8-3 m m , caducous. Lf-
formula (in uppermost lvs i—)ii—viii/21 —52; lf-stk of
major lvs below inflorescence 1-6 cm, the petiole
(consisting largely of pulvinus) 3-7 x 0.9-2.3 m m ,
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 191
M A P 45. Distribution of Calliandra virgata Bentham in southeastern Brazil.
the longer interpinnal segments 4-13 m m , the shal
low ventral groove weakly bridged; pinnae (when >1
pair) scarcely or distinctly accrescent distally, the
rachis of longer ones 4—8(-9) cm, the longer interfo
holar segments 0.9-1.6 m m , the lfts widely ascend
ing, forming when spread a lf-blade of narrowly
elliptic or oblance-elliptic outline; lft-pulvinules 0.1-
0.25 x 0.6-0.9 m m ; lfts abmptly decrescent near top
of rachis, more gradually so toward its base, the
blades linear from postically angulate base, obtuse,
straight or almost so, those near mid-rachis 6-12 x
1.1-2.5 m m , 4.6-6.7 times as long as wide; venation
palmate, the simple midrib displaced to divide blade
±1:2-3, the inner posterior and in broader blades one
anterior primary nerve ascending parallel to midrib
almost or quite to blade's apex, the 2-3(-4-) outer pos
terior primaries progressively much shorter, the vena
tion weakly prominulous or immersed on upper face,
slenderly prominulous beneath. Peduncles 9-26 m m ,
either ebracteate or charged just below the capitulum
with 1 small bract; capitula 5-12-fld, the fls homo
morphic, the receptacle clavate or hemispherical 1.8-
2.5 m m diam; floral bracts 0.8-1.8 m m , caducous;
perianth usually 4-, rarely 5-merous or the calyx ran
domly 3-merous; pedicels cryptic, 0.2-0.5 x 0.45-0.9
m m ; calyx campanulate (1.5-)1.6-3.2(-4) m m , usu
ally only thinly pubemlent, brown, nerveless or
faintly 5-15-nerved, the deltate or triangular teeth
0.3-1.2 m m ; corolla 5.2-8.2(-9.5) m m , always
silvery-pilose, rarely densely so and barbate, the
lobes 2.4-3.8 m m ; androecium 12-20(-25)-, in one
population to 40-merous, 27-50 m m , the stemono
zone 1-3 m m , the tube 1.8—3.8(-7) m m , the tassel
red. Pods in profile 5-9 x 0.95-1.15 cm, the dilated
sutural ribs 2.5-3.5 m m wide in dorsal view, the
recessed valves densely brown-papillate and thinly
pallid-pilosulous, not externally venulose; seeds
(scarcely known) ±7 m m in long diam, the close-
fitting testa ohvaceous, smooth, with well-defined
pleurogram.
About rock outcrops in savanna, on and near tepuis
of Venezuelan Guayana and adjoining Brazil, most
common on table-mountain tops at 1100-1500 m but
occurring also along rocky banks in the foothills at
100-150 m, represented by variously modified types
in different parts of a discontinuous range, described
below in more exact detail. — M a p 46. — Fl. I-VI,
very hkely at other times.
In the small Guayanan series of calhandras charac
terized by parallel primary nervation of leaflets the
polymorphic C. tsugoides is recognized by number
and size of ventrally glossy leaflets and by lack of
granular trichomes; for details see Key III. Its known
populations are ecologically restricted and mutually
distant in dispersal, and differ one from the next in
leaf-formula, pubescence, proportions of calyx to
corolla, and filament-number. The intraspecific varia
tion is analysed in the key that follows, but seems too
weak to support the recognition of formal taxa.
Key to the unnamed geographic forms of
C. tsugoides
a. Dorsal face of lfts either glabrous or finely appressed-pubescent dorsally; anterior and
posterior margins of lfts equally recurved;
calyx 1.5-2.7(-3.2) m m , thinly puberulent;
androecium 12-20(-25)-merous, the tube
1.8-3.8 m m ; T.F. Amazonas in Venezuela to n. state of Amazonas in Brazil.
b. Blade of lfts glabrous on both faces;
pinnae of larger lvs i-iv-, but mostly
ii-iii-jug.; Cerros Yapacana, Sipapo and
Autana (both cumbre and lowland) in
Venezuelan Amazonas, and C. Tunui
(on upper rio Icana in n.-w. Brazilian
Amazonas 125. alpha
b. Blade of lfts glabrous on upper faces, finely
strigulose dorsally; pinnae iv-viii-jug;
Sa. de Araca in Brazilian Amazonas 125. beta
a. Dorsal face of lfts glabrous ventrally, loosely
pilose dorsally; anterior margin of lfts abruptly
revolute, posterior margin plane; calyx 4 m m ;
androecium 40-merous, the tube ±7 m m ;
C. Guaiquinima in Venezuelan state of
Bolivar. 125. gamma
126. Calliandra vaupesiana Cowan, Bot. Mus.
Leafl. 18: 142. 1958. — T y p u s infra sub var. vaupe
siana indicatur.
192 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
M A P 46. Distribution of Calliandra tsugoides Cowan in northern South America.
Awkwardly stiffly branched, amply microphyllidi
ous, bushy shrubs ±1-2 m tall with fuscous defohate
and glabrescent older stems and trunks, notable for
long multifoliolate pinnae and silvery-pilosulous pe
rianth of fls, the new branches, lf-axes and peduncles
pilosulous-tomentulose or subappressed-pilosulous
with usually flexuous, sordid or gray hairs to 0.2-0.6
m m mixed with minute red-brown granular tri
chomes, the narrow crowded subconcolorous but ven
trally lustrous lfts either glabrous overall, or glabrous
ciholate, or thinly finely silky-strigulose dorsally, the
capitula borne, solitary or 2-4- together, in axils of
upper primary lvs and beyond these forming a short
compact, efoliate pseudoraceme commonly nestled in
fohage, sometimes barely exserted; phyllotaxy distic
hous. Stipules firm, lanceolate or narrowly ovate 2-7
m m , nerveless dorsally, deciduous. Lf-formula of
lower or of all lvs iii-vi/46-80, but in var. oligandra
pinnae of upper lvs (in some specimens of all lvs)
only i-ii; lf-stks either (var. vaupesiana) of all lvs or
(var. oligandra) of early lvs distant from inflores
cence 1.5—5(—7.5) cm, of upper lvs of var. oligandra
only 5-17 m m , the longer interpinnal segments
4—11 (-15) m m , the ventral groove bridged at inser
tion of pinnae; pinna-rachises commonly recurved,
the longer ones (5—)6—11.5 cm, the longer interfoho
lar segments 0.5-1.2 m m ; lft-pulvinules 0.2-0.35
m m ; lfts decrescent only at very ends of rachis,
otherwise subequilong, the blades linear from shortly
bluntly auriculate or obtusangulate base, obtuse,
straight or very slightly incurved, the longer ones
(6—)7—11.5 x 1-1.9 m m , 5-8.5 times as long as wide;
venation of 4—5 simple nerves from pulvinule, the
midrib forwardly displaced to divide blade 1:2-2.5,
one submarginal anterior and the inner posterior
nerve produced, parallel to midrib, to blade's apex,
the outer 1-2 posterior ones much shorter, the whole
venation delicately prominulous on both faces or
immersed on upper one. Peduncles mostly 6-22 m m ,
exceptionally to 4 cm, ebracteate; capitula densely
5-10-fld, the receptacle 1.5-2.5 x 1.5-3 m m (rarely a
fl downwardly displaced on peduncle); floral bracts
ovate or deltate-acuminate 1.5-5 x 1-2.5 m m ,
caducous; pedicels (best seen in longitudinal section
of fl) broadly turbinate or dmm-shaped 0.5-1 x 0.8-
1.7 m m ; perianth 5-merous but 1-2 pairs of calyx-
teeth sometimes adherent nearly or quite to apex,
the calyx pilosulous overall or glabrescent in patches,
the corolla of firm texture, densely white-silky-
pilosulous overall; calyx campanulate or turbinate-
campanulate 2-8 m m , the depressed-ovate or lanceo
late lobes 0.4-3.5 m m ; corolla 6-10.5 m m , the lobes
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 193
1.7-4 m m ; androecium 14-52(-57)-merous, 3.2-5.5
cm, the stemonozone 1.6-3 m m , the comeously
thickened tube 4-6 m m , the tassel either (var. vaupe
siana) red overall or (var. oligandra) pallid proxi
mally, the tassel pink-purple; no intrastaminal nec
tary; ovary subsessile, pilosulous above middle. Pod
in profile oblanceolate 7-12.5 x 0.9-1.2 cm, densely
velutinous-pilosulous overall with silvery-gray or
brownish erect hairs, the sutural keels in dorsal view
±2 m m wide, the recessed valves hgnescent; ripe
seeds not seen.
The widely divergent and reflexed leaf-axes of C.
vaupesiana contribute to a distinctive facies. The
known populations are geographically segregated
into two varieties in which flowers of different sizes
and filaments different at once in number and color
coincide.
Key to the varieties of C. vaupesiana
1. Lvs scarcely simpler upward along stem, the
pinnae of those at and near base of inflorescence
like those of main stems; calyx 4-8 m m and
corolla 8-10.5 mm; androecium (36-)40-52
(-57)-merous, red throughout; on stony savanna
and about arenitic outcrops at ±250-500 m, in
basins of upper rios Vaupes and Apaporis in
Vaupes, Colombia 126a. var. vaupesiana
1. Lvs simplified distally, the pinnae of those near base of inflorescence 1-jug.; calyx 2-3
m m and corolla 6-9 mm; androecium 14-24-
merous, the tassel pink-purple distally, pallid
proximally; savannas at 100-125 m of s.-w. state of Amazonas, Venezuela, and closely
adjoining Colombia, around the confluence of
rio Orinoco with rios Ventuari and Atabapo in
lat. 3°-4°30,N 126b. var. oligandra
126a. Calliandra vaupesiana Cowan var. vaupe
siana. C. vaupesiana Cowan, 1958, I.e., pi. XXIX.
— " C O L O M B I A : Comisaria del Vaupes, Rio
Kuduyari, Cerro Yapoboda . . . October 3-4, 1951.
Richard Evans Schultes & Isidoro Cabrera 14203."
— Holotypus, U S 21712801; isotypus, NY!.
Stipules linear-lanceolate 3-7 mm; otherwise as
described in key to varieties. Lvs sensitive (Schultes
5533, GH).
In stony savanna and around arenitic outcrops, in
habitats described by R. E. Schultes, Caldasia 3(12):
121-sequ. 1944, locally plentiful on the upper Vaupes
and Apoporis rivers in Vaupes, Colombia, within
0°-2°N, 70°30'-72°30'W. — M a p 47. — Fl. III-IX.
126b. Calliandra vaupesiana Cowan var. oligandra
Bameby, var. nov, a var. vaupesiana foliis inflores-
centiam versus paucipinnatis, floribus minoribus
(calyce 2-3, nee 4-8 m m , corolla 6-9, nee 8-10.5
m m longis), et praesertim androecii oligomeri fila-
mentis 14—24 (nee 36-57) basi pallidis supeme
roseo-purpureis (nee omnino rubris), ulterius distri-
butione disjuncta recedens. — V E N E Z U E L A .
Amazonas: Sabana de Moyo, on right bank of rio
Orinoco 10 k m above mouth of rio Ventuari, 31 Jul
1959 (fl, fr), J. J. Wurdack & L. S. Adderley 43701.
— Holotypus, NY.
Stipules narrow-ovate 2-3 m m ; otherwise as de
scribed in key to varieties.
Locally plentiful on white-sand savannas at
100-125 m along the Orinoco and immediate tribu
taries within lat. 3°-4°30'N, in s.-w. Amazonas,
Venezuela and (on lower rio Atabapo) in adj. Vaupes,
Colombia. — M a p 47. — Fl. VII-IX.
127. Calliandra pakaraimensis Cowan, Mem. New
York Bot. Gard. 10(1): 142. 1958. — "B. Maguire,
W. M. C. Bagshaw, & C. K. Maguire 40561... at Im-
baimadai, elev 550 m, Mt. Ayanganna, Pakaraima
Mountains, British Guiana, Feb. 1, 1955." — Holo
typus, NY!; isotypi, K!, US!. FiG. 30
C. resupina Cowan, Mem. New York Bot. Gard. 10(4): 65, fig. 45. 1961. "VENEZUELA. Bolivar: . . in drainage of
Rio Aponguao, alt. 1200 m, Gran Sabana . . . 27-28 Mar
1952, Bassett Maguire 33643r — Holotypus, US
22818201; isotypus, NY!.
Shrubs with slender fuscous defohate trunks, either
erect 0.5-2.5 (one record, Pipoly 7669, NY, to 7.5) m
or procumbent and potentially rooting from humistrate
nodes, the young stems, lf-stks, and inflorescence
densely silky-pilosulous or tomentulose with gray or
sordid hairs to 0.3-1 m m , the firm narrow lfts dark-
olivaceous and glabrous (but either dull or lustrous)
above, paler and either glabrous or minutely strigu
lose, or loosely pilosulous beneath, commonly ciho
late, the capitula of silky-pubescent fls forming at
anthesis an efohate terminal pseudoraceme, this some
times in fmit surpassed by a renewed fohate axis;
phyllotaxy distichous. Stipules firm, ovate-acuminate
or lanceolate 8-14 x 3-7 m, densely pubescent
dorsally, glabrous and venulose within, imbricate on
the developing inflorescence axis, deciduous. Lf-
formula i-ii(-iii)/24—37(-41); lvs widely spreading
and pinnae often arcuately recurved, the lf-stk
7-23(-35) m m , the stout petiole including pulvinus
(4—)5-ll m m , at middle (1-)1.4—2.2 m m diam, the
1(2) interpinnal segments a little longer or shorter;
rachis of longer (or only) pinna-pair 6-12 cm, the
longer interfoholar segments 1.4—2.5 m m ; lft-pulvin
ules 0.2-0.6 x 0.7 x 1.2 m m , the lft-blades essentially
194 M E M O I R S OF T H E N E W Y O R K BOTANICAL G A R D E N [VOL. 74(3)
M a p 47. Distribution of Calliandra vaupesiana Cowan
var. vaupesiana and var. oligandra Bameby in northern
South America.
sessile against rachis; lfts decrescent at each end of
rachis, contiguous or narrowly imbricate, in outhne
hnear or broad-linear from tmncate or shortly bluntly
auriculate base, straight or incipiently falcate, obtuse,
plane, or low-convex ventrally, or revolute along ante
rior margin, those near mid-rachis (12-) 13-22 x
2.4—3.6 m m , (4—)4.8-6.8 times as long as wide; vena
tion palmate, of 3-4 parallel, simple nerves ascending
nearly to apex of blade and sometimes in addition one
or more much shorter posterior ones, the midrib dis
placed to divide blade 1:2-3, the principal nerves
bluntly prominulous on both faces or (except midrib)
almost immersed beneath. Peduncles solitary or 2-3
together, 6-21 m m , either ebracteate or bracteate close
under the first fls; floral bracts rudimentary; capitula
9-13-fld, the receptacle (sometimes prolonged into a
short pedestal) ±2-4 m m ; fls homomorphic as to peri
anth, this commonly 4-, sometimes 5-merous, always
white-silky, but the corolla lobes sometimes glabres
cent toward the margin; pedicels often poorly differen
tiated externally, 0.3-0.8 x 0.5-1.1 m m ; calyx
(2.6-)3^1 x 2-2.2 m m , the triangular teeth 1-1.9 m m ;
corolla (5.6-)6-8.5 m m , the lance-ovate lobes 2.2-3.3
m m ; androecium (10-)14-22(-34)-merous, ±3.5 c m
(poorly observed, probably sometimes longer), the
tube 2.4-4.1 m m , the stemonozone (0.7-)1.2-1.7 m m ;
ovary subsessile, at anthesis densely villosulous; no
disc seen. Pods solitary and 2-4 per capitulum, erect,
in profile 5-9 x 0.8-1 cm, gray-brown-tomentulose
overall, the sutural keels in dorsal view 2.5-3 m m
wide, the stiffly hgnescent recessed valves indistinctly
transverse-venulose; seeds unknown.
In xeromorphic brush communities on and near
sandstone outcrops and on sandy savanna, 460-1385
m, localized within 5°-7°N and 60°-62°W on Gran
Sabana in e. Bolivar, Venezuela and on upper forks of
Mazaruni River in adj. Guyana. — M a p 48. — FL
I-VI, IX.
Calliandra pakaraimensis is the east Guayanan
counterpart of west Guayanan C. vaupesiana, different
in coarser growth, longer, wider and on the average
fewer leaflets per pinna, but not in any fundamental
character. The typus of C. resupina is depauperate, but
certainly, in light of contemporary collections, not tax
onomically distinct.
128. Calliandra rigida Bentham, London J. Bot. 5:
103. 1848. — "British Guiana, [Robert] Schom-
burgk," the locality furnished by Schomburgk.
1847, infra: "aus dem Flussgebiete des Carimani
oder Camarang, eine Zuflusses des Mazaruni." —
Holotypus, K (hb. Benth.)! = N Y Neg. 1964; isoty
pus, ̂ B = F Neg. 12541. — C. hookeriana Schom
burgk, Linnaea 20: 754. 1847, nom. substit. illegit.
— Feuilleea rigida O. Kuntze, Revis. Gen. PL 1:
189. 1891.
Microphyllidious shrubs 1-3 m with slender, either
erect or trailing defoliate trunks and virgate new
branches clad in upwardly imbricate lvs but lacking
axillary brachyblasts, the young stems pilosulous
with mixed gray and brown hairs to 0.3-0.7 m m , the
narrow, contiguous or imbricate lfts glabrous except
for random cilia, dark olivaceous on both faces, low-
convex and sublustrous ventrally, the capitula of
bright red fls crowded into a short terminal, efoliate
or only proximally foliate pseudoraceme; phyllotaxy
distichous. Stipules ovate-triangular 1.5-4.5 x
1.2-2.2 m m , pubescent dorsally, weakly 7-9-nerved,
deciduous. Lf-formula \l(25-)21-45; lf-stks stout
5-14 m m , the petiole including pulvinus 1-6 m m , at
middle 0.6-1 m m diam, the one interpinnal segment
not much longer; pinnae subequilong or the distal
pair somewhat longer, the rachis of longest 3.5-6 cm,
the longer interfoholar segments 0.8-1.1 m m ; lft-pul
vinules 0.05-0.15 x 0.3-0.45 m m , the lfts essentially
sessile against rachis; lfts abmptly decrescent at each
end of rachis, otherwise subequilong, the blades
linear from deltately auriculate base, obtuse or ob
scurely apiculate, those near mid-rachis 6.5-10.5 x
0.9-1.7 m m , 6-9.5 times as long as wide; venation
seemingly of 3 slender parallel nerves extending the
length of blade, the midrib displaced to divide blade
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 195
FIG. 30. Calliandra pakaraimensis Cowan.
196 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
M A P 48. Distribution of Calliandra pakaraimensis Cowan in Venezuela and Guyana. M a p 49. Distribution of Calliandra rigida in Venezuela
and Guyana.
±1:2, the 2 posterior primary nerves nearly or quite as
long, sometimes a fourth very short outermost one,
all these simple or almost so, immersed on upper
face, faintly prominulous beneath. Peduncles mostly
geminate, 1-2.8 cm, bracteate at or above middle, the
bract deciduous; capitula ±8—14-fld, the oblong
receptacle including short terminal pedestal 3.5-5 x 2
m m ; bracts 0.5-0.9 m m , tardily deciduous; fls homo
morphic except the terminal one a trifle wider than
the rest, the peripheral ones either staminate or bisex
ual, the perianth of all 4-merous glabrous; pedicels
stout 0.4—0.6 x 0.6-0.8 m m ; calyx of peripheral fls
1-1.5 x 1.2-1.3 m m , the depressed-deltate teeth 0.2-
0.3 m m ; corolla 4—5.2 m m , the ovate lobes 1.4—2.3
m m ; androecium 6-7-merous ±4-5 cm, the tube 1.4—
2 m m , the stemonozone <1 m m ; ovary sessile
glabrous. Pods in profile 6-11 x 0.8-0.9 cm, the
sutural keels in dorsal view ±3-4 m m wide, the
recessed valves sinuously obhquely venulose, the
whole becoming fuscous or nigrescent, minutely pilo
sulous overall with erect hairs; seeds (not seen fully
ripe) light brown, smooth: pleurogram 0.
In brush thickets on and around arenitic outcrops,
1000-1100 m, localized on slopes of the Pakaraima
Mts. in e. Bolivar (Gran Sabana), Venezuela, and im
mediately adj. Guyana (sources of Mazaruni R.). —
M a p 49. — Fl. II, VI-IX, perhaps at intervals
throughout the year.
Calliandra rigida has crowded, imbricately up
swept leaves that recall the Bahian C. calycina, but
the parallel leaflet-venation of compatriot Guayanan
species. It is readily distinguished from all these (ser.
Tsugoideae) by the glabrous perianth, and further
from its close neighbor C. pakaraimensis by smaller
leaves and leaflets, much smaller flowers, and 6-7-
merous androecium.
IV/D. Series C O M O S A E Barneby
Calliandra sect. Calliandra ser. Comosae Bameby,
ser. nov, ab aliis sect. Calliandrae seriebus inflo-
rescentia terminali aphylla paniculatim ramosa ab-
stans. — Sp. typica: C. comosa (Swartz) Bentham. Anneslia Comosae Britton & Rose, 1928: 52, nom. nud.
Shmbs or treelets; lf-formula (i-)ii-iii/7-16, the
longer lfts ±1-2 cm; inflorescence a terminal efoliate
panicle of pseudoracemes; capitula 3-7(-8)-fld; an
droecium ±26-34-merous, the tassel white. — Spp. 2,
endemic to Jamaica.
The condensed terminal panicle of pseudoracemes
produced above the foliate stem of the year was de
scribed by Bentham (1875: 551) as very peculiar and
is not exactly duplicated elsewhere in Calliandra.
The two described Comosae are poorly known from
few localities and not strongly differentiated.
129. Calliandra comosa (Swartz) Bentham, London
J. Bot. 5: 104. 1846. Mimosa comosa Swartz, Prodr.
85. 1788. — "Jamaica," the locality more exactly
stated by Swartz, FL Ind. Occ. 2: 980. 1798: "in
rupibus montium Jamaicae borealisV — Holo
typus, S n.v; clastotypus (If), NY!; isotypus, B M ! =
N Y Neg. s.n. — Inga comosa Willdenow, Sp. PL 4:
1026. 1804. Pithecolobium comosum Bentham,
London J. Bot. 3: 221. 1844. Feuilleea comosa O.
Kuntze, Revis. Gen. PL 1: 187. 1891. Anneslia co
mosa Britton & Rose, N. Amer. Fl. 23: 74. 1928.
Mimosa 9. Fruticosa erecta inermis ... R Browne, Civ. Nat. Hist. Jamaica 253. 1756.
Calliandra comosa sensu Bentham, 1875: 551; Fawcett &
Rendle, 1920: 144; Adams, Fl. PL Jamaica 334. 1972.
Erect slender shrubs 1-2.5 m or reportedly
arborescent to 7 m, with terete gray older branches,
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 197
densely foliate distally, glabrous or almost so, the lvs
bicolored, dark green sublustrous above, paler dull
beneath, the inflorescence a condensed but shortly
exserted, terminal efohate panicle of pseudoracemes;
phyllotaxy distichous. Stipules firm, deltate or
depressed-deltate 0.4—1.2 m m , ±7-nerved when
young, persistent. Lf-formula (i-)ii-iii/9-16; lf-stks
3-10 cm, the petiole ±1-2 cm, the longer interpinnal
segments 8-16 m m ; pinnae distally accrescent, the
rachis of further ones 4—9.5 cm, the longer interfoho
lar segments ±4—7 m m ; lfts proximally decrescent,
otherwise subequilong, the pulvinules almost 0, the
blades sessile against rachis, in outline inequilaterally
lanceolate or lance-oblong from deeply semicordate
base, obtuse, straight or incipiently falcate, the longer
ones 12-19 x 3.5-5 m m , 3.4-4.8 times as long as
wide; venation finely prominulous, palmate-pinnate,
the primary nerves 4—5, the midrib subcentric, the
inner posterior primary nerve produced well beyond
mid-blade. Primary axis of panicle 4—10 cm, errati
cally branched, the interval between successive
bracteiform stipules ±2-3 mm; peduncles solitary
6-16 mm, bracteate distally; capitula 3-6-fld, the
receptacle not over 1 m m diam, the fls sessile, homo
morphic, the glabrous perianth mostly 4-merous, the
calyx weakly striate, the greenish corolla not so, and
not thickened at base; calyx campanulate ±2 x 2.5
mm, the broad obtuse teeth ±0.5 mm; corolla ±6 mm,
the lobes 1.5 m m ; androecium (?-)34-merous, 17-20
mm, the stemonozone 1.4 m m , the tube ±6 mm, the
tassel white; intrastaminal disc ±1.2 mm, lobulate;
ovary at anthesis glabrous, gray-strigulose after fertil
ization. Pods ±9 x 0.7 cm, falcately curved, not seen
fully ripe, the sutures and valves thinly strigulose.
On limestone rocks, ?-700 m, apparently rare and
local in centr. and n. Jamaica. — Fl. I—III (-?).
130. Calliandra paniculata C. D. Adams, Phytologia
20: 310. 1970. — "C. D. Adams 11154 . . . between
Two Rivers and Gut River, Manchester Parish,
Jamaica, ... 13 May 1962." — Holotypus, U C W I
n.v; isotypus, BM!.
C. paniculata sensu Adams, Fl. PL Jamaica 334. 1972.
Slender shrubs ±1-2 m, resembling C. comosa in
habit and especially in the terminal efoliate panicles
of pseudoracemes and in falcate pods, perhaps not
specifically distinct but apparently different in evi
dent (but thin) pubescence, in manifestly pulvinulate
lfts, in proportionately wider lfts and in somewhat
smaller calyces, the lf-stks and axes of inflorescence
pilosulous with gray hairs to 0.5 m m , some lfts
remotely ciholate. Stipules lance-subulate or triangu
lar ±2-2.5 mm. Lf-formula ii/7-10; lf-stks 14-22
mm, the petiole 4—12 mm, the one interpinnal seg
ment as long or longer; rachis of distal pinnae 3.5^-
cm, the longer interfoholar segments to 5 m m ; lft-
pulvinules ±0.45-0.6 x 0.4 mm; lfts scarcely gradu
ated, the blades elliptic or oblong-elliptic from
broadly semicordate base, broadly obtuse apiculate,
the larger ones ±12-15(-18) x 4.5-6(-7) m m , 2.2-3
times as long as wide. Inflorescence scarcely 1 dm
diam; peduncles solitary and (few) geminate, the
longer ones 12-18 mm; capitula 3-6-fld, the flowers
obscurely pedicellate, the perianth greenish, glabrous
except for microciliolate calyx, this weakly ±10-
nerved; calyx shallowly turbinate-campanulate ±1.1
x 1.5 mm, the depressed-deltate teeth 0.2 mm; corolla
5.4 mm, the ovate lobes 2 mm; androecium 26-mer-
ous, 19-20 mm, the tube 6.3 mm, the stemonozone
and intrastaminal disc of C. comosa. Pods not seen
ripe, in youth gray-pilosulous, apparently otherwise
like those of C. comosa.
On rocks at ±6 m, known only from the type-local
ity on the s. coast of Jamaica near 77°30'W. — Fl.
IV-VI.
V. Sectio MICROCALLIS Barneby
Calliandra sect. Microcallis Bameby, sect, nov, in-
florescentia terminali saltern distaliter efoliata,
floribus minuscuhs vinaceis (calyce 0.8-1.4 m m ,
corolla 1.6-2.8 m m tantum longis), androecioque
minimo (5-7 m m longo) praestans; species duae,
foliorum formula stipulisque inter se quam maxime
diversis, utraque igitur seriem peculiarem effor-
manti. — Sp. typica sub ser. Microcallide indicate.
Distinguished by the syndrome of terminal pseudo
racemose inflorescence at least distally efoliate, and
minute fls (calyx 0.8-1.4 mm, corolla 1.6-2.8 m m
long) with androecium only 5-7 m m long. — Spp. 2,
of tropical extra-Amazonian Brazil, Paraguay, and n.-
e. Bohvia.
The two species of sect. Microcallis were assigned
by Bentham (1875: 413, 427) to mutually remote
series of Calliandra, the first, C. leptopoda, being
associated with long-pedicellate species of the pre
sent sect. Androcallis, and the second, C. parviflora,
tacked onto the large-flowered North American ser.
Racemosae (= ser. Calliandra), where it was certainly
misplaced. Given the great difference between them
in many morphological characters, the tiny short-
stamened flowers common to both may perhaps have
198 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
been independently derived, but from what source, or
from what sources, I cannot guess. In order to give
due emphasis to the differences, I have referred each
species to an independent monotypic series.
V/A. Series MICROCALLIS
Calliandra sect. Microcallis Bameby ser. Microcal
lis. C. sect. Microcallis Bameby, sens. str. —
Species unica: C. parviflora Bentham.
Shmbs 1-2.5 m; stipules heteromorphic, the lowest
in each growth-cycle perulate, the upper ones linear-
setiform; lf-formula x-xxxvi/30-49; longer lfts ±2-
3.5 m m ; peduncles to 6-13 mm; pedicels almost 0. —
Sp. 1, mainly of campo in tropical extra-Amazonian
Brazil, Paraguay, and n.-e. Bolivia.
131. Calliandra parviflora Bentham, London J. Bot.
3: 112. 1844. — "Brazil (Minas Geraes ?), Sello,
Pohl, Claussen, LangsdorffV Lectoholotypus,
Claussen 119, K (hb. Benth.)!. FiG. 31
Acaciapachyloma Martius, 1875: 557, in syn., nom. nud. —
Spma authentica: Martius Hb. Fl. Bras. 1108, K!, NY!. —
Feuilleea pachyloma O. Kuntze, Revis. Gen. PL 1: 186.
1891. Calliandra parviflora sensu Bentham, 1876: 427, t. CX;
Glaziou, 1905; Bernardi, 1984: 171; Lewis, 1987: 175.
Slender, elaborately microphyllidious, densely
leafy shrubs 1-2.5 m, several-stemmed from base and
freely branching distally, the older trunks defoliate
glabrate lenticellate, the young stems and all lf-axes
pilose or pilosulous with fine, widely ascending or
subappressed, white or partly brownish hairs to
0.4-1.3 m m , the dark-ohvaceous lvs concolorous, the
minute, closely imbricate lfs glabrous facially, some
times remotely appressed-ciliolate, the inflorescence
composed of axillary and terminal, efoliate or hyster-
anthously foliate pseudoracemes of small, subglo-
bose and incipiently spiciform capitula fasciculate at
each node on subfiliform peduncles, the fmits devel
oping either on efoliate axes or immersed in delayed
fohage; phyllotaxy distichous. Stipules heteromor
phic: a) the lowest of each annual growth-comple
ment forming, contemporary with the fmit, an ovoid
resting-bud axillary to a mature primary If of the pre
ceding cycle, the imbricate blades ovate or oblong-
ovate ±1.5-3 m m , striate, glabrous dorsally but often
lanulose-ciliolate, persisting at base of the new
branchlets, b) those subtending the lowest lvs of
new branchlets linear-oblong, transitional in form
between a) and c); and c) at fohate nodes of mature
branchlets and subtending hysteranthous lvs of inflo-
FlG. 31. Calliandra parviflora Bentham. Reproduced
from Bentham in Martius, Fl. bras. 15(2): t. 110. 1876.
rescence-axes, linear-lanceolate or setiform 2-9 x
0.2-0.6 m m , incurved, 1-nerved or externally nerve
less, eventually deciduous. Lf-formula (x-)xiv-
xxxvi/30-49, the leaflets of largest lvs 3000+, seldom
<1000 in any; lf-stk of longer lvs (4-)5-12(-17) cm,
the petiole, consisting largely of obese pulvinus,
2-6(-12) m m , the longer interpinnal segments 2.5-6
m m , the ventral groove continuous; pinnae sub
equilong, the rachis of longer ones 14-32(-38) m m ,
the longer interfoholar segments 0.2-0.5 m m ; lft-
pulvinules 0.1-0.25 m m , mostly yellowish, not wrin
kled; lfts subequilong, the blade hnear or linear-
lanceolate from obtusely auriculate base, acute or
mucronulate, the larger ones (1.8-)2.2-3.5 x 0.4-0.8
m m , 4.4-6.5(-7.2) times as long as wide, all weakly
1-nerved, the simple costa subcentric at mid-blade, a
short posterior primary nerve sometimes faintly per
ceptible. Inflorescence units (l-)2-8(-9) per node,
each 3-14-fld, the subfihform ebracteolate peduncles
of unequal length, the first at each node of the
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 199
pseudoraceme seldom <6 and to 13 m m , the floral
axis becoming 0.5-3.5(-5) m m ; bracts spreading-
incurved, lanceolate or ovate 0.45-0.9 m m , either
ciholate or glabrous, persistent; fls homomorphic
except some staminate, sessile or almost so, the
obconic pedicel no more than 0.2 m m ; perianth crim
son, glabrous or microscopically pubemlent distally,
both calyx and corolla submembranous 5-nerved,
bluntly pentagonal; calyx turbinate-campanulate
0.8-1.3 m m , the incurved teeth 0.2-0.5 mm; corolla
1.6-2.4 m m , the ovate lobes 0.6-1.2 m m ; androe
cium (6-)7-ll(-"12")-merous, when straightened
out 5-7 m m , the stemonozone ±0.3-0.4 mm, the
membranous tube 0.8-1.3 m m , the cupular intrasta
minal disc 0.4—0.75 m m tall; ovary (often wanting)
pubemlent in age, becoming densely brown-strigu
lose after fertilization. Pods erect on thickened
peduncle, solitary or rarely 2-3 per capitulum, in
profile linear-oblanceolate, when well fertilized
5-11 x 0.6-0.9(-1.2) cm, 5-8-seeded, the dilated
sutural ribs 2.5-4.5 m m wide in dorsal view, the
recessed hgnescent valves plane, faintly if at all venu
lose, the whole densely pilosulous with loosely
appressed-ascending, brownish and gray hairs; seeds
(few seen) in broad profile obovate ±4.5-5 x 3-3.5
m m , the testa pale brown or faintly fuscous-mottled,
smooth, the U-shaped pleurogram ±3.5 x 2 mm.
In cerradao, in campo cerrado, on bush-islands in
palm-savanna, at gallery margins and in other types
of brush-woodland, sometimes surviving disturbance
in hedgerows, 170-800, infrequently up to 1000 m,
locally plentiful over much of the Brazilian Planalto,
from the upper Rio S. Francisco valley in Minas
Gerais and s.-w. Bahia w. to the upper Paraguai basin
and n.-e. Bohvia, thence n. in Brazil to the middle
Araguaia valley in s.-e. Para and to the upper Par-
naiba valley in Maranhao, within latitude 6°-24°S
and 44°-62°W. — Map 50. — Fl. X-III(-IV). —
Angiquinho.
Calliandra parviflora is notable for the syndrome
of: high leaf-formula and proportionately minute
leaflets; perulate resting buds; efoliate (or some
hysteranthously foliate) pseudoracemes of tiny
carmine flowers; short, mostly 6-11-merous androe
cium; and a cupular intrastaminal nectary in all
flowers, whether staminate or bisexual. Similar rest
ing buds and similarly decompound leaves are com
mon in the family Mimosaceae; but all such differ
at anthesis from C. parviflora either by extrafloral
nectaries, or by white to yellow perianth and androe
cium, or by free stamens, or by some combination
of these characters.
V/B. Series L E P T O P O D A E Barneby
Calliandra sect. Microcallis ser. Leptopodae Bar
neby, ser. nov. monotypica, stipuhs omnibus magnis
foliaceis, foliorum formula i/3-5, foliolis majoribus
17-31 m m usque longis, pedicelhs productis um-
bellatim radiantibus ±1-2 cm longis, flosculisque
minutis brevistamineis fere C. parviflorae persingu-
laris. — Sp. unica: C. leptopoda Bentham.
Subshrubs and small shmbs to 1.5 m; stipules foli
aceous, homomorphic; lf-formula i/3-5; longer lfts
1.7-3.1 cm; pedicels to 1-2 cm, radiately spreading;
perianth and androecium of C. parviflora. — Sp. 1,
mostly of caatinga in tropical e. Brazil, from Piaui
and Pemambuco s. through interior Bahia to n.-e.
Minas Gerais.
132. Calliandra leptopoda Bentham, London J. Bot.
3: 101. 1844. — "Brazil, Serra Acuma, Blanchet, n.
2833, between Boa Esperanza and Santa Anna das
Merces [near Jaicos, Piaui], Gardner, n. 2138." —
Lectotypus, Blanchet 2833, K (hb. Benth.)! = N Y
Neg. 7955; isotypi, BM!, K (hb. Hook.)! = IPA Neg.
1465, N Y (2 sheets)!, P!; paratypi, Gardner 2138,
BM!, K!. — Feuilleea leptopoda O. Kuntze, Revis.
Gen. PL 1: 188. 1891. FiG. 32
C. leptopoda sensu Bentham, 1875: 544; 1876: 413, t. CVI; Lewis, 1987: 174, fig. HE; Rizzini & Mattos Filho, Min-istro Ambiente (Rio de Janeiro) ser. Estudos e Con-tribucoes9: 11. 1992.
Slender macrophyllidious several-stemmed shrubs
or subshrubs 6-15 dm with conspicuously dilated
foliaceous stipules, either glabrous throughout except
for a tuft of pallid hairs in the axil of some older lvs,
or for sometimes remotely cili(ol)ate lfts, or thinly
pilose faces of some lfts, the ample thin-texmred lfts
either green or glaucescent, a little paler dorsally, the
small umbellate crimson-red fls borne on subfihform
pedicels of variable length, the umbels at first axillary
to lvs but by suppression of distal lvs forming a
broad-bracteate pseudoraceme projected well above
fully expanded foliage. Stipules broadly reniform or
suborbicular from shallowly cordate, semiamplexi-
caul base, obtuse or apiculate, the larger ones 12-26
x 11-35 mm, all flabellately 7-11-nerved from base
and openly reticulate-venulose. Lf-formula i/3-5; lf-
stks (12-) 15-28 mm, at middle 0.4-0.8 m m diam;
rachis of longer pinnae 2-4.5 cm, the longer interfo
holar segments 7-12 m m ; lft-pulvinules 0.6-1 m m ,
cross-wrinkled; lfts little graduated, the blade broadly
oblong-elhptic or elhptic-obovate from shallowly
broadly semicordate base, obtuse or depressed-deltate
200 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
M A P 50. Distribution of Calliandra parviflora Bentham in South America
at apex, the larger ones 1.7-3.1 x 0.8-1.5 cm, 14-22
times as long as wide; venation palmate-pinnate, the
slightly excentric midrib giving rise on each side to
±3-6 major (and random intercalary) secondary
nerves, brochidodrome well within the plane margin,
the inner posterior primary nerve incurved-ascending
to or somewhat short of mid-blade, the whole venation
finely sharply prominulous and palhd beneath, usually
a httle less so above. Peduncles (1—)2—4 per node,
ebracteolate, the longer ones of a fascicle 2.5-4 c m x
0.2-0.3 m m , the floral receptacle globose, <1 m m
diam; umbels 11-21-fid, the fls homomorphic, the fil
iform pedicels ± unequal, the longer ones of any umbel
±1-2 c m x 0.1 (-0.15) m m , the 4—6 outermost ones
subtended by an involucelliform ring of ovate acute
green but submembranous bracts 1-4 m m , the inner
ones ebracteate; perianth glabrous, green or red, the
calyx weakly 5-nerved, the corolla externally nerve
less; calyx campanulate 0.8-1.4 m m , the obtuse teeth
0.3-0.7 m m ; corolla 2.5-2.8 m m , the erect ovate lobes
±1.2 m m ; androecium 27-30-merous, 6-7 m m , the
stemonozone 0.3-0.6 m m , the campanulate tube 1.2-
1.5 m m , the filaments dark wine-red, uniseriate, the
intrastaminal disc 0.65-0.8 m m tall; ovary substipitate,
glabrous. Pods 1-2 per umbel, pliantly geotropic, in
broad view oblanceolate ±6-8 x 0.9-1.2 cm, the
sutural ribs relatively slender, <1 m m wide in dorsal
view, the lustrous papery, pale brown valves bullately
distended over each of 5-8 seeds; dehiscence not seen;
seeds (httle known) in broad view 6 x 3.5 m m , the testa
pale brown, dull, the shadowy pleurogram not incised.
In the caatinga formation, surviving in roadside
capoeira, and about outcrops on morros, ±300-500
m, apparently local in e. tropical Brazil within lat.
7°-17°S: e-centr. Piaui, w. Pemambuco, and Alagoas,
s. through the Chapada Diamantina of interior Bahia
just into n.-e. Minas Gerais. — M a p 51. — Fl.
(XI-)I-IV.
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 201
FiG. 32. Calliandra leptopoda Bentham. Reproduced from Bentham in Martius, Fl. bras. 15(2): t. 106. 1876.
A delightfully idiosyncratic plant, resembhng C.
parviflora in size and stmcmre of perianth and an
droecium, but in other respects extremely different,
and isolated in the genus.
Acknowledgments
It is a pleasure to express my gratitude to friends,
colleagues, and foundations that have contributed to
preparation and editing of this book. In particular to:
Melissa Luckow (BH) and James W . Grimes (MEL,
formerly N Y ) for careful peer reviews and valuable
professional advice. Dr. Grimes, furthermore, put sev
eral versions of the original typescript into electronic
form, mounted the distribution maps, and helped
M A P 51. Distribution of Calliandra leptopoda Bentham
in eastern Brazil.
organize the List of Exsiccatae. His aid was precious
and indispensible.
William S. Moye HI, for 25 evocative portraits of
Calliandra species that inject a lively reality into the
technical descriptions.
William R. Buck (NY), meticulous editor, who pa
tiently and cheerfully piloted m y manuscript through
the straits of review and revision.
Curators of herbaria A, ARIZ, B H , B M , BR, C,
CAS, CEPEC, COL, F, Fl, G, GH, GOET, H A M B ,
HUEFS, K, M A , M B M , MICH, M O , OXF, P, POM,
R, RB, S, TEX, UB, UC, US, VEN, and W for loan
of specimens and/or permission to consult the collec
tions in their care.
Correspondents M. M. Arbo (CTES), S. G. Beck
(LPB), A. de Carvalho (CEPEC), A. G. Fernandes
(EAC), G. Hatschbach (MBM), L. R. Landrum (ASU),
G. P. Lewis (K), D. J. Macqueen (OXF), L. P. de
Queiroz (HUEFS), B. Stergios (PORT), T R. Van
Devender (ARIZ), and R. O. Vanni (CTES) for gifts of
select specimens.
E. Forero and S. Mori, former and current directors,
respectively, of the Institute of Systematic Botany at
NY, for support of long-term research in Mimosaceae
tribe Ingeae.
J. E. Runyon and colleagues in Scientific Pubhca
tions at N Y for skilled and devoted attention to tech
nical aspects of publication.
The Director and staff of the Library at N Y
for procuring copy of rare pubhcations bearing on
Calliandra.
D. Brunner ( M O ) and E. Forero (COL, formerly
N Y and M O ) for loan of an important manuscript
paper on Calliandra of Venezuela Guayana.
202 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
The curatorial staff at NY, particularly S. Mi sir,
who over the years have handled and recorded a brisk
traffic of gifts and loans.
Finally, the National Science Foundation, throught
grant # 9420283 to The New York Botanical Garden,
and the Eppley Foundation, for generous funding of
research on Mimosaceae.
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204 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Species Incertae vel M i n u s Cognitae
Acacia selloi Sprengel, Syst. Veg. 3: 137. 1826. — "Brasil,
Sello." — No typus known to survive and the protologue
taxonomically ineffective. Surmised by Bentham to be
equivalent to C. brevipes. — Calliandra selloi (Sprengel)
Macbride, Contr. Gray Herb. 59: 5. 1919.
Calliandra grandifolia P. H. Allen, Rain Forests of Golfo
Dulce 148, 409. 1956. — "[Costa Rica.] Esquinas Forest,
between Golfo Dulce and Rio Terraba, prov. Puntarenas,
100 ft., Skutch 5362." — Holotypus, EAP n.v. — Closely
related to C. brenesii, but leaves supposedly larger.
Calliandra grisebachiana var. carolae Spegazzini, Revista
Argent. Bot. 1: 79, 187. 1926. — Described as closely
resembhng var. grisebachiana, but of dwarf stature. Not
seen, perhaps Cojoba sp.
Calliandra kuhlmannii Hoehne, Com. Linh. Telegr., Bot.
8: 20, est. 134. 1919. — "... nas matas que margeiam o
rio Arinos. . . . Nos. 461-463 do Sr. J. G. Kuhlmann." —
Described as related to Calliandra filipes (= Zapoteca fil-
ipes (Bentham) H. Hernandez), but perhaps close to C.
laxa var. stipulacea (Bentham) Bameby.
Calliandra orinocensis Pittier, 3rd Conf. Interam. Agric.
Caracas 354. 1945, nom. nud.
Calliandra pubens Renvoize, Kew Bull. 36: 79, fig. 4(34),
5D. 1981. — "BAHIA. Sera [sic] do Curral Feio, Lagoinha
to Minas do Mimoso, Harley & al. 16694A." — Holotypus,
K; isotypus, CEPEC, neither seen, perhaps overlooked.
The cited figure suggests a form of C. bahiana Renvoize,
sympatric in Serra do Curral Feio.
Calliandra rondoniana Hoehne, Com. Linh. Telegr., Bot.
12: 6, est. 179. 1922. — "No. 2015 Kuhlmann, margem do
Cautario Grande, Rondonia, no noroeste do Estado de
Matto-Grosso, em Janeiro de 1919." — No type seen; the
protologue suggests C. laxa.
Calliandra seleri Harms, Repert. Spec. Nov. Regni Veg. 17:
90. 1921. — "[Mexico.] Chiapas, distr. Comitan . . . bei
Zapaluta . . . Seler n. 2512. . . ." — No typus found; de
scribed as resembling C. humilis. — Anneslia seleri
(Harms) Britton & Rose, N. Amer. Fl. 23: 62. 1928.
Calliandra stenophylla Wawra, Oesterr. Bot. Z. 29: 215.
1879. — "Brasilia: comm. Schwarz." — Holotypus not lo
cated at W by Dr. B. Wallnofer in July 1994.
Calliandra tamarindifolia Ettingshausen, Blatt-Skel. Dikot.
224, fig. 275. 1861. — "Cultiviert im K.-K. Hofgarten im
Schoenbriinn." — No typus found; only the nervation of
leaflets is described.
Calliandra urbanii Alain, Brittonia 20: 157. 1968. — Holo
typus consisting of 2 leafy twigs, one of them in young
bud, NY!. — The petiolar nectaries exclude the specimens
from genus Calliandra.
Calliandra viscidula var. leucantha Bentham in Martius,
Fl. Bras. 15(2): 423. 1876. — "[Brasil. Mato Grosso:] . . . in campis arenosis Serra da Chapada et Rio Pardo: Riedel
(Herb. Acad. Petropolit.)." — No typus found at K or L.
Calliandra yucunensis N. Mattos, Loefgrenia 71: 3, fig.
1977. — "Estado de Rio Grande do Sul: Tenente Portela,
Parque Estadual do Turvo, no Salto Yacuna, 10-1-1977, J.
Mattos 1644 & N. Mattos." — Holotypus, IPRN n.v. —
The protologue, with crude figure, suggests C. brevipes.
Numerical List of Taxa
I. sect. ANDROCALLIS Bameby A. ser. Androcallis
1 C. pittieri Standley
a. var. pittieri
b. var. polyphylla (Harms) Bameby
2. C. purdiaei Bentham
3. C. glomerulata Karsten
a. var. glomerulata
b. var. parvifolia (Bentham) Bameby
4. C. cruegeri Grisebach
5. C. tolimensis Taubert 6. C. laxa (Willdenow) Bameby
a. var. laxa b. var. stipulacea (Bentham) Bameby
c. var. urimana (Brunner & Forero, ined.)
Bameby 7. C. rubescens (Martens & Galeottii)
Standley 8. C. belizensis (Standley) Standley
9. C. goldmanii Rose ex Bameby
10. C. bijuga Rose
11. C. molinae Standley
12. C. hirsuta (G. Don) Bentham
13. C. peninsularis Rose
14. C. sesquipedalis McVaugh
15. C. californica Bentham
16. C. a. b.
17. C. a. b. c.
18. C. 19. C. 20. C.
20a. C.
21. C. 22. C. 23. C. 24. C. 25. C.
a. b. c. d.
26. C. 27. C.
a. b.
28. C. 29. C.
eriophylla Bentham
var. eriophylla
var. chamaedrys Isely
humilis Bentham var. humilis
var. gentryana Bameby
var. reticulata (A. Gray) L. Benson
tumbeziana Macbride
taxifolia (Kunth) Bentham
parvifolia (Hooker & Arnott) Spegazzini
carrascana Bameby foliolosa Bentham
tweedii Bentham
bella Bentham
subspicata Bentham
dysantha Bentham
var. dysantha
var. macrocephala (Bentham) Bameby var. opulenta Bameby
var. turbinata (Bentham) Barneby gardneri Bentham
macrocalyx Harms
var. macrocalyx
var. aucta Bameby
fernandesii Bameby
ulei Harms
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 205
30. C. pilgerana Harms 31. C. umbellifera Bentham 32. C. imperialis Barneby 33. C. concinna Bameby 34. C. squarrosa Bentham 35. C. glaziovii Taubert 36. C. depauperata Bentham 37. C. silvicola Taubert
38. C. surinamensis Bentham 39. C. samik Bameby
40. C. purpurea (Linnaeus) Bentham
41. C. riparia Pittier
42. C. magdalenae (de Candolle) Bentham a. var. magdalenae
b. var. colombiana (Britton & Killip) Bameby 43. C. caeciliae Harms
44. C. chulumania Bameby 45. C. carcerea Standley & Steyermark 46. C. pityophila Bameby 47. C. colimae Bameby
48. C. hintonii Bameby 49. C. conferta Bentham 50. C. brevipes Bentham 51. C. staminea (Thunberg) Bameby 52. C. sessilis Bentham
53. C. spinosa Ducke 54. C. duckei Bameby 55. C. blanchetii Bentham
56. C. aeschynomenoides Bentham B. ser. Biflorae Bameby
57. C. biflora Tharp C. ser. Chilenses Bameby
58. C. chilensis Bentham D. ser. Pauciflorae Bameby
59. C. pauciflora (A. Richard) Grisebach
60. C. enervis (Britton) Urban
E. ser. Ambivalentes Bameby
61. C. medellinensis Britton & Kilhp 62. C. mollissima (Willdenow) Bentham 63. C. guildingii Bentham
64. C. falcata Bentham 65. C. haematocephala Hasskarl
a. var. haematocephala
b. var. boliviana (Britton) Bameby
66. C. erythrocephala H. Hernandez & Sousa
67. C. rhodocephala J. D. Smith
F. ser. Macrophyllae Bentham 68. C. trinervia Bentham
a. var. trinervia b. var. carbonaria (Bentham) Bameby
c. var. peruicola Bameby d. var. pilosifolia (Cowan) Bameby
e. var. paniculans Bameby
f. var. stenocylix Bameby g. var. arborea (Standley) Bameby
69. C. bombycina Spruce ex Bentham
70. C. glyphoxylon Spruce ex Bentham
71. C. jariensis Barneby 72. C. coriacea (Willdenow) Bentham
73. C. antioquiae Bameby
74. C. angustifolia Spruce ex Bentham
75. C. harrisii (Lindley) Bentham 76. C. tergemina (Linnaeus) Bentham
a. var. tergemina b. var. emarginata (Willdenow) Bameby
77. C. macqueenii Bameby 78. C. brenesii Standley
79. C. laevis Rose
G. ser. Hymenaeodeae Bameby
80. C. hymenaeodes (Persoon) Bentham H. ser. Longipedes Bameby
81. C. longipes Bentham
II. sect. ACISTEGIA Bameby
82. C. haematomma (de Candolle) Bentham a. var. haematomma
b. var. colletioides (Grisebach) Bameby
c. var. correllii Bameby
d. var. rivularis (Urban & Ekman) Bameby
e. var. tortuensis (Alain) Barneby f. var. glabrata Grisebach
g. var. locoensis (Garcia & Kolterman)
Bameby 83. C. pedicellata Bentham
III. sect. ACROSCIAS Bameby 84. C. brevicaulis M. Micheh
a. var. brevicaulis b. var. glabra Chodat & Hassler
IV. sect. CALLIANDRA A. ser. Calliandra
85. C. lintea Bameby 86. C. nebulosa Bameby 87. C. bahiana Renvoize
a. var. bahiana
b. var. erythematosa Bameby 88. C. lanata Bentham 89. C. fuscipila Harms 90. C. feioana Renvoize
91. C. asplenioides (Nees) Renvoize 92. C. viscidula Bentham 93. C. fasciculata Bentham
a. var. fasciculata
b. var. bracteosa (Bentham) Bameby 94. C. linearis Bentham
95. C. elegans Renvoize 96. C. mucugeana Renvoize
97. C. calycina Bentham 98. C. x cumbucana Renvoize
99. C. hirsuticaulis Harms 100. C. crassipes Bentham
101. C. hirtiflora Bentham
a. var. hirtiflora
b. var. ripicola Bameby 102. C. sincorana Harms
103. C. stelligera Bameby 104. C. coccinea Renvoize
a. var. coccinea
b. var. trimera Bameby
105. C. involuta Mackinder & Lewis
106. C. santosiana Glaziou ex Bameby
107. C. renvoizeana Bameby 108. C. longipinna Bentham
109. C. debilis Renvoize 110. C. iligna Bameby
111. C. paterna Bameby 112. C. ganevii Bameby
113. C. erubescens Renvoize
114. C. semisepulta Bameby 115. C. germana Barneby
116. C. luetzelburgii Harms
117. C. hygrophila Mackinder & Lewis
206 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
118. C. houstoniana (Miller) Standley
a. var. anomala (Kunth) Bameby
b. var. colomasensis (Britton & Rose)
Bameby
c. var. acapulcensis (Britton & Rose) Bameby
d. var. calothyrsus (Meisner) Bameby
e. var. houstoniana
119. C. juzepczukii Standley
120. C. palmeri S. Watson
121. C. physocalyx H. Hernandez & Sousa
122. C. wendlandii Bentham
123. C. quetzal (J. D. Smith) J. D. Smith
B. ser. Virgatae Bameby
124. C. virgata Bentham
C. ser. Tsugoideae Bameby
125. C. tsugoides Cowan 126. C. vaupesiana Cowan
a. var. vaupesiana
b. var. oligandra Bameby
127. C. pakaraimensis Cowan
128. C. rigida Bentham
D. ser. Comosae Bameby
129. C. comosa (Swartz) Bentham
130. C. paniculata C. D. Adams
V sect. MICROCALLIS Bameby
A. ser. Microcallis
131. C. parviflora Bentham B. ser. Leptopodae Bameby
132. C. leptopoda Bentham
List of N u m b e r e d Exsiccatae
The numbers in parentheses refer to those in the Numerical List of Taxa.
Abbott, W. L., 1308 (82b). Acevedo-Rodriguez, P., 2398 (65a); 3950 (82b). Acosta P., A., 534 (76b). Acuiia, J., 16664 (65a).
Adams, C. D., 6335 (82f). Adams, D. J., 209 (130). Aguilar H., M., 356 (76b). Alain, Bro., see Liogier, A.
Albuquerque, M. G., 35 (38). Alencar, L., 388 (68d). Alexander, E. J., 871, 1901, 1948 (118a).
Allen, C, 563 (40). Allen, P. H., 1958, 3666 (42a). Almeda,E, 3363(118d). Almeida, J., 104 (23). Alonso, F, 5660 (41); 5620, 5675 (38). Alston, A. H. G., 6049 (41); 6989 (lb); 7075 (63).
Alvarado Flores, A., 161 (118a). Alvarenga, D., 117, 438 (25a). Alvarez, A., 620 (la). Amaral, I. A. de, 1518(125).
Amorim, A. M. A., 1022 (91); 1288 (23); 1681 (97); 1683 (85). Anderson, W. R., 3757 (118e); 6271 (25a); 6823, 6920
(131); 6954 (25a); 6954A (26); 8372, 8715 (91); 8902 (33); 9105 (25a); 9128 (131); 9161 (20); 9720, 10203, 11215 (131); 13134 (118a); 35451 (93b); 35621 (91); 36302 (94); 36432, 36662, 37097 (25a); 37184 (20).
Andrade Lima, D. A., 68-5356 (36). Andre E., 151 Ibis (42a); 1519 (la); 1555 (72); 2755 (68b).
Andrews, L. M., 221, 222 (118a); 279, 279a (16a); 772,
772a (82f). Anhanguera, U. L. de, 5 (25a). Araujo, A. P. de, 19 (91); 319 (27a). Arbo, M. M., 3415 (124); 3561 (26); 3666 (20); 3783 (81);
4236 (94); 4317 (93a); 4569 (25a); 4677 (93b); 4696
(94); 5005 (110); 5685 (100); 5736 (99); 5768 (92).
Archer, W. A., 166 (25d); 178 (la); 2188 (72); 4984 (93b);
4985 (25a); 7268, 7527, 7869 (38).
Argent, G., 6502, 6747 (81). Aristeguieta, L., 1622 (38); 3835, 3972 (64); 5909 (6b);
6412 (4); 6836 (42b).
Arnoldo, M., 3024 (65a). Arsene, Bra, 2159 (118a).
Asplund, E., 5600 (65b); 19399 (74).
Ataide, M., 1 (53); 586 (101b); 587 (56); 593, 599 (101a); 600 (97); 604 (52); 607 (36).
Atta, A. L., 7 (25a). Atwood, N. D., 2183a, 17364 (16a).
Aulistia, M., 407 (74); 2885 (68e).
Aymard, G., 7690 (6b); 7729, 8735 (6a); 8736 (4); 8750
(6a); 9283 (63); 9906 (68a).
Bailey, L. H. & E. Z, 296 (76a). Baker, M. A., 10014 (17a). Balansa, B., 1436 (84a).
Baldwin, J. T., 4513(38). Balick, M. J., 3494, 3590 (118e). Balls, E. K. & Everett, 22923 (16a). Balslev, H., 4359 (74). Bang, M., 1586 (65a); 1603 (65b). Barclay, A. S., 556 (125a). Barkley, F. A., 16012 (16a); 30C293 (42a). Bameby, R. C, 2504 (16a). Bartlett, H. H., 11406 (118a); 11448 (118d); 13123 (76b). Basler Lobo, C. M., 41 (101b).
Bautista, H. P., 856, 906 (132). Bautista, H. & Orlando, 991 (92).
Bautista H. P. & Salgado, 843 (27a). Beard, J. S., 457 (40). Beck, S. J., 370, 1758, 6631 (65b); 8159 (74); 8597 (44);
9268 (65b); 12067, 12162, 12166 (44).
Belem, R. P., 146 (25a); 395 (52); 438 (25a); 1162 (23); 1325, 1806(24).
Belem, R. P. & Magalhaes, 96 (131); 707 (24).
Belem, R. P. & Pinheiro, 2369 (23). Belanger, C. P., 77 (40); 297, 1046 (50). Benitez de Rojas, E., 1021 (3b).
Bercot, A. C. S., 13 (25a). Berlin, B., 554 (39).
Bernal, R., 239, 248 (la); 1552 (38).
Bemardi, L., 1793 (3a); 2497 (4); 2516 (6a); 2727, 3706
(6b); 18311 (84a); 18861 (50); 20510 (20). Bertoni, M.S., 1017 (22); 3768 (21).
Betancour, J., 693 (38); 2956, 3443 (la).
BHCB, 7138 (25a); 8918 (20); 10643 (93a); 10797, 14242,
17560 (20); 18701 (93b); 18852, 18937 (25a); 22241, 24228 (20); 24741 (25b); 26539, 26540 (20); 26543, 26544(91).
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 207
Black, G. A., 40-7012 (16a); 48-2682 (125). Blackmore, S. & Heath, 1755 (66); 2041 (76b). Blanchet, J. S., 2105 (24); 2584 (55); 2620 (92); 2816 (52);
2833 (132); 3315 (52); 3896 (56). Blumer, J. C , 1677 (17a).
Boom, B. M., 5719 (72); 7782 (68e); 9421 (125); 10541 (6b). Bordas, - & Schinini, 20557 (81).
Box, H. E., 1724, 1997 (76a).
Brace, L. K. J., 1477 (82a); 1695, 3695, 3956, 4030 (82c). B R A D E , 39283 (24).
Brandegee, T. S., 194, 4722 (15)
Bredemeyer, F, 15 (6a).
Breedlove, D. E., 9789 (118e); 20768 (118d); 20923 (9); 22255 (118d); 23585, 25277 (76b); 28553, 25950 (42a);
31263 (76b); 35769 (14); 36688 (9); 42197 (76b); 47004
(16a); 50390 (43); 52779 (119); 54121, 56339 (9); 60645 (12); 60781 (15); 61597 (66); 62282 (15).
Brenes, A. M., 4424 (78); 4443, 6508 (118d); 13503,13680
(78); 15134a (118d); 15677(78). Breteler, F. J., 3244 (lb); 3428, 3670 (38). Brett, J., 538(118e). Blinker, J., 5178 (50).
Brito, H. S., 295 (36); 330 (25a). Britton, N. L., 814 (82f); 1921, 1999 (82a); 8284, 9107 (82b).
Britton, N. L. & Cowell, 12545 (82a). Britton, N. L. & Hazen, 431, 727 (63). Britton, N. L. & Millspaugh, 6024 (82a); 6065 (82c). Broadway, W E., 61, 184 (6a); 5041, 9226 (41).
Brooks, R. R., 298 (25a). Bruijn, J. de, 1361 (3a); 1434 (2).
Buchtien, O., 762 (68e). Bunting, G. S., 1112 (118d); 1437 (2); 5186, 5335 (41); 5381
(42b); 6456, 6702 (3a); 6793, 6861 (2); 7431, 8020 (42b); 8345, 8446 (2); 8852 (41); 9793 (2); 9826 (3a); 10140 (4); 10939 (2); 12571, 12776 (41); 13400, 13424 (63).
Burch, D., 4367 (83c). Burchell, W. J., 5909, 6053 (124); 6081 (131); 6538, 7188
(81); 7232 (37); 7499 (81); 7526 (37); 8003 (26).
Burger, W., 11171 (78). Burkart, A., 16382 (lb); 16578 (76b); 16648 (3b).
Caballero, Z., 153 (118c).
Caballero M., R., 76 (21). Cabanillas S., J., 494 (19).
Cabrera, A. L., 1572(20).
Cabrera, E., 9859 (8).
Cabrera, I., 3517 (78b).
Caceres, S., 362 (84b). Callejas, R., 4324 (38); 4512 (73); 7285 (la); 9354 (76b);
9596 (63); 10289 (la). Calzada, J. I., 885 (118e); 1068 (65a); 9055 (119). Camp, W H., 2362, 2377 (118a); 2752 (12); 3177, 3539
(70). Caraiba, J. P., 3320 (59). Carauta, J. P. P., 778(131).
Cardenas, K., 1739 (74); 2011 (65a). Cardenas de Guevara, L., 1262 (42b); 1333 (lb); 1343 (3b).
Cardona, F, 2135 (Ba); 2763 (6c). Carlson, M., 3044 (76b); 3388 (41).
Carreira, L., 432 (6b); 636 (25d). Carvalho, A. M. de, 1675 (36); 1758 (52); 1763,1884, 1910
(27a); 2006 (23); 2153 (131); 2371 (92); 2523 (96); 2930
(97); 2970(92); 2990(111); 2998 (91); 3229 (87b); 3241
(113); 3686 (52); 3695 (86); 3742 (27a); 3940 (23); 3980
(25a); 4086 (23); 4158 (52); 4208 (92); 4258 (23).
Castillo, A., 2182 (6b); 2886 (38).
Castillo, J. J., 1048 (118a). Cazalet, P. C. D., 7744 (68a); 7781 (74). Cer6n, C. C, 733, 2637 (68e); 4675 (68a); 9376 (38).
Ceruti, T. M., 326 (92).
Cervi, A. C, 4173 (50). CFCR, 860 (91); 876 (52); 1203 (23); 1306 (108); 1692
(116); 1696 (86); 2136 (99); 2312, 2695 (91); 3529 (93b) 4101, 6159, 6272, 6597 (91); 6795 (104a); 6895 (116) 6896 (89); 6963 (96); 7018 (92); 7039 (117); 7069 (109)
7095 (117); 7100 (52); 7128 (97); 7248 (105); 7262 (99)
7320 (97); 7395 (104a); 7398 (88); 9615 (91).
Chiang, F, 1734, 1814 (16a). Chisaki,F, 519(15).
Chow, K. S., 78217 (41). Churchill, J. A., 72058 (16a). Cid F, C. A., 1255 (72); 1550, 1552 (38); 2103, 2245 (72);
3285, 6867 (38); 7354 (68f); 8024 (38); 9214 (6b). Clausen, R.T., 6082 (118a). Claussen, P., 41 (25a); 119 (131); 122, 820 (25a); 837 (131);
902, 910 (25a); 971 (131). Clemens, J. & A., 21805, 22525 (65a). Clemente & Alain, Bros., 4007 (60). Coker, W. C, 484 (82c).
Colella, M., 1308 (82b); 1340 (82d); 1410 (6a); 1725 (63).
Contreras, E., 1614 (8). Contreras, J. L., 1216 (66); 1732 (76b); 1733 (66).
Combs, R., 724 (59). Constance, L., 3123 (15). Conzatti, C, 1424, 1607 (118a).
Cook, O. F, 269 (76b). Cooper, G. P. & Slater, 318 (42a). Coradin, L., 918 (4); 5775 (25a); 6388 (91). Core, E. L., 1120 (68b). Correll, D. S., 21932 (49); 43777 (82a); 44596, 45031,
46248, 47644 (82c); 47912 (41); 48830 (82c).
Correll, D. S. & Johnston, 20200, 21509 (17a); 21564 (16a). Cory,VL., 52063 (17a). Cott, J. W. van, 1565 (12). Coulter, J.M., 511 (17a). Courbon, - ,80 (20). Cowan, C, 2024(118e).
Cowan, R. S. & Wurdack, 31533 (72). Cremers, C, 9966 (38).
Croat, T. B., 14144 (6a); 21501 (3a); 21637 (76a); 21650
(lb); 23531 (76b); 24488 (118a). Croizat, L., 605 (6b).
Cross white, F. S., 460 (160).
Cruger, H., 266, 1008 (4). Cruxent, J. M., 339 (6b).
Cuadros V, H., 1674, 1718 (38); 1885 (42b); 1861 (40);
2243 (41); 2377 (42b).
Cuatrecasas, J., 6570 (72); 8056, 8113 (la); 11211 (68a); 13847 (68b); 16412 (72); 19620 (la).
Cuming, H., 1248 (6a).
Curran, H. M., 5 (3a); 139, 140, 221 (6a); 265 (27a); 444 (3b); 575 (3a).
Curran, H. M. & Haman, 1017 (38). Curran, M. F, 1916 (72).
Cutler, H., 4746 (16a).
Czerwenka, K., 397 (83).
Dahlgren, B. E., 875 (32); 191 (53).
Daly, D. C, 953 (52); 5310 (40); 6408 (65a). Damazio, L. B., 2034 (25a).
208 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Daniel, T. R, 2161 (16a).
Davidse, G., 5038 (6a); 10708, 12113 (25a); 13642 (lb); 15261(4); 17974(38).
Davidson, C , 4891 (68e); 6475 (17a).
Davis, E.W., 1194 (65b).
Dawe, M. T., 635 (40).
Dawson, E. Y, 14736 (25a).
Deam, C. C, 6015 (67); 6093 (76b); 6102 (118a); 6288 (76b). Degener, O., 26647 (118a); 36711 (65a).
DeHaas & Belem, 176 (25b); 300 (25a). Delascio, F 7016 (6a).
Delgado, L. 850 (72); 1659 (38).
Delgado S., A. 23 (118a); 158 (118e); 352 (79). Devard, -, 144 (38).
Dezzeo, N., 396 (72).
Diaz, M., 518 (76a). Diederichs, E., 91 (63).
Diggs, G., 3990 (118a).
Dillon, M. O., 3936 (19).
Dodson, C. H., 15449 (74); 15734 (68e). Dorantes, J., 1467 (7).
Dorr, L. J., 2344 (16a); 4727 (38); 4793 (3a); 7597 (lb). Dressier, R. E., 1551 (118e); 3914 (6a).
Duarte, A. P., 223 (75); 468, 1405 (31); 2279 (93b); 6772
(23); 7474 (52); 7483 (131); 8949 (93b); 9095 (106); 9354 (85); 10633 (97).
Duarte, L., 152 (25a); 803 (20); 840 (21).
Ducke, A., 301 (68a); 496, 1400, 1679 (38); 1810 (68a); 2117(53); 15650(20).
Dudley, T. R., 11485 (68e).
Dugand,A., 6955 (la). Duke, J. A., 3654 (6a); 9955 (6b); 9955 (6a); 11952 (72);
13053 (6a).
Dunn, D., 30, 17502 (118a); 20120 (16a). Duque-Jaramillo, J. M., 2624 (la). Dusen, P., 9574 (50); 10569, 17393 (25b).
Duss, Pere A., 1164 (40); 1165 (76a); 2637 (40). Dwyer, J. D., 11520 (118e). Dziekanowski, C, 56 (118a); 3411 (118c).
Earle, F. S. & Tracy, 314 (17a). Edwards, J. H., 711 (118e).
Eggleston, W. W , 17236 (17c); 20273A, 20400 (17a). Egler, F. E., 39-74 (40); 39-284 (76a). Ehrenberg, C, 563 (17c).
Eiten, G. & L., 2217 (20); 3922,4013 (131); 4604 (52); 4862 (131); 6776 (94); 6842 (93b); 9455,10172,10408,10482, 10499, 10545, 10547 (131); 10863, 10873 (36).
Ekman, E. L., 985 (83); 1730 (84b); 3325 (82b); 3814 (60); 6042 (82d); 7530 (82a); 9599 (59); 9948 (82d); 12623
(82b); 19148 (59). Elcaro, S., 4A, 73 (6b).
Elias, Bro., 1576 (40). Elias, T., 8379 (16a).
Ellis, C.C., 471 (17a). Enriquez, A., 192 (17c). Ertter, B., 2610 (16a); 8057 (15).
Estrada C, A. E., 1524, 1525,1607 (16a); 1831 (17a); 2345, 2357 (118a); 2462 (16a); 2836 (17a); 3344 (17c); 3463
(17a); 3581 (17c).
Eustachio, N. A. N., 2 (25a).
Fanshawe, D. B., 2163, 2826 (38).
Fassett, N. C, 25939 (41). Feddema, C , 59, 166 (118a); 799 (118e); 916 (118a).
Felix, M., 23 (25a). Fendler, A., 350 (63); 352 (64); 354 (6a); 2255 (3b).
Fernandes, A., 2310, 2985, 3520 (28); 8208, 9691, 11081 (20); 11109, 12297, 15046 (36); 17884 (28).
Fernandez Alonso, J. L., 5575 (2); 5696 (65a); 5929 (lb);
6091 (la).
Fernandez Casas, J., 3831 (84a); 3855 (81); 5981 (84a);
6322 (25c); 6364 (84a). Fernandez N., R., 2987 (16a); 3320 (118a).
Ferreyra, R., 4792 (69); 16495, 16823 (19).
Ferris, R. S., 5990 (120); 7181 (16a); 8570 (15); 8788 (16a). Feuerer, T., 4634a (65b).
Fiebrig, K., 297 (20); 4862 (84b). Fiedler, L., 31 (25a). Fishbein, M., 1733 (17b).
Fisher, G. L., 35249 (7). Flores M., A., 2702 (12); 2967 (76b).
Fonnegra G., R., 1570 (42b).
Fonseca, S., 1517 (131).
Fonseca, W. A. de, 366 (36). Forero, E., 743 (72); 1078 (la); 1506 (73); 1543 (65a); 1754
(6a); 7729 (93a); 9326, 9382 (5); 9414, 9433 (2); 9716
(74); 9717, 9718, 9725, 9731 (la); 9797 (74); 9921 (40);
9955 (2); 10028, 10100, 10101, 10102, 10106, 10109,
10110, 10112 (5); 10154 (2); 10167 (12); 10179, 10184 (la); 10185 (41); 10188, 10189 (38); 10191 (2); 10192 (41); 10193 (65a); 10194(64); 10200(38); 10201,10202, 10241, 10243 (41); 10244, 10247, 10249, 10253 (38);
10256 (lb); 10257 (38); 10258 (la). Fors,A., 19609 (65a).
Fosberg, F R., 23209 (19). Foumet, A., 536 (65b). Franco, E. M., 29 (25a).
Franklin, B., 4521 (16a); 5217 (17a). Franklin, M. A., 5316 (17a); 5801 (15). Friend, E. A., 90(41).
Friedrichs, -, 29742 (22). Friedrichstal, E. Ritter von, 1210 (76b). Fr6es, R de L., 20119 (23); 20120 (92); 20168 (96); 20978
(15a); 30260 (3a). Frye, T. C, 2260 (16a); 2596 (118a). Fuertes, Padre, 1927 (82b). Funck, - & Schlim, 305 (4); 370, 456 (64).
Gandara, J. M., 138 (7).
Galeano, G., 2250 (40). Galeotti, H. G., 3314 (7). Ganev, W., 294 (112); 472 (89); 506 (115); 677 (104a); 769
(88); 793 (104a); 804 (99); 822 (96); 864 (101a); 917
(103); 980, 1055 (86); 1076 (95); 1148 (86); 1205 (91); 1319,1416(20); 1676(112).
Garcia, C, 183 (118e). Garcia, M. & Camineros, 3425 (82g). Garcia, R., 63, 751, 2311 (82b).
Garcia Barriga, H., 11705, 12312 (5); 16045 (126a); 18217 (2); 18668 (68e).
Gardner, G., 23 (75); 1581 (31); 2138 (132); 2555 (31); 3129
(25b); 3130 (131); 3703 (26); 4523,4524 (93b); 4525 (21).
Gaumer, G. F, 24240 (76b). Geisse, G., 143 (58).
Gentle, P. H., 1008 (118e); 1609 (76b); 2301 (118d); 2488, 2486 (76b).
Gentry, A., 8429 (72); 9721 (74); 14770 (76a); 17482a (72);
26797 (68e); 28793 (la); 29696 (67); 36617 (68a);
47567 (42b); 48185 (la); 50564A (42b); 70747 (65b).
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 209
Gentry, H. S., 1218 (17b); 1470 (76b); 2628 (17b); 4860, 5374, 5467 (76b); 5628, 6330a, 6576 (118a); 10669 (79).
Gibbs, P., 2772 (25b). Gillespie, L. J., 2832 (38).
Gillett, J. M., 17023 (16a).
Gillis, W. T., 5260 (82c).
Ginzbarg, S., 874 (52).
Glassman, S. F, 1776 (11); 1872 (118e); 1923 (118d).
Glaziou, A. F. M., 1946 (50); 3794a (21); 7587 (50); 9403
(75); 11195 (25b); 12633 (22); 12640,13787 (35); 14648 (20); 14655 (93a); 15929 (81); 19112 (131); 19113
(106); 19114 (99); 19115 (25a); 21036 (124); 21037, 25038 (25a).
Goldman, E. A., 850 (9).
Gomes-Santis, F, 136, 179 (118e). Goncalves, L. M., 209 (27b). Gonzalez, F, 3251 (la).
Gonzalez-Villarreal, L. M., 3342, 3826 (76b); 3903 (118e). Goodding, L. N., 402-45 (16a). Gorts-van Rijn, A. R. A., 219 (6b).
Goftsberger, I. & G., 36-23771 (25a); G18-12888 (74). Gould, F. W., 3685 (17c). Gregory, D. P., 148 (12).
Griffiths, D., 1674, 2450 (16a).
Grimes, J. W., 2821 (68g); 2822 (42a); 2826 (68g). Groger, A., 669 (4).
Grubb, P. J., 8 (74).
Guedes, M. L., 645 (85). Guedes, T. N., 423 (53). Guppy, N., 382 (38).
Gutierrez, G. & Schultes, 698 (6b). Guyana Forest Dept., 7512 (72).
Hage, J. L. & dos Santos, 1860 (41). Hagen, C. & W. von, 1034, 1201 (118e).
Hahn, L., 1121 (40).
Hahn, W , 1265, 1311 (131); 1520 (75); 1815 (131); 4974
(3a); 5608 (6b); 5848 (38). Hancock, W., 1882 (118e).
Handro, O., 352 (25b). Hanson, H. C, 58 (17a); A/522 (16a). Harley, R. M., 10993 (131); 15089 (88); 15101 (104a); 15416
(113); 15521 (100); 15532 (108); 15687 (100); 15843 (52); 15931, 15932, 15933 (92); 15934 (98); 15937 (97); 15978 (52); 16061 (107); 16095 (96); 16321 (36) 16540 (97); 16684 (87a); 16971 (97); 16972 (90); 16985 (87a); 18630 (52); 18676 (109); 18698 (92); 18745 (97); 18762
(96); 18860 (97); 18973 (132); 19227 (113); 19526 (89);
19717 (87b); 19822 (88); 19928 (95); 19971 (116); 19978 (52); 19980 (104a); 20073 (116); 20091 (87b); 20092
(86); 20137 (87b); 20542 (101b); 20568 (92); 20633 (96);
20644 (92); 20653 (112); 20654 (97); 20726 (100); 20731
(52); 20777 (108); 20784 (91); 21016 (92); 21041 (97);
21042 (92); 21050 (96); 21051 (92); 21055 (117); 21116A
(25a); 21158 (27a); 21178 (25a); 21183 (27a); 21518
(132); 21564 (17a); 22254 (97); 22288 (91); 22423 (111)
22493 (91); 22537 (99); 22537a, 22541 (91); 22544 (92)
22838 (113); 24138 (99); 24427,24619 (89); 25312 (116)
25349 (95); 25350 (99); 26102 (114); 26150 (116); 26157
(99); 27362, 27538 (88); 27611 (20); 27746 (87a); 27753
(45); 27817 (114).
Harriman, N. A., 17629 (65).
Harris, J. A., 16492 (16a).
Harris, W., 8921, 9318 (82f).
Harrison, G. J., 6539 (16a).
Harrison, H. G., 1557 (41).
Harrison, S. G., 1460 (38). Harshberger, C. W., 1042 (22).
Hartweg, T., 964 (68b). Hassler, E., 995 (21); 1628 (84b); 1773 (131); 2974, 3292
(84b); 3556 (131); 4126 (20); 4312 (81); 4490 (21); 4560
(84a); 4566 (84b); 4980 (84a); 5943 (131); 6723 (20);
6991 (84b); 7005 (81); 7765, 8915 (84a); 9303 (81); 9350
(20); 9730 (84a); 9737 (25c); 10135 (84a); 10145, 10818,
10818a (25c); 10983 (131); 11048a (20); 11387 (84b). Hatschbach, G, 1509 (21); 6260 (25b); 6369 (20); 8861 (22);
12886 (20); 17012 (21); 18303 (50); 19803 (20); 22620
(21); 22664 (22); 23247 (50); 26049 (131); 26999, 27467, 27487 (93a); 27516 (22); 27784 (25a); 29628 (93a); 29861 (93b); 32100, 32331 (25d); 33876 (50); 34647 (25d); 35091 (124); 35259 (25a); 39013 (20); 39533 (132); 39662 (52); 40781 (41); 41251 (91); 42053, 42309 (25a); 42436 (113); 42842 (91); 44142 (25a); 44838 (22);
46312 (20); 46466 (104a); 46501 (116); 46515 (87b); 46569 (27a); 46602 (25a); 47424 (52); 47469 (92); 47474
(96); 47511 (117); 47515 (97); 47876 (91); 47910 (92);
47990 (107); 48062 (101b); 48273 (97); 48323 (99); 48347 (108); 48350 (20); 48374 (75); 48422 (84a); 49994
(132); 50029 (26); 50476 (25a); 50503 (27a); 50624 (21);
51126 (91); 51524 (84a); 53394 (86); 53395 (88); 53986, 54144 (25a); 54204 (91); 54373 (124); 54419 (50); 54423
(21); 54682 (25b); 55429 (22); 55768 (21); 56035, 56251 (131); 56716 (113); 56908 (91); 58669 (25c); 58919 (131); 59214, 59409 (21); 60030 (25a); 60415 (26); 61833, 62027 (52); 62032 (86); 64538 (50).
Haught, O., 1429 (2); 3557 (40); 6246, 6305 (42a). Hauthal, R., 12 (21). Helmreichen, -, 7 (75).
Henkel, T. W., 1679, 1974, 2224 (38); 3121 (6b); 5404 (38); 939 (41).
Heredia, M. D., 337, 366, 516 (la).
Heringer, E. P., 782 (131); 2479 (25a); 2544 (124); 3287
(131); 5369 (25a); 6446, 6805 (20); 6912 (25a); 7292 (94); 8668 (25a); 8682 (20); 9009 (124); 11225 (131);
11341 (52); 14213 (131); 15512 (50); 18276 (41); 31900 (52).
Hernandez, H. M., 258 (16a); 262 (12); 263 (16a); 276 (118a);
318 (119); 452 (121); 493 (119); 518 (118d); 758 (72). Hernandez, L., 170 (6b).
Hernandez Garcia, L., 347 (12); 298 (118a). Hernandez M., R., 1798 (7); 8013 (17a). Herrera, C. & Murillo, 23993 (9).
Higgins, L. C, 6292 (16a); 8715, 8794 (17a); 11623, 11684, 11725 (16a).
Hill, S. R., 2389 (82c); 10656 (49); 12124, 14677, 14825 (17a); 20303 (118d).
Hinckley, L. C, 433 (17a); 1654, 3100 (49); 3157 (17a).
Hinton, G. B., 2772 (118a); 4099 (48); 6317 (43); 7806 (10);
9154 (43); 10183 (76b); 10503 (118a); 11044 (118c);
11457, 11685 (118e); 11877, (118a); 12476 (118e); 12712
(118a); 12726 (10); 13931 (79); 14124 (76b); 14377 (46); 14808, 15384 (118a); 15553 (118e); 16524 (16a).
Hioram, Bro., 2298 (82a).
Hitchcock, A. S., 20664 (70); 21324 (19).
Hitchcock, C. L., 22223, 23207, 25518, 25732 (16a).
Hoehne, F. C, 23432 (50); 28403 (22). Hoff, M., 5865 (38).
Hoffman, B., 375 (6b); 1121 (38); 1617, 1679 (127); 2281 (38); 3049 (127); 3757 (61).
Holdridge, L. R., 525, 1075 (82b).
210 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Holmes, W. C, 4490 (118a). Holmgren, N. H., 7052 (16a). Holt, E. G., & Gehriger, 68 (6b). Holton, L. F, 999 (72).
Hostmann, F. W., 171 (38).
Howard, R. A., 625 (118d); 5539 (59); 8371 (83); 10908
(40); 10102 (41); 19613 (40); 19946 (76a); 20011 (65a). Huashikat, V, 876 (63).
Huber, O., 385 (lb); 1663, 1766 (125); 2480, 2648, 3728,
3961 (126b); 5364 (125); 6322 (lb); 6640 (6b); 7439
(127); 8164 (6b); 10625, 11733 (128).
HUEFS, 6239 (56); 10077 (96); 13684, 13685 (27a).
Hughes, C. E., 1235 (76b); 1268,1271 (118e); 1272 (118d);
1287, 1351 (118e); 1490 (118d); 1493 (9); 1503 (12); 1541, 1542, 1546 (15); 1675 (119); 1696 (118d); 1749
(76b); 1786(10); 1807 (16a). Humbert, H., 27612 (82b).
Humboldt, F. W. H. A. von & Bonpland, 260 (40). Hunt, D. R., 5492 (25a). Hutchison, P. C, 1562, 5869 (62).
Hutchison, P. C. & Wright, 5073 (19).
Ibarra M., C, 2361 (118e). Ibarrola, T., 1960, 2164, 2200 (84b).
ICN, 8549, 20622 (22).
Idrobo, J., 11013 (42a). IPA, 22552 (24).
Irwin, H. S., 8328 (124); 8370 (25a); 8805, 10103 (124); 10477 (25a); 11061 (124); 13318 (91); 13861, 14472 (25a); 15011, 15112 (131); 15372 (25a); 16680 (131);
19070 (37); 20010 (93b); 20047 (94); 20367 (25a); 20495 (93b); 20610 (94); 21318, 21640 (131); 21822, 22615 (93a); 22789 (93b); 22946 (52); 22987 (91); 22989 (25a); 23548 (93b); 23702 (52); 23997 (26); 24500 (25b); 24665 (131); 26592 (124); 26871 (131); 27178, 27457 (93a); 27564 (33); 27595 (93a); 28008 (91); 30870 (52); 30888
(99); 30996 (101b); 31459 (25a); 31596 (131); 32275 (113); 32570 (52); 32571 (87a); 37301 (82b); 54088 (38); 55163 (72); 55414 (38).
Irwin, H. S. & Soderstrom, 6204 (37). Isely, D., 10733, 10765 (49); 10878 (17a); 10913 (17c);
10943, 10944 (17a).
Itapu International, 173 (84a).
Jansen-Jacobs, M. J., 76, 2033 (6b); 3137 (38).
Jaramillo, J., 94 (38). Jardim, J. G., 624 (23). Jativa, C. & Epling, 908 (la); 1165 (74).
Jesus, J. O. de, 80(131). Jimenez, J. de J., 5574 (41); 5575 (65a); 5842 (41).
Jimenez M., A., 1004 (7); 2439 (118d).
Johnson, C. D., 267-78 (118a). Johnston, A. R., 27 (6a). Johnston, M. C , 10447, 10530, 11418 (16a); 11758 (49);
11767 (17a); 12218 (16a). Jones, M. E., 186 (76b); 17176 (13); 23001 (118a); 24101,
24243 (15); 27175 (120). Jones, S.G., 5354 (118a). Jorgensen, P., 3630 (84b); 4404 (21); 4819 (84a).
Joyal, E., 1985 (16a).
Judd,A. F, 132(41).
Juzepczuk, S., 1382(119).
Karling,J.T.,46(118c).
Kearney, T. H. & Peebles, 14003 (17a).
Keck, D. D., 6214 (16a).
Keil, D., 1373 (16a). Kellerman, W. A., 7287, 8037 (118e); 8092 (118a).
Killeen, T., 887, 1415, 1610 (131). Killip, E. P., 11260 (la). Killip, E. P. & Smith, 14982, 16177 (2); 16205 (42a); 20922
(3a); 20926 (41); 23611, 25122, 26347 (74).
King, R. M.,486 (119); 581 (118d); 1805 (76b); 2730(119);
2767, 2836 (118d); 2907, 3026 (118a); 4299 (118e);
5059 (118a).
Kirkbride, J. H., Jr., 175 (42a); 2641 (42b); 3009 (131);
3530 (25a); 3677 (124); 5101 (131).
Klug, G., 152 (63); 421 (68a); 1716 (68e); 2349 (68a); 2679,
3765 (74); 4354 (69).
Knapp, S., 7045 (69). Knight, G. H., 697 (68e). Koch, S. D., 83281 (12). Koming, J., 58620 (63). Krai, R., 70733 (41); 72800 (23); 75239 (124); 75524 (20);
75614 (85); 75630 (92); 75631 (97).
Krapovickas, A., 32845, 32871 (25c); 32945 (25d); 33193 (124); 36255 (131); 44443 (84b); 44480 (21); 44972
(84a); 45151 (84b). Krukoff, B. A., 8670, 10144 (68a); 10220, 10220a (65b).
Kubitski, K., 84/66 (38). Kuhlmann, M., 31706 (65a). Kummrow, R., 792, 2078 (50).
Kuntze, O., 1685 (6a). Kvist, L. P., 351 (38); 40556 (74).
Langlasse, E., 26 (la); 41 (10); 1042 (76b). Larpin, D., 704 (38). LaRue, E., 91-32 (16a). Lasseigne, A., 4330 (25a): 4452 (6a); 4899 (118e). Lasser, T., 1640 (6b). Lavastre, Bro., 24 (82b); 1518, 2049 (83).
Lavin, M., 4921 (17c). Lawrence, A., 141 (74). Leavenworth, W. C, 389 (118e).
Ledingham, G. F, 3182 (16a). LeDoux, 2206(119). Lehmann: see Plantae Lehmannianae
Lehr, J. H., 2192 (16a). Lehto, E., 10874B (16a); 14468, 20612 (17c). Leite, E., 554, 1696 (22).
Leite, F. E., 2228 (50). Leme, R. de O., 19 (25a). Lemke, T. O., 5 (41).
Lemmon, J. G., 2662 (17a). Lent, R.W., 2228 (118d).
Leon, Bro., 5921, 8234, 9339 (59); 12393 (82a). Leonard, E. C, 2883, 7295 (82b). Leonard, E. & G., 11088 (82b); 11565 (82e).
Lewis, G. P., 852 (99); 853 (97); 872, 874 (85); 880 (111);
908 (97); 930 (101b); 958 (20); 960 (111) 998 (41); 1075,1132, 1153(36); 1812(59); 1869(36); 1884, 1927
(132); 1939 (36); 1949 (87b); 1967 (36); 1986 (87b);
1990 (36); 2010 (132); 2025 (23). Liesner, R. L., 837 (42b); 5316 (6a); 8282 (42b); 10427
(38); 11690(63); 12042(40). Lima, A., 50542 (34).
Lima, D. A., 53-1589 (3a).
Lima, H. de, 3883 (113); 3896 (52); 3953 (30); 3971 (85). Lindeman, C. A. M., A261 (22).
Lindeman, J. C. & de Haas, 1873 (22); 2210, 3412 (21).
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 211
Linden, J., 43 (64); 696 (3a) Liogier, A., 13923 (83); 16475 (82b); 17561 (83); 17902,
21170 (82b); 25968 (83); 27183 (82b); 27700 (41); 31440 (65a); 31448 (41); 33359 (65a); 33594, 35431 (41); 36152 (82a).
Lisboa, F, 1252 (52). Little, E. L., 663 (68e); 4002 (21); 7082 (la); 7173 (72);
8443 (5); 26005 (41). Lloyd, F. E., 42 (16a); 593, 782 (76a). Loaiza, C. A., 126 (38).
Lopes, R. D., 5 (26). Lopez, A., 196 (82b).
Lopez Perez, J., 105, 513 (118a). Lowry, P. P. Ill, 1193 (16a).
Luetzelburg, P. von, 3 (91); 152 (116); 245 (89); 376 (132); 382 (27a); 12299(116).
Lugo S., H., 4297, 4382, 4519, 4692, 5052 (68e); 5914 (68a); 6072 (68e).
Luiz Emygdio, 3586 (25a).
Lundell, C. L., 148 (8); 6610 (72). Luteyn, J. L., 1561 (42a); 9258 (41). Lyonnet, E., 4 (118a).
Maas, J. P. M., 10926 (72).
McAdams, C. S., 351 (41). Macbride, J. F, 2419 (19).
MacDougal, T. D., 387 (17a).
McDowell, T., 3897 (38). Macedo, A., 856(131). Machuca Nunez, J. A., 6769 (118c).
MacQueen, D. J., 133, 143 (118a); 162, 163 (118c); 170 (118e); 185, 189 (118c); 191 (118e); 203 (118a); 207 (120); 208 (118e); 209 (120); 211, 212, 216 (118e); 217, 218 (118a); 228 (119); 229 (118a); 230 (118e); 234 (118a); 255 (118); 278, 281 (118c); 282 (119); 285
(118a); 293,296 (119); 302, 307, 308 (118d); 311 (118e); 315 (118d); 318 (118e); 327 (68g); 335, 345, 349 (118d); 355 (118a); 358, 362, 363, 365 (118d); 369, 371, 379 (118e); 380 (118d); 390 (42a); 411 (76b); 414 (77); 421,
435 (68g); 445, 454 (76b); 462 (118c); 463 (76b); 464 (68g); 469 (118e); 492 (118d); 493 (118a); 505 (118e); 514 (118a); 516 (118d); 518 (7); 519, 521, 523 (118d);
526 (67); 532, 533 (118d); 534 (68g); 555, 556, 558, 560,
561, 563B (118e); 564(118a); 572(118e); 581, 587, 590, 597, 601 (118d); 604 (6a); 614, 615 (118d); 617 (79);
618, 619 (68g); 621 (6a); 623 (72). McVaugh, R., 12149 (79); 13703 (118a); 13795 (14); 13992
(118a); 15152 (118e); 15550 (47); 16213 (118d); 16387
(118); 116475 (79); 16590 (120); 16868 (17c); 18592
(120); 20301 (79); 21323 (118a); 21467 (79); 21769
(120); 23283, 23520 (79); 24843 (118a); 25851 (16a).
McVaugh, R. & Koelz, 914 (118a).
Magalhaes, M., 18936(91).
Magallanes, A. S., 4389 (43). Maguire, B., 10064, 10153, 10232, 10559, 10725, 10865,
11167 (16a); 25068 (72); 27303 (3a); 28721 (68d); 30341
(6b); 30365 (28); 30390 (72); 30633 (114); 30671, 30705
(125); 32180, 32189 (127); 32299, 32447, 32448 (38)
33643 (127); 35024 (38); 35862,35870 (3a); 37510 (68d)
40172, 40199, 40263 (6b); 40561 (127); 41451 (126b)
44140 (68a); 44633 (94); 44661 (93b); 49105 (93a) 49134 (93b); 49142 (25a); 49152, 49211 (91); 49283
(25a); 54139 (72); 56192 (25a); 56245 (25d); 56905 (25c);
56942 (25d); 56986 (25c); 57152 (25a); 65703 (125).
Maguire, B. & Politi, 28421, 28629 (68d).
Mahler, W. R, 3090 (16a); 9188 (57).
Makrinius, E., 613 (42a).
Marcano Berti, L., 1002, 12-6-28, 78-980 (38); 317-979,
324-979 (69); 433-579 (40); 435-975 (4); 982-138 (63).
Marquez R., W., 405 (118c).
Marshall, N.T., 325(118). Martin, R. T. & Plowman, 110 (la).
Martinelli, G., 5399 (92); 11044 (23); 11346 (93b).
Martinez S., E., 4307 (118e); 4416 (118d); 5009, 6808 (118e); 14407 (76b).
Martinez, E. & S. & R Barrie, 5763 (121). Martinez, E. & Villasenor, 5237 (66).
Martinez-Calderon, G., 1528 (119).
Martins, G. J., 201 (118a).
Martins, P., 8620 (20). Martins, T. L. R, 31 (25a). Martius, C. R P. von Herb. Fl. Bras., 1100 (50); 1108 (131);
Obs. 1516 (91); 1882 (86); 1985 (108).
Marulanda, O., 2004 (76b). Mason, C.T., 2970 (118a).
Mattos, J. R., 8529, 8553 (131). Mattos Silva, L. A., 382, 827 (23); 1603 (92); 1618 (96);
2600 (23); 2816 (96). Matuda, E., 1342 (118a); 1888 (12); 2608 (42a); 3340 (76b);
4132, 4267 (12); 4394 (76b); 4717 (118d); 26982, 27468, 28931 (118a); 38392 (118e).
Maxon,W.R., 3218(118e).
Medina-Cota, M., 3049 (118a). Medrano, F.G., 1416 (118a). Meier, W., 851 (64).
Mejia , M., 107 (83); 1108 (82b); 7004 (83); 8330 (82b); 23504 (83).
Mell, C. D., 2237 (68g). Mello, C. M. S., 25 (25a). Mello Barreto, M., 6407 (93a). Mello-Silva, R., 612 (93a); 697 (91); 795 (100); 753 (91). Melo, M. M. R, 538 (50). Melo, N. J., 16 (41). Mennega, A. M. W., 397 (72). Metcalfe, O. B., 294, 1382 (17a); 8824 (17c).
Mexia, Y, 1634a (10); 1818 (79); 2705 (118a); 5523 (131); 5620 (25a); 5858 (93a); 6313 (74); 6336 (68e); 8464 (74).
M G , 2483 (31). Michel, R., 327 (75).
Miller, O. O. & Johnston, 58 (6a). Millspaugh, C. R, 9068 (82c).
Miranda, M. R, 5790 (16a).
Montero O., G., 2860 (58).
Montes, J. E., 396 (22).
Moore, S. L., 160 (25d); 737 (131). Moraes R., M., 765 (65b). Morawetz, W., 17-7288 (38).
Moreira, A. G., 9 (25a).
Mori, S. A., 9913 (23); 11316 (20); 11681 (23); 12033 (41);
12297 (86); 12383 (87b); 12430 (88); 12465 (89); 12548
(107); 12597, 12624, 12668 (92); 13091 (85); 13131
(96); 13139 (117); 13173 (92); 13312 (96); 13318 (91);
13346 (97); 13463, 13470 (27a); 13532 (104b); 13611
(89); 14285 (97); 14341, 14353 (96); 14399 (97); 14400
(85); 14454, 14495 (1.13); 16692, 16898, 16949 (25a).
Mori, S. A. & Kallunki, 9259 (41).
Morillo, G., 9217 (2).
Morong, T., 412 (84b).
Mostacero C, J., 1567 (19).
Muller,-, 252, 1594 (118a).
212 MEMOIRS OP THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Molina R., A , 10163 (76b); 10829 (68g); 11001, 11891
(118d); 11906 (43); 11913 (76b); 12349 (123); 12411
(118a); 12896 (118e); 13531 (118a); 13673, 13821 (68g);
14020 (118a); 14240 (7); 14392 (76b); 14399 (11); 14401
(7); 14442 (43); 14457 (11); 14487 (118d); 14567 (11);
15661 (76b); 16015 (118a); 18633 (7); 18706 (11); 21095
(118a); 21558 (76b); 21923 (11); 22289 (118a); 22435
(43); 22654, 22701 (7); 22808 (11); 22933 (118d); 23011
(43); 24029 (76b); 24148, 25757 (11); 25779 (7); 26058
(68g); 26194, 31551 (76b).
Mutis, J. C, 441, 3542 (6b); 5611 (5).
Nash, C. V. & Taylor, 1423 (82c); 1465 (82b); 1543 (83).
Navarro de Andrade, -, 861 (22).
Nee, M., 17524 (lb); 26071 (118e); 26957 (16a); 30287
(58e); 30951 (6b); 35363 (65b); 37700 (74); 40520,
40605, 40849, 41069 (65b); 42432 (41); 42708 (38);
44233 (65a).
Negrete, M., 13 (118a).
Neill, D., 6061 (74); 6140, 6163, 6434, 6888, 7389 (68e);
7652 (68a); 7797, 7997 (74).
Neinstedt, E. P., 36(131).
Nelson, A. & R., 1262 (16a).
Nelson, E. W., 2782 (76b); 2795 (7); 3258 (118d); 3396
(68g); 3838 (42a); 4095 (10); 4696 (16a); 4975 (17a); 6183 (17c); 6863 (118e).
Nelson, E. W. & Goldman, 7455 (13). Nevling, G. & Gomez-Pompa, 1916 (16a).
Nicholls, H. A., 25 (76a).
Noblick, L. R., 2810 (91); 2841 (20); 2842 (101b); 2886 (96); 3146 (36); 3770 (25a); 3775 (86).
Noblick, L. R. & Pinto, 2808 (97); 2880 (107).
Norris, D. N. & Taranto, 13555 (79); 13662 (76b); 13855 (79).
Oldeman, R. A. A., 1314 (68a); 2499, 2714, 3531 (38).
Oldenburger, P. H., 1542 (124); 1597, 1630 (25a); 1815, 1879(124).
Oliveira, E., 1928 (20); 4603 (71).
Oliveira, P. I., 512(50). Onishi, E., 26 (25a); 715, 728 (131).
Opler, P. A., 404 (7); 1781 (76b). Orlandi, R. P., 390 (36). Orozco, C. J., 1134(72); 1143(74); 1144 (la); 1145, 1165,
1186 (74); 2361 (la).
Ortega, J. C, 5640 (76b).
Pabst, G., 2870 (91); 3761 (25a); 5184 (22); 7097 (25a).
Pacheco, M., 201 (38).
Padilla, S. A., 89(118e).
Palacios, W., 845, 1135 (68e); 3490 (63); 6795 (la).
Palmer, E., 14 (16a); 22 (13); 59 (118c); 134 (76b); 138
(10); 174 (17a); 266 (118a); 279 (120); 293, 318 (16a);
330, 370 (17a); 590 (16a); 790 (15); 1200 (118e).
Palmer, E. J., 21353, 34041 (49).
Parish, S. B. & W. R, 762 (16a).
Parish, W. P., 52 (16a).
Parry, C.C, 317a (16a). Parry, C. C. & Palmer, 212 (16a).
Paz, N., 198 (41); 203 (65a).
Peckolt,T., 135,399(20). Pedersen, T. M., 3007 (84b); 5250 (84a); 9384 (121); 15613
(84b). "Pedra do Cavalo," 337 (55). Peebles, R. H., 292, 1035 (16a); 2314 (17c).
Peebles, R. H. & Kearney, 2623 (17a); 10951 (16a).
Pennell, P. W. & Killip, 5978 (la).
Pereira, B. A. S., 1340 (81); 2600 (131). Pereira, B. 2184 (101b); 2925 (25a); 3706 (91); 8849 (93b);
9114 (124); 9962 (93b); 10067 (85); 10240 (124).
Pereira, E. M., 16 (25a).
Perez Arbelaez, E., 195 (38); 8338 (5).
Perez Arbelaez, E. & Cuatrecasas, 5797 (68b).
Perez Asso, A., 1712 (82a).
Perez M., A., 139, 202 (118e).
Phelps, K. P. & Hitchcock, 305 (6c).
Philcox,D., 3180,4163(131).
Picarda, 1. 869 (82b).
Pickel, D. B., 3218 (54); 3726, 3758 (24).
Piedade, M. L., 14 (25a).
Pimentel, J., 308 (82b); 373 (83); 538 (82b).
Pinheiro, R. S., 282 (24).
Pinkava, D. J., 9958 (16a); 12446 (17a); 14663 (17c); 14491
(16a).
Pinkley, H. V, 395 (74).
Pinkus, A. S., 10 (38).
Pipoly, J. J., 7669, 7907 (127); 8382 (38); 10251 (127);
10325 (38).
Pirani, J. R., 1985 (91); 11887 (25a).
Pires, J. M., 880, 10370, 10527, 10721, 10832 (38); 17257
(131); 52483 (38); 58133 (25a).
Pires, J. M. & Black, 2491 (131). Pires, J. M. & Silva, 596, 881 (38); 1053 (71). Pittier, H., 2721 (7); 6374 (76a); 6509 (42a); 8784 (64); 8816,
8822, 8834 (76a); 8861 (lb); 8866, 8927 (76a); 9018 (3b);
9055, 9166 (3a); 9275 (6a); 10219 (64); 10221 (lb);
10935 (2); 11658 (64); 11784 (6a); 11940 (76a); 12224
(4); 12309 (41); 12407 (3b); 12529 (4); 13114 (3a); 13925
(lb); 14148,14150 (3a); 15204 (64); 15208 (41).
Pittier, H. & Tonduz, 891 (118d).
Plantae Lehmannianae, 365 (68b); 4580, 5376 (la).
Plowman, T., 8629 (131); 9775, 9809 (52); 13357 (lb).
Pohl, J. E., 605 (25b); 692 (131); 1362 (25b); 1433 (75);
1454 (20); 1455 (50); 1456 (93a); 1474 (124); 2550 (25a); 3460 (93a).
Poole, J. M., 2046 (68a).
Prance, G. T., 7273 (68a); 8929 (38); 9217, 9278 (6b); 14192, 14876,14913,17851 (68a); 18694 (6b); 18702 (41); 18880
(25d); 20811, 21801, 21945 (68a); 22386 (72); 24001 (68a); 25044 (6b); 26134 (131); 28958 (125); 29534 (6b).
Prance, G. T. & Silva, 59595 (21).
Pringle, C. G., 44, 915 (16a); 2371, 2426 (118a); 3713 (12);
4093 (16a); 4096 (118a); 6351 (16a); 8163 (118e); 8671 (43); 8745 (17c); 9736 (12).
Proctor, G. R., 16179 (129); 31085 (82f).
Purpus, C. A., 135 (7); 487 (118e); 1184 (118a); 1574 (16a);
1876 (7); 2335 (76b); 3186 (16a); 3642 (7); 2666 (118a);
7199, 7505 (76b); 7643 (7); 8391 (76b); 8393 (7); 8402,
8597, 8736 (76b); 10159 (7); 10240, 10346, 10554 (12);
10582, 10762 (7); 11000, 11000a (76b); 11001 (7); 10214(43); 13028(7).
Pursell, R., 1361 (3a); 8898, 8912, 9000 (6a).
Purseglove, J. W., 6396 (63).
Queiroz, L. P. de, 348 (56); 600 (85); 628 (111); 769
(99); 1692 (101a); 1922 (99); 2031 (25a); 2534 (20); 2745
(92); 2751, 2752 (86); 3334 (99); 3359 (91); 3385 (52);
3471, 3488 (92); 3517 (113); 3540 (87a); 3579 (25a);
3584 (27a); 3595 (86); 3674 (20); 3678 (88); 3684 (86).
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 213
Rabelo, B. V, 2949, 2951, 2953 (38). Ramella, L., 29527 (75). Ramia, M., 8467 (41). Ramirez, N., 3087 (41); 2478 (6a); 3095, 3222 (3b); 4023
(127); 7160 (6b). Ramirez, G., E., 640 (118e). Ramirez P., B. R., 82 (68a); 93 (74); 1045 (la). Ramos, J. E., 1740 (la). Ratter, J. A., 801, 1066, 1898, 2293 (131); 2661 (20); 3282
(25d); 5240 (131). Rauh-Hirsch, P1577 (74). Rebolledo V, A., 300 (118e). Regnell, A. P., 1.99 (50); III.516 (20).
Reitz, PR., 2232, 6851 (50).
Reitz, P. R. & Klein, 2453 (50); 3729 (22); 7372 (50); 7382 (22); 17270, 17830 (50).
Reko,B. P.,4130(68g).
Renvoize, S. A., 3001 (21); 3083 (22); 3205 (21).
Restrepo, D., 707 (6b).
Reveal, J. L., 3622 (118a).
Revilla, J., 250, 2125 (68a).
Reyes Garcia, A., 1818 (43).
Reznicek, A. A., 225 (118a).
Rico A., M. de L., 192, 209, 874 (118a).
Riedel, L., 20 (68a); 307, 918 (131); 1118 (75); 1247 (94);
2033 (131); 2483 (25a); 2527 (81).
Riley, L., 95 (63).
Rimachi Y, M., 2852, 8411, 3412 (68a).
Ripley, H. D. & Bameby, 13457 (17c); 14839 (118e).
Rivas, R. M., 148 (41).
Rivera R., J., 1777 (118e).
Roca,Y, 211(41).
Rodrigues, W., 7853 (38). Rodriguez, H., 1396 (64).
Rodriguez, M., 683 (13).
Roe, K., 769 (118a); 888, 1047 (118e).
Rohwer, J. G., 101 (25a).
Roldan, P. J., 1833 (42b). Rombouts, H. E., 455 (38); 617 (72).
Romero, G. A. & Guanchez, 1803 (68a).
Rommer, D. F. R., 36(131).
Rosa, N. A. & Lima, 2276 (125).
Rosales, J., 288 (6a). Rosas R.,M., 1338(119). Rose, J. N., 1378, 1380, 1504 (76b); 1660 (118b); 1753
(79); 1927 (118e); 2254 (118a); 2674 (17c); 2874, 2963
(16a); 3036 (118a); 3365 (79); 3844 (83); 4209 (16a);
7525 (118e); 9538 (16a); 14155 (76b); 15185 (16a);
16470 (15); 18532 (65a); 22255 (70); 23040,23322 (19);
23834 (70). Rose,J. N. & Hay, 5909(12).
Rose, J. N. & Hough, 4580 (12).
Rose, J. N. & Standley, Russell, 12743 (16a).
Rose, K., 1686 (118a).
Rothrock, J. T., 302 (16a).
Rowell, C. M., 145 (118a).
Rubio, D., 538, 612 (la).
Ruiz-Teran, L., 10678 (41). Rusby, H. H., 22 (118a); 118 (16a); 394, 654 (65b); 724
(68a); 1314, 1315 (65b). Rusby, H. H. & Pennell, 249 (72); 1153 (42a); 1156 (72).
Saer d'Heguert, J., 726 (40); 869 (3a). SagasteguiA., A., 8656, 11741, 12324, 15374(19).
Saint-Hilaire, A. de, CV643 (124).
Saldias P., M., 401 (65b).
Salgado, O. A., 337 (36).
Salsedo, C. A., 119 (65a).
Sanchez Vega, I., 6643(62).
Sanchez Vega, J. G., 422, 874, 2582, 5848 (19).
Sandeman, C, 4157 (19).
Sandoval, E., 690 (76b).
Sano,T., 182 (84a).
Santana Michel, R J., 2809 (43).
Santiago, A. R, 8 (25a).
Santis Cmz, E., 530 (118e).
Santos, E. B. dos, 274 (91).
Santos, R. S., 24191 (25a). Santos, R. S. & Castellanos, 24104 (21).
Santos, T. S. dos, 682, 1673 (23); 1698 (24); 3014 (23).
Saunders, A. C, 3371 (15); 9960 (8); 13249 (16a).
Saunders, J., 274, 727 (67).
SawadaR, M., 122(19).
Schaffher, J. G., 596/626 (16a).
Schaller,G., 114(131). Schatz, G. E., 816 (38).
Schiede, C. J. W., 678 (76b).
Schinini, A., 12553 (84b); 21356 (84a); 21925 (84b).
Schipp, W. A., 369 (118d); 657 (118e).
Schnell,R., 11454(38). Schomburgk, Robert, 582 (6b); 806B (3a); 1542 (128).
Schott, H. W. d, 1433 (75).
Schucht,J., 1427(131). Schultes, R. E., 5553, 5612, 5819, 14203 (126a); 15782
(38); 17036 (6b). Schultes, R. E. & Cabrera, 12787, 13568 (68a); 14203
(126a); 15329, 15419 (76b); 16407 (40); 19152, 19915,
19985 (126a).
Schultze, A., 553(411).
Schunke, J. M., 258 (68a).
Schunke V, J., 874, 4991, 6824 (68e).
Schwarz,G..J.,4750(21).
Sehnem, A., 10638 (22).
Seidel, R., 2550 (74); 3095 (65b).
Seidenschwartz, -, 166/1 (74).
Seigler, D., 13216 (1186); 13550 (119).
Seler, E., 1734 (118a); 1988 (76b); 3006 (43); 3195 (118e).
Sello, R, B94 (25a); 8169 (50).
Semir, J., 595 (93b).
Shafer, A. J., 426 (40); 1181 (82a); 3710 (59); 4055 (60);
7626 (82a); 8252 (60).
Shelton, M. G., 96 (17c); 252 (17a).
Shepherd, G. J., 4119 (131); 7580 (25d); 10219 (93b).
Sherman, C, 106(118e).
Shunsuke, T., 8.46 (50).
Sieber, P. W., 455 (76a).
Silva, A., 86 (38).
Silva, R C. da, 102 (131); 163 (25a).
Silva, J. A., 284 (6b).
Silva, L. A. M., 60 (25a).
Silva, M., 2600 (38).
Silva, M. B. da, 25 (38).
Silva, M. R F , 1200(131).
Silva, M.G., 658(131).
Silva, N. T., 1807, 1031, 2353 (71); 2495, 60869 (38);
60970 (6b).
Silva, S. B. da, 179 (86); 347 (25a).
214 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Silverstone-Sopkin, P., 615, 3727 (la). Simpson, D. R., 435 (68c). Singh, U., 40 (65a). Sintenis, P., 4916 (82b). Slane,V, 133 (76a). Skutch, A. P., 1371 (42a); 2900, 5154 (118d). Smith, A. C, 2378, 2505 (6b); 3156 (3a); 3168 (40); 3268
(6b); 10186, 10490 (40). Smith, C. L., 323 (118a); 1849 (118e). Smith, D. N. & Garcia, 13844 (68e). Smith, E. G., 14164 (17c). Smith, G. L., 10011 (82b). Smith, H. H., 32 (42b). Smith, H. H. & G. W., 1000 (40). Smith, J. D., 2836 (118a). Smith, L. B., 6930 (25a); 9271 (21). Smith, L. B. & Klein, 12244 (50); 13836 (21); 14091,14946
(50). Smith, L. B. & Reitz, 12764, 12778, 12871 (22). Sneidern, K. von, 425 (68b); 1157, 2284, 5037, 5536 (la). Sobral, M., 4615 (50); 5532 (23). Sodiro, A., 392 (74); 393 (70). Soejarto, D. D., 3579 (lb); 4154 (74). Solis, I., 518 (118a); 595 (118e). Soria, N., 4500(131). Sousa S., M., 3124 (118a); 4560 (12); 4653 (16a); 5208
(76b); 5587 (66); 7006 (121); 7137 (66); 8511 (121); 8584 (7); 8766 (42a); 12130 (8); 12521 (121).
Spellenberg, R., 3882 (17c). Spichiger, R., 368, 395 (75). Spruce, R., 389 (38); 4235 (69); 4466 (74); 5571 (70). Standley, P. C, 5809 (76b); 6856 (7); 14602 (11); 20396,
23846 (118d); 23877 (118e); 41212, 53888 (118d); 56397 (118e); 73144 (67); 90049 (118a).
Standley, P. C. & L. Williams, Allen, 571 (11). Stannard, B., 6551 (25a). Stehle H., 1643 (40); 3519, 3529 (76a). Stehle H. & Quentin, 5546 (40). Steiger, T. L., 2102 (17a). Stein, B., 2153 (74). Stein, B. A., 2771 (68e); 3057 (74); 3377 (63). Steinbach, J., 5612, 7154 (65b). Stergios, B., 3479 (4); 8168 (72); 11685 (6b); 13085 (64). Stevens, W. D., 8353 (67). Stevenson, D. W , 1137 (118e). Steward, W. C, 160 (6b). Steyermark, J. A., 42359 (118a); 42985 (45); 50601 (118a);
53738, 54830 (19); 60998, 61795 (lb); 62226 (63); 74637 (33); 86250 (6a); 86583 (6b); 87856, 87962, 89282 (3a); 90505 (6b); 94701 (40); 96444 (4); 94723 (64); 97017 (42b); 97623 (63); 97809 (6b); 99057 (3b); 101482 (38) 102978 (125); 105299 (64); 105360 (76a); 105879 (lb); 106787 (63); 106889 (6b); 107850 (4); 108034 (40); 108062, 108232 (4); 108233 (40); 108710 (6a); 111504 (63); 113605 (41); 114062 (6a); 116347 (63); 121219 (64); 121835, 122835 (6a); 125706 (6b); 130957(4).
Steyermark, J. A. & Wurdack, 38, 52 (6c). Stiibel, A., 239 (5). Stutz de Ortega, L. C , 1560 (21).
Tamayo, R, 3400, 3756 (64).
Tavares, A. S., 116(125). Taylor, J. & C , 29419 (16a).
Taylor, R. J., 4406 (7).
Teilher, S., 1058 (58). Teixeira, L. O. A., 1254 (68a). Tellez, O., 3902 (121). Tessmann, G., 4101 (68e). Teyssman, -, 1565 (21).
Tharp, B. C, 44419 (57).
Theckery, P. A., 57 (16a).
Thieu, L. B. 1562 (la). Thomas, W. W , 3224 (72); 3698 (12); 4458 (131); 9386
(23); 9621 (36); 9787, 9848, 10711 (23).
Thomber, J. J., 5203 (16a).
Thome, R. P., 57580, 60175 (16a).
Thurber, G., 360 (16a).
Tillett, S. S., 45519 (38).
Ton, A. S., 1405 (118d); 2697 (118a); 3503 (118d).
Tonduz, A., 4544 (42a). Torke,A.,441 (118a). Toro,RA., 109(61).
Toro, R. A., 25 (la); 109A (73).
Torres C, R., 4757 (119). Townsend, C. H. T. & Barber, 7 (17c).
Tressens, S. G., 4401 (20). Triana, J., 467, 4478 (la); 4479 (42a); 6837 (72).
TRIN, 5026 (63).
Trott, S., 254 (119); 256 (42a).
Trujillo, B., 13370 (64). Tryon, R. & A., 6814(94).
Tuerckheim, H. von, 1324 (123); 11/1366 (118d); 11/1953
(118a); 11/2294 (123). Tun Ortiz, R., 489 (118d). Turner, B. L., 3642 (16b). Tweedie, J., 1218(75).
Ule, E., 2833 (25b); 2834 (37); 2835 (131); 3326 (50); 7133, 7310 (102); 7311 (100); 7385 (132); 7386 (27a); 7439 (52); 7440 (29); 7530 (30); 7573 (34); 7660 (22); 7931
(3a). Uribe U., L., 633 (76b); 2326 (5). Utley,J. P., 3132 (76b).
Valera, A., 457 (6b). Valeur, E. J., 205 (82b).
Valverde, L,. 1271 (41). Valverde, L. & Calderon, 81 (38).
Van Devender, T., 92-227 (17b); 94727 (118a); 94-778 (118e); 94-974 (76b); 95-22 (16a).
Vanni, R., 237 (25c); 2116 (65b); 3060 (21). Vargas C.,C, 10285(19).
Vasquez, M., 2256 (118e). Vasquez, R. & Jaramillo, 11842 (68a).
Vauthier, -, 102 (25a); 110 (91). Velasco, J., 772 (126b). Ventura A., A., 2110, 2892 (118a).
Ventura, E., 606, 827, 1678 (42a).
Ventura, P., 2583 (76b). Vidal, L. A., 108 (5).
Villasenor, J. L., 831 (118a). Votava, P., 92 (82b).
Wagenknecht, R., 18417 (58).
Wagner, R. G., 741 (65a).
Walker, S., 74H09 (12); 75H14 (76b); 78H38 (17a). Wallace, - & Dunn, LeDoux, 224 (120). Walmor, -, 274 (97). Wallnofer, B., 11/10488(38).
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 215
Wanderley, M. G. L., 777 (25a); 940 (23); 952 (96). Ward, D. B., 7769 (57). Waterfall, U. T., 3483, 4747, 6205 (49). Watkins,-, 704 (118a). Weberbauer, A., 3260 (19); 6238, 7123 (62). Webster, G. L., 23610 (3a); 31278 (la). Weddell, H. A., 2789 (25b). Wendt, T., 3671 (68g); 10006 (16a). Werdermann, E., 2278, 2279 (65b). Werff, H. van der, 5093 (63). Wessels Boer, J. G., 1327 (38). White, A., 736 (la). White, O. E., 443 (65b); 997 (65a). Whitefoord, C, 5783 (76a). Wiggins, I. L., 4369 (15); 4454 (14); 5675 (15); 8557 (16a);
10848 (19).
Wilbur, R. L., 1497 (16a); 8356 (76a). Wilbur, R. L. & C. R., 2125 (79).
Williams, L. O., 5383 (50); 6918, 6883 (91); 10414 (64); 21857, 25165 (118a); 26363 (7); 28368 (118d); 42800
(11). Williams, L. O. & Assis, 6972 (94). Williams, L. O. & Molina, 20215 (7). Williams, LL, 6821, 6833 (69); 10228 (6a).
Williams, LL & Alston, 65 (3a). Williams, R. S., 141 (72); 707 (la).
Wilson, P., 7295, 7633 (82c). Wilson-Brown, Fr., 93 (6b). Winzerling, -, VII-2 (8). Wolf, G. P. de, 2010(41).
Wood, C. W , 452 (64).
Woolston, A. L., 452 (131); 778 (84b).
Worth, C. & Morrison, 15748 (19). Worthington, R. D., 9318 (118a); 10340, 11331 (17a); 11787
(16a); 12488(41); 13075 (16a); 13330 (17a); 13454 (16a);
14347 (17a); 20647 (16a); 23489 (17a) 24059 (8). Wright, C, 153 (82a); 168, 315 (17a); 601 (49); 1043 (16b);
1045 (17c); 3544 (59). Wurdack, J. J., 397 (6a); 2168 (68e); 2322 (74); 34326A,
34326B (38). Wurdack, J. J. & Adderley, 43015 (125); 43594 (72); 43701
(126b). Wurdack, J. J. & Monachino, 39843, 40884 (4); 41228 (3a).
Wynde, F. L. & Muller, 14 (16a).
Yanes, E., 7 (3a). Yip, H. G., 71 (65a). Yuncker, T. G., 5585 (118e); 5622 (7); 5681 (76b); 6047
(68g); 6128 (118a); 8586 (67); 8723 (118d); 8833 (67);
17390 (82f).
Zak,V, 3141 (68e). Zambrano, O., 1383 (2). Zanoni, T., 11862 (82b); 18068 (83); 20789 (82b); 24637
(83); 25408 (82b); 26074, 26289, 27703, 28803 (83);
28855 (82b); 28857 (83); 32503 (82d); 32584, 34559, 34635, 34647 (82b); 35116 (83); 37332 (82b); 41196
(82d). Zardini, E., 12628, 14704 (21). Zarucchi, J. L., 3285, 6636 (lb).
Zollner, O., 8095 (58). Zuluaga R., S., 219 (41); 366, 399 (6a).
I n d e x to Scientific N a m e s
Synonyms are in italics, new names and new combinations in boldface. The symbol ° indicates a homotypic syn
onym; the symbol =, a taxonomic synonym. A n asterisk (*) indicates an illustration or map. The generic name
Calliandra is abbreviated to C. Combinations in Zapoteca are indexed in various papers by H. Hernandez, hsted
above on page 202; names in various genera attributed to Bameby and Grimes are indexed in parts 1 and 2 of the
present volume 74 of the Memoirs of The N e w York Botanical Garden.
Acacia Linnaeus asplenioides Nees = C. asplenioides caracasana (Jacquin) Willdenow = Zapoteca caracasana
(Jacquin) H. Hernandez
fasciculata (Willdenow) Poiret = C. surinamensis
formosa Kunth = Zapoteca formosa (Kunth)
H. Hernandez subsp. formosa gracilis Martens & Galeotti = Zapoteca formosa (Kunth)
H. Hernandez subsp. rosei H. Hernandez
grandiflora (L'Heritier) Willdenow = C. houstoniana
anomala haematomma de Candolle = C. haematomma
haematomma
haematostoma Sprengel = C. haematomma
haematomma houstoni (L'Heritier) Willdenow = C. houstoniana
houstoniana
humilis Schlechtendal = C. humilis humilis
lambertiana G. Don = Zapoteca lambertiana (G. Don)
H. Hernandez
laxa Willdenow = C. laxa laxa magdalenae de Candolle = C. magdalenae magdalenae
media Martens & Galeotti = Zapoteca media (Martens &
Galeotti) H. Hernandez
metrosiderifolia Schlechtendal = C. houstoniana houstoniana
mollicula Martens & Galeotti s Zapoteca formosa
(Kunth) H. Hernandez subsp. mollicula (Martens & Galeotti) H. Hernandez
nigra Clos = C. chilensis
pauciflora A. Richard = C. pauciflora
pilosa Bertero ex de Candolle = C. haematomma glabrata
portoricensis Jacquin = Zapoteca portoricensis (Jacquin)
H. Hernandez
rubescens Martens & Galeotti = C. mbescens selloi Sprengel, 204
squarrosa Martius = C. squarrosa
tetragona Willdenow = Zapoteca tetragona (Willdenow)
H. Hernandez
Anneslia Salisbury, nom. rejic. = Calliandra
acapulcensis Britton & Rose = C. houstoniana
acapulcensis
alamosensis Britton & Rose = C. houstoniana houstoniana
216 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Anneslia Salisbury (continued)
albanensis Britton & Rose = C. houstoniana anomala
albescens Britton & Rose = C. houstoniana anomala
belizensis Britton & Rose ex Standley = C. belizensis
bijuga (Rose) Britton & Rose = C. bijuga
brandegeei Britton & Rose = C. brandegeei
bullata (Urban) Britton & Rose = C. enervis
caeciliae (Harms) Britton & Rose = C. caecihae
callistemon (Schlechtendal) Britton & Rose =
C. houstoniana anomala
calothyrsus (Meissner) Britton & Rose = C. houstoniana
calothyrsus
canescens (Chamisso & Schlechtendal) Britton & Rose
= C. tergemina emarginata
capillata (Bentham) Britton & Rose = Zapoteca formosa
(Kunth) H. Hernandez subsp. formosa
caracasana (Jacquin) Britton & Rose = Zapoteca
caracasana (Jacquin) H. Hernandez
centralis Britton & Rose = C. centralis
chiapensis Britton & Rose = C. magdalenae colombiana
chihuahuana Britton & Rose = C. houstoniana anomala colletioides (Grisebach) Britton = C. haematomma
colletioides colomasensis Britton & Rose = C. houstoniana
colomasensis comosa (Swartz) Britton & Rose = C. comosa
compacta Britton & Rose = C. rubescens
confusa (Sprague & Riley) Britton & Rose =
C. houstoniana calothyrsus conzattiana Britton & Rose = C. houstoniana anomala
cookii Britton & Rose = Zapoteca formosa (Kunth)
H. Hernandez subsp. formosa
costaricensis Britton & Rose = Zapoteca costaricensis (Britton & Rose) H. Hernandez
coulteri (S. Watson) Britton & Rose = Zapoteca media (Martens & Galeotti) H. Hernandez
cruziana Britton & Rose = C. tergemina emarginata
cubensis Britton & Rose = Zapoteca formosa (Kunth)
H. Hernandez subsp. gracilis H. Hernandez
cumingii (Bentham) Britton & Rose = C. laxa laxa
deamii Britton & Rose = C. tergemina emarginata
densifolia (Rose ex Harms) Britton & Rose = C. caeciliae
diquetii Britton & Rose emend. McVaugh = C. hirsuta
emarginata (Willdenow) Britton & Rose = C. tergemina
emarginata
enervis Britton = C. enervis eriophylla (Bentham) Britton & Kearney = C. eriophylla
eriophylla etzatlana Britton & Rose = C. houstoniana houstoniana falcifolia Salisbury = C. houstoniana houstoniana
fasciculata (Willdenow) Kleinhoonte = C. surinamensis
formosa (Kunth) Britton & Millspaugh = Zapoteca formosa (Kunth) H. Hernandez subsp. formosa
fulgens (Hooker) Britton & Rose = C. falcata
goldmanii Rose = C. goldmanii gracilis (Grisebach) Britton & Rose = Zapoteca formosa
(Kunth) H. Hernandez subsp. gracilis H. Hernandez
grandiflora (L'Heritier) Britton & Rose =
C. houstoniana anomala
grisebachii Britton & Rose = Zapoteca formosa (Kunth)
H. Hernandez subsp. gracilis H. Hernandez
guildingii (Bentham) Britton & Rose = C. guildingii
haematocephala (Hasskarl) Britton & Wilson =
C. haematocephala haematocephala
herbacea (Engelmann) Britton & Rose = C. humilis
humilis houghiana Britton & Rose = C. hirsuta houstoni (L'Heritier) Sweet = C. houstoniana
houstoniana humilis (Schlechtendal) Britton & Rose = C. humilis
humilis
izalcoensis Britton & Rose = Zapoteca formosa (Kunth)
H. Hernandez subsp. formosa
juchitana Britton & Rose = C. tergemina emarginata
laevis (Rose) Britton & Rose = C. laevis
lagunae Britton & Rose = C. peninsularis
lambertiana (G. Don) Britton & Rose = Zapoteca
lambertiana (G. Don) H. Hernandez
langlassei (Harms) Britton & Rose = C. tergemina
emarginata
leucotricha Britton & Rose = C. tergemina emarginata
lucens Britton = C. houstoniana houstoniana
magdalenae (de Candolle) Britton & Rose =
C. magdalenae magdalenae
marginata (R. O. Williams) Moldenke = Zapoteca
formosa (Kunth) H. Hernandez subsp. formosa
media (Martens & Galleoti) Britton & Rose =Zapoteca media (Martens & Galleoti) H. Hernandez
mexicana (T. Brandegee) Britton & Rose = C. tergemina emarginata
minutifolia Britton & Rose = C. haematomma
haematomma
mixta Britton & Rose = C. californica
mollicula (Martens & Galeotti) Britton & Rose =
Zapoteca formosa (Kunth) H. Hernandez subsp. mollicula (Martens & Galeotti) H. Hernandez
mollis (Standley) Britton & Rose = Zapoteca mollis
(Standley) H. Hernandez
mucronulata Britton & Rose = C. californica nervosa Britton & Rose = Calliandropsis nervosa
(Britton & Rose) H. Hernandez & Guinet nicaraguensis (Taubert & Loesner) Britton & Rose =
Zapoteca portoricensis (Jacquin) H. Hernandez
subsp. portoricensis
nipensis Britton & Rose = C. pauciflora
oaxacana Britton & Rose = Zapoteca media (Martens &
Galeotti) H. Hernandez
orientalis Britton & Rose = Zapoteca formosa (Kunth)
H. Hernandez subsp. formosa
pallida Britton & Rose = C. rufescens
papillosa Britton & Rose = C. rufescens
parvifolia (Hooker & Arnott) Britton = C. parvifolia
pauciflora (A. Richard) Britton & Rose = C. pauciflora pedicellata (Bentham) Britton & Rose = C. pedicellata
penduliflora (Rose) Britton & Rose = Zapoteca formosa
(Kunth) H. Hernandez subsp. formosa
peninsularis (Rose) Britton & Rose = C. peninsularis
pilosa (Bertero) Britton & Rose = C. haematomma glabrata
pittieri (Standley) Britton & Rose = C. pittieri pittieri
portoricensis (Jacquin) J. D. Smith = Zapoteca
portoricensis (Jacquin) H. Hernandez subsp. portoricensis
pubiflora Britton & Rose = C. hirsuta
pueblana Britton & Rose = C. houstoniana anomala
purpurea (Linnaeus) Britton = C. purpurea
purpusii (T. Brandegee) Britton & Rose = C. tergemina emarginata
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 217
quetzal (J. D. Smith) J. D. Smith = C. quetzal
rekoi Britton & Rose = C. trinervia arborea
reticulata (A. Gray) Britton & Rose s C. humilis reticulata
rhodocephala (J. D. Smith) Britton & Rose =
C. rhodocephala
rubescens (Martens & Galeotti) Britton & Rose = C. rubescens
rupestris (T. Brandegee) Britton & Rose = C. tergemina emarginata
rusbyi Britton & Rose = C. houstoniana anomala
salvadorensis Britton & Rose = Zapoteca formosa
(Kunth) H. Hernandez subsp. salvadorensis (Britton & Rose) H. Hernandez
schottii Britton & Rose = Zapoteca formosa (Kunth)
H. Hernandez subsp. schottii (Britton & Rose) H. Hernandez
seemannii (Bentham) Britton & Rose = C. tergemina emarginata
seleri (Harms) Britton & Rose = C. seleri
simulans Britton & Rose = C. rubescens
sinaloana Britton & Rose = C. tergemina emarginata
socorrensis (I. Johnston) Britton & Rose = Zapoteca
formosa (Kunth) H. Hernandez subsp. rosei (Wiggins) H. Hernandez
speciosa (Martens & Galeotti) Britton & Rose = C. hirsuta
strigillosa Britton & Rose = C. houstoniana anomala
surinamensis (Bentham) Kleinhoonte = C. surinamensis
tergemina (Linnaeus) Britton & Rose = C. tergemina
tergemina
tetragona (Willdenow) J. D. Smith = Zapoteca tetragona (Willdenow) H. Hernandez
tonduzii Britton & Rose = C. magdalenae colombiana unijuga (Rose) Britton & Rose = Zapoteca formosa
(Kunth) H. Hernandez subsp. mollicula (Martens
& Galeotti) H. Hernandez
xalapensis (Bentham) Britton & Rose = C. rubescens yoroensis Britton = Zapoteca formosa (Kunth)
H. Hernandez subsp. formosa
yucatanensis Britton & Rose = C. tergemina
emarginata
Calliandra Bentham, 3
sect. Acistegia Bameby, 139
sect. Acroscias Bameby, 146
sect. Androcallis Bameby, 21
ser. Ambivalentes Bameby, 103
ser. Androcallis Bameby, 21
ser. Biflorae Bameby, 98
ser. Chilenses Bameby, 100
ser. Hymenaeodeae Bameby, 134
ser. Longipedes Barneby, 137 ser. Macrophyllae Bentham, 111
ser. Pauciflorae Bameby, 101
sect. Calliandra, 148
ser. Calliandra, 149
ser. Comosae Bameby, 196
ser. Tsugoideae Barneby, 190
ser. Virgatae Barneby, 189
sect. Caulanthon Grisebach = Zygia P. Browne
sect. Eucalliandra Grisebach = sect. Calliandra, 148
sect. Formosae Britton & Rose = Zapoteca
H. Hernandez
sect. Laetevirentes Bentham = Zapoteca H. Hernandez
sect. Microcallis Bameby, 187
ser. Leptopodae Bameby, 199
ser. Microcallis Bameby, 198
sect. Portoricensis Britton & Rose = Zapoteca
H. Hernandez
sect. Pseudacacia Grisebach = Zapoteca H. Hernandez
sect. Saman Grisebach = Samanea Merrill
ser. Laetevirentes Bentham = Zapoteca H. Hernandez
ser. Nitidae Bentham = ser. Androcallis, 21
ser. Pedicellatae Bentham = sect. Acistegia, 139
ser. Racemosae Bentham = ser. Calliandra, 149
abbreviata Bentham = C. dysantha dysantha
aculeata Spmce ex Bentham = Zapoteca aculeata (Bentham) H. Hernandez
aeschynomenoides Bentham, 98
affinis Pittier = C. cruegeri
amazonica Bentham s Zapoteca amazonica (Bentham) H. Hernandez
amblyophylla Harms = C. falcata
angelica Bentham = Zapoteca media (Martens &
Galeotti) H. Hernandez
angusta Renvoize = C. calycina angustidens Britton & Killip = C. surinamensis
angustifolia Spmce ex Bentham, 125, 127*
anomala (Kunth) Macbride = C. houstoniana anomala var. longipedicellata McVaugh = C. houstoniana
anomala
anthoniae Grimes = C. coriacea
antioquiae Bameby, 123
arborea Standley = C. trinervia arborea
aristulata Rizzini = Chloroleucon foliolosum (Bentham) G. P. Lewis
asplenioides (Nees) Renvoize, 155 axillaris Bentham = C. sessilis bahiana Renvoize, 151
var. bahiana, 151
var. erythematosa Bameby, 151 belizensis (Standley) Standley, 34, 36* bella Bentham, 58, 60*
var. trianae Bentham = C. pittieri pittieri bicolor Bentham = C. parvifolia
biflora Tharp, 86*, 99, 99*
bijuga Rose, 36*, 37
blakeana Pittier = C. glomerulata parvifolia blanchetiana Bentham, 97
boliviana Britton = C. haematocephala boliviana bombycina Spruce ex Bentham, 118, 120*
brachyandra Spegazzini = Chloroleucon tenuifolium (Bentham) Barneby & Grimes
bracteosa Bentham = C. fasciculata bracteosa
brandegeei (Britton & Rose) Gentry = C. peninsularis brenesii Standley, 132, 134*
brevicaulis M. Micheli, 146, 147*
var. brevicaulis, 148, 149*
var. genuina Chodat & Hassler = C. brevicaulis brevicaulis
fma. robusta Chodat & Hassler = C. brevicaulis brevicaulis
var. glabra Chodat & Hassler, 148, 149*
fma. grandiflora Hassler = C. brevicaulis glabra
fma. parviflora Hassler = C. brevicaulis glabra
fma. roseiflora Chodat & Hassler = C. brevicaulis glabra
218 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Calliandra Bentham (continued) var. puberula Hassler = C. brevicaulis brevicaulis
fma. hirsutula Hassler = C. brevicaulis brevicaulis
var. pubescens Hassler = C. brevicaulis brevicaulis
fma. intermedia Hassler = C. brevicaulis brevicaulis
var. tomentosa Hassler = C. brevicaulis brevicaulis
brevipes Bentham, 90, 91*, 93*
bullata Urban = C. enervis
caeciliae Harms, 84, 86*
californica Bentham, 42, 43*
callistemon Schlechtendal = C. houstoniana anomala
calothyrsus Meissner = C. houstonianus calothyrsus
canescens (Chamisso & Schlechtendal) Bentham = C. tergemina emarginata
capillata Bentham = Zapoteca formosa (Kunth)
H. Hernandez subsp. formosa caracasana (Jacquin) Bentham = Zapoteca formosa
(Kunth) H. Hernandez subsp. caracasana
(Jacquin) H. Hernandez
carbonaria Bentham = C. trinervia carbonaria
carcerea Standley & Steyermark, 87
carrascana Bameby, 55
catingae Harms = C. squarrosa
centralis (Britton & Rose) Standley = C. trinervia
arborea
chamaedrys Engelmann = C. eriophylla eriophylla
chapadae S. Moore = C. dysantha turbinata
chiapensis (Britton & Rose) Lundell = C. magdalenae
colombiana
chilensis Bentham, 51*, 100
chotanoana Harms = C. mollissima chulumania Bameby, 85
cinerea Taubert = C. staminea
clavellina Karsten = C. purdiaei
coccinea Renvoize, 167
var. coccinea, 168
var. trimera Bameby, 168, 168*
codonandra Bentham = C. magdalenae magdalenae
colimae Barneby, 88
colletioides Grisebach = C. haematomma colletioides
subsp. gonavensis (Urban & Ekman) Bassler = C. haematomma haematomma
colombiana Britton & Killip = C. magdalenae
colombiana comosa (Swartz) Bentham, 196
comosa Grisebach = Zapoteca formosa (Kunth)
H. Hernandez subsp. gracilis H. Hernandez
concinna Bameby, 71, 71*
conferta Bentham, 86*, 89, 90*
confusa Sprague & Riley = C. houstoniana calothyrsus
conzattiana (Britton & Rose) Standley = C. houstoniana
grandiflora
conzattii Standley = Zygia conzattii (Standley) Britton
& Rose
cookii (Britton & Rose) Standley = Zapoteca formosa
(Kunth) H. Hernandez subsp. formosa
coriacea (Willdenow) Bentham, 122, 124*
var. aquae-nigrae Bameby = C. coriacea
coroensis Karsten = C. purpurea
costaricensis (Britton & Rose) Standley = Zapoteca
costaricensis (Britton & Rose) H. Hernandez
coulteri S. Watson = Zapoteca media (Martens &
Galeotti) H. Hernandez
crassipes Bentham, 164
cruegeri Grisebach, 26, 27*, 29* cruziana (Britton & Rose) Standley = C. tergemina
emarginata cubensis (Macbride) Leon = Zapoteca formosa (Kunth)
H. Hernandez subsp. gracilis H. Hernandez
x cumbucana Renvoize, 162
cumingii Bentham = C. laxa laxa
var. galeottii Bentham = C. hirsuta
cylindrocarpa Bentham = C. harrisii
cynometroides Beddome, 3
deamii (Britton & Rose) Standley = C. tergemina
emarginata
debilis Renvoize, 171 decrescens Killip & Macbride = C. guildingii
dendroides Renvoize = C. asplenioides densifolia Rose ex Harms = C. caeciliae
depauperata Bentham, 73, 74*, 75*
diademata Lemaire = C. foliolosa
diversifolia Britton & Killip = C. laxa stipulacea
duckei Bameby, 95
dysantha Bentham, 60
var. dysantha, 62, 64*
var. macrocephala (Bentham) Bameby, 63, 65*
var. opulenta Bameby, 63, 65* var. pilosa Bentham = C. dysantha dysantha
var. turbinata (Bentham) Bameby, 64, 65*
elegans Renvoize, 159 emarginata (Willdenow) Bentham = C. tergemina
emarginata enervis (Britton) Urban, 102, 102*
eriophylla Bentham, 44
var. chamaedrys Isely, 43*, 45
var. eriophylla, 43*, 44
erubescens Renvoize, 174
erythrocephala H. Hernandez & Sousa, 110
expansa Bentham = C. taxifolia
exsudans Harms = C. sincorana
falcata Bentham, 105, 107*
falcifera Ducke = C. parvifolia
fasciculata Bentham, 158
var. bracteosa (Bentham) Bameby, 158 var. fasiculata, 158
fasciculata (Willdenow) Bentham = C. surinamensis
feioana Renvoize, 154
fernandesii Bameby, 67, 68*, 69*
filipes Bentham = Zapoteca filipes (Bentham)
H. Hernandez
flavida Urban = Zapoteca portoricensis (Jacquin)
H. Hernandez subsp. flavida (Urban) H. Hernandez
foliolosa Bentham, 55, 57*
formosa (Kunth) Bentham = Zapoteca formosa (Kunth)
H. Hernandez subsp. formosa
fulgens Hooker f. = C. falcata
fuscipila Harms, 152
ganevii Bameby, 173
gardneri Bentham, 60*, 64
geminata Bentham, 3
gentryi Standley = Zapoteca formosa (Kunth)
H. Hernandez subsp. formosa
germana Bameby, 175
gilbertii Thulin & Hunde, 2
glaberrima (Bentham) Britton & Killip = C. coriacea
glabra (Chodat & Hassler) Hassler = C. brevicaulis
glabra
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 219
glaziovii Taubert, 73
glomerulata Karsten, 25, 25*
var. glomerulata 25
var. parvifolia (Bentham) Bameby, 26
glyphoxylon Spruce ex Bentham, 119, 120*
var. glaberrima Bentham = C. coriacea
goldmanii Rose ex Bameby, 35, 36*
gracilis Klotzsch = Zapoteca formosa (Kunth)
H. Hernandez subsp. gracilis H. Hernandez gracilis Renvoize = C. renvoizeana
grandiflora (L'Heritier) Bentham = C. houstoniana anomala
grandifolia P. H. Allen, 204
grisebachiana Spegazzini = Chloroleucon foliolosum (Bentham) G. P. Lewis
var. carolae Spegazzini, 204
grisebachii (Britton & Rose) Leon = Zapoteca formosa
(Kunth) H. Hernandez subsp. gracilis H. Hernandez
guildingii Bentham, 104, 105*
haematocephala Hasskarl, 108
var. boliviana (Britton) Barneby, 107*, 109
var. haematocephala, 107*, 109
haematomma (de Candolle) Bentham, 140
var. colletioides (Grisebach) Bameby, 141, 143*
var. correllii Bameby, 142, 143*
var. glabrata Grisebach, 143, 143*, 144*
var. haematomma, 141, 143*
var. locoensis (Garcia & Kolterman) Bameby, 144
var. pubescens (Urban) Macbride = C. haematomma haematomma
var. rivularis (Urban & Ekman) Bameby, 142
var. tortuensis (Alain) Bameby, 143 haematostoma Sprengel = C. haematomma
haematomma var. genuina Urban = C. haematomma haematomma
var. minutifolia Urban = C. haematomma
haematomma
harrisii (Lindley) Bentham, 126, 127*, 128*
herbacea Engelmann = C. humilis humilis
hintonii Bameby, 88
hirsuta (G. Don) Bentham, 38, 40* var. sancti-pauli (Hasskarl) Macbride = C. foliolosa
hirsuticaulis Harms, 163
hirticaulis. Harms = C. hirsuticaulis
hirtiflora Bentham, 165
var. hirtiflora, 165
var. ripicola Bameby, 165
hookeriana Schomburgk = C. rigida houstoni (L'Heritier) Bentham = C. houstoniana
houstoniana
houstoniana (Miller) Standley, 177 subsp. alamosensis (Britton & Rose) Macqueen &
H. Hernandez = C. houstoniana houstoniana
subsp. stylesii Macqueen = C. houstoniana
houstoniana var. acapulcensis (Britton & Rose) Bameby, 180,181*
var. anomala (Kunth) Bameby, 179, 181*
var. calothyrsus (Meissner) Bameby, 180, 183*
var. colomasensis (Britton & Rose) Bameby, 180
var. houstoniana, 182, 184* var. longipedicellata MacVaugh = C. houstoniana
houstoniana var. pedicellata de Candolle = C. houstoniana anomala
humilis Bentham, 45
var. gentryana Bameby, 47, 49*
var. humilis, 47, 49* var. reticulata (A. Gray) L. Benson, 48, 48*, 50*
humilis Schlechtendal = C. humilis humilis
hygrophila Mackinder & Lewis, 176
hymenaeodes (Persoon) Bentham, 134, 136*, 138*
hystrix (A. Richard) Grisebach = Pithecellobium hystrix
(A. Richard) Bentham
iligna Bameby, 171, 172*
imperialis Bameby, 69*, 70 inaequilatera Rusby = C. haematocephala
haematocephala
inermis Druce = C. houstoniana houstoniana
involuta Mackinder & Lewis, 168 izalcoensis (Britton & Rose) Standley = Zapoteca
formosa (Kunth) H. Hernandez subsp. formosa
jacobiana Renvoize = C. calycina
jariensis Bameby, 120, 120*, 121* juzepczukii Standley, 182, 185*, 186*
kingii Prain, 3
kuhlmannii Hoehne, 204
laevis Rose, 133, 134*, 135* lambertiana (G. Don) Bentham = Zapoteca lambertiana
(G. Don) H. Hernandez
lanata Bentham, 152, 153*
langlassei Harms = C. tergemina emarginata
lasiopus (Bentham) Grisebach = Zygia latifolia
(Linnaeus) Fawcett & Rendle var. lasiopus
(Bentham) Bameby & Grimes latifolia (Linnaeus) Grisebach = Zygia latifolia
(Linnaeus) Fawcett & Rendle laxa (Willdenow) Bentham, 29
var. laxa, 31, 32*
var. parvifolia Bentham = C. glomerulata parvifolia
var. stipulacea (Bentham) Bameby, 32, 32*
var. urimana Bameby, 33
lehmannii Harms = C. pittieri pittieri
leptopoda Bentham, 199, 201*
leucothrix Standley = C. houstoniana anomala
linearis Bentham, 157, 159*
lintea Bameby, 149
locoensis Garcia & Kolterman = C. haematomma
locoensis
longipes Bentham, 137, 138*, 139*
var. valenzuelensis Chodat & Hassler = C. longipes fma. nana Chodat & Hassler = C. longipes
longipinna Bentham, 170
lucens (Britton) Standley = C. houstoniana houstoniana luetzelburgii Harms, 176
macqueenii Bameby, 131
macrocalyx Harms, 66
var. aucta Bameby, 66, 67*
var. macrocalyx, 66, 67*
magdalenae (de Candolle) Bentham, 82
var. colombiana (Britton & Killip) Bameby, 83, 85* var. magdalenae, 84, 85*
malacophylla Bentham = Zapoteca formosa (Kunth)
H. Hernandez subsp. formosa
marginata (R. O. Williams) Moldenke = Zapoteca
formosa (Kunth) H. Hernandez subsp. formosa matisiana Uribe = C. tolimensis
matudai Lundell = C. hirsuta hirsuta
medellinensis Britton & Rose ex Britton & Killip, 103
220 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Calliandra Bentham (continued) media (Martens & Galeotti) Standley = Zapoteca media
(Martens & Galeotti) H. Hernandez mertensioides (Nees & Martius) Bentham =
C. asplenioides var. debilis Bentham = C. silvicola
metrosiderifolia (Schlechtendal) Bentham ex Jackson =
C. houstoniana houstoniana
mexicana T. Brandegee = C. tergemina emarginata
michelii (Britton & Rose) Standley = C. bijuga
microcalyx Harms = C. parvifolia
microcephala Britton & Killip = Zapoteca microcephala
(Britton & Killip) H. Hernandez
minutifolia Pittier = C. glomerulata parvifolia molinae Standley, 36*, 37
mollicula (Martens & Galeotti) Standley = Zapoteca
formosa (Kunth) H. Hernandez subsp. mollicula
(Martens & Galeotti) H. Hernandez
mollis Standley = Zapoteca mollis (Standley) H.
Hernandez
mollissima (Willdenow) Bentham, 103, 105*
moritziana Cardenas = Zapoteca formosa (Kunth)
H. Hernandez subsp. formosa
mucugeana Renvoize, 160, 161*
myriophylla Bentham = C. parvifolia nebulosa Barneby, 150
nervosa Urban & Ekman = Zapoteca nervosa (Urban &
Ekman) H. Hernandez
nicaraguensis Taubert & Loesner = 2&poteca
portoricensis (Jacquin) H. Hernandez subsp.
nicaraguensis (Taubert & Loesner) H. Hernandez
nitida S. Watson = C. hirsuta
nogalensis Lundell = Zapoteca portoricensis (Jacquin)
H. Hernandez subsp. portoricensis
novaesii Hoehne = C. haematocephala haematocephala oaxacana Rose = Zapoteca media (Martens & Galeotti)
H. Hernandez
obovata Bentham = Abarema obovata (Bentham)
Bameby & Grimes obtusifolia (Willdenow) Karsten = C. purpurea
orientalis (Britton & Rose) Leon = Zapoteca formosa
(Kunth) H. Hernandez subsp. formosa
orinocensis Pittier, 204 pacara Grisebach = Enterolobium contortisiliquum
(Velloso) Morong
pakaraimensis Cowan, 193, 195*, 196*
pallatangensis Harms = C. glyphoxylon
pallens Bentham = Havardia pallens (Bentham) Britton
&Rose
pallida Standley = C. rufescens
palmeri S. Watson, 184, 186*
paniculata Adams, 197 panlosia J. R. Johnston = C. laxa laxa
papillosa (Britton & Rose) Standley = C. rubescens
parviflora Bentham, 198, 198*, 200* parvifolia (Hooker & Arnott) Spegazzini, 52, 54*
paterna Bameby, 172
patrisii Sagot = C. hymenaeodes pauciflora (A. Richard) Grisebach, 101
peckoltii Bentham = C. parvifolia
pedicellata Bentham, 144, 145* penduliflora Rose = Zapoteca formosa (Kunth)
H. Hernandez subsp. formosa
peninsularis Rose, 40 physocalyx H. Hernandez & Sousa, 186 picardae Alain = C. haematomma haematomma
pilgerana Harms, 69, 69* pilosa (Bertero) Urban = C. haematomma glabrata
pilosifolia Cowan = C. trinervis pilosifolia
pittieri Standley, 22
var. pittieri, 22, 24*
var. polyphylla (Harms) Bameby, 23, 24*
pityophila Bameby, 87
polyphylla Harms = C. pittieri polyphylla
porphyrea Pittier = C. pittieri polyphylla
portoricensis (Jacquin) Bentham = Zapoteca
portoricensis (Jacquin) H. Hernandez subsp.
portoricensis
var. major Sprague = Zapoteca portoricensis (Jacquin)
H. Hernandez subsp. portoricensis
var. multijuga M. Micheli = Zapoteca tetragona
(Willdenow) H. Hernandez
prehensilis C. Wright = Sphinga prehensilis (C. Wright) Bameby & Grimes
prostrata Bentham = C. taxifolia
pubens Renvoize, 204 pubiflora (Britton & Rose) Standley = C. hirsuta
purdiaei Bentham, 23, 25*
purpurea (Linnaeus) Bentham, 78, 79*
var. dussiana Stehle" = C. purpurea var. quentiniana Stehle = C. purpurea
purpusii T. Brandegee = C. tergemina emarginata
pyrophila Fernandez Casas & Schinini = C. longipes
quetzal J. D. Smith, 187, 188* redacta (J. H. Ross) Thulin & Hunde, 2
rekoi (Britton & Rose) Standley = C. trinervia
arborea
renvoizeana Bameby, 169
resupina Cowan = C. pakaraimensis
reticulata A. Gray s C. humilis reticulata
revoluta Grisebach = Abarema obovalis (A. Richard)
Bameby & Grimes rhodocephala J. D. Smith, 111, 112*
rigida Bentham, 194, 196*
riparia Pittier, 80, 81*, 82*
rivalis Lundell = C. coriacea rivularis Urban & Ekman = C. haematomma rivularis
robusta Renvoize = C. calycina
rondoniana Hoehne, 204
rosei Wiggins = Zapoteca formosa (Kunth) H.
Hernandez subsp. rosei (Wiggins) H. Hernandez
rotundifolia Killip & Macbride = C. trinervia trinervia
rubescens (Martens & Galeotti) Standley, 33, 35* rupestris T. Brandegee = C. pittieri polyphylla
saman (Jacquin) Grisebach = Samanea saman (Jacquin)
Merrill
samik Bameby, 77
sancti-pauli Hasskarl = C. foliolosa santanderensis Britton & Killip = C. magdalenae
colombiana
santosiana Glaziou ex Bameby, 169
schottii (Britton & Rose) S. Watson = Zapoteca formosa
(Kunth) H. Hernandez subsp. schottii (Britton & Rose) H. Hernandez
schultzei Harms = C. riparia
schwackeana Taubert = Zygia cataractae (Kunth) Rico
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 221
scopulina T. Brandegee = Zygia scopulina
(T. Brandegee) Britton & Rose
scutellifera Bentham = Zapoteca scutellifera (Bentham) H. Hernandez
seemannii Bentham = C. tergemina emarginata seleri Harms, 204
selloi (Sprengel) Macbride, 204
semisepulta Bameby, 174
sericea Renvoize = C. lanata
serjanioides Urban = C. falcata
sessilis Bentham, 92, 95*
siltepecensis Lundell = Zapoteca portoricensis (Jacquin)
H. Hernandez subsp. portoricensis silvicola Taubert, 75
similis Sprague & Riley = C. houstoniana calothyrsus sinaloana (Britton & Rose) Standley = C. tergemina
emarginata
sincorana Harms, 166
slaneae Howard = C. purpurea
socorrensis I. Johnston = Zapoteca formosa (Kunth)
H. Hernandez subsp. rosei (Wiggins) H. Hernandez sodiroi Harms = C. angustifolia
speciosa (Martens & Galeotti) Standley = C. hirsuta spinosa Ducke, 93*, 94, 96*
splendens (Britton & Rose) Standley = C. trinervia
arborea spraguei (Britton & Rose) Lundell = Zapoteca
portoricensis (Jacquin) H. Hernandez subsp.
portoricensis spruceana Bentham = Chloroleucon mangense (Jacquin)
Britton & Rose
squarrosa (Martius) Bentham, 72
var. crassifolia Bentham = C. nebulosa
staminea (Thunberg) Bameby, 92
stelligera Bameby, 166
stenophylla Wawra, 204
stipulacea Bentham = C. laxa stipulacea
stricta Rusby = C. angustifolia strigillosa (Britton & Rose) Standley ex Leavenworth =
C. houstoniana anomala
suberifera Rizzini = C. spinosa subnervosa Bentham = C. angustifolia
subspicata Bentham, 59, 60*
surinamensis Bentham, 76, 78* tamarindifolia Ettingshausen, 204
tapirorum Standley = C. rubescens
taxifolia (Kunth) Bentham, 50, 51*
tenuiflora Bentham = C. surinamensis
tergemina (Linnaeus) Bentham, 127 var. emarginata (Willdenow) Bameby, 129, 132*
var. tergemina, 131, 132* tergemina (Linnaeus) Standley s C. tergemina tergemina
tetragona (Willdenow) Bentham = Zapoteca tetragona
(Willdenow) H. Hernandez
tetraphylla (G. Don) Bentham = C. tergemina
emarginata
tocantina Ducke = C. sessilis
tolimensis Taubert, 25*, 28, 30* tolimensis Uribe = C. tergemina emarginata
tonduzii (Britton & Rose) Standley = C. magdalenae
colombiana
toroana Britton & Killip = Zapoteca tetragona
(Willdenow) H. Hernandez
tortuensis Alain = C. haematomma tortuensis
trijugata Schery = C. laxa laxa
trinervia Bentham, 113
var. arborea (Standley) Bameby, 117, 119*
var. carbonaria (Bentham) Bameby, 115, 16*
var. paniculans Bameby, 116*, 117
var. parvifolia Huber = C. trinervia trinervia
var. peruicola Bameby, 115, 116* var. pilosifolia (Cowan) Bameby, 116, 116*
var. stenocylix Bameby, 116*, 117
var. trinervia, 114, 116*
tsugoides Cowan, 190, 192* tubulosa Bentham = Samanea tubulosa (Bentham)
Bameby & Grimes tumbeziana Macbride, 49, 51* turbinata Bentham = C. dysantha turbinata
tweedii Bentham, 56, 59* var. sancti-pauli (Hasskarl) Bentham =
C. foliolosa
ulei Harms, 68, 69* umbellifera Bentham, 69*, 70
umbrosa Wallich, 3
var. griffithii (Baker) Paul, 3 unijuga Rose = Zapoteca formosa (Kunth)
H. Hernandez subsp. mollicula (Martens &
Galeotti) H. Hernandez
urbanii Alain, 204
uribei Kilhp & Dugand = C. tergemina emarginata vaupesiana Cowan, 191
var. oligandra Bameby, 193, 194*
var. vaupesiana, 193, 194* vespertina (Macfadyen) Bentham = Zapoteca
portoricensis (Jacquin) H. Hernandez subsp.
portoricensis
villosiflora Harms = C. macrocalyx macrocalyx
virgata Bentham, 189, 190*, 191*
viridiflora Bentham = Zapoteca formosa (Kunth)
H. Hernandez subsp. formosa
viscidula Bentham, 156
var. leucantha Bentham, 204
weberbaueri Harms = Zapoteca formosa (Kunth)
H. Hernandez subsp. weberbaueri (Harms) H. Hernandez
wendlandii Bentham, 186
xalapensis Bentham = C. rubescens
yoroensis (Britton) Standley = Zapoteca formosa
(Kunth) H. Hernandez subsp. formosa
yucatanensis (Britton & Rose) Standley = C. tergemina
emarginata
yucunensis N. Mattos, 204
yunckeri Standley = C. rubescens
Calliandropsis H. Hernandez & Guinet, 2
nervosa (Britton & Rose) H. Hernandez & Guinet, 2 Clelia Casaretto = Calliandra
ornata Casaretto = C. harrrisii
Codonandra Karsten = Calliandra
purpurea Karsten = C. magdalenae magdalenae Cojoba Britton & Rose
siltepecensis (Lundell) Rico = C. hirsuta
Desmanthus Willdenow, 2
Dichrostachys Wight & Arnott
kirkii Bentham, 2
222 MEMOIRS OF THE N E W YORK BOTANICAL GARDEN [VOL. 74(3)
Feuilleea O. Kuntze
abbreviata (Bentham) O. Kuntze = C. dysantha dysantha
aeschynomenoides (Bentham) O. Kuntze = C.
aeschynomenoides
amplebracteata O. Kuntze = C. fasciculata bracteosa
angelica (Bentham) O. Kuntze = Zapoteca media
(Martens & Galeotti) H. Hernandez
angustifolia (Bentham) O. Kuntze s C. angustifolia
asplenioides (Nees) O. Kuntze = C. asplenioides
axillaris (Bentham) O. Kuntze = C. sessilis
bella (Bentham) O. Kuntze = C. bella
benthamiana O. Kuntze = C. fasciculata fasciculata
blanchetii (Bentham) O. Kuntze = C. blanchetii
boliviana (Britton) O. Kuntze = C. haematocephala
boliviana
bombycina (Bentham) O. Kuntze = C. bombycina
brevipes (Bentham) O. Kuntze = C. brevipes
calothyrsa (Meissner) O. Kuntze = C. houstoniana
calothyrsus
calycina (Bentham) O. Kuntze = C. calycina
canescens (Chamisso & Schlechtendal) O. Kuntze =
C. tergemina emarginata
caracasana (Jacquin) O. Kuntze = Zapoteca caracasana
(Jacquin) H. Hernandez subsp. caracasana
carbonaria (Bentham) O. Kuntze = C. trinervia
carbonaria
cearana O. Kuntze = C. umbellifera
chilensis (Bentham) O. Kuntze = C. chilensis
codonandra (Bentham) O. Kuntze = C. magdalenae
magdalenae
colletioides (Grisebach) O. Kuntze = C. haematomma
colletioides
comosa (Swartz) O. Kuntze = C. comosa
crassipes (Bentham) O. Kuntze s C. crassipes cruegeri (Bentham) O. Kuntze = C. cruegeri
cumingii (Bentham) O. Kuntze = C. laxa laxa depauperata (Bentham) O. Kuntze s C. depauperata
dimidiata O. Kuntze = Zapoteca filipes (Bentham)
H. Hernandez dysantha (Bentham) O. Kuntze = C. dysantha dysantha
emarginata (Willdenow) O. Kuntze = C. tergemina
emarginata expansa (Bentham) O. Kuntze = C. taxifolia
fasciculata (Willdenow) O. Kuntze = C. surinamensis
foliolosa (Bentham) O. Kuntze = C. foliolosa
formosa (Kunth) O. Kuntze = Zapoteca formosa (Kunth)
H. Hernandez subsp. formosa
gardneri (Bentham) O. Kuntze = C. gardneri
glomerulata (Karsten) O. Kuntze = C. glomerulata
glomerulata
glyphoxylon (Bentham) O. Kuntze = C. glyphoxylon
guildingii (Bentham) O. Kuntze = C. guildingii
haematocephala (Hasskarl) O. Kuntze =
C. haematocephala haematocephala
harrisii (Lindley) O. Kuntze = C. harrisii
hirsuta (G. Don) O. Kuntze = C. hirsuta
hirtiflora (Bentham) O. Kuntze s C. hirtiflora
humilis (Schlechtendal) O. Kuntze = C. humilis humilis
hymenaeodes (Persoon) O. Kuntze = C. hymenaeodes
lambertiana (G. Don) O. Kuntze = Zapoteca lambertiana
(G. Don) H. Hernandez
lanata (Bentham) O. Kuntze = C. lanata
laxa (Willdenow) O. Kuntze h C. laxa laxa
leptopoda (Bentham) O. Kuntze = C. leptopoda
linearis (Bentham) O. Kuntze = C. linearis
longipes (Bentham) O. Kuntze = C. longipes
longipinna (Bentham) O. Kuntze = C. longipinna
macrocephala (Bentham) O. Kuntze = C. dysantha
macrocephala magdalenae (de Candolle) O. Kuntze = C. magdalenae
magdalenae
multifoliolata O. Kuntze = C. parvifolia
mutica O. Kuntze s C. sessilis
myriophylla (Bentham) O. Kuntze = C. parvifolia
pacara (Grisebach) O. Kuntze = Enterolobium
contortisiliquum (Vellozo) Morong
pachyloma (Martius) O. Kuntze = C. parviflora
peckoltii (Bentham) O. Kuntze = C. parviflora
portoricensis (Jacquin) O. Kuntze = Zapoteca
portoricensis (Jacquin) H. Hernandez subsp.
portoricensis
prostrata (Bentham) O. Kuntze = C. taxifolia
purdiaei (Bentham) O. Kuntze = C. purdiaei
purpurea (Linnaeus) O. Kuntze = C. purpurea
quitaro O. Kuntze = C. laxa stipulacea
reticulata (A. Gray) O. Kuntze = C. humilis reticulata
rigida (Bentham) O. Kuntze = C. rigida
scutellifera (Bentham) O. Kuntze = Zapoteca scutellifera
(Bentham) H. Hernandez
squarrosa (Bentham) O. Kuntze = C. squarrosa subnervosa (Bentham) O. Kuntze = C. angustifolia
subspicata (Bentham) O. Kuntze = C. subspicata tenuiflora (Bentham) O. Kuntze = C. surinamensis
tergemina (Linnaeus) O. Kuntze = C. tergemina
tergemina tetragona (Willdenow) O. Kuntze = Zapoteca tetragona
(Willdenow) H. Hernandez
tetraphylla (G. Don) O. Kuntze = C. tergemina
emarginata texana O. Kuntze = C. conferta
trinervia (Bentham) O. Kuntze = C. trinervia trinervia turbinata (Bentham) O. Kuntze = C. dysantha turbinata
tweedii (Bentham) O. Kuntze = C. tweedii
virgata (Bentham) O. Kuntze = C. virgata
viscidula (Bentham) O. Kuntze = C. viscidula
Inga Miller
sect. Annesleya G. Don = C. sect. Calliandra
anomala Kunth = C. houstoniana anomala
var. pedicellata de Candolle = C. houstoniana
anomala
bauhiniifolia Poeppig = Zapoteca amazonica (Bentham)
H. Hernandez
canescens Chamisso & Schlechtendal = C. tergemina
emarginata
caripensis Willdenow = C. tergemina tergemina
comosa (Swartz) Willdenow = C. comosa
coriacea Willdenow = C. coriacea
emarginata Willdenow = C. tergemina emarginata fasciculata Willdenow = C. surinamensis
harrisii Lindley = C. harrisii
hirsuta G. Don = C. hirsuta
houstoni (L'Heritier) de Candolle = C. houstoniana
houstoniana
hymenaeodes (Persoon) Desvaux = C. hymenaeodes
mertensioides Nees & Martius = C. asplenioides
1998] SILK TREE, GUANACASTE, MONKEY'S EARRING 223
mollissima Willdenow = C. mollissima
obtusifolia (Willdenow) Karsten = C. purpurea
parvifolia Hooker & Arnott = C. parvifolia
pulcherrima Cervantes ex Sweet = C. tweedii
purpurea (Linnaeus) Willdenow = C. purpurea
semicordata Bertoloni = C. tergemina emarginata taxifolia Kunth = C. taxifolia
tergemina (Linnaeus) Willdenow = C. tergemina
tergemina
tetraphylla G. Don = C. tergemina emarginata
Jacqueshuberia Ducke, 2
Mimosa Linnaeus
caripensis Willdenow = C. tergemina tergemina
comosa Swartz = C. comosa
emarginata (Willdenow) Poiret = C. tergemina emarginata
grandiflora L'Heritier = C. houstoniana anomala
houstoni L'Heritier = C. houstoniana houstoniana
houstoniana Miller = C. houstoniana houstoniana
hymenaeodes Persoon = C. hymenaeodes
laxa (Willdenow) Poiret = C. laxa laxa
mollissima (Willdenow) Poiret = C. mollissima
obtusifolia (Willdenow) Poiret = C. purpurea portoricensis Jacquin = Zapoteca portoricensis (Jacquin)
H. Hernandez subsp. portoricensis quadrangularis Poiret = Zapoteca tetragona (Willdenow)
H. Hernandez
spartioides Vahl = C. haematomma haematomma
staminea Thunberg = C. staminea
tergemina Linnaeus = C. tergemina tergemina
yaguaronensis Larranaga = C. tweedii
Viguieranthus Villiers, 3
