Monograph of Andira (Legimnosae-Papilionoideae)

Monograph of Andira (Leguminosae-Papilionoideae)
Author(s): R. Toby Pennington

Source: Systematic Botany Monographs, Vol. 64, Monograph of Andira (LeguminosaePapilionoideae) (Mar. 31, 2003), pp. 1-143
Published by: American Society of Plant Taxonomists
Stable URL: http://www.jstor.org/stable/25027903 .
Accessed: 03/09/2014 14:45
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .
http://www.jstor.org/page/info/about/policies/terms.jsp
.
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of
content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms
of scholarship. For more information about JSTOR, please contact support@jstor.org.
American Society of Plant Taxonomists is collaborating with JSTOR to digitize, preserve and extend access to
Systematic Botany Monographs.
http://www.jstor.org
This content downloaded from 158.39.37.3 on Wed, 3 Sep 2014 14:45:31 PM

All use subject to JSTOR Terms and Conditions
MONOGRAPH OF ANDIRA
(LEGUMINOSAE-PAPILIONOIDEAE)
R. Toby Pennington
Botanic
Garden Edinburgh
Royal
20a Inverleith Row
Edinburgh EH3 5LR, United Kingdom
The principally Neotropical
29 species; one
genus Andira Lam. (Leguminosae)
comprises
two of which occur in Africa. Chloroplast DNA restriction site
includes three subspecies,
analysis allows phylogenetic
placement of 21 species o? Andira,
including five species that were not previously
studied. These data are also used to investigate
and to identify ac
intraspecific chloroplast DNA polymorphism
Abstract.
species,
cessions
A.
inermis,
of A. humilis with plastomes

characters provides
12 morphological
of cpDNA
restriction
A cladistic analysis of
from probable introgressive hybridization.
little phylogenetic
resolution in comparison with a simultaneous
analysis
some of the clades discovered
site data and morphology.
Although
by this analysis are
derived
characters and have no morphological

the
"cryptic," i.e., they are supported only by molecular
synapomorphies,
results provide a reasonable estimate of the phylogeny
of Andira and are used as the basis of discussions
of char
acter evolution,
and biogeography.
Previous
of Andira
classification,
infrageneric
infrageneric classifications
were based on the nature of the indumentum of the gynoecium,
but glabrous ovaries have evolved
in parallel at
least three times. Detailed
informative characters and

analysis of fruit wall anatomy provides phylogenetically
biology. There have been at least five instances of independent evolution of large, puta
fruits and three of large, persistent stipules. The frequent changes in stipule morphology
tively rodent-dispersed
are perhaps due to mutation
in a single gene. Evidence
radi
points to a relatively recent (possibly Pleistocene)
insights
into dispersal
ation of species of Andira

are illustrated. Five new
macrocarpa,
A. praecox,
are fully described and their ranges are mapped,

and 15
are proposed: A. chigorodensis,
A. jaliscensis,
A.
subspecies
and A. inermis subsp. glabricalyx.
in eastern Brazil. All
species
and one new
species
A. taurotesticulata,
INTRODUCTION
is a genus of 29 woody species (including three subspecies of A. inermis) dis
tributed throughout tropical America; A. inermis also occurs in Africa. Andira is most
Benth. (Bentham 1860; Polhill 1981), a genus of 10-12
closely related toHymenolobium
species confined to South America. They are similar florally and vegetatively, and they
Andira
share a root nodule morphology

that is very rare in the Papilionoideae
(de Faria et al.
1987; Sutherland et al. 1994). The principal difference between the two genera is in fruit
has samaras and Andira drupes. The two genera also differ in their
type; Hymenolobium
mode of germination: Andira is cryptohypogeal, whereas Hymenolobium
is phanerohy
a
Andira
is
considered
with
group
pogeal. Thus,
monophyletic
synapomorphies of a dru
paceous fruit and cryptohypogeal germination (Pennington 1995, 1996).
The close relationship of Andira and Hymenolobium
is confirmed by recent chloro
trnL
intron
et
nucleotide sequence data (Pennington
al. 2001). The two genera form
plast
a well-supported monophyletic
group, but their phylogenetic position is unresolved, and
no clear sister group has been discerned; however, it is clear that they are
distantly related
to the other genera of tribe Dalbergieae
to which they are currently assigned. Dalbergieae
comprises 19 genera and is centered in tropical America; only Inocarpus J. R. Forst. &
G. Forst, is endemic to the Old World
(in tropical Asia; Polhill 1981). Other recent
l

SYSTEMATICBOTANYMONOGRAPHS
VOLUME 64
systematic studies (Doyle et al. 1997; Lavin et al. 2001) indicate that Dal
This confirms earlier suggestions, based upon consideration
bergieae is not monophyletic.
that Dalbergieae are not a natural group (de Lima
of fruit, seed, and seedling morphology,
molecular
1991).
This taxonomic revision
is based upon examination of ca. 3,000 herbarium specimens

in five Neotropical
countries from 1991 to 1997. Species de
inAndira has been regarded as problematic, and the only recent revision (Mat
and field studies conducted
limitation
tos 1979), covering the Brazilian species, is inadequate for species identification, particu
coastal rain forest and coastal restinga forest of
larly for taxa from the Atlantic
southeastern Brazil (Lewis 1987), where Andira is both diverse (eight species) and abun
dant. Three new species have been described recently from eastern Brazil (Pennington &
de Lima 1995) and the Venezuelan Guayana (Pennington et al. 1997). Four of the five new
species included in this monograph
area of diversification for Andira.
grow around the fringes of Amazonia,
which
is an
frameworks for Andira based upon both chloroplast DNA (cpDNA) re

Phylogenetic
striction site data and morphology
have been presented by Pennington (1995, 1996). The
are drawn from these studies, but
in
this monograph
phylogenetic
analyses presented
in coding of some characters are proposed based upon new observations of
modifications
pollination syndromes and fruit anatomy. Furthermore, new cpDNA restriction site data
have provided phylogenetic placements for five more species and helped resolve some of
the problems of intraspecific cpDNA polymorphism
discussed by Pennington
(1995,
are used to re-evaluate infra
The
refined
here
1996).
phylogenetic hypotheses presented
in Andira, to present the first biogeographic
generic classification
the evolution and diversification of the genus, and to investigate
phological features, such as stipules, flowers, and fruits.
hypotheses to explain
the evolution of mor
TAXONOMICHISTORY
to the attention of European science through the use of extracts
from the bark and seeds as anthelminthic drugs by indigenous peoples in the Neotropics.
as an anthelminthic in Per
Piso (1648) described the use of the seeds of A. fraxinifolia
Andira
first came
nambuco, Brazil, and named the species "Andira Ibaiariba." The first species descriptions
were included in accounts of the medicinal use of the bark and seed of A. inermis in Ja
maica (Wright 1777) and A. surinamensis in Suriname (Bondt 1788). Both authors placed
their species in Geoffroea Jacq., which shares Andira's unusual drupaceous fruit, but is
distinct inmany floral and vegetative characters.
the first valid publication of the name Andira has been problematic. It
Determining
had generally been assumed to be by Lamarck (1783), and as such was proposed (Harms
1904) and accepted (Briquet 1906) for conservation against Vouacapoua Aubl. Although
these two genera are now considered taxonomically quite distinct, Lamarck had included
Vouacapoua in his original concept of Andira, making the latter name illegitimate. Yet, as
part of the Rickett and Stafleu revision of the lists of Nomina Conservanda, Cowan (1959)
noted that Lamarck's
(1783) description of Andira racemosa provided no generic diagno

sis, and since Lamarck cited Vouacapoua americana Aubl. (1775) in synonymy his genus
and species could not be treated as new. Cowan decided that the long-standing provisions
of the International Code of Botanical Nomenclature
(ICBN) permitting a "descriptio
to validate simultaneously
the generic and specific names could not
generico-specifica"

ANDIRA
2003
apply. He therefore concluded that neither Andira nor the species name, A. racemosa,
were validly published by Lamarck, and proposed that the conservation of Andira be from
its next usage, that by A. L. de Jussieu (1789), with A. racemosa Lam. ex J. St.-Hil., ap
parently the first validation of that name, as type.
R. K. Bmmmitt
(pers. com.) noted that Lamarck did indeed validly publish Andira
but as an illegitimate replacement (a nomen novum) of Vouacapoua Aubl., because he
treated
Vouacapoua
as a synonym
of his
generic
name
Andira.
His
species
name
A.
race
is also illegitimate, as a nomen novum for V. americana. Under Art. 7.5 of the cur
rent ICBN (Greuter et al. 2000), the type of Andira Lam. is that of Vouacapoua (i.e., the
type of V. americana Aubl.), but because Vouacapoua is now recognized as a distinct cae
salpiniaceous genus, this typification would be extremely disruptive.
mosa
The current conservation is of "Andira Juss.," not "Andira Lam." Jussieu (1789) cited
no species in his account, and Cowan (1959) considered Andira Juss. to be typified by A.
racemosa Lam. ex J. St. Hil.; however, Jaume St. Hilaire (1804) was merely using
Lamarck's A. racemosa, so there is no case for treating his name as a later homonym
under Art. 48.1 of the ICBN. If "Andira Juss." were to be maintained as the conserved
name, the type requires correction to A. racemosa Lam., which is automatically typified
by V. americana. Therefore, themaintenance of Andira Juss. would defeat the purpose of
the conservation of the name Andira.
For these reasons, a formal proposal to change the authorship of Andira to Andira
Lam. nom. cons., and to provide it with a conserved type, A. inermis, was submitted and
accepted (Pennington 2002). The name Andira inermis (Wright) DC. has been widely ap
plied to the species that includes A. racemosa Lam., the type of the conserved Andira (i.e.,
excluding the Vouacapoua element). Thus, itwas appropriate to propose, under Art. 14.9
of the ICBN, that the type of A. inermis be the conserved type of Andira Lam. If the name
A. racemosa Lam. were conserved with a new type, itwould displace A. inermis, the name
traditionally used for the most
common, widespread,
and frequently
collected
species of
Andira.
Infrageneric
Bentham
classification
1839, 1860, 1862) features prominently in the taxonomic history of

studying the specimens of Martius, Pohl, and Schott from eastern Brazil, he
the
first major account of Andira (1837, reprinted in 1839). He placed the
produced
two
in
sections: sect. Lumbricidia, defined by a hairy ovary on a relatively short
species
(1837,
Andira. After
stipe, and sect. Euandira, defined by a glabrous ovary on a relatively long stipe. Walpers
(1843) followed Bentham. In his later works (1860, 1862), Bentham abandoned both the
indicate that the ratio of stipe length to ovary length
stipe character (my measurements
across all species of Andira varies very little) and the formal sections, but he did group
species based upon the presence of hairy or glabrous ovaries.
The next attempt at infrageneric classification was by Mattos
(1979). She recog
two sections, one with two subsections. Section Lumbricidia
is defined by the
presence of more than five leaflets (i.e., more than two pairs of leaflets). Subsection
Lumbricidia
includes species with a hairy ovary and subsect. Glabratae
those with a
nized
sections. Section Paucifoli

glabrous ovary; these subsections correspond to Bentham's
olatae is defined by presence of 1-3 leaflets, and therefore contains only A. unifoliolata
and A. trifoliolata.

VOLUME 64
MORPHOLOGY
Habit. Species of Andira grow in a wide variety of habitats and have varied life forms
(character 1 in cladistic analysis). Rain forest species, such as A. macrothyrsa, A. parvi
flora, A. cori?cea, and A. anthelmia, are trees that can reach 40 m. Several species (in
are small trees and shrubs abundant in
cluding A. nitida, A. carvalhoi, and A. fraxinifolia)
the sandy coastal restinga scrub and forest of eastern Brazil. Andira humilis is a geoxylic
suffrutex, defined by White (1976) as plant with massive, woody, underground axes but
only annual or short-lived shoots above ground. Its aerial shoots rarely exceed 50 cm. The
species appears to be well adapted to survive the regular fires in the seasonally dry cer
rado woodland of central Brazil by an ability to sprout from its underground rhizomes
after burning. This growth form is unique to this species of Andira, but several other
are able to
species (A. verm?fuga, A. carvalhoi, A. legalis, A. nitida, and-A. fraxinifolia)
to
A.
often
This
enable
carvalhoi
survive on
may
root-sprout (pers. obs.),
extensively.
steep and easily eroded white sand slopes in restinga scrub.
Life form provides a useful field character for distinguishing A. ormosioides from A.
fraxinifolia. The latter species has a broad, spreading crown and short bole when in open
situations, whereas A. ormosioides has a long bole and small crown. Polhill (1969) indi
cated that growth form may also be useful in distinguishing A. inermis subsp. rooseveltii.
Field notes (from herbarium specimen labels) indicate it to have a narrow, conical crown,
which contrasts with the broad, spreading crown of A. inermis subsp. inermis.
Bark and slash. All species of Andira produce a little red ex?date when the bark is
cut, a characteristic also of some other woody papilionoids (e.g., Dussia Krug & Urb. ex
Taub., Geoffroea, Pterocarpus
Jacq.). Bark color, bark texture, and slash color vary, al
too
few
been made to assess their taxonomic significance. Pale
observations
have
though
grey-brown bark color characterizes A. fraxinifolia, whereas other eastern Brazilian tree
species (A. ormosioides, A. legalis, A. anthelmia, and A. nitida) have darker bark (pers.
slash of A. parviflora and A. cori?cea contrasts with the more
obs.). The orange-brown
reddish brown observed in other Brazilian species, such as A. cordata and A. surinamen
sis. The thick, corky barks of A. cordata, A. cujabensis, and A. verm?fuga probably serve
as protection against fire in the cerrado woodlands of central Brazil.
Wood. The wood anatomy of eleven species of Andira (A. anthelmia, A. fraxinifolia,
A. legalis, A. humilis, A. nitida, A. carvalhoi, A. verm?fuga, A. inermis, A. surinamensis,
A. cori?cea, and A. cordata) was studied by Herridge (1992). All these species have very
similar wood anatomical features, and any interspecific differences noted are quantitative
and may be due to the nature of the sample (from a branch or the trunk), its age, or vari
ation in the growth rate of different species. These difficulties in interpretation mean that
these data are of little relevance to the species-level
taxonomy of Andira.
The wood is diffuse-porous and without growth rings. The storied vessels, which are
of up to four, are large (98-234 mm), widely
solitary or clustered in radial multiples
spaced, and often contain deposits. The intervessel pitting is vestured, small, and alternate.
The axial parenchyma is storied, aliform (winged to lozenge-shaped),
and in tangential
bands ranging from 3-20 cells wide. There are gelatinous prismatic crystals in chambered
axial parenchyma cells. The rays range from 1-3 to 1-6 cells wide and 12-19 cells high,
and are irregularly storied or non-storied. Multiseriate
rays aremostly heterocelluar (A. in
ermis is homocellular).
The fibers are non-septate
and usually very thick-walled.

ANDIRA
2003
Indumentum. When present, hairs are always simple and generally either flexuose and
erect or short and appressed. The indumentum of different structures (twigs, stipules,
rhachis, petiolules, leaflet undersurfaces, inflorescence, calyx, and gynoecium) varies in
dependently, and the variation for each part is discussed separately below. This variation
for species delimitation,
(characters 5, 8, 10).
is useful
and provides
three characters
for cladistic
analysis
Stipules. Four species (A. multistipula, A. grandistipula, A. anthelmia, and A. legalis)

have large (to 7.5 cm long, 6 cm wide in A. grandistipula), persistent stipules, which are
crowded at the shoot apices and around the base of the inflorescence (Fig. 1A; character
3 in cladistic analysis). These contrast with the more typical small stipules of all other
Andira species (Fig. IB). This difference in stipule size distinguishes A. multistipula from
the otherwise similar A. inermis, and A. anthelmia and A. legalis from A. ormosioides. The
function of the large stipules is presumably to protect the shoot apices. There is no obvi
ous correlation of the occurrence of large stipules with geography, habitat, or life form.
Leaves. The leaves of all species of Andira are imparipinnate. The leaflets are gener
ally opposite with their petiolules subtended by stipels. Leaf length varies from 2.5 to 60
cm and is an important character in species delimitation. The number of leaflets varies
to up to 19 (nine pairs) in A. chigorodensis
from one in A. unifoliolata
(character 4 in
cladistic analysis; Fig. 1C, D). Unifoliolate
leaves are unique to A. unifoliolata (Fig. 1C).
Andira
trifoliolata is the only species with uniformly trifoliolate leaves; A. tervequinata

has both trifoliolate and 5-foliolate leaves. In the other species the number may be con
stant (e.g., A. micrantha, 2 pairs; A. cori?cea, 3 pairs) or vary within certain limits (e.g.,
A. chigorodensis, 7-9 pairs).
Leaflet shape provides characters that are diagnostic of certain species. For example,
the obovate, blunt-ended leaflets of A. marauensis distinguish it from the otherwise simi
larA. nitida. The most critical character of the leaflets for species delimitation is the na
ture and density of the abaxial indumentum (character 5 in cladistic analysis). Some
species, such as A. inermis (Fig. 2A), have glabrous leaflets. In the other species, three in
dumentum types are present (Fig. 2B, C, D). Eleven species (including A. surinamensis,
A. macrothyrsa, A. praecox) have an indumentum of short, tightly appressed hairs (Fig.
2B), which contrasts with the varying densities of vestures composed of longer, erect hairs
these two
(Fig. 2C) of six species (such as A. legalis and A.fraxinifolia).
Distinguishing
indumentum types (short appressed versus longer erect hairs) is useful for species identi
fication, but variation between the two is continuous, and they are lumped into a single
character state in the cladistic analysis. Yet, two species, A. cujabensis andA.jaliscensis,
have leaflet undersurfaces with a distinct indumentum of dense, fine, pale, tangled, erect
hairs (Fig. 2D), which is the character that separates them from otherwise similar species
(A. cordata and A. inermis; Fig. 2A) that are glabrous. There is no association of indu
mentum type with habitat. For example, A. micrantha (glabrous), A. macrothyrsa
(short,
tightly appressed hairs), and A. parviflora
(long, erect hairs) are all rain forest species.
Similarly A. cordata (glabrous), A. humilis (short, tightly appressed hairs), A. verm?fuga
(long, erect hairs), and A. cujabensis (long, fine, pale, tangled hairs) are all species of the
seasonally dry cerrados of central Brazil.
The primary vein is prominently raised on the leaflet undersurface (Fig. 2C), and ei
ther channelled (Fig. 2E) or plane (Fig. 2F) on the leaflet uppersurface (character 6 in
cladistic
analysis). Venation
pattern varies continuously
from entirely eucamptodromous

FIG. 1. Stipules and leaves

= ca. 5 cm;
(scale bar
photo G. P.
scale = inches; right-hand
scale =
leaves of A. legalis. (Based on: A,
in Andira
(characters 3 and 4). A. Large, persistent
stipules of A. anthelmia
Lewis). B. Small stipules ("s" at arrows) of A. surinamensis
(ruler left-hand
leaves of A. unifoliolata.
D. Large, imparipinnate
cm). C. Small, unifoliolate
R. T. Pennington
183; B, R. T. Penningfon
463; C, W. Rodrigues
11182; D, R.
T Penningfon
305.)

VOLUME 64
ANDIRA
2003
y.y
FIG. 2. Indumentum
and venation of leaflets of Andira
abaxial leaflet
(characters 5 and 6). A. Glabrous
surface of A. inermis, showing plane secondary and tertiary veins. B. Abaxial
leaflet surface of A. surinamensis
with short, straight, tightly appressed hairs, prominently
raised secondary vein, and slightly raised tertiary veins.
leaflet surface of A. fraxinifolia
erect hairs, prominently
C. Abaxial
with long, flexuose,
raised primary and sec
ondary veins, and slightly raised tertiary veins. D. Pale, long, fine, tangled hairs of the abaxial leaflet surface of
A. cujabensis. E. Adaxial
leaflet surface of A. surinamensis with channelled primary vein. F. Adaxial
leaflet sur
face of A. unifoliolata with plane primary vein. P = primary vein; s = secondary vein; t = tertiary vein. Scale bars
= 1mm.
463; C, G. Hafschbach
13558; B, R. T. Penningfon
(Based on: A, T. D. Pennington
55043; D, C. A. Cid
Ferreira 6259; E, R. T. Penningfon
463; F, A. Ducke 35078; all from herbarium material except E, leaf from live
plant growing
at Royal
Botanic
Gardens,
Kew.)

VOLUME 64
to entirely brochidodromous
tomixed eucamptodromous
(terminology of Hickey, 1979).
The number of secondary veins, their angle of divergence from the primary vein and then
nature of curvature varies and can be useful for distinguishing
species. For example, the
contrast with those of A. inermis,
uniformly curving secondary veins of A. macrothyrsa
curve only as they approach the margin. The secondary and tertiary veins also
vary from plane to sunken on the leaflet uppersurface, and plane (Fig. 2A) to raised (Fig.
2B, C) on the undersurface, but this variation is continuous and not useful for cladistic
which
analysis.
Inflorescences are terminal and axillary panicles, which vary in size,

Inflorescence.
shape, compactness, number of branches, angle at which the branches are held, and the
and
density of the flowers. Intraspecific variation is often extreme (e.g., in A. fraxinifolia
A. inermis), and this limits the use of these inflorescence characters for taxonomic pur
poses. Terminal inflorescences may not be obvious on herbarium sheets, but have been ob
served in all species seen flowering in the field.
Bracts
and
bracteoles.
Bracts
and
bracteoles
ducous. There is little variation in theirmorphology

paired at the base of the calyx.
are
small,
narrow,
and
often
early
throughout Andira. The bracteoles
ca
are
length varies continuously from 5 to 23 mm (Fig. 3).Within species,

the range of variation is relatively narrow, approximately 2 mm for small-flowered species
and 6 mm for large-flowered species. Flower length is a useful character for separating
similar pairs of species, such as A. inermis and A. cubensis, A. fraxinifolia
and A. ormo
Flowers.
sioides,
Flower
and A. macrothyrsa
and A.
chigorodensis.
The angle and depth of the calyx lobes varies widely in several species and is thus of
limited utility in species delimitation; however, the calyx indumentum (character 8 in
cladistic analysis) provides more useful characters. For example, the densely hairy calyx
of A. cubensis distinguishes
this species from A. inermis. In A. inermis, A. multistipula,
and A. humilis, the calyx is either sparsely hairy (the hairs short and appressed) or
and this variation shows
of
scription
subspecific taxa inA.
The corolla o? Andira consists
but are firmly attached because of
geographical correlation, which is the basis for the de

inermis (see discussion under the individual species).
of five glabrous petals. The keel petals are not united
two interlocking folds. The wing petals are free from
the keel. Petal color (character 7 in cladistic analysis) is an important phylogenetic
char
acter. Species with flowers of less than 8 mm (with the exception of A. cubensis) have
white to yellowish petals with the standard marked with red or purple. The larger-flow
ered species have pink to purple petals with the standard bearing a central white mark.
glabrous,
Lamellate
wing petal sculpturing (Fig. 4A; character 9 in cladistic analysis) is present in

all species with flowers greater than 9 mm long, and also in A. cubensis where the flow
ers are 7 to 8.5 mm long. Lamellate wing petal sculpturing is absent from the other small
flowered species (Fig. 4B).
In all species of Andira, the stamens are diadelphous (9 + 1); the vexil
Androecium.
and A. macrothyrsa
lary stamen is free (Fig. 3). Very rarely, individuals of A. fraxinifolia
have one side of the vexillary filament united to the other nine. The filaments are unequal
in length; those closest to the standard are shorter and united for one half to two-thirds of
their length. The ratio of the length of the united and the length of the free filaments varies

ANDIRA
2003
FIG.
of Andira
3. Floral morphology
(characters 8, 9, and 10). For each species, from left to right: full
and gynoecium;
showing calyx indumentum and wing petal sculpturing; dissection
showing androecium
C. A. taurotestic
longitudinal section of gynoecium
showing ovule number. A. A. cordata. B. A. chigorodensis.
v = vexillary
ulata. D. A. nitida. E. A. jaliscensis.
F. A. carvalhoi. G. A. fraxinifolia.
H. A. ormosioides.
stamen,
= 5 mm; all drawn to same scale.
in
in
H.
all
but
marked
Scale
bar
present
species,
only
flower

10
VOLUME 64
B(i)
in Andira. Species with wing petal sculpturing: A, wing petal of A. ver

FIG. 4. Wing petal sculpturing
= lamellate
S
scale
bar = 1mm; A(i) wing petal of A. fraxinifolia
(16 mm long); A(ii) wing
m?fuga,
sculpturing,
(15 mm long); A(iii) wing petal of A. nitida (11 mm long); A(iv) wing petal of A. jalis
petal of A. carvalhoi
censis (12 mm long). Species without wing petal sculpturing: B, wing petal of A. trifoliolata,
scale bar = 1mm;
B(i) wing petal of A. cordata (5.5 mm long); B(ii) wing petal of A. chigorodensis
(5 mm long); B(iii) wing petal
of A. taurotesticulata
228; A(ii) M. P. M. de
(8 mm long). (Based on: A, J. Ratter 3595; A(i) R. T Pennington
Lima et al. 20; A(iii) A. J. Ribeiro 62; A(iv) E. J. Lott 2544; B, J. J. Wurdack & L. S. Adderley 43368; B(i) G.
.
Hatschbach
39477; B(ii) /. Brand & A. Cogollo
114; B(iii) G. Lozano & G. Galeano
3963.)
little across all Andira species, which is further proof of the highly conservative floral or
in Andira that appears to reflect a general bee pollination mechanism.
ganization
Pollen. The pollen morphology

of two species, A. inermis and A. macrothyrsa,
has
been found to be uniform by B. Klitgaard (unpubl. data). This is unsurprising, given the
morphological
uniformity of pollen in the many tropical woody papilionoids
(Ferguson &
Skvarla 1981; Ferguson et al. 1994). It therefore appears unlikely that pollen characters

11
ANDIRA
2003
will provide taxonomic characters within Andira. The pollen grains of the two species ex
amined have an even, minutely reticulate tectum, are 3-colporate, with the colpi extend
ing only partly to the poles, and have granular colpal membranes.
Gynoecium. The ovary of Andira is borne on a stipe and contains 1-8 ovules (Fig. 3).
The fruit of Andira is usually single-seeded
(see "Fruit" below), the result of ovule abor
tion. The style is curved and is included within the keel petals. The stigma is ciliate (sensu
Lavin and Delgado,
1990). The proportions of the stipe, style, and ovary are more or less
invariant. The extent of the indumentum of the gynoecium is, however, very variable (Fig.
3; character 10 in cladistic analysis) and is a critical character for species delimitation.
are single-seeded
2-3-seeded)
(occasionally
drupes with
or
and
hard
thin
exocarps. They are diverse
mesocarps,
endocarps, fibrously fleshy
Fruit.
woody
Fruit of Andira
in the color
of
exocarp
and mesocarp,
scent,
mesocarp
structure,
and
endocarp
structure.
There are two general size classes of fruit: 6 cm long or less, weighing ca. 10-20 g when
to 40-300 g when dry. These
dry, and much larger fruits up to 12 cm long and weighing
in dispersal biology
size differences
reflect probable differences
(see "Dispersal
Syndromes").
Bentham
in
(1860) and Lewis (1987) point to the probable utility of fruit morphology
species delimitation and the lack of adequate fruit collection. Field observations
and the inferences made from alcohol preserved fruit confirm this. I have made field ob
servations of fruit of seven species, which suggest that coloration of themesocarp and ex
ocarp are useful characters. For example, the green mesocarp of A. anthelmia and A. frax
Andira
inifolia contrasts with the pale green-white mesocarp of

distinctness of A. ormosioides. Andira anthelmia and A.
is green.
ocarps, whereas that of A. fraxinifolia
Detailed field notes of exocarp and mesocarp color
fruits in alcohol should be a priority for future collectors
A. ormosioides, and confirms the

ormosioides have dark brown ex
and structure, and preservation of

of Andira. Better field data might
some
in Andira (such as that
of
of
delimitation
the
clarify
outstanding problems
species
of A. nitida; see notes under that species), would benefit studies of dispersal biology in
Andira, and provide suitable material for studies of fruit anatomy.
A preliminary study of Andira fruit anatomy (Pennington 1994) revealed two distinct
types of endocarp in Andira; one composed of woody fibers, the other of stone cells. The
endocarp of stone cells appeared to be a possible synapomorphy for a well-supported
clade of Andira
species discovered by cpDNA data (Pennington 1995). Pennington and

Gemeinholzer
(2000) examined the pericarp anatomy of 25 species in search of further
taxonomic characters, and a summary of this work is provided here.
The mesocarp of all Andira species examined comprises parenchyma cells in which
stone cells are embedded, either individually or in patches. The mesocarp of seven species
is homogeneous, whereas in 20 species it is heterogeneous, having either patches of stone
cells close to the exocarp, fibers near the endocarp, and/or ergastic materials distributed
unevenly throughout the parenchyma. Despite this variation, itwas decided thatmesocarp
structures could not be used cladistically, because intraspecific variation was too great.
Starch grains are abundant, occurring either scattered throughout the tissue, as greyish cell
content or as "tyloses-like"
features (probably tightly packed starch grains in highly
packed parenchyma cells with squashed cell walls).
At maturity, the endocarp is divided into a thicker hard layer and a thin papery layer
within it, which is the remnant of the "seed cushion." This latter structure, described by

12
VOLUME 64
(1914), is characteristic of legume fruits and consists of thin-walled, juicy

parenchyma cells inwhich the developing seeds are embedded. At fruit maturity it shrinks
and dries out, forming flake-like shreds.
The hard, thick part of the endocarp is variable and provides phylogenetically
useful
information. Three main structures are present: (i) dominated by parenchyma and/or col
lapsed cell tissue; (ii) dominated by fibers; (iii) dominated by stone cells.
This variation can be coded as amultistate cladistic character (Pennington & Gemein
holzer 2000; character 10). This is a refinement of the coding of Pennington (1996), where
Fucsk?
the distinct
was
not
state of the endocarp dominated
by parenchyma
and/or collapsed
cell tissue
recognized.
formed by parenchyma and/or col

species have an endocarp predominantly
most
to
cell
often
tannin. Twelve species have an en
which
is
brown
due
tissue,
lapsed
dominated
fibers.
Small
cells
and
sometimes squashed cell tissue
by
parenchyma
docarp
occur between the fibers. Air gaps were often observed, probably due to ripening or de
in layers, but in A. ver
generation processes. The fibers can be arranged predominantly
Five
m?fuga and A. carvalhoi they form sub-rounded patches surrounded by parenchyma cells.
Andira galeottiana, A. legalis, and A. anthelmia are characterized by a mixture of these
arrangements. Nine species have an endocarp that predominantly comprises stone cells.
This endocarp type is the hardest and most dense, and it is often very difficult to cut with
a sliding microtome. The densely packed tissue, lacking intercellular space, consists ei
ther of stone cells alone (three species) or of stone cells interspersed with small patches
of parenchyma (six species). In a single species, A. jaliscensis,
the hard layer of the en
an
two
of
thickness:
inner
of
equal
layer
parenchyma and col
docarp comprises
layers
an
outer
stone
cell
of
is
with an unique
and
cells.
This
coded
tissue,
lapsed
layer
species
(autapomorphic) state for this character.
The thicker, fibrous endocarps, and the endocarps of stone cells may offer greater me
chanical protection against seed predators (Pennington & Gemeinholzer
2000). Janzen
in A. inermis, and I have seen fruit of
(1982) noted seed pr?dation by Cleogonus weevils
A. macrothyrsa on a herbarium specimen (A. Gentry 37168) that have been attacked by
weevils. Both of these species have weak endocarps composed
Andira humilis has an endocarp of woody fibers, but this layer
species examined, perhaps making it less of a barrier against
Handro (1969) reported seed pr?dation by Cleogonus weevils
of parenchymatous
tissue.
is thinner than in any other
burrowing seed predators;
in this species. Field obser

of preserved fruit revealed no seed pr?dation inA. fraxinifolia, A.
anthelmia, A. ormosioides, A. nitida, and A. carvalhoi, which have thicker, tougher,
woody fibrous endocarps and in A. cujabensis, which has an endocarp of stone cells.
vation and examination
These data suggest
that greater seed protection may be provided by these harder, thicker
endocarps.
Seeds of species of Andira can be considered "overgrown" (sensu Comer,

because
1976),
they have an undifferentiated testa and contain no endosperm. They fill the
entire cavity of the fruit, and are more or less globose tomore elongated, 2-8 cm long and
1.5-6 cm wide. The testa is thickly chartaceous, generally dark reddish brown, and often
remains attached to the walls of the seed cavity (overlying the remains of the seed cush
Seeds.
ion) when the seed is removed. The cotyledons are virtually united, and a thin line can be
seen in transverse section where they meet. The hypocotyl-radicle
axis is small and ap
parent as a fold at the apical end of the seed.

2003
ANDIRA
13
(de Lima 1991; pers. obs. for A. carvalhoi,

Seedlings. Germination is cryptohypogeal
A. cujabensis, A. fraxinifolia, A. nitida, A. surinamensis, A. cori?cea, A. anthelmia). Nu
merous spirally arranged scale-like prophylls are produced before the first true leaves,
which are trifoliolate or with two or more pairs of leaflets (even in the unifoliolate A. uni
was not observed in
pers. obs.). Leaflet nyctinasty (evening leaf movements)
foliolata,
seven
of
the
of
Andira
this
any seedling
grown during
species
study.
It appears possible that polyembryony?the
presence of more than one embryo in an
occur in Andira, because more than one shoot can be pro
(Richards 1997)?may
duced from a single seed o? A. fraxinifolia
(P.Griffiths, pers. comm.). This raises the pos
that
the
variation
patterns of this species might be ex
sibility
complex morphological
ovule
plained
in part by apomixis.
studies of this and other species are
Further developmental
necessary.
POLLINATIONBIOLOGY
Frankie et al. (1976) recorded numerous species of bees visiting the flowers of A. in
ermis in Costa Rica in large numbers. Other observations of floral visitors are anecdotal.
I have made similar observations of a variety of bee species visiting A. fraxinifolia
and A.
carvalhoi (R. T. Pennington, unpubl.) in Bahia, Brazil. These are two species with pink
purple flowers, less than 18mm long. Although the larger (19-24 mm long) flowers of A.
and are not significantly larger in terms
visited by large xylocopid bees, which
suggests a different pollination biology. This also appears to be reflected in the phenology
of A. anthelmia, where a high percentage of individuals were observed to be synchro
nously flowering in one area (vicinity of Ilh?us, Bahia, Brazil, October 1994; R. T. Pen
nington, unpubl.) in contrast to smaller-flowered
species, such as A. fraxinifolia, A. car
anthelmia
of flower
have superficially similar morphology

length, I have only seen these flowers
valhoi, and A. verm?fuga, where

T. Pennington,
unpubl.;
J. A.
few individuals
Ratter,
pers.
comm.).
in a population
Two
other
flower synchronously
species,
A.
legalis
and A.
(R.
or
mosioides, have flowers of a correspondingly

large size to those of A. anthelmia and may
share the same pollination biology, but direct pollination observations are lacking.
The completely different flower color of other species o? Andira (white to yellow, the
standard petal with red or purple markings) suggests
tion biology, although there are no field observations
that they, too, have distinct pollina

to confirm this. James Ratter (herb,
label) has observed large numbers of bees visiting the small, white-flowered A. cujaben
sis. This suggests that the pollinators of the small-flowered species may simply be smaller
bees. The absence of wing petal sculpturing suggests a difference in pollinator of the small
flowered species, because wing petal sculpture is thought to provide a foothold for floral
visitors (Stirton 1981). Further evidence that these small flowers may attract different pol
linators comes from the general correlation of the characters flower size, flower color, and
presence of wing petal sculpturing.
Bee pollination of all species o? Andira might provide an explanation for the covari
ance in size of the floral parts throughout the genus. Yet, the
probable pollination of A. an
thelmia by different vectors (xylocopid bees), demonstrates
that care must be taken in
generalization. For example, all the small flowers may not be as similar as they appear; A.
Ducke (herb, label) noted that the small flowers of A. macrothyrsa smell fetid, which con
trasts with the sweet scent of the flowers of A. cujabensis noted by J. Ratter.

14
FIG. 5. A. Small,
fruits of A.
legalis
fruits of A. fraxinifolia
(R. T. Pennington
bat-dispersed
(G. P. Lewis & H. C. de Lima 1196; photo G. P. Lewis.)
VOLUME 64
213). B. Large,
rodent-dispersed
These observations of pollination biology have influenced the coding of character 7,

which now has three states, rather than the two described by Pennington (1996).
DISPERSAL SYNDROMES
Janzen et al. (1976) observed dispersal of the fruits of A. inermis by bats of the fam
in Costa Rica. It seems reasonable to assume thatmost Andira species
ily Phyllostomidae
also have bat-dispersed fruits, because their fruits are morphologically
similar to those of
A. inermis (Pennington & de Lima 1995; Fig. 5A). The fruits of these bat-dispersed
species do not exceed 6 cm in length, weigh 10-20 g when dry, are green or occasionally
yellow (A. humilis) when ripe, have a strong, sweet scent and a fibrous mesocarp. The bat
dispersal of Andira fruits is well known by local people in South America; "Andira"
means bat in the Tupi Amerindian
language (Milliken et al. 1992).
of
Andira
possess much larger, heavier fruits, up to 12 cm long and
Eight species
to
500
g when fresh (Fig. 5B; dry fruit of A. macrocarpa weigh
weighing probably up
300 g). These fruit are too large and heavy to be carried by South American fruit bats,
which can carry a maximum of 100 g (Fleming 1986). It is probable that they are adapted
for dispersal by large rodents (van Roosmalen
1985; Pennington & de Lima 1995), such
as paca (Agouti paca), agouti (Dasyprocta spp.), and acouchy (Myoprocta spp.), which are
(Emmons 1990). Of these large
important dispersers of large fruit in the Neotropics
fruited species, I have only seen fresh fruit of A. carvalhoi, which are brown and odorless

ANDIRA
2003
15
when ripe, with a hard, non-fibrous, pale green to green-white mesocarp. When dry, the
mesocarp becomes hard and finely granular. Dried fruit of the other large fruited species
also have these characters, which suggests that they may be dispersed by similar means.
No direct observations of rodent dispersal have been made, but if these large fruits are ro
dent dispersed, then this represents a considerable novelty in legumes. The only reported
are of Parkia mul
cases of rodent dispersal in the estimated 650 genera of Leguminosae
tijuga Benth. (Mimosoideae; Hopkins & Hopkins 1983) and Hymenaea courbaril L. (Cae
1986). Some species appear to be secondarily dispersed by
salpinioideae; Hallwachs
water. The Mexican endemic A. galeottiana has a large fruit of which the dry mesocarp is
soft and spongy with air cavities, apparently adapting this species to dispersal by water;
its fruits are regularly found on Mexican
coastal beaches (Rovirosa 1890; Gunn & Den
nis 1976). Andira anthelmia and A. surinamensis are small-fruited species that show river
ine distributions and may also be secondarily dispersed by water (Pennington & Gemein
holzer 2000). It also seems probable that small-fruited
species may be secondarily
dispersed by rodents, because their fruits fall to the ground (pers. obs.).
Pennington and de Lima (1995) noted thatmode of dispersal appeared to have an ef
fect on the distributions of Andira species, with the putatively bat-dispersed species gen
erally more widespread. The best example of a wide distribution of a bat-dispersed species
isA. inermis (see Figs. 14,15). Putatively rodent-dispersed species are often restricted en
is confined to the vicinity of Manaus in central Amazonia, A.
demics, e.g., A. micrantha
to
is
the Pakaraima Mountains
in Guyana, A. carvalhoi is endemic
endemic
grandistipula
to southern Bahia in Brazil. These distributional differences may reflect the restricted
home ranges of seed-dispersing
rodents (Emmons 1990; Hallwachs
1986) and the long
distances flown by Neotropical fruit bats from their sleeping roosts to feeding areas (up to
10 km; Williams & Williams
1967).
Standard analyses of the composition of the dried mesocarps of five Andira species
were made using the methods of AOAC (1990). These confirm that starch, sugars, pro
tein, and fat are important nutritional components of these fruits (Table 1). It was hoped
that the probable differences in dispersal of large and small Andira fruits might be re
flected in the composition of their mesocarps. For example, Pannell and Koziol (1987)
found wide differences in the nutritional composition of fruits of Aglaia (Meliaceae) dis
the data in Table 1 are somewhat inconclu
persed by birds and primates. Unfortunately,
sive, and it should be noted that A. carvalhoi was the only large-fruited species for which
material was available. First, the difference in fat content between the two accessions of
A. fraxinifolia
suggests that small interspecific differences in values must be treated with
Table
1.Mesocarp
of five species of Andira. All figures are % by weight of the air-dried
composition
the maturity of the fruit was proven by the subsequent germination
ex
of seed from all accessions,
mesocarp;
All vouchers are collections
of R. T. Pennington
and deposited at CEPEC, FHO, and K.
cept A. ormosioides.
Fruit
Speciespresumed
A. surinamensis
A. fraxinifolia
A. fraxinifolia
RTP
364
RTP 202
RTP
213
size and
disperser
Starch
Sugars
6.6
3.4 32.6
7.3
7.23.3
9.5
5.3
11.6 2.8
5.3
2.0
6.8
A. anthelmia
RTP 227
A. carvalhoi
RTP 232
large, rodent
RTP 306
Fat
small, bat
small, bat
small, bat
small, bat
small, bat
A. ormosioides
Protein
8.9
8.6
1.62.2 5.6
2.3
26.2
13.7
0.6

28.7 8.8
16
VOLUME 64
caution. Second, although A. carvalhoi has a particularly high starch content, which re
the sim
flects the large numbers of starch grains in the parenchyma cells of itsmesocarp,
ilar value for A. anthelmia indicates that high starch content is not a unique feature of large
fruit. Overall, there is little difference in composition between the two fruit types, and the
inclusion of more rodent-dispersed species and more accessions of individual species are
necessary to further investigate the small differences found.
A notable aspect of these data is the high sugar content of the mesocarp of A. suri
namensis. This sample was analyzed four weeks after collection in Guyana, whereas the
other accessions were a year old; however, the sugars should remain more or less stable
unless dried fruit suffer microbial contamination, of which there was no evidence. It ap
pears that these fruit are much sweeter than the others. It is possible that these sweet fruits
are attractive to different bat species from those that disperse other species of Andira. The
fruit of A. humilis has a completely different scent from the fruit of the five other bat-dis
persed species that I have collected, which may also be attractive to alternative bats.
Andira fruit generally contain a single seed, so the large, putatively rodent-dispersed
fruit contain a correspondingly
large seed. It is possible that large seeds allow seedlings
to tolerate shade for long periods in the rain forest understory (Leishman & Westoby
1994), which may explain why several rain forest species o? Andira (e.g., A. micrantha,
A. cori?cea, A. taurotesticulata) have large fruit. In restinga vegetation, the large seed of
A. carvalhoi may permit the immediate development of an extensive root system, which
helps the seedlings to survive in the easily eroded, rapidly drained, white sandy soils. Seed
of this species, grown at the Royal Botanic Gardens, Kew, produced extensive roots, and
little above-ground growth, in contrast to themuch stronger shoot growth of the other four
bat-dispersed
species germinated.
CHROMOSOMENUMBERS
of chromosome numbers in Andira is restricted to those reported by
Knowledge
Goldblatt (1981). Both n = 10 and n = 11 have been reported in A. inermis. One of these
counts (Fritsch 1970; n = 11) is from a tree from Cuba, where A. inermis is absent, and is
therefore more likely to be of A. cubensis, the only Andira species in Cuba. Two other
= 10.
species (not named by Goldblatt) have n
Further chromosome counts were attempted for A. anthelmia {R. T. Pennington 282)
and A. fraxinifolia
{R. T. Pennington 236), two species for which living plants were avail
able at the Royal Botanic Gardens, Kew and Edinburgh. Root tips were collected and
in distilled water before pre-treatment in a saturated solution of paradichloroben
zene for four hours to overnight at room temperature. They were
then fixed in
ethanol:acetic acid (3:1 vol:vol). The root tips were hydrolyzed for 40 minutes in 5N hy
washed
acid at room temperature before staining with Feulgen (prepared after Fox,
two
for
hours. They were then mounted
in 45% acetic acid and observed under
1969)
drochloric
phase
contrast.
Perhaps because the root tips were not collected from entirely healthy, vigorously
growing plants, the preparations showed few cells inmetaphase. Thus, the counts made
should be regarded as preliminary. Counts for A. fraxinifolia
show 2n = 22, and for A. an
=
21 or 22. These results, together with those reviewed by Goldblatt (1981),
thelmia, 2n
indicate that there may be two basic chromosome numbers in Andira: n = 10 and n = 11.

ANDIRA
2003
17
AND
INTRASPECIFTC
cpDNA POLYMORPHISM
POTENTIALHYBRIDIZATIONINANDIRA
Introduction
Intraspecific cpDNA variation is often low due to slow mutation rates, small effective
population size, and the possibility of "selective sweeps" caused by the non-recombining
nature of this genome (Ennos et al. 1999); however, Pennington (1995) reported intraspe
in A. inermis, A. humilis, A. carvalhoi, and A. verm?fuga.
cific cpDNA polymorphism
inA. verm?fuga was dismissed, be
Subsequently, the intraspecific cpDNA polymorphism
cause the accession R. T. Pennington 250 does not belong to this species, but is an indi
or perhaps a new species (Pennington 1996; see notes under A.
vidual of A. fraxinifolia
fraxinifolia). The intraspecific variation inA. inermis involves few restriction site changes
and is likely to be due to processes of mutation within this species (Pennington 1995). In
contrast, the haplotypes inA. humilis and A. carvalhoi are phylogenetically
disparate. This
can
be explained by two processes: introgres
type of intraspecific cpDNA polymorphism
of an ancestral polymorphism. Unfortunately,
sive hybridization and/or the maintenance
the most basic information necessary to confirm introgression as a possibility inAndira?
whether different species are interfertile?is missing. Yet, hybridization and introgression
is perhaps a more
likely explanation than the maintenance of ancestral polymorphism be

cause of the large number of restriction sites (7-10) that differentiate the haplotypes in A.
carvalhoi and A. humilis, and the lack of intermediate haplotypes in these or other extant
then this polymorphism
has
species. If ancestral polymorphism were the explanation,
been successfully maintained over remarkably long periods, and perhaps we might
of the intermediate haplotypes.
expect to discover maintenance
also
Whatever
the explanation for this infraspecific cpDNA polymorphism
involving phy
logenetically disparate haplotypes, it is vital to assess whether it is widespread, because
processes of hybridization with subsequent introgression and lineage sorting (where an
ancestral species is polymorphic for cpDNA, and these cpDNA types sort in subsequent
species) can result inmisleading
interpretations of phylogenetic relationships based upon
cpDNA data (Doyle 1992). Fieldwork in Central Brazil, Costa Rica, and Ecuador pro
vided a valuable opportunity to extend this survey to other species from different geo
graphical areas. In particular, cpDNA variation was examined inmore populations of A.
humilis in an attempt to discover how widespread
is
intraspecific cpDNA polymorphism
in this species.
Methods
All methods
followed
(1995). Fourteen restriction site mutations

Pennington
(Ap
for
different
clades of species (Figs. 7, 9) were selected for study, a
pendix 1) diagnostic
and Brunsfeld
technique recommended by Doyle et al. (1990) and Rieseberg
(1992).
DNAs of accessions included in this previous study (A. humilis RTP 239, 269; A. ver
m?fuga RTP 265; A. fraxinifolia RTP 213; A. inermis C. E. Hughes 1673) were included
to facilitate identification of banding patterns. Forty-one accessions of eight species were
screened (Appendix 2). These included 14 accessions from three populations of A. hu
milis, 11 accessions from three populations of A. verm?fuga, and eight accessions from
two populations of A. cujabensis.

18
VOLUME 64
Results
The results are summarized in Appendix 2, which also includes data reported by Pen
nington (1995) in order to provide an overall summary of intraspecific cpDNA polymor
involving
phism in Andira. These data confirm that intraspecific cpDNA polymorphism
phylogenetically
disparate haplotypes is confined to A. humilis and A. carvalhoi. Wide
geographic sampling of several species (e.g., eight populations o? A. fraxinifolia, five pop
ulations of A. verm?fuga) has revealed no further instances of such polymorphism. This
corroborates the conclusion (Pennington 1995, 1996) that intraspecific cpDNA polymor
phism in Andira involving phylogenetically
disparate haplotypes is restricted enough to
treat the phylogenetic hypothesis derived from cpDNA restriction site data as an approx
imation of the true species phylogeny.
in seasonally dry cerrado vegetation in central Brazil,
Andira humilis is widespread
where it is sympatric with A. verm?fuga (Piastome Group I), A. cordata, and A. cujaben
sis (both Clade II). Every accession sampled from the center of its distribution (Fig. 6)
possessed the plastome type of Plastome Group I, indicating that the true phylogenetic po
sition of A. humilis is in Plastome Group I; however, the accessions of A. humilis RTP 239,
246, and 247 have the plastome type of Clade III. These were sampled from populations
in the state of Bahia at the edge of the species' range, where the cerrado vegetation meets
vegetation of Atlantic coastal Brazil (Fig. 6); there are seven species of
Andira, all with Clade III plastomes. It is only here, where these vegetation types meet,
that A. humilis comes into close contact with several species of Clade III (e.g., A. fraxini
the more mesic
folia, A. nitida, A. anthelmia, A. legalis), which are common elements of vegetation in

coastal southeastern Brazil. It is thus tempting to hypothesize
that hybridization and sub
area.
in
This
have
occurred
this
sequent introgression
hypothesis is supported by the ob
servation that the individuals RTP 246 and 247 were
small trees, rather than geoxylic suf

of
A.
the
form
humilis.
frutices,
typical growth
Although A. surinamensis, a species of
that border the northern
Clade III, is distributed along the southern fringes of Amazonia
cerrado (see Fig. 25), it does not occur at high density, and the records of A. humilis from
the northern area of the cerrado are few (see Fig. 23). Introgression here seems less likely
because of the low probability of contact between species, and itwould be interesting to
screen populations of A. humilis from this area to test this.
The case of A. carvalhoi ismore complex. Pennington (1995) screened two accessions
informative restriction sites). One had the plastome of
fully (i.e., for 37 phylogenetically
Plastome Group I {RTP 229), and the other that of Clade III {RTP 233; see Fig. 6). Another
accession screened for only two diagnostic mutations had one characteristic of Clade III
and the other for Plastome Group I, indicating a third plastome type (Pennington 1995).
Only one further accession of A. carvalhoi was available for the study reported here. This
comes from the same locality as RTP 229, is identical to it in all mutations screened, and
thus unequivocally has a plastome type of Plastome Group I. Therefore, the restricted data
indicate A. carvalhoi has three different plastome types.
Andira carvalhoi has a restricted coastal range in Bahia (Fig. 6) and is entirely sur
rounded by species with the Clade III plastome type. If the true phylogenetic affinity of A.
carvalhoi is with species of Plastome Group I and its "true" cpDNA type is that of Plas
tome Group I, it is possible to explain the presence of the Clade III plastome type in ac
cession RTP 233 by introgressive hybridization with a species of Clade III. If A. carval
available
hoi's true phylogenetic position is in Clade III and its "true" cpDNA type is that of Clade
in A. carvalhoi by recent
III, it is difficult to explain the plastome type polymorphism

2003
andira
19
PLASTOMES:
CLADEI
CLADE II
CLADE III(one species)
PLASTOME
GROUP I (one species)
-10?
PLASTOMESr
CLADE II
PLASTOME
GROUP I
H 20?
PLASTOME:
CLADE IIIONLY
50?
Amazonian
Rain Forest
Populations of A. humi/is with

plastome type of Clade III
Seasonally
Dry Vegetation
Range
FIG.
6. Location
polymorphism.
of populations
The plastome
of Andira
humilis
types present
carvalhoi
A Populations of A. humilis with

plastome type of Plastome Group
Atlantic Coastal
Rain Forest
cpDNA
of A.
and A. carvalhoi
in each of the different
sampled
vegetation
in a study of intraspecific
types are indicated.
hybridization, because its coastal range is ca. 200 km from the closest localities of A. hu
milis and A. verm?fuga, species with the cpDNA type of Plastome Group I. Itmight have
been in closer proximity and hybridized with these cerrado species in times of drier cli
mates during the Pleistocene,
when species of both cerrado and restinga, which are
to
water
have
been more widespread. That the cpDNA type of acces
stress, may
adapted
sion A. carvalhoi RTP 229 differs from those of A. humilis or A. verm?fuga by three re
striction sites also indicates that recent introgression with these two species is unlikely to

VOLUME 64
20
have been the source of its plastome. To unravel the reasons for the complex intraspecific
inA. carvalhoi clearly requires sampling of more populations and
cpDNA polymorphism
full characterization of plastome types.
It is tempting to ascribe some of the difficulties in species delimitation inAndira, such
some specimens as A. fraxinifolia
as the difficulties in distinguishing
and A. ormosioides,
of hybrids. Ifmorphological
intermediacy may be taken as evidence for
see
are
Bennett 1994, McDade
1990), then hy
frequent exceptions; e.g.,
hybridity (there
evidence
bridization may be possible, but rare. I reported having seen no morphological
of hybridity amongst sympatric species of Andira in southeastern Brazilian restinga
forests (Pennington 1995). Subsequently, I have observed a single plant that appeared to
to the occurrence
between A. anthelmia
be intermediate in leaf and stipule morphology
in the Poco das Antas Reserve in the State of Rio de Janeiro.
and A. fraxinifolia
CLADISTICANALYSES OF SPECIESRELATIONSHIPS
Introduction
Cladistic analysis of chloroplast DNA restriction site characters for species o? Andira
are presented by Pennington
and two outgroup species of Hymenolobium
(1995). Pen
site
and
that
both
restriction
directly combining
cpDNA
morpho
nington (1996) argued
logical characters in a single cladistic matrix and analyzing both simultaneously provided
the best phylogenetic hypothesis for Andira. Cladistic analysis of cpDNA restriction site
characters alone failed to resolve relationships between closely related species. Cladistic
characters produced a highly unresolved result because of the
analysis of morphological
lack of characters with discrete states that are suitable for cladistic analysis in Andira. In
contrast, combining these data sets directly produced good phylogenetic
resolution, be
cause the different character sets appear to be providing resolution at different hierarchi
cal levels: the cpDNA characters appear to be evolving slowly and delimit groups of
characters provide resolution within these groups. The
species, whilst the morphological
same view is taken here: that a combined cladistic analysis of all available data represents
the best phylogenetic
sions of infrageneric
hypothesis for Andira, the hypothesis that will be used for discus
classification, character evolution, and biogeography.
Species
Concepts
were delimited largely using the framework of the phylo

1990; Luckow 1995) and die characters
genetic species concept (e.g., Nixon & Wheeler
and character states listed below. They are diagnosable entities, characterized by constant
or consistent differences.
Species
in this monograph
Methods
Three cladistic analyses are presented that are based upon Pennington's studies (1995,
These are: 1)morphological
1996), with small modifications.
analysis; 2) combined mol
ecular and morphological
and
combined
molecular
and morphological
3)
analysis;
analy
a
of
subset
restriction
site data for species not included by Pen
sis, including
cpDNA
nington (1995, 1996). It should be noted that "Andira sp. nov. 1" and "Andira sp. nov. 2"
(Pennington 1995, 1996) have subsequently been named A. carvalhoi and A. cordata, re
spectively (Pennington & de Lima 1995). "Andira sp. nov. 3" (Pennington 1995, 1996) is

2003
21
ANDIRA
Table
2. Data matrix
of morphological
characters
of Andira.
Character
flavum
nitidum
Hymenolobium
Hymenolobium
A. inermis subsp.
inermis
0
0
0
subsp. rooseveltii
subsp. glabricalyx
0
A. jaliscensis
A. multistipula 0
A.
inermis
A.
inermis
A. cubensis
A. galeottiana
A. macrothyrsa
0
0
A. verm?fuga 1
A. humilis 2
A. anthelmia
A. carvalhoi
A. fraxinifolia
A. legalis 0
A. marauensis
A.
/u'f?fti 1
A. ormosioides
A. surinamensis
A. cordata
A. cori?cea
0
A. cujabensis
A. grandistipula
A. micrantha
A. parviflora
0
A. praecox
0
A. trifoliolata
A. unifoliolata
0
0
A. chigorodensis
A. taurotesticulata
A. tervequinata
A. macrocarpa
0
0
0
0
here included inA. nitida, which is now polymorphic for several character states (see dis
cussion under that species), and "Andira sp. nov. 4" (Pennington 1996) is included in
A. fraxinifolia
(see notes under that species).
Data matrices were handled using Dada (Nixon 1993). Parsimony analyses were car
ried out using Hennig86
(Farris 1988), using the mh*bb* command, and PAUP* 4.0, beta
2 (Swofford 1999). Character optimization and tree printing was achieved using Clados
1.2 (Nixon
1992).
Morphological
analysis. Characters and character states. The data matrix is shown in
Table 2. The criteria for selecting morphological
characters used in the cladistic analysis
follow those of Lavin (1993), and include the assumption that they are functionally and
developmentally
independent of one another, and that they are intrinsic attributes of the
taxa. The characters should also show uniform and consistent occurrence or absence
related.
among the terminal taxa, which implies that they are not environmentally

22
VOLUME 64
The view taken here is that characters with states that show overlapping variation are
of doubtful cladistic significance or utility, because there is no objective means of delim
1987; Farris 1990; Stevens 1991; Gift &
iting states within them (Pimentel & Riggins
Stevens 1997; for alternative viewpoints see: Archie 1985; Chappill 1989; Thiele 1993).
For this reason, characters such as leaf and leaflet size, which are useful for species de
limitation, were excluded from the phylogenetic analyses.
characters are included. These were
treated as unordered, thus min

imizing the assumptions built into the analysis.
In the list of characters and character states, notes have only been added where
characters were not presented or where their coding differs from that described by Pen
nington (1996). These differences are: (i) anatomical studies of fruit have refined the
coding of the endocarp character; (ii) observation of the pollination of A. anthelmia has
suggested a re-coding of the petal color character based upon presumed pollinator; and
Several multistate
(iii) including all species (rather than just those for which cpDNA data are available) al
lows a character state to be added to the leaflet indumentum character (5) and two new
characters to be included: leaflet number (4) and nature of the primary vein (6). The fol
lowing conventions are adopted: 1) The first number of each character refers to its po
sition in the data matrix for the morphological
cladistic analysis; the second number (in
to
refers
its
in
the
data
matrix
of the combined analysis. 2) The
parentheses)
position
states of each character are described and assigned a code (the number in parentheses
after
each
Vegetative
character
state).
characters.
1 (39). Tree (0); able to root-sprout and form amultistemmed

shrub (1); geoxylic suf
frutex (2).
2 (40). Germination phaneroepigeal
(0); germination cryptohypogeal
(1).
3 (41). Stipules small and caducous (0); stipules large and persistent (1).
4. Leaves with two or more pairs of leaflets (0); leaves trifoliolate only (1); leaves
unifoliolate only (2).?This
character was used by Mattos (1979) to define her sections
Lumbricidia
(defined by possession of five or more leaflets) and Paucifoliolata
(defined
by possession of trifoliolate or unifoliolate leaves). The character was run ordered and un
ordered in separate analyses. The justification for treating it as ordered is ontogeny;
seedlings of A. unifoliolata have multifoliolate
eophylls (R. T. Pennington, unpubl.). Ap
of Nelson's
(1988) modification
plying Weston's
(1978) ontogenetic rule indicates that
unifoliolate
leaves are less general (i.e., derived) relative to multifoliolate
leaves; how
ever, until the ontogeny of A. trifoliolata is studied, the exact order of the character states
cannot be entirely certain.
5 (42). Leaflet undersurface glabrous (0); leaflet undersurface with indumentum of short
(<0.2 mm), straight, tightly appressed hairs or indumentum of long (>0.2-1.0 mm), flexuose
? appressed to ? erect hairs (1); leaflet undersurface of
pale, tangled, erect hairs (2). [Fig.
erect
hairs
2A, B, C, D]?The
fine, pale, tangled,
(state 2) of A. cujabensis and A. jaliscen
sis are most distinct from the indumentum of all other species of Andira
(Fig. 2D).
6. Primary vein channelled above (0); primary vein plane above (1). [Fig. 2E, F]?
State 1 is found only in A. unifoliolata and A. trifoliolata.
Floral
characters.
7 (43). Flowers yellow

flowers
(0);
pink to purple,
to white,
the standard generally with red or purple markings

the standard generally with a pale central marking, less than

ANDIRA
2003
23
18mm
long, pollinated by numerous species of small bees (1); flowers pink to purple, the
standard with a pale central marking, 18-23 mm long, pollinated by large xylocopid bees
(2).?The
coding of this character reflects inferences of pollination biology described
length inAndira varies continuously (Fig. 3), and alone it does not provide
a good basis for the delimitation of character states.
8 (44). Calyx with indumentum (0); calyx glabrous (1).
9 (45). Wing petal sculpturing present (0); wing petal sculpturing absent (1).
10 (46). Gynoecium with indumentum (0); gynoecium totally glabrous or the ovary
above. Flower
with very few (1-3)
scattered hairs (1).
Fruit characters
11 (47). Fruit a samara (0); fruit a small, bat-dispersed drupe that dries smooth (1);
fruit a small, bat-dispersed drupe that dries wrinkled (2); fruit a large, putatively rodent
dispersed drupe, with non-fibrous mesocarp
(large drupes with mesocarp drying finely
granular) (3); fruit a large, putatively rodent-dispersed
taining air spaces (A. galeottiana only) (4).
con
drupe, with fibrous mesocarp
12 (48). Endocarp absent (0); endocarp dominated by parenchyma and/or collapsed

cell tissue (1); endocarp dominated by fibers (2); endocarp dominated by stone cells (3);
character presents
endocarp of fiber and stone cells (A. jaliscensis
only) (4).?This
an
of
because
the
lack
outgroup species
Hymenolobium
problems,
endocarp. Hawkins
et al. (1997) argued that the most theoretically consistent way of coding such characters
is by re-conceptualizing
them as two characters (the first coding the presence and ab
sence of the structure, the second coding the different types of structure, with the ter
minal taxa that lack the structure scored as missing data). This is not a perfect solution,
because it introduces problems of optimization?for
the second character, species of
for states of an endocarp, a structure that they do not
is coded here as a single multistate character with ab
coding as two characters, as recommended by Hawkins et
become optimized
Hymenolobium
have. For this reason the character
sence as one state. Moreover,
al. (1997), does not change
lack the structure.
the outcome
of any analyses,
because
only the outgroup
taxa
and morphological
analyses. Species for which cpDNA data
are lacking (e.g., A. cubensis, A. chigorodensis) were not included in these analyses. The
first analysis is identical to that presented by Pennington (1996) but includes the re-cod
ing of the fruit characters and flower color character described above. The accession A.
Combined
molecular
humilis RTP 239 has been identified as possessing a "foreign" plastome, probably derived
from introgressive hybridization (see above), and is thus excluded from this and the fol
lowing cladistic analysis. The data matrix is presented in Table 3.
The second analysis includes a subset of cpDNA restriction site data for species not
included by Pennington (1995, 1996). These are accessions o? A. jaliscensis, A. multistip
ula,
A.
taurotesticulata,
A.
cujabensis,
and A. marauensis
that were
screened
for
the di
(Appendix 1) as part of the study of intraspecific

and
cpDNA polymorphism
potential hybridization. Where cpDNA data were not avail
were
states
character
scored
with question marks. The data matrix is presented in
able,
Table 3.
agnostic mutations
described
above

24
VOLUME 64
data for Andira. Characters

1-38 are re
3. Matrix of cpDNA restriction site and morphological
a
for
accessions
included
in
the
molecular
sites; characters 39-48 are morphological.
species
Multiple
are abbreviated: AMC = A. M. de Carvalho; CEH = C.
study are indicated by collector and number. Collectors
de Lima; MC = M. Cheek; MS = M. Sugiyama; RTP = R. T. Pennington; TDP
E. Hughes; HCL = H. Cavalcante
= T. D.
Pennington.
Table
striction
Character
Hymenolobium
flavum
nitidum
Hymenolobium
A. inermis CEH 1673
A.
inermis TDP
A.
inermis MC
A. macrothyrsa
A. galeottiana
13558
3579
0
0
A. humilisRTP 269
A. fraxinifolia
RTP 250
A. verm?fuga 0
A. anthelmia RTP 227,282
A. carvalhoi
RTP 229
A. carvalhoi
RTP 233
A. fraxinifolia
A. fraxinifolia
A.
A.
RTP 213
MS
889
legalis RTP 307

legalis HCL s.n.
A. nitida RTP
300
A. nitida RTP 292

A. nifwfa RTP
301
A. ormosioides
A. surinamensis
A. cordata
A. grandistipula
A. parviflora
A. unifoliolata
inermis KY?
A.
512
A.
inermis RTP 580
A.
inermis RTP 589
A. multistipula
A. jaliscensis
A. taurotesticulata
A. marauensis
A. cujabensis
A. carvalhoi AMC
A
carvalhoi
RTP 217
12
3 4 5 6 7
1
1
1 1
1 1
?
1
0
0
0 0
0 0
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
?
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
1
0
0
0
0
0
0
0
0
1
0
1
0
1
1
1
1
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
1
0
0
1
1
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
?
1
1
?
1
0
0
0
?
1
1
1
1
1
1
1
1
0
1
0
1
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
?
1
1
1
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
1
1 1
? 1
0 ?
1
90123456789012345
1
1
1
1
1
1
1
1
0
1
0
1
0
0
0
0
0
0
0
0
0
0
1
1
1
1
1
?
1
0
1
?
1
1
1
1
1
1
1
0
0
0
?
?
1
1
1
1
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
0
0
0
0
1
1
1
1
?
0
0
1
?
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
?
0
0
0
0
0
1
1
1
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
?
?
?
1
1
1
0
0
0
1
0
1
0
1
1
1
1
1
1
1
1
1
1
1
0
1
0
0
0
0
? 1 0
0 1 1
0 1 0
0 1 1
0 1 0
0 1 0
0 1 0
0 1 0
0 1 0
0 1 0
0 1 0
0 1 0
0 1 0
0 1 0
0 0 1
0 1
0 1
0 1
? ?
1 0
1 0
? ?
9 9
1 0
? 0
? ?
9 9
? 0
1 0 1 ? 9 9?
1 ? 9 9 9 9
0 1 0 ? ? ?
1 ? ? ? ? ?
? 0 9 9 ? 0
?
0
0
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
?
1
1
0
1
1
1
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
?
0
0
0
?
0
1
1
1
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
?
1
1
1
1
1
?
?
?
?
?
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
?
0
?
0
1
1
1
1
0
0
0
?
?
0
0
1
?
0 1
0 1
1 1
1 1
1 1
0?
0 1
0 1
0 1
0 1
0 1
0 1
0 1
0 1
0 1
0 1
0 1
0 1
? 1
0 1
0 1
0 1
0 0
0 0
0 0
0 0
1 ?
1 ?
1 ?
1 ?
1 ?
1 ?
0?
0 ?
0 ?
?
9
?
9
?
?
9 ?
? ?
? ?
? 9
1
1
1
0
0
1
1
1
1
1
1
1
1
1
1
1
1
1
1
1
?
1
1
1
0
0
1
1
1
0
0
0
0
0
0
0
0
0
0
?
0
0
0
0
?
0
1
1
? ?
?
? ?
9
? ?
9
? ?
? ?
9
Results
Morphological
analysis. Pennington (1996) showed that a cladistic analysis of mor
characters
of a subset o? Andira species provided little phylogenetic resolution.
phological
A cladistic analysis of all species, using the twelve morphological
characters listed above,
also produced little phylogenetic
resolution. This lack of resolution reflects the low num
ber of characters and the high levels of homoplasy. With all characters run unordered, the
trees (the limit that the computer
analysis resulted in 2966 equally most-parsimonious
could store; CI = 0.57, RI = 0.79, length = 38). The strict consensus tree is entirely unre

ANDIRA
2003
Table
25
3 continued.
Character_67890123456789012345678
Hymenolobiumflavum
Hymenolobiumnitidum
A. inermisCEH 1673
A. inermisTDP 13558
A. inermisMC 3579
A. macrothyrsa
A. galeottiana
A. ?urni/frRTP 269
A. fraxinifolia RTP 250
A. verm?fuga
A. anthelmia
A. carvalhoi
RTP 227,282
RTP 229
A. carvalhoiRTP 233
A.yhmm/o/??RTP213
A.yhmm/0/uzMS889
A. legalisRTP 307
A.
legalis UCLs.n.
A. nitidaRTP 300
A. mttfo RTP 292
A. n?tidaRTP 301
0 1 ? 1 10?00010?0001100000
?0?0001110000
01??????01
10010000100100011
010001
01000100010000100100011
10010000100100011
010001
1000101000011
10001
07001
1000101100142
10001
00001
10011
100210?1?0122
00001
1 101100012
?00?1
10701
101
00001100111001101100722
00001101011010111200012
00001100011001100100032
00001101011011100100032
00001101011011101100012
70077777011011101100012
70071107011010111200032
00001101011010111200032
00001
10101
07701
10701
00001101011011107170772
101
101
A. ormosioides
7077777701
1010101200012
A. surinamensis
00001101011010101100022
A. cordata
10101100010100100011113
A. grandistipula
11101100010100111717033
A.parviflora
70177107017100101001013
A. unifoliolata
70101100010100101071013
A. inermisRTP 5\2
A. inermisRTP 5S0
A. intrm?RTP 589
77777777070700100100011
77777777070700100100011
77777777070700100100011
A. multistipula
77777777077700110100011
A. jaliscensis
????????0???00102100014
A.
77777777077700101001031
taurotesticulata
A. marauensis
1107170772
1 10?1?0??2
7777777707770010111017?
A.cujabensis
77777777070700102001013
A. carvalhoiAMC
77777777077701
A. ozraz//i0/RTP217
77777777777771100100032
100100032
solved, except for a clade comprising A. cordata, A. cori?cea, and A. micrantha. With the
character of leaflet number ordered, the analysis resulted in the same number of equally
trees (2966, CI = 0.57, RI = 0.80, length = 38), but the strict consen
most-parsimonious
sus tree resolved another clade comprising A. unifoliolata and A. trifoliolata.
Combined molecular and morphological
analyses. The first analysis resulted in five
trees of length = 76; CI = 0.72; RI = 0.88. The strict consen
equally most-parsimonious
sus is shown in Fig. 7. Of the equally most-parsimonious
trees, two can be rejected be
cause they result from incorrect optimization of missing values in accessions of A. nitida
for character 47. These accessions are assigned a character state for large, rodent-dis
for the two
persed fruit, when they do not have this state (this species is polymorphic
states of bat-dispersed fruit). Rejection of these topologies does not change the topology

26
VOLUME 64
-Hymenolobium
flavum
-Hymenolobium
n'rtidum
-A.
inermis
CEH
1673
-A.
inermis
TDP
13558
-A.
inermis MC
CLADE I
3579
-A. parviflora
-A.
cordata
-A.
grandistipula
-A.
unifoliolata
-A.
carvalhoi
CLADE II
RTP 229
-A. macrothyrsa
PLASTOME
GROUP I
-A.
galeottiana
-A.
humilis
-A.
verm?fuga
-A.
surinamensis
-A.
ormosioides
-A.
anthelmia
-A.
legalis
-A.
legalis HCL sn
-A.
fraxinifolia
-A.
carvalhoi
-A.
fraxinifolia
-A.
fraxinifolia
-A.
nitida
RTP 300
-A.
nitida
RTP 292
RTP 269
RTP 227,282
RTP 307
CLADE III
RTP 250
RTP 233
RTP 213
MS
889
-A. nitida RTP 301

FIG. 7. Strict consensus
cladogram of five equally most-parsimonious
resulting from cladistic
cladograms
data. Collector
and number are indicated
analysis of combined Andira cpDNA restriction site and morphological
when more than one accession
for a single species was included in the molecular
study.

27
ANDIRA
2003
of the strict consensus tree. When the same analysis was run on PAUP 4.0b2, where the
fruit character (47) can be coded as polymorphic for A. nitida, 26 equally most-parsimo
nious trees resulted, with an identical strict consensus. The greater numbers of equally
trees reflect different arrangements of the accessions of A. fraxinifo
most-parsimonious
A.
A.
and
carvalhoi {RTP 233). The semi-strict consensus tree ismore resolved,
lia,
nitida,
with the accessions of A. nitida grouping with A. carvalhoi RTP 233.
The only difference between this analysis and that presented by Pennington (1996)
lies in the extra resolution in the strict consensus tree in Clade III (Fig. 7). Andira ormo
sioides, A. anthelmia, and A. legalis form amonophyletic
group, supported by the synapo
morphy of larger flowers pollinated by xylocopid bees. Andira fraxinifolia, A. nitida, and
A. carvalhoi are a monophyletic
group diagnosed by the ability to root-sprout and form a
multistemmed
shrub.
The combined analysis that included a subset of cpDNA restriction site data for
species not included by Pennington (1995, 1996) resulted in ca. 2700 trees (the limit that
the computer could store; length = 87; CI = 0.65; RI = 0.87). The strict consensus is shown
in Fig. 8 and demonstrates that A. taurotesticulata, A. multistipula, and A. jaliscensis are
members of Clade I, which previously only comprised accessions of A. inermis (Pen
nington 1995, 1996). The affinities of A. multistipula and A. jaliscensis with A. inermis
were suggested by their close overall similarity in morphology. Andira taurotesticulata,
with its white-yellow
flowers and large ridged fruit, is more divergent morphologically.
Yet, it does have in common with A. inermis and A. multistipula an endocarp of parenchy
matous
tissue, shown in the equally-most parsimonious tree (Fig. 9, character 48, state 1)
to be the plesiomorphic
state for Andira. Similarly, Pennington (1994) and Pennington and
Gemeinholzer
(2000) predicted that all species with an endocarp of stone cells would be
members of Clade II. This is corroborated by the placement of A. cujabensis in this clade,
and it is likely that A. cori?cea, A. trifoliolata, A. tervequinata, A. micrantha, and A. prae
cox also belong here.
Andira marauensis
is a member of Clade III, placed as sister species to all other
of
this
because
it has the plesiomorphic
state of restriction site characters
clade,
species
33 and 38. Clade III remains "cryptic" in the sense of Wojciechowski
et al. (1993), be
cause it is not marked by any morphological
characters but well supported by cpDNA re
striction
Andira
site
characters.
jaliscensis,
A.
multistipula,
A.
taurotesticulata,
A.
cujabensis,
and
A.
ma
rauensis have incomplete restriction site data, and therefore many missing values. Termi
nal taxa with missing values are liable to occupy different topological positions in indi
vidual equally most-parsimonious
trees (Nixon & Davis 1991; Platnick et al. 1991), thus
trees and decreasing the resolution
increasing the numbers of equally most-parsimonious
of consensus trees. Therefore, in an attempt to gain a more accurate idea of the relation
ships of each of these species, separate analyses were run where all except one was ex
cluded. In only one case, that of A. taurotesticulata, did this result in a different placement
for the species in comparison with Fig. 8. In the analysis where A. taurotesticulata was in
cluded alone, it is resolved as a sister species to the different accessions of A. inermis in
Clade I in a strict consensus cladogram, because it possesses the plesiomorphic
state for
character 42 (leaflet undersurface indumentum).
The results of this combined analysis that includes five extra species is used as the
basis for the following discussion of character evolution and biogeography,
despite the
problem of the large number of equally-most parsimonious trees.

28
VOLUME 64
-Hymenolobium
flavum
-Hymeno/obium
nrtidum
-A.
inermis
CEH
1673
-A.
inermis
TDP
13558
3579
-A.
inermis MC
-A.
inermis
RTP 512
-A.
inermis
RTP 580
-A.
inermis
RTP 589
CLADE I
-A. mu/tistipula
-A.
jaliscensis
-A.
taurotesticulata
-A.
parvif/ora
-A.
cujabensis
-A.
cordata
-A.
grandistipu/a
-A.
unifoliolata
-A.
carvalhoi
RTP 229
-A.
carvalhoi
AMC
CLADE II
PLASTOME
GROUP I
-A. macrothyrsa
-A.
galeottiana
-A.
humilis
-A.
verm?fuga
-A. marauensis
-A.
surinamensis
-A.
ormosioides
-A.
anthelmia
RTP 227,282
-A.
legalis
RTP 307
-A.
legalis
HCL sn
-A.
fraxinifolia
RTP 250
-A.
fraxinifolia
RTP 213
-A.
fraxinifolia
MS
CLADE III
889
-A.
n?tida RTP 300
-A.
n?tida RTP 292
-A.
n?tida RTP 301
-A.
carvalhoi
RTP 233
-A.
carvalhoi
RTP 217
FIG. 8. Strict consensus

cladogram of 2996 equally most-parsimonious
cladograms
resulting from cladis
tic analysis of combined Andira cpDNA restriction site and morphological
of species
data, including accessions
not studied previously by Pennington
Collector
and number are indicated when more than one ac
(1995,1996).
cession for a single species was included in the molecular
study.

ANDIRA
2003
29
CHARACTEREVOLUTION
Stipules (Fig. 9, character 41). Large persistent stipules have
pendently in Andira, and are autapomorphic for A. grandistipula
synapomorphic for A. anthelmia and A. legalis. In Pisum sativum
mutant, stipules-reduced
{st) causes a marked reduction in stipule
arisen three times inde

and A. multistipula and
L. (pea), a single gene
size from the large, fo
liaceous wild-type
stipules (Marx 1987). It is tempting to ascribe the apparent lability of
this character in Andira to a similarly simple genetic switch. Indeed, Andira could be a
good model system to investigate the evolutionary significance of such a developmental
mutation, especially in the case of A. multistipula, where it is the presence of these large,
this species from A.
persistent stipules that is the diagnostic character that distinguishes
inermis. From the standpoint of the phylogenetic
(Nixon & Wheeler
concept
species
1990), it was the fixation of the trait of large stipules in a population that represented the
speciation
event.
Flowers
(Fig. 9, character 43). White petals are shown to have arisen at least three
times. They are a synapomorphy for Clade II, and an autapomorphy diagnosing both A.
macrothyrsa and A. taurotesticulata. Other species not sampled in themolecular study (A.
cori?cea, A. micrantha, A. praecox, A. tervequinata, A. trifoliolata) are known to have an
endocarp composed of stone cells and to lack wing petal sculpturing, both synapomor
phies for Clade II. Hence, all these species probably belong in this clade, which would
then be defined by white flowers. Andira chigorodensis,
the remaining species with white
most
to
is
A.
similar
and may be its sister species.
petals,
morphologically
macrothyrsa
Loss of wing petal sculpturing is a synapomorphy for Clade II, and an autapomorphy for
A. taurotesticulata.
Itmay have also been independently
most
is
species
closely allied with A. macrothyrsa.
lost by A. chigorodensis,
if this
Fruit type and dispersal (Fig. 9, character 47). The new restriction site data for A. tau
rotesticulata demonstrate that it is a member of Clade I, and that there have been at least
five independent origins of rodent dispersal from bat dispersal inAndira. This is one more
independent origin than reported by Pennington
(1996), confirming the prediction that
four origins was an underestimate. Of the remaining species with large fruit, A. cori?cea
and A. micrantha are likely to be members of Clade II based upon their floral characters
of stone cells, but they share no great morphological
similarity with A.
the other species of that clade with large fruit. Andira macrocarpa
has a
woody endocarp, and its affinities are hard to guess, because it has never been collected
in flower. Again, I suggest that further independent origins of rodent dispersal may have
and endocarps
grandistipula,
occurred during the diversification
o? Andira.
Endocarp (Fig. 9, character 48). Examining the sequence of character state change on
the cladogram (Fig. 9) shows a transition from weak endocarps of parenchyma to
woody
fibrous endocarps to endocarps composed of stone cells. This represents a sequence of in
2000; discussion above), and seed
creasing seed protection (Pennington & Gemeinholzer
have
the
been
this
pr?dation may
pressure driving
evolutionary sequence.

30
r-fl-[]?Hymenolobium
?|?Hymenolobium
VOLUME 64
f/avum
nitidum
?A.
1 7 13202530323642
fOttHf?
10 1 110
inermis
1673
CEH
1232
1824 rff--A.
10 0
4XH
0 ?
inermis
00
?-A.
CLADE I
13558
TDP
3579
inermis MC
parvif/ora
4246
I-A.
3 1012192122283637434548
ttOHUOHHH
000100101013
A.
cordata
CLADE M
16274147
0 113
A. grandistipu/a
A. unifoliolata
0 1111
394247
S-S?
A.
RTP 229
carvalhoi
1? 3
4348
1423
0 1
4647
if 1 2
-A. macrothyrsa
-??A.
3439
/H//W//S /?7P 269
-??A.
ft 1 1
HHKHKtH
0 0 0 10 2
PLASTOME
GROUP I
galeottiana
?A.
verm?fuga
A. surinamensis
A.
ormosioides
A.
anthelmia
RTP 227r282
/?7P 307
4 6 8 3338
A.
/e#3//s
LKK}ft
10 0 11
A.
legalis HCL sn
A.
fraxinifolia
RTP 250
A.
fraxinifolia
RTP 213
-A. fraxinifolia
47
?A,
MS
carvalhoi
CLADE III
889
RTP 233
A. n?tida RTP 300

A. n?tida RTP 292
A. n?tida RTP 301
FIG. 9. An equally most-parsimonious
cladogram
resulting from cladistic analysis of combined Andira
data. The bars representing
character changes are labelled above with
cpDNA restriction site and morphological
the character number and below with the character state. Solid black bars = unique changes, stippled black bars
=
and number are indicated when more than one accession
for a single species
changes. Collector
homoplasious
was included in the molecular
study.

2003
ANDIRA
31
CRYPTICCLADES
Pennington (1995) reported that none of themonophyletic
groups, even the well-sup
ported clades, discovered in themolecular analysis o? Andira had been recognized by pre
vious workers. They lack major morphological
innovations and hence are "cryptic" clades
sensu Wojciechowski
et al. (1993). The combined analysis demonstrates that new micro
characters provide unambiguous support for some of these cryptic groups.
morphological
Clade II is supported by two unique micromorphological
character states: lack of wing
an
and
of
stone
cells
petal sculpturing (character 45)
(character 48). All species
endocarp
of Clade I have an endocarp of parenchyma, but this maps as a plesiomorphy for Andira
rather than a synapomorphy. At a higher taxonomic level, micromorphological
characters
have also proved congruent with other clades that are well supported by molecular data in
(Rudall 2000). In both cases, reciprocal illumination provided by new
monocotyledons
hypotheses of grouping from molecular data is important. For example, itwas only when
the cpDNA restriction site data suggested the Clade II grouping of Andira species that I
noticed
that endocarps of these species appeared somewhat paler and harder than in other
species, which prompted the anatomical study reported above.
Other clades, such as clade III, remain cryptic, supported by restriction site data but
not by morphological
characters. This raises the question of whether they should be rec
ognized in formal classifications (Wojciechowski et al. 1993; Pennington & Gemeinholzer
2000), such as an infrageneric classification o? Andira.
INTRAGENERICCLASSIFICATIONSIN
THE LIGHTOF THE CLADISTICANALYSES
The cladistic analyses show that the infrageneric classifications proposed for Andira
Bentham
(1860, 1862) and Mattos (1979) are flawed. The only section that has been
by
that
is sect. Paucifoliolata Mattos, comprising only A.
may be monophyletic
proposed
A.
amonophyletic
and
This
is
cladis
unifoliolata
trifoliolata.
group in themorphological
tic analysis, if the character of leaflet number is treated as ordered. The
gynoecium indu
mentum character used by Bentham to define his sections Lumbricidia and Euandira, and
by Mattos to define corresponding subsections in her section Lumbricidia is homoplasious
is paraphyletic and
(Fig. 10); thus Bentham's section (Mattos's subsection) Lumbricidia
section Euandira (Mattos's subsection Glabratae) polyphyletic.
the cladistic framework presented here does not provide a clear route
Unfortunately,
to a new infrageneric classification o? Andira based upon
monophyly. There are four well
supported clades (Fig. 9): Clade I, Clade II, Clade III, and the group of Clade II, Clade III
and Plastome Group I; however, giving these clades formal taxonomic
recognition leaves
the weakly supported "Plastome Group I," supported as monophyletic
by a single, homo
restriction
site
character.
both
Clade
III
and
I remain cryptic,
Clade
Moreover,
plasious
restriction
site
but
not
data
characters. Whilst there is no
supported by
by morphological
reason to doubt that these cryptic clades reflect true
phylogeny because they lack mor
ultrastructural
and
characters may well
phological
support (indeed,
micromorphological
be found to support them, as was the case for Clade II), I question the
practical utility of
giving them formal taxonomic recognition. Because of the lack morphological
characters,
identification. Hence, I propose that sectional
they cannot be keyed out for conventional
classification within Andira
should be abandoned. A further factor supporting this

32
?Hymenolobium
-Hymeno/obium
flavum
nitidum
VOLUME 64
= 0
= 0
-A.
= 0
taurotesticulata
?A.
= 0
jalisensis
-A.
1-A -A.
inermis
TOP
-A.
inermis
-A.
inermisRTP
CEH
1673=
inermis
RTP 580=
-A.
inermis
RTP 589
?A.
= 0
512=
-A.
13558=
3579
inermis MC
0
0
= 0
= 0
multistipu/a
-A.
= 0
cujabensis
= 0
-A. parviflora
cordata
-A.
grandistipula
A.
A.
A.
carvalhoi
RTP
A.
carvalhoi
AMC=
-j?A.
i-A.
humilis
.
RTP
=
verm?fuga
ormosioides=
A.
A.
RTP
legalis
RTP 307
legalis
HCL
fraxinifolia
RTP
250
fraxinifolia
RTP
213=
fraxinifolia
MS
A.
n?tida RTP
A.
carvalhoi
A.
0
0
<?
0 <?
A. n?tida RTP 301=
sn = 0
889=
292=
= 0
= 0
A.
n?tida RTP 300=
227,282=
A.
A.
anthelmia
-r ??A.
?A.
269=
1 <?
= 0
surinamensis
A.
marauensis=
A.
= 0
A. galeottiana
??A.
unifoliolata
= 0
229=
A. macrothyrsa=
<?
RTP
carvalhoi
233=
RTP
217=
0
0
FIG. 10. An equally most-parsimonious

cladogram
resulting from cladistic analysis of combined Andira
data showing character state changes for indumentum of the gynoe
cpDNA restriction site and morphological
taxa marked
Terminal
cium. The bars represent character state changes from a hairy to a glabrous gynoecium.
"1" have a glabrous gynoecium.
Andira nitida and A. ver
terminal taxa marked
"0" have a hairy gynoecium;
and number are indicated when more than one acces
m?fuga have both hairy and glabrous gynoecia. Collector
sion for a single
species was
included
in the molecular
study.

ANDIRA
2003
33
argument is thatmolecular data are not available for some species whose relationships
hard to estimate based upon morphology
alone (e.g., A. cubensis, A. macrocarpa).
are
BIOGEOGRAPHY
Distribution
and Habitats
The majority o? Andira species are endemic to South America. The exceptions are A.
and A. galeottiana
and
(endemic to Cuba), A. jaliscensis
(endemic toMexico),
A. inermis (widespread throughout the entire Neotropics and also present in Africa).
cubensis
Most species grow in rain forests. Four species, A. cordata, A. cujabensis, A. humilis,
and A. verm?fuga grow in the savanna woodlands
("cerrados") of central Brazil. Andira
inermis subsp. rooseveltii occurs in a similar wooded savanna habitat in Africa. Andira
carvalhoi is entirely restricted to the sandy coastal restinga scrub and forests of eastern
Brazil. Andira fraxinifolia, A. legalis, A. anthelmia, A. ormosioides, and A. nitida grow
both in restinga and adjacent rain forests in eastern Brazil. Andira jaliscensis
is a species
of seasonally dry tropical forests, as is A. inermis subsp. glabricalyx. Andira inermis
subsp. inermis also occurs in seasonally dry tropical forest in Central America and on the
Caribbean coasts of Venezuela and Colombia, but elsewhere in the Neotropics,
and in
Africa, it grows in rain forests.
Origin
and Transatlantic
Distribution
lacks a fossil record, though it is reasonable to hypothesize

that it originated
early in the Tertiary, because there is relatively extensive fossil record of related genera of
Sophoreae and Dalbergieae from middle Eocene deposits in north America (Herendeen et
al. 1992). It is not clear whether the relationships of Andira and its sister genus Hy
menolobium are with New or Old World legumes, because their phylogenetic position is
Andira
(Pennington et al. 2001). Furthermore, the lack of resolution in the basally di

Andira
clade (clade I, Fig. 8), inwhich all distribution areas (South America, Cen
vergent
tralAmerica, the Caribbean, and Africa) are represented, means that little can be inferred
about the early diversification o? Andira. The placement of A. inermis and A. jaliscensis
in this basal lineage is intriguing and not inconsistent with a northern Boreotropical ori
unresolved
gin (cf. Lavin and Luckow 1993), but no firm conclusions can be drawn until relationships
are better resolved. The phylogenetic position of theMexican A. galeottiana suggests that
it is derived from South American progenitors, which is not a Boreotropical pattern.
The transatlantic distribution of A. inermis might be the result of ancient vicariant
events or of more recent long-distance dispersal. Lavin et al. (2000) described vicariant
relationships between Africa and North America in two legume groups
in the Tertiary in seasonally dry tropical vegetation. This vicariance ex
planation would predict higher levels of cpDNA restriction site divergence between
American and African accessions of A. inermis than are observed (Fig. 9). The ability of
A. inermis to disperse over water barriers makes more recent long-distance dispersal ap
pear to be amore likely explanation. The distribution of A. inermis in the Caribbean gives
indications of the ability of its fruits to be dispersed over water, presumably by floating.
Its presence in Jamaica indicates that the species has the capability of crossing short
stretches of ocean. Jamaica appears to have been completely submerged during a period
in the Oligoc?ne
1988, as cited in Lavin, 1993), and therefore its
(Buskirk 1985; Donnelly
biogeographical
that diversified

34
VOLUME 64
current biota must have the ability to disperse over water. The distance of Jamaica from
the nearest island where A. inermis is present (Haiti; ca. 200 km) is, of course, an order of
shorter than that between South America and Africa.
magnitude
Nothing is known of the potential of Andira fruits for long-distance dispersal by float
ing in sea water. Four observations are pertinent: (i) the distributions of A. surinamensis
and A. anthelmia often appear to follow river courses, a pattern seen inmany taxa known
to be dispersed by water (data from herbarium labels; pers. obs.; H. C. Lima, pers.
comm.); (ii) the fruits of A. galeotiiana are often found washed up on beaches inMexico
(Gunn & Dennis
1976), and this species is characteristic of river and lake banks, and
flooded areas (Pennington & Sarukh?n 1968); (iii) Andira seeds, protected by their hard
endocarps, have remarkable viability; seeds of A. carvalhoi that were not excised from
their endocarps germinated at Kew Gardens 14 months after collection; (iv) Andira iner
mis grows on the strand line in Cameroon (M. Cheek, pers. comm.; see notes onM. Cheek
it often grows along rivers and can survive inmangrove swamps
3579); in the Neotropics
(in Colombia; notes on H. Murphy & G. Parra 723).
The low frequency of long distance dispersal events may be indicated by the absence
of A. inermis from Cuba (where only A. cubensis occurs) despite the proximity toHaiti (ca.
100 km at the closest point). Given the presence of A. inermis on the islands of the Lesser
and all the other major islands of the Greater Antilles, this absence from Cuba is
puzzling, although a similar pattern occurs in Coursetia caribaea (Jacq.) Lavin van carib
aea (Lavin 1988). No other species of Andira have ranges as wide as that of A. inermis. For
example, in the Caribbean, A. surinamensis (widespread in South America) is present only
on Trinidad and Tobago, which are extremely close to the South American mainland. It is
Antilles
enigmatic that long-distance dispersal of these species is less effective, because there are
no apparent differences in their dispersal biology in comparison with that of A. inermis.
The explanation may lie in the ecology of A. inermis, such as its ability to survive inman
grove swamps and on beach strand-lines, which appears unique in the genus.
Recent
Patterns
in South America
The lack of cpDNA divergence within the species of clades I, II, and III (Fig. 9) is con
sistent with the origin of most South American species in recent times, possibly as late as
the Pleistocene. Morphological
change has occurred, but the slowly evolving cpDNA has
accumulated few, or in the case of Clade III, no differences among species. Such patterns
are characteristic of recent radiation on oceanic islands (reviewed by Bateman, 1999).
All
(an Amazonian
species of Clade III, with the exception of A. surinamensis
occur
seems to have
in
moist
the
forests
of
coastal
southeastern
Brazil.
there
Thus,
species),
been a recent burst of speciation in this area. Andira surinamensis and the other species of
Clade III are separated by the wide "dry diagonal" of cerrado and caatinga vegetation that
runs across central and northeastern Brazil. The close relationship of A. surinamensis and
the species of Clade III is suggestive of previous links between the mesic vegetation of
southeastern Brazil and Amazonia, which are more often evidenced by disjunctions of the
same species between these two areas (Mori et al. 1981; Prance 1985, 1987).
The species of Clade II are from the central Amazonian
region (A. parviflora, A. uni
the Guianas (specifically Guyana; A. grandistipula),
and the central Brazilian
(A. cordata, A. cujabensis). The lack of divergence between the plastomes of
these species also suggests a relatively rapid radiation of these species. The morphologi
foliolata),
cerrados
cal characters
supporting
this clade (lack of wing
petal sculpturing,

endocarp composed
2003
ANDIRA
35
H oc
H 20e
CERRADO
BIOIVJE
50?
80?
-Contact
line of Andira
cujabensis/cordata
-Contact
line of harry & glabrous
Andira humilis
.Miiimmi.
inContact
line of
Pterodon
hairy
& glabrous
forms of
forms
of
emarginatus
Lines dividing
distinct
northern and
south-eastern
cerrado
sites
FIG.
11. Cerrado biogeography.

Dashed
line: line of contact between
the parapatric distributions
of Andira
and A. cujabensis
line with dots: line of contact between hairy and glabrous forms
(see Fig. 39). Dashed
of A. humilis. Comb: line of contact between hairy and glabrous forms of Pterodon
lines:
emarginatus. Dotted
cordata
lines dividing the distinct northern and southeastern cerrado

tion of 98 areas of cerrado vegetation
(Ratter et al. 1996)
sites identified
by analysis
of the floristic
composi
of stone cells) suggest that this clade comprises nine species (those listed above plus A.
trifoliolata, A. micrantha, A. praecox, and A. cori?cea), all of which are confined to these
areas, so the five species represented in themolecular and combined analyses probably are
a representative geographic sample.
The parapatric distributions of the Clade II species A. cujabensis and A. cordata, which
are found in the Brazilian cerrado vegetation, are especially interesting (see Fig. 11). The

36
VOLUME 64
line of contact of these species' distributions through the states of Goi?s and Tocantins
closely matches the lines dividing the distinct northern and southeastern sites in the cerrado
discovered by TWINSPAN
(two-way indicator species analysis) of the floristic
composition of 98 areas of cerrado and Amazonian savanna vegetation (Fig. 11 ;Ratter et al.
1996). This is also the line of contact for the glabrous and hairy forms of A. humilis (Fig. 11;
see account under that species), and similar distributions are shown by hairy and glabrous
forms of Pterodon emarginatus Vogel (Leguminosae, Papilionoideae; Fig. 11; R. T. Pen
nington, unpubl.). The subspecies of Diptychandra aurantica Tul. also have similar distrib
vegetation
utions, with
subsp. aurantica occurring in the southeast of the cerrado biome, and subsp.
epunctata (Tul.) Lima, Carvalho & Costa occurring in the north, though in this case, the
ranges only abut at their southern ends inMinas Gerais (de Lima et al., 1990). These con
gruent patterns are suggestive of a common underlying historical factor. A possibility is re
cent separation in the late Quaternary of the now continuous cerrado vegetation by more
mesic vegetation, such as that suggested by the palaeopalynological
data of Ledru (1993).
TAXONOMY
Note: The following terms are used to describe the placement of indumentum on
ovary, which is approximately oval in cross section. The "upper surface" refers to
areas on and adjacent to the adaxial suture, and the "lower surface" refers to the areas
and adjacent to the abaxial suture; the "sides" refer to the two regions bounded by
the
the
on
the
adaxial and abaxial sutures.

Andira
Lamarck, Encycl. 1: 171. 1783, nom. conserv. Andira sect. Euandira Bentham,
Comm. legum. gen. 45. 1837. Andira sect. Lumbricidia subsect. Glabratae N. F.
58: 2. 1973, nom. superfl.?Type:
Andira
inermis (W.
Mattos, Loefgrenia
Wright)
Lumbricidia
DC.
Vellozo, Fl. flumin. 305. 1829. Andira sect. Lumbricidia (Vellozo) Ben
Comm.
tham,
legum. gen. 43. 1837. Andira sect. Lumbricidia subsect. Lumbri
cidia (Vellozo) N. F.Mattos, Loefgrenia 58: 2.1973.?Type:
Lumbricidia legalis
Vellozo
[=Andira legalis (Vellozo) Toledo].
Andira sect. Paucifoliolatae
N. F. Mattos, Loefgrenia 58: 3. 1973.?TYPE: Andira
unifoliolata
Ducke.
Trees, shrubs, or rarely geoxylic suffrutices. Bark often producing small amounts of
red ex?date when cut. Indumentum of simple, red-brown or occasionally pale whitish
hairs, or absent. Stipules large and persistent or small, narrow and early caducous. Leaves
spirally arranged, imparipinnate, with up to 9 pairs of leaflets, occasionally 3-foliolate or
1-foliolate; rhachis canaliculate; leaflets subtended by stipels (occasionally absent); peti
olules stout, swollen, canaliculate; leaflets glabrous adaxially, glabrous or hairy abaxially,
primary vein channelled or plane adaxially, prominently raised abaxially, secondary veins
or mixed eucamptodromous-brochidodromous.
brochidodromous
Inflorescences axillary
and terminal, paniculate, with pale brown, brown, or red-brown, simple indumentum, the
branches subtended by caducous bracts. Flowers pedicellate or occasionally
sessile, each
subtended by a caducous bract; paired caducous bracteoles at base of calyx. Calyx shal
lowly to deeply and subequally 5-lobed, the upper lobes broader than the lower lobes, all
lobes with simple, red-brown indumentum, or glabrous. Petals 5, free, clawed, purple,

ANDIRA
2003
37
pink, or white; wing petals with or without lamellate sculpturing; keel petals overlapping,
firmly attached, but not united. Stamens 10, the filaments united for at least half of their
length, the vexillary stamen free. Ovary distinctly stipitate, with simple hairs or glabrous;
ovules 1-8. Fruit a 1 (-2-3)-seeded
drupe, ? globose to elongated; sutures (along abaxial
and adaxial surfaces of the fruit) raised or sunken or indistinct; stylar remnant present or
indistinct; exocarp glabrous, green, brown, or occasionally
yellow when ripe and fresh;
sweet
and
mesocarp fibrously fleshy
smelling (drying hard, fibrous and granular) or hard,
non-fibrous, and odorless (drying hard and finely granular); endocarp very hard, woody,
or woody and fibrous. Seeds white, filling the entire cavity, the hypocotyl-radicle
axis a
small fold at the apex; testa dark reddish brown, adhering to the endocarp. Seedlings cryp
number: x = 10, 11.
tohypogeal. Chromosome
The order of species reflects the groups discovered by the combined analysis of
The relationships
of species not
cpDNA restriction site characters and morphology.
characters. The
included in this analysis are estimated based upon their morphological
A. multistipula,
groups are (refer to Fig. 8): (i) "Clade I": A. inermis, A. jaliscensis,
A. taurotesticulata,
and A. cubensis (the last species not included in the combined analy
to A. inermis);
similarities
sis but placed in this group because of its morphological
A.
A.
"Plastome
I":
A.
A.
humilis, and
(ii)
macrothyrsa,
galeottiana,
verm?fuga,
Group
not
included
in
A. chigorodensis
last
the
combined
(the
analysis but placed
species
to A. macrothyrsa);
similarities
here because of its morphological
(iii) "Clade III":
A.
A.
surinamensis,
anthelmia,
A.
legalis,
A.
A.
ormosioides,
fraxinifolia,
A.
n?tida,
A. carvalhoi, A. marauensis,
and A. macrocarpa
(the last species not included in the
of
combined
but
here
because
its
similarities to A. suri
analysis
morphological
placed
namensis);
(iv)
"Clade
II": A. praecox,
A. micrantha,
A.
cori?cea,
A.
trifoliolata,
A.
ter
vequinata (these species not included in the combined analysis but included here be
cause they share two features that define this clade: an endocarp of stone cells and wing
A. cujabensis,
petals without sculpturing), A. parviflora, A. cordata, A. grandistipula,
and A.
unifoliolata.
Keys
to the Species
of Andira
of Andira are easy to identify if complete material

of leaves, flowers,
and fruit is available,
Specimens
one emphasizing
to distinguish
when sterile. Two keys are presented:
but they are often very difficult
vege
use of the keys, especially
of the first key, re
tative characters and the other flowers
and fruits. Successful
the three different
among
quires distinguishing
leaflets (refer to Fig. 2 and Fig. 12). Understanding
ical (refer to Fig. 13).
used to describe
leaf morphology
terminology
is measured
from the joint of the stem and petiole
The
length
surface of the
found on the abaxial
types of indumentum
of the terminology
used to describe fruit shape is also crit
FIG.
follows
the guidelines
of Hickey
(1979). Leaf
to the joint of the rhachis and terminal petiolule.
long), dense,
fine, pale,
All
types of the abaxial leaflet surface o? Andira. A. Short (ca. 0.1 mm long), appressed
erect hairs of A. fraxinifolia.
B. Long (ca. 0.5 mm long), flexuose,
C. Long (ca. 1mm
(See also Fig. 2B, C, D.)
tangled hairs of A. jaliscensis.
12. Indumentum
hairs of A. surinamensis.
axis

38
VOLUME 64
of Andira. A. ? Globose,
FIG. 13. Fruit morphology
smooth, dried fruit (ca. 3.5 cm long) of A. jaliscen
C. Elongated, wrinkled,
sis. B. Elongated,
dried fruit (ca.
smooth, dried fruit (ca. 6 cm long) of A. chigorodensis.
4.5 cm long) of A. nitida.
of petiolule
length is
length and leaflet size and shape are for the terminal pair of leaflets. Leaflet
to the tip of the lamina and does not include the length of the petiolule. Acumen
length is mea
the acumen is broad, this point is
sured from the point where the leaflet margin curves towards the apex. Wlien
measurements
from the base

ill defined,
Flower
wing
petals
sculpturing
Many
"elongated"
13C). Fruit
lengths must be treated as approximate.

and ismeasured
from the base of the calyx to the end of the
length is reported for dried specimens,
in fully open flowers. A fully open flower is one in which the standard petal is reflexed. Wing petal
is described following
the terminology
of Stirton (1981).
so acumen
have more or less globose fruits (Fig. 13A). Other

(Fig. 13B, C). Fruit of some species dry smooth (Fig. 13A,
from the tip of the stipe (where it joins
length ismeasured
from the abaxial to adaxial
is the maximum measurement
species
fruit height
surement between
the fruit walls.
I. Key emphasizing
1. Leaves
vegetative
all uni- or trifoliolate;
2.
Leaves
trifoliolate.
2.
Leaves
unifoliolate.
1. Leaves
3.
Geoxylic
3.
Tree or shrub.
5.
vein plane on adaxial
primary
surface.
28.
A. unifoliolata.
pairs of leaflets, or sometimes
A.
irifoliolala.
also some
leaves
trifoliolate;
primary
shrub-like,
to 2 m
suffrutex
forming mats
more
up to 10 m
in diameter
(occasionally
than 1.7 cm long and 0.5 cm wide
at base, persistent,
crowded
glabrous
Leaflets
Leaflets abaxially
lens or microscope);
6. Leaflets abaxially
visible
1.
at in
3. A. multistipula.
(use lens or
tiny (<0.1 mm long) appressed straight hairs beneath (use

25. A. grandistipula.
leaflets up to 4 pairs; the Guianas.
with erect or ? appressed, often flexuose hairs (0.2-1.0 mm long;
to the naked eye); leaflets 3-9 pairs; eastern Brazil.
6.
7.
tall).
10. A. humilis.
and shoot apices.
with sparse, erect hairs); Amazonia.

(very occasionally
indumentum
abaxially hairy, sparsely hairy, or with tiny appressed
Guianas
and eastern Brazil.
microscope);
Leaflets
vein
surface.
Stipules generally
florescence
bases
5.
characters.
29.
two or more
all with
sunken on adaxial
4.
species have fruits that are referred to as

B) and of others distinctly wrinkled
(Fig.
the body of the fruit) to the tip of the fruit;
suture; fruit width is the maximum mea
with
at maturity;
leaflets hairy abaxially,
Stipules red-brown pubescent,
glabrescent
over 100 g when dry, pale
hairs red-brown, erect; fruit 5.6-12 cm long, weighing
14.
A.
brown when fresh and dry, odorless when fresh.
legalis.
when
(red-brown appressed-pubescent
Stipules generally
glabrous at maturity
to
leaflets sparsely to very sparsely hairy abaxially, hairs red-brown
young);
ca. 20 g when dry,
whitish, ? appressed to ? erect; fruit 3-6.2 cm long, weighing
13. A. anthelmia.
dark brown when fresh and dry, sweet smelling when fresh.
4.
sometimes
less than 1.7 (-2.0) cm long and 0.5 cm wide at base, often caducous,
entirely absent (rarely persistent and then paired only at leaf bases and not crowded at inflo
rescence bases or shoot apices).
Stipules
8.
Leaflets
with
indumentum
abaxially
(use lens or microscope).

ANDIRA
9.
39
undersurfaces
(particularly primary and secondary veins) with erect, gener
visible with the
hairs (>0.2-1.0 mm
generally
ally flexuose
long); indumentum
to touch.
naked eye and perceptible
Leaflet
tan
red-brown),
abaxially with dense indumentum of pale (occasionally
leaflets appear pale abaxially (seen with
gled hairs that ? obscure the epidermis;
the naked eye).
to cordate; flowers
11. Leaflets
base rounded
5.5-7 mm
coriaceous,
long;
21. A. cujabensis.
Brazil.
10. Leaflets
to subcoriaceous,
to rounded; flowers
base obtuse
Mexico.
2.
A.
jaliscensis.
long;
abaxially with indumentum of red-brown hairs (or if pale, then indu
leaflet often appearing red
sparse) that do not obscure the epidermis;
11. Leaflets
chartaceous
9-1 lmm
10. Leaflets
mentum
brown
is sparse).
abaxially (to the naked eye; if pale then indumentum
12. Secondary
veins plane or slightly raised (occasionally
raised)
prominently
indumentum
abaxially, tertiary veins plane or very slightly raised abaxially;
of abaxial
suture;
leaflet
surfaces
in montane
forests
caducous; fruit distinctly ribbed from suture to

at 700-2000
in lowland forest
m, occasionally
(Ecuador).
12. Secondary
veins
raised abaxially;
ribbed from
prominently
indumentum
5. A.
taurotesticulata.
raised
abaxially,
tertiary veins prominently
of abaxial leaflet surface persistent; fruit not
suture to suture, or with
broad,
indistinct ribs;
lowland
forests
and savanna.
13. Leaflets
2-3 pairs, the apex rounded, often slightly retuse; fruit 10-11
cm long, weighing
300 g when dry.
H.A. macrocarpa.
13. Leaflets
(2-) 3-7 pairs, the apex obtuse to rounded, often acuminate,
cm long,
sometimes
retuse; fruit 2.4-6
(-10 [A. galeottiana
only])
10-20
(-125 [A. galeottiana

only]) g when dry.
cm long, 40-125
6-10
g when dry, dried mesocarp
8. A. galeottiana.
structure; Mexico.
relatively soft with sponge-like
14. Shrub or tree; fruit 2.4-6 cm long, weighing
10-20 (-30) g when
weighing
14. Tree;
fruit
hard (difficult to insert the point of a needle),

dry, dried mesocarp
with hard, fibrous or granular structure; South America.
15. Secondary
veins
flowers
6-7.5 mm
10-15;
long, whitish,
standard with purple markings; Amazonia.
20. A. parviflora.
15. Secondary
veins 5-11; flowers
12.5-23 mm long, pink to pur
ple, the standard with a central white or cream marking; eastern
and central Brazil (occasionally Amazonia).
16. Stipels 1-2 mm long; stipules <5 mm long, caducous; gy
or with
noecium
sparse hairs along upper and
glabrous
lower surfaces of ovary; dry fruits distinctly
wrinkled;
Central
Brazil
and
Bolivia
(cerrado),
occasionally
Amazonia.
9. A. verm?fuga.
1-9 mm long; stipules to 16 mm long, caducous or

moderately
persistent; upper and lower surfaces and walls
of ovary hairy; dry fruits smooth; eastern Brazil (restinga,
rain forest, campo rupestre, and secondary vegetation).
16. Stipels
17. Shrub or small
tree to 12 m, with
flowers
spreading
crown
in
13-17 mm
long; fresh fruit

A. fraxinifolia.
16.
green with green mesocarp.
17. Tree to 30 m, tall with small crown in open situations;
open
situations;
18-23 mm long; fresh fruit very dark brown

15. A. ormosioides.
pale greenish-white
mesocarp.
tiny (<0.2 mm long), tightly appressed hairs; indumentum
to the naked eye or perceptible
by touch (use lens or
flowers
with
9.
Leaflets
generally
abaxially with
not visible
microscope).
18. Tertiary
impressed
raised on abaxial
leaflet surface,
distinctly
or slightly impressed on adaxial leaflet surface.
veins

veins
secondary
12. A. surinamensis.
40
VOLUME 64
18. Tertiary veins plane or slightly raised on abaxial leaflet surface,

plane or slightly raised on adaxial leaflet surface.
19. Shrub.
20. Leaflets
1-2 pairs; flowers
20. Leaflets
2-4
9 mm
long; Venezuelan
secondary
Guayana.
27. A.
10-13 mm
(-5) pairs; flowers
veins
long; eastern Brazil.
tervequinata.
17. A. n?tida.
19. Tree.
21. Leaflets
(1-) 2 (-3) pairs.

22. Secondary veins 6-7; leaflets widely elliptic to widely
pairs; small tree to 7 m tall; Venezuelan
Guayana.
22. Secondary
veins
to 10 mm
23. Stipules
long; stipels
1-2 mm
long; flowers
26.
long;
Amap?.
to 2.5 mm
23. Stipules
long; stipels absent

9-11 mm long; Bahia.
1-2
27. A. tervequinata.
obovate
(occa
elliptic to narrowly
(1-) 2 (-3) pairs; tree to 35 m
leaflets
8-10;
obovate),
sionally widely
Brazil.
obovate,
(perhaps
tall; eastern
6.5-7 mm
A. praecox.
early cadu
18. A. marauensis.
cous); flowers
3-9 (-10) pairs.
24. Leaf axis 37-55 cm long; leaflets 7-9 pairs.
7. A. chigorodensis.
24. Leaf axis 6-33 (-40) cm long; leaflets 3-8 (-10) pairs.
25. Leaflets coriaceous; flowers red to purple; Cuba.
4. A. cubensis.
21. Leaflets
25. Leaflets
marked
flowers white to yellowish,

the standard
subcoriaceous;
red or purple; Panama and South America.
with
26. Stipules absent (or early caducous; never seen), stipels to 2

mm long, caducous; fruit 5-5.5 cm long, smooth,
weighing
10-20 g when dry.
6.
A. macrothyrsa.
to 12 mm long, stipels 1-3 mm long, generally per
cm long, distinctly ribbed from suture
sistent; fruit 5.4-7.2
to suture, weighing
100-200 g when dry.
5. A. taurotesticulata.
(use lens or microscope).
26. Stipules
8.
Leaflets
glabrous abaxially
27. Leaflet base truncate or cordate
27. Leaflet
base obtuse
eastern Brazil,
Guianas,
28. Shrub
22. A. cordata.
(rarely rounded); Brazil, in cerrado.
?
or
truncate
the
Amazonia,
cordate);
(rarely
slightly
in rain forest and restinga.
to rounded
(occasionally
small tree to 10 m).
19.
A.
carvalhoi.
28. Tree.
in 2-9 pairs (some leaves on a plant always with more
of leaflets); Neotropics
and Africa.
1.
inermis.
A.
29. Leaflets
in 2-3 pairs; South America.
29. Leaflets
30. Stipules
to 5 mm
quite persistent,
wide
at base,
than 3 pairs
to 15 mm
Guianas.
30. Stipules caducous, often

long; Central Amazonia,
entirely absent, <1 mm wide

eastern Brazil.
at base,
long; the
24. A. cori?cea.
to 6 mm
31. Stipules early caducous, probably

small and narrow (never seen);
leaflets 2 pairs; petals 6.5-7 mm long, white to cream with the stan
dard marked with lilac; fruit 91-0 cm long, weighing
up 300 g when
23.
A.
micrantha.
dry; central Amazonia.
31. Stipules
at shoot apices
present
generally
(though caducous);
leaflets 2-4 (-5) pairs; petals 10-13 mm long, pinkish white to pur
coastal Brazil.
ple, the standard with a central white spot; Atlantic
17. A. n?tida.
II. Key emphasizing

1. Leaves
flower and fruit characters.
all uni- or trifoliolate; primary vein plane on adaxial

trifoliolate.
28. A. trifoliolata.
2.
Leaves
2.
Leaves
unifoliolate.
surface.
29. A. unifoliolata.

2003
Leaves
all with
two or more
sunken on adaxial
3.
41
ANDIRA
pairs of leaflets, or sometimes
also some leaves
Stipules generally more than 1.7 cm long and 0.5 cm wide

florescence
base and shoot apex.
4.
19-24 mm
Flowers
5.
trifoliolate;
primary
vein
surface.
cm
at the base, persistent,
crowded
long; eastern Brazil.

over
long, weighing
100 g when dry; stipules red-brown

at maturity.
14. A. legalis.
young, glabrescent
cm long, weighing
ca. 20 g when
Fruit 3-6.2
dry; stipules red-brown
13.
pubescent when young, generally glabrous at maturity.
Fruit
5.6-12
at in
pubescent
when
5.
4.
Flowers
6.
11-15 mm
and Guianas.
long; Amazonia
in (2-) 3-4 pairs, apex rounded (rarely obtuse) with an acumen to 3 mm long,
tiny (<0.1 mm long) appressed hairs abaxially (use lens or microscope).
25. A. grandistipula.
Leaflets
with
pairs, apex acute to obtuse with an acumen to 15 mm long, glabrous

A.
with few, erect hairs).
3.
multistipula.
(very
occasionally
abaxially
sometimes entirely
Stipules less than 1.7 (-2) cm long and 0.5 cm wide at base, often caducous,
base
absent (rarely persistent and then paired only at leaf bases, but not crowded at inflorescence
6.
3.
appressed
A. anthelmia.
in (5-) 6-9
Leaflets
or shoot apex).
7. Fruit 5-11 cm long, weighing
(40-)
8. Leaflets glabrous abaxially.
9.
Flowers
9.
Flowers
14-15 mm
g when
dry.
long; shrub or small tree to 10 m; eastern Brazil,
in restinga.
19. A. carvalhoi.
6-7 mm
10. Ovary
10. Ovary
8.
100-300
and the Guianas,

in rain forest.
long; trees to 40 m; Amazonia
mm
mm
to
15
5
wide.
24. A. cori?cea.
glabrous; stipules quite persistent,
long,
23. A. micrantha.
sparsely pubescent;
stipules early caducous (not seen).
Leaflets
with indumentum abaxially.

11. Fruits ? globose; petals white, the standard marked with red; tertiary veins plane on
A.
abaxial surface of leaflet.
5.
taurotesticulata.
11. Fruits
elongated; petals
tiana; flowers unknown
pink to violet, the standard marked with white (A. galeot

for A. macrocarpa);
tertiary veins distinctly raised on abax
ial surface of leaflet.

12. Dried mesocarp
relatively soft (the point of a needle is easily inserted), fibrous,
and spongy; fruits weighing
40-125
8. A. galeottiana.
g when dry; Mexico.
12. Dried mesocarp
not
hard (the point of a needle is inserted only with difficulty),
1.
300 g when dry; Ecuador.

fibrous; fruits weighing
10-20 (-30) g when dry.
(-7) cm long, weighing
13. Petals white to yellow
(occasionally
pale pinkish or pale
H.A.
macrocarpa.
Fruit 2.4-6
lilac),
the standard generally
red or purple markings.

14. Ovary glabrous.
22. A. cordata.
with
14. Ovary pubescent.

15. Leaflets with
on abaxial
erect indumentum
16. Secondary
veins
epidermis
clearly
Para), in rain
surface.
leaflets abaxially
tree to 20 m,
visible;
forest.
10-15;
with
red-brown
the
indumentum,
smooth; Brazil
(Amazonas,
A.
20.
parviflora.
with
generally
pale indumentum,
bark
veins 8-10; leaflets abaxially

16. Secondary
often so dense that the epidermis
a
is hidden; tree to 12 m (occasionally
do Sul,
shrub), bark thick, fissured; Brazil
(Mato Grosso, Mato Grosso
21.
A. cujabensis.
Goi?s, Para), in cerrado and gallery forest.
glabrous or with appressed indumentum
17. Leaflets 4-9 pairs; endocarp of fruit fibrous
15. Leaflets
inserted).
18. Flowers
9-11 mm
18. Flowers
5-6
17. Leaflets
a needle
long;
(-7) mm
leaf axis
on abaxial
surface.
(the point of a needle
10-33
long; leaf axis 37-55
is easily
cm
long; leaflets 4-8

6. A. macrothyrsa.
pairs.
cm long; leaflets 7-9 pairs.
(-40)
1-2 (-3) pairs; endocarp of fruit very hard, amorphous

is inserted only with difficulty).

7. A. chigorodensis.
(the point of
42
19. Flowers
VOLUME 64
6.5-7 mm
ally with
long; secondary veins 8-12 per leaflet; leaves gener

tree
2 pairs of leaflets (occasionally with 3 pairs or trifoliolate);
to 30 m. 26. A. praecox.
9 mm long; secondary veins 6-7 per leaflet; leaves with 1-2

27. A. tervequinata.
pairs of leaflets; shrubs or small trees to 7 m.
red or pink to purple, the standard generally with a white or cream central mark
19. Flowers
13. Petals
ing.
20. Dry fruits
distinctly wrinkled
(Fig.
with erect indumentum
13).
on abaxial
surface.
9.
A. verm?fuga.
glabrous or with appressed indumentum on abaxial surface.
22. Geoxylic
suffrutex forming mats up to 10 m in diameter
(occasionally
to 2 m tall); fresh mature fruit yellowish;
central Brazil and
shrublike,
21. Leaflets
21. Leaflets
10.
A.
humilis.
Paraguay, in cerrado.
22. Shrub or tree to 40 m; fresh mature fruit green; Amazonia,
eastern Brazil.
the Guianas,
23. Secondary
veins plane to slightly raised abaxially,
tertiary veins
17.
A.
flowers 10-13 mm long; eastern Brazil.
and
plane;
n?tida.
raised abaxially; flowers

23. Secondary veins and tertiary veins prominently
12. A. surinamensis.
13-18 mm long; Amazonia
and the Guianas.
not
and
but
20. Dry fruits smooth (the surface sometimes
wrinkled)
warty,
rough
(Fig.
13).
24. Flowers
7-8.5 mm
24. Flowers
9-23 mm
A.
4.
cubensis.
long; Cuba.
Caribbean
(excluding Cuba), and Africa.
long; Neotropics,
25. Leaflets with erect, spreading indumentum on abaxial surface.
26. Flowers 9-11 mm long; only the upper surface of ovary hairy; leaflets
abaxially densely hairy and the epidermis often not visible; Mexico.
26. Flowers
13-23 mm
pubescent;
Brazil.
27. Flowers
greenish
27. Flowers
leaflets
2. A. jaliscensis.
long; upper and lower surfaces and sides of ovary
abaxially hairy but the epidermis
always visible;
18-23 mm
long; fresh fruit very dark brown with pale
tree to 30 m, with a small crown in open
white mesocarp;
situations.
15. A. ormosioides.
13-17 mm
shrub or small
long; fresh fruit green with green mesocarp;

tree to 12 m, with a broad spreading crown in open
situations.
16. A. fraxinifolia.
indumentum.
glabrous or abaxially with appressed
1. A. inermis.
28. Leaflets glabrous abaxially.
28. Leaflets with appressed hairs abaxially.
25. Leaflets
29. Calyx with

leaflet base
leaflets 2-4 (-5)

indumentum;
sparse appressed
obtuse, or rounded to slightly cordate; flowers
mm long; ovules (1-) 2-4.
17.
A.
29. Calyx glabrous except around margins of lobes; leaflets (1-)
pairs; leaflet base acute to obtuse; flowers 9-11 mm; ovules
pairs;
10-13
n?tida.
2 (-3)
1-2.
18. A. marauensis.
1. Andira
inermis (W.Wright) DC, Prodr. 2: 475. 1825. Geoffraea

inermis W. Wright,
Lond. Med. J. 8: 256. 1787. Geoffrea acutifolia Stokes, Bot. mat. med. 4: 46.
1812, nom superfl. Vouacapoua inermis (W.Wright) Lyons, PL nam. 396. 1900.
Andira jamaicensis Urban, Symb. antill. 4(2): 298. 1905, nom. superfl.?TYPE:
Jamaica. W. Wright s.n. (holotype: BM!).
Tree to 35 m tall with broad spreading crown (in open situations in the Neotropics;
trees in the African savannas may have ascending branches and a pyramidal crown; see
Polhill, 1969); buttresses slight; bark rough, scaling, grey to dark brown; slash very pale

2003
ANDIRA
43
5-10 mm thick; twigs sparsely hairy when young, the hairs red-brown, erect,
glabrescent. Stipules to 17 mm long, to 1mm wide, occasionally persistent; leaf axis 6-34
cm long; rhachis sparsely to very sparsely hairy, hairs red-brown, erect; stipels 1-9 mm
long, often caducous; petiolules 2-7 mm long, sparsely to very sparsely hairy, hairs red
brown, erect; leaflets 2-9 pairs, 3.5-13.5 cm long, 1.4-6 cm wide, narrowly elliptic, el
subcoriaceous to thick-chartaceous, base obtuse
liptic, narrowly obovate to oblanceolate,
an acumen to 15 mm long, glabrous (occa
to
rounded
with
obtuse
acute),
apex
(rarely
brown,
sionally very sparsely hairy abaxially, the hairs erect) except the primary vein adaxially
and abaxially very sparsely hairy, the hairs red-brown, erect, >0.2-1.0 mm long; primary
vein channelled adaxially, raised abaxially, secondary veins 10-14 (-16), plane adaxially
and abaxially or slightly raised abaxially, pattern brochidodromous,
often with the basal
few veins eucamptodromous,
tertiary veins plane adaxially and abaxially. Panicles termi
cm long, hairy to sparsely hairy at branch tips, becom
10-40
nal, occasionally
axillary,
the
less
hairs
towards
red-brown, ? erect; bracts 2.5-3 mm long, with red
base,
ing
hairy
brown appressed hairs; pedicels to 4 mm long or flowers subsessile; bracteoles 1.5 mm
long, indumentum like that of bracts. Flowers (9-) 10-12 (-19) mm long. Calyx 4-7 mm
long, with appressed, red-brown indumentum or glabrous, except at the base and margins
of lobes; lobes 0.3-1 mm long. Petals pale pink to purple; standard blade 7-14 mm high,
9-11 mm wide, claw 1-3 mm long; wing 6-13 mm long, 3^.5 mm wide, claw 2.5-5.5
mm
long, lamellate sculpturing present; keel 6-13 mm long, 3^.5 mm wide, claw 2.5-6
mm long. Stamens 10-18 mm long, filaments united for the basal 5-11 mm, free for the
distal 3-7 mm, vexillary stamen 7-13 mm long. Gynoecium 9-20 mm long, usually with
sparse, red-brown, ? appressed hairs only on the upper surface of the ovary, but more oc
casionally on both the upper and lower surfaces of the ovary, base of the style, and upper
surface of the top of the stipe; stipe 3-10 mm long, ovary 3-6 mm long; style 3-5 mm
long; ovules 2-4. Fruit 3-6 cm long, 2.5-^.3 cm high, 2-4.3 cm wide, ? globose to ? elon
gated, weighing ca. 20 g or less when dry, green, drying dark brown to black, smooth or
somewhat rough and warty, occasionally
slightly wrinkled; suture slightly raised adaxially
and below; stylar remnant insignificant or absent; mesocarp
1-3 mm thick, hard, granu
lar; endocarp 2-5 mm thick, brown, woody, fibrous. Chromosome number: n = 10.
Andira inermis, occurring in both the Neotropics and in Africa, is a widespread and
variable species.The key characters that differentiate the subspecies are the presence or
absence of indumentum on the calyx and the size of the flowers. The most widespread
subspecies, subsp. inermis, has a calyx with indumentum covering its entire outer surface,
whereas subspecies glabricalyx and rooseveltii have calices that are glabrous, except at
the base and on the margins of the lobes. Similar intraspecific variation in calyx indu
mentum is found in A. humilis and A. multistipula, but in these species the extremes are
linked by intermediates, and therefore subspecies are not recognized. Flower size inA. in
ermis varies continuously from 9-19 mm, and alone forms no basis for the distinction of
intraspecific taxa; however, theMexican
specimens with a glabrous calyx (subsp. glabri
calyx) have flowers from 10-11 mm long, whereas the African specimens with a glabrous
calyx (subsp. rooseveltii) have larger flowers (13-14 mm long). Flower length therefore
provides the diagnostic character to separate these two subspecies.
There is clear geographic partitioning of variation inAndira inermis, with subsp. roo
seveltii restricted to wooded savanna in Africa, and subsp. glabricalyx found in Nayarit
and Sinaloa at the edge of the neotropical distribution of A. inermis inMexico.
inermis jamaicensis.
Subse
Wright (1777) first described this species as Geojfroea
quently
(Wright 1787), he reported the medicinal
uses of the tree and used the binomial

44
VOLUME 64
inermis. Though he provided no botanical description in 1787, the name G. in

Geoffraea
ermis is validly published, because Wright cited his previous work (1777).
Key to the
1. Calyx
inermis
Andira
with
glabrous except
2. Flowers
10-11 mm
2. Flowers
inermis
of Andira
la.
indumentum.
appressed
at the base and margins of the lobes.
long; calyx 2.5-3 mm long; Mexico.
13-14 mm long, calyx 4-5 mm long; Africa.
covered
1. Calyx
Subspecies
lb. Andira
inermis
lc. Andira
inermis
subsp.
inermis.
subsp. glabricalyx.
subsp. rooseveltii.
la. Andira
inermis subsp. inermis.

MEXICO. Guer
Andira excelsa Kunth inH. B. K., Nov. gen. sp. 6: 385.1824.?TYPE:
rero: Prope La Venta de Tierra Colorada, Bonpland 3901 (holotype: P!).
Andira riparia Kunth in H. B. K., Nov. gen. sp. 6: 386. 1824.?Type:

COLOMBIA.
Santander: Prope La Boca del R?o Opon, in ripa fluminis Magdalenae, Bonpland
1599 (holotype: P!).
Pterocarpus sapindoides DC, Prodr. 2: 419. 1825. Amerimnum affine Sprengel, Syst.
veg. 3: 192. 1826, nom. superfl. Andira sapindoides
(DC) Bentham, J. Proc.
Linn. Soc, Bot. 4, Suppl. ("A Synopsis of the Dalbergieae"):
123. 1860. Voua
Bertero
capoua sapindoides (DC) Kuntze, Revis, gen. pi. 1: 212. 1891.?Type:
s.n. (holotype: G-DC!; isotype: TO).
Andira grandiflora Guillemin, Perrottet & A. Richard, FI. Seneg. tent. 254. 1832.
Andira inermis subsp. grandiflora (Guillemin, Perrottet & A. Richard) Gillett ex
Gambia. Albreda, Perrottet 31 (holo
Polhill, Kew Bull. 23: 490. 1969.?Type:
not
located; isotype: BM!).
type: P,
Glycyrhiza undulata Ruiz & Pav?n ex G. Don, Gen. syst. 2: 227. 1832.?Type:
Peru. Ruiz & Pav?n s.n. (holotype: BM!).
acuminata Bentham, Comm. legum. gen. 45. 1837.?Type:
"In
Brazil.
sylvis ad flumen Solimaen provinciae Rio Negro," Martius 1157 (holotype: K!;
isotype: M).
Andira inermis var. riedelii Bentham, J. Proc. Linn. Soc. Bot. 4, Suppl. ("A Synopsis
of the Dalbergieae"):
123. I860.?Type:
Brazil.
Mato Grosso: Rio Coxim,
Riedel 402 (lectotype, here designated: K!).
Andira
Lonchocarpusl
Cameroon.
staudtii Harms
Kumba
in Engler, Bot. Jahrb. Syst. 26: 301. 1899.?Type:

Staudt 912 (holotype: B, de
[=Johann-Albrechtsh?he],
stroyed; isotype: BM!).

Andira chiricana Pittier, Contr. U.S. Nati. Herb.
Chiriqu?: David,
NY!).
Flowers
Fig. 2A.
(9-)
10-12
30 Mar
(-19) mm
1911, H. Pittier
long. Calyx
18(6): 235. 1917.?TYPE: PANAMA.

3372 (holotype: US!; isotypes: GH!
covered with
appressed
indumentum.
in Central America and the Caribbean coast of South

Phenology. Dry vegetation
America: flowering concentrated February-May;
other records scattered throughout the
but scattered through the year.
year. Caribbean: flowering concentrated May-September,
Guianas and Amazonia: flowering records scattered throughout the year. Africa: in rain
forest (Cameroon, Nigeria, Femando Po), flowering March-May;
in wooded
savanna
(Senegal, Ivory Coast, Mali, Central African Republic), flowering March-April.
Distribution
(Fig. 14, 15). Caribbean (Jamaica, Dominican Republic, Haiti, Puerto

ANDIRA
2003
110
105
100
95
90
85
80
75
70
45
65
60
55
50
45
40
35
Rico, Virgin Islands, Antigua, Guadeloupe, Dominica, Martinique, St. Vincent, St. Lucia,
Grenada, Trinidad, Tobago); Mexico and throughout Central America; Venezuela, Colom
bia, Ecuador, Peru, Bolivia, the Guianas, Paraguay, and northern Argentina (Corrientes);
Africa (Gambia, Senegal, Ivory Coast, Nigeria, Cameroon, Equatorial Guinea). In the dif
ferent parts of its range, A. inermis subsp. inermis grows in various habitats. In Africa, it
occurs in wooded savanna and rain forest. In Central America and the Caribbean coast of
it is common in seasonally dry tropical forests, whereas else
northern South America
it grows in rain- and gallery forest, including inundated areas and
where in the Neotropics
even
mangrove.
Black plum (Tobago); l'angelin, angelim, bois palmis (Mar

palo maco (Dominican Republic); cabbage bark, bastard cabbage
tinique, Guadeloupe);
cuilimbuca, maquilla
cuautololote,
(Mexico); cab
(Jamaica); tololote, quiringuacua,
almendro
cirvelillo,
(El Salvador,
(Belize);
frijolillo
bage bark, barley wood,
Guatemala, Honduras, Nicaragua);
cabbage bark, quira, harino, cocu (Panama); congo,
Vernacular
names.
(Colombia);
majagua gallina, palmillo, ciruelo de playa, bolombolo, mot?n
(Venezuela); barbasco caspi, montera
caspi, numi,
pil?n, lumbricero, peloto, mot?n
pisho, sacha ushuu (Peru); koraro, bat seed (Guyana); reddie kabbes, vremoessoe noto
rouge (French Guiana);
sapupira
(Suriname); lebi kiabici, St. Martin
lombrigueira,
(Brazil); mendubira
(Argentina).
dividivi,

46
FIG.
15. Distribution
Representative
Massif
Nord-Ouest:
of Andira
Specimens.
inermis
Jamaica.
subsp.
inermis and A.
Manchester:
du Nord,
Petit Ford, Ekman

Port-de-Paix,
Domingo:
vie. Ciudad Trajillo, Allard 13367 (US).?SAMAN?:
Las Terrenas, Zanoni & Mejia
Loma
17748 (NY).?DAJAB?N:
inermis
VOLUME 64
subsp. rooseveltii
in Africa.
Donnell
Smith 2389 (US).
Haiti.
Mandeville,
4326 (NY, US).
Santo
Dominican
Republic.
Finca Hacienda Nydia,
3.5 km E of village of
de Cabrera, banks of R?o de Masacre,

Zanoni &
on road to Cotui, Zanoni et al. 16093
RAM?REZ: 7 km from center of Pimentel
(NY).?S?NCHEZ
Puerto Rico. Ca. 2 km N Hwy 3, just E town of Rio Grande, Boom 6783 (GH, NY).
Islands.
(NY).
Virgin
St. Croix: Belvedere
Box 1385 (A, US).
estate, D'Arcy 4727 (GH, MO).
Antigua. Wallings,
Guadeloupe.
Pointe Noire, Duss 3231 (NY, US).
Dominica.
Sastre
Batalle, Ramage 221 (K). Martinique.
Basse-Pointe,
17899
Mej?a
7731 (GH).
St. Lucia. Anse Louvet, Slane 803 (A).
St. Vincent.
Kingstown,
Grenada.
Trinidad.
Balthazar, banks of Great River, Beard 157 (GH).
US).
s.n. (F, NY).
ton 2906 (GH, NY, US).
Tobago.
Scarborough, Broadway
carretera Esc?rcega-Villahermosa,
Mexico.
CAMPECHE: Km 30 desviaci?n
Beard
1336
Pitch Lake,
(F, GH, MO, NY,

vie. Brighton, Brit
rumbo a Palizada, Chan & Lira

sobre la
(MEXU).?Chiapas:
Mpio. Angel Albino Corzo, 500 m W de la salida de la colonia Quer?taro,
terraceria a Finca Prusia, Reyes Garc?a 1724 (MO).?Guerrero:
de los Bravos, 4 km SE
Mpio. Chilpancingo
de Tlahuizapa,
camino El Ocotito-Jalenca,
R?o Potrero, Mart?nez & T?llez 170 (MO).? MlCHOAC?N: El Hue
4707
outskirts of San Pedro de Tepantepec,

close to main
tamo, arroyo de Chapa, Soto N??ez 93 (MO).?Oaxaca:
road running ENE towards Tuxtla Guti?rrez, Hughes
1673 (E, FHO, K, MEXU, MO, NY).?SlNALOA:
Sierra
Madre foothills, between Rosario and Colonias, Rose 1639 (NY, US).?Tabasco:
camino Playa Azul, 8 km de
3046 (MO).
Para?so, Cowan & Maga?a
Belize.
BELIZE: Belize-Sibun
hwy, Croat 24861

mingbird
ten River, 2 km N of Victoria
with
Road,
Gentle
(MO, US).?Stann
15 (F, NY, US).?Cayo:

along
Creek:
Cockscomb Mountains,
Sibun River, just W of Hum

trib. of Cocoa Branch of Sit
Peak, Gentry 7887 (MO, US).?TOLEDO:

hwy to Punta Gorda, 1mi E of junction
Croat 24471 (MO, US).
Alta Verapaz:
Rio Sebol, downstream
Guatemala.
from
44779
Smith 2823 (GH, US).?IZABAL:
Escuintla, Donnell
Maricos,
(US).?ESCUINTLA:
road to San Antonio,
Carrizal,
Steyermark
near Jutiapa, Standley 60583 (F).?PETEN: Laguna
7600 (MO).?JUTIARA:
bordering Lake Izabal, E. Contreras
S of Sayaxch?, Steyermark 46202
Petexbat?m
Marcos:
Palmatto Flats, 1-2
(F, US).?San
(Laguna M?xico)
mi N of Oc?s, Steyermark
37865
(F).?SANTA ROSA: Chiquilmulilla,
Standley 79182 (F).?SUCHITEP?QUEZ:
vie. Tiquisate,
and Santa Cruz, Standley
47462
between R?o Honda
74130
(F).?Zacapa:
(F).
Steyermark
en la vega de Quebrada Tolobre,
ATLANT?DA: vic. Tela, Standley 53570
(GH, US).?Choloteca:
entre Morolica
et
y Tolobre, Standley 14219 (NY).?COL?N:
Trujillo, R?o Negro, Clewell
al. 4339 (MO).?COMAYAGUA:
Pito Solo, Lake Yojoa, Edwards
1 km N
429P (GH).?CORTES:
Matorrales,
DE LA Bah?a: West End, Isla de Roat?n, Nelson & Romero 4519
Villa Nueva, Molina R. 6790 (GH, US).?ISLAS
Honduras.
matorrales
y bre?ales
(MEXU).?Olancho:
Mpio.
San Esteban,
Santa Mar?a
del Carb?n,
30 km al NE
de San Esteban,

Sousa
et al.
ANDIRA
2003
47
R?o
R. 8633 (MO).
El Salvador. Ahuachap?n:
13369 (MEXU).?Valle:
Bah?a de San Lorenzo, Molina
vie. La Libertad, Standley 23206 (US).?
Libertad:
Paz, near Paso de Santa Cruz, Standley 20333 (US).?La
La Uni?n:
Miguel:
vic. La Uni?n, Standley 20861 (GH, US).?San
Standley 20991 (GH,
Laguna de Olomega,
Vicente:
vic. San
Salvador:
Finca Altamira, hills S of San Salvador, Alien 7190 (US).?San
MO, NY).?San
vic. Sonsonate, Standley 21796 (GH, MO, US).
Nicaragua.
3 km S El Sanee on banks of a small dry river
(F).?LE?N:
camino a Salamina, hacienda Santa Cruz, a 6 km de la car
R?o Lim?n, Lago da Granada, Shannon 5025
(MEXU).?RlVAS:
Standley 21277 (GH, US).?SONSONATE:

11189
Chinandega:
vic. Chichigalpa,
Standley
452 (FHO, MEXU).?MANAGUA:
course, Hughes
Vicente,
retera a Montelimar,
& Castro
Guzman
138
San
Brenes,
(F). Costa Rica. ALAJUELA: Parque Alberto Manuel
at R?o Abangartos,
Croat 61182 (MO).?
Carvajal 263 (F,MO).?GUANACASTE:
hwy Esparza-Ca?as
Rio Colorado
10047 (MO).?LLM?N:
HEREDIA: Finca La Selva, OTS field station on Rio Puerto Viejo, Folsom
Cant?n de Osa,
between Islas Buena Vista and Cerro Coronel, Davidse & Herrera 31242 (MO).?PUNTARENAS:
El Recreo,
(US).?Zelaya:
229
Long
Ram?n,
R?o Terraba, Alien 5220 (F,MO, US).?San

Jos?: 10 km NW Gu?piles, L. D.
vic. El
Panama.
CfflRlQU?: Progreso, Cooper & Slater 265 (F, GH, NY, US).?Cocl?:
(WO).
Alto de Las
Rio Ucurganti, Bristan
1127 (NY, US).?HERRERA:
Valle, Allen 1770 (GH, NY, US).?DARBEN:
256 (F).?Panam?:
R?o San Juan D?az, above Juan D?az, Alien 938 (GH).?SAN
BLAS:
Minas,
Carrasquilla
trail Palmar Norte-Ca?ablancal,
18517
G?mez
mainland
opposite
Play?n
Chico,
0-3 mi
from Caribbean,
Gentry
6373
(F).?VERAGUAS:
vic. Santa F?, Allen
4435 (GH).
DELTA Amacuro:
Serra Imataca, Cerro La Paloma, E side R?o Cuyubini,
Venezuela.
S?eyermark 87604
FEDERAL: E of Caraballeda, Hacienda
Juan D?az, Steyermark 62930 (F,MO).?FALC?N:
Dist.
(U).?DISTRITO
113652 (MO).?Miranda:
R?o Chico, Jahn 1239
Colina, R?o Ricoa, Dos Bocas, Steyermark & A. Gonzales
between La Pica and Ca?o Colorado, E Marur?n 6 km W La Ormega, Wurdack &
(GH, NY, US).?Monagas:
Monachino
Davidse
Cerro Las Minas, S main road from Santa Ana, 17 km SE Santa Ana, Stey
(NY).?T?CHIRA:
of R?o Catatumbo
and La Fria-Maracaibo
Distrito Col?n,
intersection
road,
re
(MO).
Guyana. West Bank Demerara River, Boom 7198 (NY, US); Pomeroon-Supernaam
39495
et al.
ermark
(MO).?Zulla:
18831
Between mouths
& Persaud
1217 (MEXU, US).
Suriname.
Gillespie
gion, Adventure,
and Coronie, Lanjouw & Lindeman
1460 (MO, U).
French Guiana.
Vicinity Cayenne,
Amazonas:
P. N. Amacayac?,
Colombia.
Cabana Pamat?, Narv?ez
& Olmos 69
US).
2 km from Barraquillita,
lands of
95 (COL, JAUM, MO).?BOL?VAR:
Brand & Cogollo
Curran
Loba,
23
(US).?Caldas:
R?o Magdalena,
cerca
a La Dorada,
of rivers Coppename
611 (GH,
Broadway
(COL).?Antioquia:
Loba, San Mart?n
de
del R?o Purnio, Idrobo & R.

la Yuca, M. S?nchez & G. Mahecha
confl.
2269 (COL, F, MO, NY, US).?CAQUET?:

Florencia, Quebrada
9a (UDBC).?CAUCA:
camino a Pablo Sexto y Cabo de Hornos, Lozano et
Mpio. Guap?, P. N. de Isla Gorgona,
al. 5245 (COL).?CHOC?:
Archer 2050 (NY,
Llor?, 50 km S Quibd?, at june. R?o Atrato and R?o Andagueda,
Planeta Rica, S. L?pez 4005 (COL).?Guajira:
Sierra Nevada de Santa Marta, Sierra de San
US).?C?rdoba:
Jaramillo
de La Cueva, Cuadros & Gentry 2965 (MO, NY).?MAGDALENA:

valle inferior del Mag
Bosque
841 (F, US).?Meta:
R?o Umea, Gavia 5122 (US).?Narl?O:
dalena, Dugand
Tamaco, R?o Rosario, 6 kms ar
near Bonda, H. H. Smith 18
del Caunap?, Romero 5218 (MO, NY).?Santa
riba de la desembocadura
Marta:
Puerto Araujo, Renter?a e? al. 1887 (JAUM, MO).?Valle:
Isla del
(F, GH, MO, NY, US).?Santander:
Antonio,
en el desembocadura
del R?o Cajambre, Cua?recasas
16224 (F).?Sucre:
around Verrugas, Finca La
Guayabal
Esmeraldas:
Ecuador.
San Lorenzo, 0.5 km S on walk to R?o Nadadero, Little
Aguada, Bernai 158 (COL).
Jr. 6341 (F, US).??apo:
La Joya de los Sachos Cant?n, Pompeya, R?o Indillana, between
its mouth at the R?o
et al. 2125 (QCNE).
and the MAXUS
Peru. AMAZONAS: Quebrada Huampami,
road, Gudi?o
?apo
Kayap
1061
(F,MO).?Cuzco:
Pilcopta, Atalaya, Paucartambo, N??ez
de Iparia a lo largo de R?o Pachitea,
Bosque Nacional
Croat
GH,
NY,
US).?LORETO:
(F,
Quebrada Cuninico,
Honoria,
1958
6851 (MO).?Hu?NUCO:
Prov. Pachitea, Dtto.
1 km arriba del pueblo de Tournavista,
Schunke
DE DIOS: Prov. Manu,
17759 (MO).?MADRE
10-15 km NNW Shintuya, Foster et al. 11027 (F).?PASCO: Prov. Oxapampa,

between Puerto Bermudez
and Paujul, Gentry et al. 42145
Bermudez,
(F,
Mart?n:
1-2 km E Tabalosos, Belshaw 3370 (F, GH, MO, NY, US).?
Prov. Lamas, Distr. Lamas,
MO).?SAN
Prov. Coronel Portal, Yarinacocha,
Ucayali:
Bolivia. BENI: Mamore,
Gentry & Horna 29375 (MO).
Tyson &
Cerro de Pantiacolla, R?o Palotoa,
1 hour below Puerto
R?o Pichis,
Kuns
Prov. N Su?rez, road Cobija-Porvenir

Km 2, Dom?nguez & E. Gonzales 57 (NY).?
(MO).?PANDO:
Cruz: La Guardia, W.M.A.
Brooke 5825 (F, NY).
Brazil. Acre: Mpio. Tarauac?, ?rea urbana, M. C.
Ferreira & A. P. Araujo 61 (CEPEC).?AMAP?:
Rabelo et al. 2238 (MO,
Igarap? do lago do Marac?, Mazag?o,
Serra
NY).?AMAZONAS:
Mpio. Mara?, Rio Japur?, afl. do Rio Solim?es, Cid Ferreira 3380 (F,MO).?Goi?S:
1003
Santa
de Caldas
da Pousada
do Rio Quente, margem

do Rio Quente, Heringer
12228 (MO, UB).?
Una do Trauira, Froes
1836 (GH, NY).?Mato
GROSSO: Mpio.
Region,
11292 (US).?Mato
GROSSO DO SUL: Pantanal do Rio Negro, Fazenda Salina,
Coxim, Rio Taquar?, Anderson
Dubs 475 (NY).?PARA:
lina de Arana, Mpio.
Santarem, Curral Grande, Black & Ledoux 50-10366
(GH).?
Maranh?o:
Novas,
Parque
Maracassum?
River

48
VOLUME 64
& Hass
15795 (SPF).?ROND?NIA:
Forte
PARAN?: Mpio.
Icaraima, Rio Paran?, Barra Rio Ivai, Hatschbach
MISIONES:
Schinini
da
local
&
Wilson
4273
W.
Beira,
(US).
Paraguay.
Ayolas,
Concei?ao,
Rodrigues
Principe
Corrientes:
& Vanni 25958
(GH, SPF).
Argentina.
Ituzaing?, Isla Apip? Grande, Puerto San Anto
Depto.
et al 24452 (MO).
nio, Krapovickas
449 (K). Gambia.
Mali. K?m?bra, Roberty 17062 (K).
Senegal. Koudougou
(Saraya), Nongonierma
Prov. Calabar, Dist. Itu,
380 (K).
Galam, Heudelot
Ivory Coast. Near Kpakobo, Ak? Assi s.n. (K). Nigeria.
25 km NNW Douala, Letouzey
14743 (K);
Atau Eki Beach, Iyizoba s.n. (K-2 sheets).
Cameroon.
Mayaen,
near Victoria, Maitland
1064 (K); Marienberg,
Polhill et al. 5214 (K); Douala-Edea
Reserve, bank of
D.
W.
313
Bioco
Km 5, J.
Thomas
Guinea.
River,
Malabo-Rebola,
(K).
P?),
(Fernando
Equatorial
Sanaga
et al. 2823 (K).
Fern?ndez
Victoria,
(1860) recognized A. sapindoides based on two specimens from Antigua

These and other collections from the Lesser Antilles (Antigua, Dominica,
and St. Lucia) have much larger flowers, up to 19
Guadeloupe, Martinique, Monserrat,
mm long, than specimens from other parts of the range. The existence of specimens with
flowers of intermediate size, such as H. E. Box 1385 (Antigua), appears to confirm Ben
Bentham
and Dominica.
tham's (1860) doubts that A. sapindoides may only represent a well-marked variant of A.
inermis, and the name is therefore placed in synonymy.
The type of A. chiricana {Pittier 3372; Panama: Chiriqui) has hairy ovaries and
leaflets that are sparsely hairy abaxially, but it otherwise matches A. inermis subsp. iner
mis. Ovary and leaflet indumentum appear sporadically in populations
throughout the
range of subsp. inermis.
Polhill (1969) provided an excellent summary of the variation displayed by A. iner
mis in Africa. He recognized two subspecies, which are geographically
separate and have
a calyx bearing indumentum: subsp. inermis and subsp. grandiflora. If the African mate
rial is considered in isolation, then these morphological
differences appear to merit sub
inermis
in
rain
forest
around the Gulf of Guinea in
grows
specific recognition. Subspecies
Cameroon and Nigeria, and has flowers of 12-13 mm long; subspecies grandiflora, from
savannas inGambia, Senegal, Mali, Central African Republic, and Ivory Coast, has larger
(to 15 mm long). These two variants agree in all other respects with Neotropical
populations of subsp. inermis. Because the range of flower size in the Neotropics encom
passes the range of variation in Africa, subsp. grandiflora cannot be considered distinct,
and the name is therefore placed in synonymy.
flowers
Andira inermis subsp. inermis has hard, heavy timber, which is used in the Neotrop
ics for furniture and cabinetwork, and also for construction, railway sleepers, fence posts,
etc. Because the wood has virtually no resonance, it is particularly suitable for radio and
television cabinets (Weaver 1989). It is also used as an ornamental street tree inVenezuela
and Costa Rica. The bark may be used as a vermifuge but is poisonous
in high doses
It
is
also
taken
for
intermittent
in
fevers
Mexico and Brazil (Morton 1981).
(Weaver 1989).
lb. Andira
inermis subsp. glabricalyx R. T. Pennington,

Mexico.
subsp. nov.?Type:
Nayarit: Mpio. Nayar, Arroyo de San Pablo, margen derecha del R?o Santiago,
frente al poblado de Colorado de laMora, 20 Jun 1992, A. Ben?tez-Paredes 3796
(holotype: MEXU!).
Ab A.
inermi subsp. inermi calycibus glabris differt. Ab A. inermi subsp. rooseveltii

(10-11 mm longis, non 13-14 mm) differt.
Flowers 10-11 mm long. Calyx 2.5-3 mm long, glabrous except at base and margins
of lobes.
floribus minoribus

2003
ANDIRA
49
Phenology. Flowering May and June, fruiting August

fruiting records in December and March.
and September,
with
single
Distribution
(Fig. 14). Mexico
(Nayarit and Sinaloa); deciduous and semi-deciduous
tropical forest. No flowering specimens have been collected in Sinaloa, but the remains of
are glabrous.
the calyx on the fruiting specimens Rose 1639,1782
Additional
de
Specimens
las Ventanas,
Examined.
Ben??ez-Paredes
Mexico.
3658
Nayartt:
(MEXU); Arroyo
between Zopilote
La Nanchflera
and Arroyo
ca. 10 km E la
P. H. Aguamilpa,
near junction for San
and Santa Cruz de Guaybel,
NE
km
San
Pedro
19
Ixcat?n, 2 km NE El
Nayar,
Mpio.
Brasil,
Nayar,
Embalse
between
et al. 18688 (MEXU);

et al 5474 (MEXU); Mpio.
del
Miguel
Zapote, Ch?zaro
terracer?a San Pedro Ixcat?n-Santa
Cruz Guaybal, G. Flores e? al. 1841 (MEXU); along dirt road be
Naranjo,
tween Pochitit?n and Francisco Madero, Miller & T?llez 3181 (MEXU, MO); Mpio. Tepic, 4.3 km E junction to
la Escondida,
R. Ramirez & G. Flores 674 (MEXU); Mpio.
old road Tepic-Mazatl?n,
Tepic, 5-7 km W.
35 km W Tepic, T?llez 9076 (MEXU); 4 km NE Pochotit?n,
road to Mojarritas,
Pochotit?n,
T?llez & Miller
cortina, Calzada
10478
(MEXU); Mpio. Tepic, Paso de Bueyes, 40 km SE Tepic, Tenorio e? al. 16869

tween Rosario and Colomas, Rose 1639 (US); near Colomas, Rose 1782 (US).
It is tempting to speculate that morphological

populations at the northern edge of the Neotropical
mis, producing both A. inermis subsp. glabricalyx
Jalisco and Michoac?n,
separating the ranges of A.
accurate information on
More
subsp. glabricalyx.
(MEXU).?Slnaloa:
be
divergence has occurred in isolated

distribution of A. inermis subsp. iner
and A. jaliscensis. The latter occurs in
inermis subsp. inermis, and A. inermis
the phylogenetic
relationships of these
taxa is necessary to corroborate this hypothesis, especially for A. jaliscensis, whose affini
ties are unresolved (Fig. 8). A similar geographic pattern is seen in Lennea viridiflora
Seem. (Leguminosae, Papilionoideae,
tribe Robinieae), where L. viridiflora var. novo
galiciensis Lavin & M. Sousa is restricted to Colima, Jalisco, and adjacent Michoac?n,
whereas L. viridiflora var. viridiflora is widespread throughout Central America (Lavin &
Sousa 1995).
lc. Andira
(de Wildeman) Gillett ex Polhill, Kew Bull. 23:

subsp. rooseveltii
Millettia rooseveltii de Wildeman, PL Bequaert. 3: 353.1925.?Type:
Mearns 3004 (holotype: BR;
Equatoria Province: Nimule-Gondokoro,
inermis
490.1969.
Sudan.
isotype: BM!).
brownii Hoyle, Kew Bull. 1932: 262. 1932.?Type:
Region: Jema, Feb 1931, W. T. Brown 2163 (holotype: K!).
Ostryoderris
13-14 mm
Flowers
long. Calyx 4-5 mm
long, glabrous
Ghana.
Ashanti
except at base and margins
of lobes.
Phenology.
Flowering
April-June.
Distribution
in wooded
February
(Fig. 15). Nigeria,
and March,
with
one
record
Chad, Central Africa Republic,
in May,
Togo, Uganda,
fruiting
Sudan;
savanna.
names. Gwaska
Vernacular
Additional
(Nigeria); Weri-dee
(Central African Republic).
Specimens
Examined.
Togo. Without
locality, Kersting 554 (K). Nigeria. Gangoro Forest
168 (K); Bauchi, Lely P192 (K); Sokoto, Uly
832
(K); northern Nigeria, Yola, Dalziel
54 (K). Chad. N of Nd?ll?, between
(K); Abakaliki,
Opara 609 (K); Plateau Prov., Selfwe River, Thornewill
Golo and Mansaca,
Chevalier
7773 (K).
Sudan. Yirol District,
around Koudogoi,
F. W. Andrews 464 (K);
Reserve,
Chapman
Imatong Mts, Kinyeti

cality, Schweinfurth
4203
lo
Valley, between Imeila and Hiliu, ca. 5 km S Hiliu, Friis & Vbllesen 1042 (K); without
1875 (K). Central Africa Republic.
Near Bonasse
lu, 20 kms from center, Fay s.n. (K);

VOLUME 64
50
2 km S Camp Koumbala
along Koumbala river, Fay 4321 (K); Bouar, Mildbraed
7252 (K).
Leya and Aiyu, river junction, Greenway & Eggeling
9422
(BM, K).
Uganda.
Madi,
Andira inermis subsp. rooseveltii ismore or less parapatric with the African savanna
form of A. inermis subsp. inermis (Fig. 15) in the wide belt of wooded savanna running
across central Africa from Gambia to Sudan and Uganda. Their ranges abut between Ivory
Coast (the most eastern record of the savanna form of A. inermis subsp. inermis) and
Ghana (the most western record of A. inermis subsp. roosveltii). Hopkins (1983) demon
strated that variation of leaf characters in Parkia biglobosa is clinal in these woodlands,
and thus did not provide a basis for recognition of intraspecific taxa. From the paucity of
herbarium specimens A. inermis appears a rare species throughout these African savan
nas, and Boulvert (1977) reported it as very rare in the Central African Republic. It is pos
sible that further collections from the savanna zone between the ranges of subsp. inermis
and subsp. rooseveltii (in southern Mali and Burkina Faso) might reveal intermediates,
and thus there may be a clinal situation similar to that found in Parkia biglobosa.
Andira inermis subsp. rooseveltii is reported to be used for children's gera, or sever
ance illness, by washing
2. Andira
babies with a decoction
(herb, label, Central African Republic).
Jalisco: Mpio. Talpa de

jaliscensis R. T. Pennington, sp. nov.?Type: Mexico.
camino a la mina del Cuale, 7.6 km E de la carretera Puerto Val
Allende,
iso
larta-Barra de Navidad, 30 May 1985, E. J. Lott 2544 (holotype: MEXU!;
type: MO!).
A.
inermi primo aspectu maxime similis sed foliolis subter dense pubescentibus.
Tree to 25 m tall; presence of buttresses unknown; bark and slash unkown; twigs
brown to dark brown with numerous pale elongated lenticels, sparsely hairy, the hairs
pale, erect, glabrescent. Stipules to 10 (-20) mm long, to 1mm wide, caducous, densely
erect; leaf axis 13-25 cm long; rhachis densely hairy, hairs white, erect,
tangled, glabrescent, especially towards the base; stipels minute (0.5 mm long), caducous;
petiolules 2-4 mm long, indumentum like that of rhachis; leaflets in 3-5 (-6) pairs, 3-10
cm long, (1.5-) 2.2-3.5
to
{-A) cm wide, narrowly obovate or elliptic, thick-chartaceous
hairy, hairs white,
subcoriaceous, base obtuse to rounded, apex obtuse to rounded, often with a short acumen
to 4 mm long, glabrous adaxially, densely hairy abaxially and the epidermis often not vis
ible, hairs >0.2-1.0 mm long, erect, tangled, pale (the leaflets appear white abaxially);
primary vein channelled adaxially, raised abaxially; secondary veins 10-12, ? plane adax
ially, slightly raised abaxially, pattern brochidodromous with the first 1-2 eucamptodro
mous, tertiary veins plane adaxially and abaxially. Panicles terminal, 20-35 cm long,
densely hairy at branch tips, becoming less hairy towards the base, the hairs red-brown
becoming white with age (or drying?), erect, tangled; bracts 2 mm long with pale brown
caducous hairs; bracteoles 1mm long with pale brown caducous hairs. Flowers 9-11 mm
long, subsessile. Calyx 5 mm long, hairy, the hairs tangled, pale brown; lobes 0.5-0.75
mm
long, acute to obtuse. Petals pink to purple; standard blade 10mm wide, 7 mm high,
claw 2.5 mm long; wing 7.5-8 mm long, 3.5 mm wide, claw 3 mm long, lamellate sculp
turing present but poorly developed; keel 6 mm long, 4 mm wide, claw 4 mm long. Sta
mens 10 mm long, the filaments united for the basal 4.5-5 mm, free for the distal 3.5-5
mm. Gynoecium
10-10.5 mm long, the upper surface of the ovary hairy, hairs red-brown,
? erect; stipe 3.5-4 mm long, ovary 3 mm long, style 3.5 mm long; ovules 2-3. Fruit
3.2-3.5 cm long, 2.8-2.9 cm high, 2.7-3 cm wide, ? globose, weighing ca. 20 g or less

2003
ANDIRA
51
when dry, green, drying brown, appearing smooth but minutely wrinkled (best seen with
suture slightly raised adaxially with a narrow groove, indistinct abax
lens or microscope);
remnant
tiny; composition of mesocarp and endocarp unknown. Chromosome
ially; stylar
number unknown. Figs. 3E, 4(iv), 5A, 16.
Phenology. Flowering May to June with occasional records in January and March.
in deciduous or semi-deciduous
Distribution
(Jalisco, Michoac?n);
(Fig. 17).Mexico
tropical forest, often by seasonal water courses.
Vernacular names. Manzanita
(Jalisco); garrapata (Michoac?n).
Examined.
Mexico.
Jalisco:
Additional
Specimens
Arroyo Chamela, Bullock 910 (MEXU); Mpio. La
Chamela,
Huerta, Est. de Biolog?a Chamela, Arroyo Chamela, Bullock 1313 (MEXU, MO); Estaci?n Biol?gica
A. Delgado
et al. 480 (MEXU); Mpio. La Huerta, 7 km al N del Puente Chamela, camino a Nacastillo,
pasando
et al. 4795 (MEXU); Mpio. La Huerta, Ma
Arroyo Colorado, por los terrenos de la U. de G., G. Flores-Franco
en el Lindero de la Universidad
de Guadala
(MEXU); Mpio. La Huerta, camino al E de Nacastillo,
W
1008
km
de
camino
585
3
Chamela,
Macuautitl?n,
(MEXU);
(MEXU);
Magallanes
jara, Magallanes
Arroyo
1044 (NY); Mpio. La Huerta, Chamela,
camino a Nacastillo, Magal
de Allende, Magallanes
Tomatl?n-Talpa
14 km N Santiago, Magallanes
1669
lanes 1383 (MEXU); Mpio. La Pe?a, camino Santiago Chacala-Colima,
gallanes
421
de Navidad, Magallanes
1710 (MEXU);
Tomatl?n, Las Jarillas, carretera Puerto Vallarta-Barra
(F); Mpio.
La Huerta, Est. de Biolog?a Chamela, Magallanes
2273 (US); Mpio. El Tuito, Las Gu?cimas,
camino El
a
La
4197
3556
camino
Tuito-Chacala,
Huerta,
Nacastillo,
(MEXU);
(MEXU);
Magallanes
Magallanes
Mpio.
4404 (NY); Chamela, L. A. P?rez 182 (MEXU); Chamela, L. A.
Mpio. La Huerta, Arroyo Chamela, Magallanes
Mpio.
P?rez & M. Hern?ndez
Vallar?a in direction
(MEXU, NY); Mpio. El Tuito, 14 km N road El Tuito-Puerto
2148 (MEXU); Mpio. Cuautitl?n,
3 km N Teqesquitl?n,
2950
Solis-Magallanes
4 km de Aquila, Guerrero 766 (MEXU); Mpio. Aquila,
(MEXU).?MICHOAC?N:
Mpio. Aquila, El Tenamaste,
Los Tanamastes,
Guerrero et al. 1262 (MEXU); Mpio. Chinicuila,
10 km NW Villa Victoria, Soto N??ez et al.
of El Cuale,
8187
848
Solis-Magallanes
(MEXU); Mpio.
Tiquicheo,
Paraje
la Escondida
el Lim?n,
Verduzco
s.n. (MEXU).
is clearly distinct from A. inermis in its abaxially densely hairy

Andira jaliscensis
leaflets. In specimens from Michoac?n
this indumentum is less dense than the material
from Jalisco, but they are separated from specimens of A. inermis from this area, which
have entirely glabrous leaflets, even as very young seedlings (e.g., Soto N??ez ?tal. 8016).
come from the southwestern part of
All the collections of A. jaliscensis from Michoac?n
this state, near Jalisco, except one, A. A. Verduzco s.n., which was collected near the bor
der with Guerrero and M?xico. This raises the possibility that A. jaliscensis may be dis
tributed across Michoac?n. Andira inermis and A. jaliscensis certainly appear to be sym
and future fieldwork should concentrate here to see whether any
patric in Michoac?n,
intermediate
3. Andira
forms exist.
Brazil.
Ducke, Bol. T?cn. Inst. Agron. N. 2: 30. 1944.?Type:
multistipula
Amazonas: S?o Paulo de Oliven?a, 2 Nov 1942, A. Ducke 1035 (lectotype, here
designated: RB!; isotypes: K! R! RB! US!).
Tree to 20 m
tall; buttresses absent; bark brown, smooth; slash pale orange-brown,

4
oxidizing darker, mm thick; twigs sparsely hairy, rapidly glabrescent, hairs red-brown,
appressed; older twigs buff, occasionally brown. Stipules to 5 cm long, 1.5 cm wide at the
base, persistent, with sparse red-brown appressed hairs, glabrescent and the hairs becom
ing paler; leaf axis 13-42 cm long; rhachis very sparsely hairy, hairs red-brown, rapidly
glabrescent; stipels 1.5-10 mm long, quite persistent; petiolules 3-5 mm long, indumen
tum like that of rhachis; leaflets in (5-) 6-9 pairs, 5.5-13.5 cm long, 2-4.5 cm wide, nar
rowly obovate, oblanceolate, elliptic to narrowly elliptic (proximal leaflets occasionally

52
FIG.
16. Andira
surface. E. Standard
cium. I. Gynoecium,
S. Magallanes
4404.)
VOLUME 64
A. Habit B. Abaxial
leaflet surface. C. Flower. D. Calyx, opened to show inner
jaliscensis.
petal, inner surface. F. Wing petal, outer surface. G. Keel petal, inner surface. H. Androe
also shown above in longitudinal
section. J. Fruit. (Based on: A-I, E. J. Lott 2544; J, J. A.

ANDIRA
2003
110
100
105
FIG.
narrowly
ovate
or
95
90
85
17. Distribution
lanceolate),
80
of Andira
chartaceous
75
70
jaliscensis,
53
65
60
A. multistipula,
to thick-chartaceous,
55
50
andA.
base
45
40
35
cubensis.
obtuse,
rarely
acute,
often slightly decurrent, apex acute to obtuse, with an acumen to 15 mm long, glabrous
(one specimen seen with very sparse erect hairs abaxially); primary vein channelled adax
ially; secondary veins 10-14, plane adaxially, ? plane to slightly raised abaxially, pattern
brochidodromous,
tertiary veins plane adaxially and abaxially. Panicles terminal and ax
illary (generally a large, terminal, ? conical inflorescence with some smaller axillary in
at the base, forming a single "attractive unit"), 15-45 cm long, hairy to
florescences
sparsely hairy, hairs red-brown, ? appressed, glabrescent at base; bracts 2.5-4 mm long,
with red-brown appressed hairs; pedicels to 0.5 mm long or flowers subsessile; bracteoles
1.5-2 mm long, with red-brown appressed hairs. Flowers 11-12.5 mm long. Calyx green
lobes, 4-5 mm long, with sparse red-brown appressed hairs
yellow with red wine-colored
to glabrous except at the base and margins of the lobes; lobes 0.4-0.7 mm long, 90? to ob
tuse. Petals pink to deep red-purple, the standard with a central white or white-yellow
marking; standard blade 9-9.5 mm wide, 8-9 mm high, claw 2 mm long; wing 8.5-9 mm
long, 3.2-3.5 mm, claw 3 mm long, lamellate sculpturing present but poorly developed;
keel 8-8.5 mm long, 3.5-4 mm wide, claw 3.5-4 mm long. Stamens 12 mm long, fila
ments white-pink, united for the basal 4.5-8.5 mm, free for the distal 2.5-5 mm, the vex
11-12 mm long, the upper surface of the ovary
illary stamen 8.5 mm long. Gynoecium
sparsely hairy, the hairs red-brown, ? erect, occasionally extending to the base of the style;
stipe 4-4.5 mm long, ovary reddish, 3.5-4 mm long, style 3^4.5 mm long; ovules 3. Fruit

54
VOLUME 64
cm long, 2-2.5 cm high, elongated, weighing ca. 20 g or less when dry, green, dry
dark brown to black, slightly wrinkled; stylar remnant barely discernible; meso
very
ing
mm thick, greyish brown, hard, granular; endocarp 1-1.5 mm thick, brown,
1.5-2
carp
2.5-3.5
Chromosome
fibrous.
woody,
number
unknown.
the year (March, April, June, October to

Phenology. Flowering
December).
Distribution
(Amazonas), Ecuador
(Fig. 17). Brazil (Amazonas, Acre), Colombia
(Napo, Sucumbios), Peru (Amazonas, Loreto, San Mart?n, Hu?nuco), Bolivia (Pando); in
seasonally flooded forest and on river banks, with occasional records from secondary
records throughout
forest.
Vernacular
names. Pisho, barbasco
caspi, arco caspi, arco sacha (Peru).
Amazonas:
Bocas del R?o Yari en el R?o Caquet?, Pab?n
La
de
los
Sachos
Cant?n, Pompeya, R?o Indillama, between its
(COL).
Joya
et al. 2145 (QCNE); Parque Nacional Yasun?, MAXUS
mouth at the Rio Napo and the MAXUS
road, Guai?o
road Km 24, by bridge over R?o Payamino, R. T. Pennington
& M. Aulestia 537 (E, K, QCA, QCNE); Parque
Nacional Yasun?, A?angu, SEF 8533 (QCA).?SUCUMBIOS:
Reserva Faunistica Cuyabeno, R?o Cuyabeno
above
Additional
& Mahecha
Examined.
Specimens
482
Ecuador.
Colombia.
?apo:
Peru. Amazonas:
Prov. Bagua, Dist. Imaza, R?o Cenepa
Laguna Grande, Balslev e? al. 97505 (QCA, QCNE).
& Gorham
1 km debajo de La Poza, bande del R?o San
162 (E, NY);
region, Comunidad Yamayakat, Hodges
49 (MO); R?o Santiago, 2 km atr?s de la Comunidad
925 (MO); Prov.
tiago, Huashika?
Caterpiza, Huashikai
de Yamayakat, R?o Mara??n,
Comunidad
Jaramillo 575 (E, MO).?
Bagua, Dist. Imaza, Regi?n del Mara??n,
Prov. Maynas,
Jacintillo, near Tingo Mar?a, Schunke 5725 (F,MO, US).?LORETO:
et al. 3102 (MO); Quebrada Cuninico,
Croat 17753 (MO); Quebrada Yanayac?,
GH, MEXU, MO, US); Prov. Coronel Portillo, Carretera Federico Basadre Km 99, Arboretum
HU?NUCO:
Itaya, Ayala
C. D?az
et al. 670
C. D?az
1315
Santa Rosa, R?o

Croat
20320
(F,
Von Humboldt,
orillas del R?o Pastaza, entre Rimachi

(F,MO, NY); Prov. Alto Amazonas,
y R?o Witoyacu,
(F,MO); Prov. Maynas, Moena Ca?o between Iquitos and R?o Itaya, Gentry et al. 15668 (F,MO);
Prov. Maynas, R?o Yarapa, Quebrada Fillico, Gr?ndez
1433 (E,MO); Prov. Maynas, Recreo, R?o Maniti, NE Iq
DE Dios: Prov. Manu, Parque Nacional Manu, Zona Reservada
1142 (F, MO).?MADRE
R?o
uitos, V?squez
Prov. Mariscal
Mart?n:
Manu, Cocha Otoronga, Foster & Vivar 13259 (F, K).?San
Caceres, Dtto. Tocache
camino
a Shunte,
Schunke 3859 (COL, GH, NY).

PANDO: Prov. Manuripi,
Bolivia.
along
from Chiba, Nee 31530 (MO).
Brazil. Acre: Mpio. Sena
(by air) downstream
Km 1-3 on road Sena Madureira
to Rio Branco, Prance e? al. 7968 (COL, GH).?AMAZONAS:
Rio
do RADAM,
Cavalcante
3164 (RB, ?B); Mpio. Nova Japura, between Tamandar? and Manquari,
Nuevo,
de Dios,
80 km NE
R?o Madre
Madureira,
Javar?, Pt. 2
Rio
Japura,
do Juami-Japura, Cid Ferreira et al. 7250
(K, NY); Mpio. Limoeiro, Est. Ecol?gica
(F, K, NY, US); Alto Solim?es, Rio Bora, afluente do Rio Jutai, L. Co?lho et al 394 (INPA); Rio Javar?, Esper
an?a, Ducke 1822 (F, GH, NY, US); ?cima do lago Jacapar?, left bank Rio Solim?es, H. C. de Lima & J. Guedes
Cid Ferreira
2731
& Lima 3611
(GH, K, NY); Alto
(GH, K, NY);
Rio
Solim?es,
Juru?, Pena
608
Mpio.
S?o Paulo
de Oliven?a,
S?o Luis do Pass?, H. C. de Lima
et al. 2767
(NY).
The flowers and fruit of A. multistipula are very similar to those of A. inermis; how
ever, it is clearly distinct because of its large, persistent stipules. Ducke (1944) claimed
that these stipules distinguish A. multistipula
from any other species of the entire Ama
zonian
flora.
All
the specimens from Peru have a calyx with appressed indumentum, in contrast to
the majority of the Brazilian material in which the calyx is glabrous, except at the base
and on the margins of the lobes. The calyx is sparsely hairy in two specimens from Brazil
(Prance et al. 7968, Acre; Cid Ferreira et al. 7250, Amazonas). The lack of other char
acters differentiating
these two groups and the presence
into two taxa.
splitting A. multistipula
Two
specimens
specimen with more
of intermediates
of the type collection (Ducke 1035) were

stipules was chosen as the lectotype.
argue against
found at RB. The

larger
'
55
ANDIRA
2003
4. Andira
Bentham, J. Proc. Linn. Soc., Bot. 4, Suppl. ("A Synopsis of the Dal
bergieae"): 121. 1860. Vouacapoua cubensis (Bentham) Kuntze, Revis, gen. pl.
Cuba. Oriente: Tinal de Nimanima, Apr 1844, J. Linden
1: 212. 1891.?Type:
2160 (holotype: K!; isotypes: G-2 sheets! K-2 sheets! NY-2 sheets!).
cubensis
Grisebach, PL wright. 179. 1860.?TYPE: Cuba. Oriente: "In

praeruptis prope Monte Verde," Jan-Jul 1865, C. Wright 1187 (lectotype, here
designated: GH!; isolectotypes: G-3 sheets! GH! K! MO! NY-3 sheets! US!).
Andira microcarpa
Tree to 20 m tall, crown often described as spreading; presence of buttresses unknown;

bark grey; slash unknown; twigs dark brown with pale raised lenticels, sparsely hairy,
mm long, to 1mm wide, very
glabrescent, hairs brown, ? appressed to ? erect. Stipules to 3
seen
at shoot apices); leaf axis 6-22 (-32 cm long on sterile branches);
early caducous (only
rhachis sparsely hairy, glabrescent, the hairs ? erect, brown, becoming or drying pale;
stipels minute, very early caducous; petiolules 1.5-5 mm
hairs ? erect, brown, becoming or drying pale; leaflets
2.3-5.5 cm wide, narrowly ovate, elliptic (rarely obovate
base obtuse, rounded to ? cordate, apex obtuse and often
often retuse (apex of the terminal leaflet tends to be more
long, sparsely hairy, glabrescent,

in 3-4 (-5) pairs, 5-13 cm long,
or broadly obovate), coriaceous,
slightly acuminate or rounded and
rounded), occasionally acute and
slightly acuminate with an acumen 7 (-10) mm long, glabrous adaxially, very sparsely hairy
abaxially, glabrescent, hairs pale (probably brown on young foliage), appressed, <0.2 mm
long; primary vein channelled adaxially, raised abaxially; secondary veins 9-12, plane
tertiary veins plane
adaxially, ? plane to slightly raised abaxially, pattern brochidodromous,
or
to
terminal, 8-30 cm long,
adaxially and plane
slightly raised abaxially. Panicles axillary
brown indumentum at branch tips, glabrescent towards base, hairs ? erect to ? appressed;
bracts 2 mm long, early caducous, densely covered with brown hairs; pedicels 1-1.5 mm
long; bracteoles 1mm long, very early caducous, densely covered with brown hairs. Flow
ers 7-8.5 mm long. Calyx 4 mm long, densely hairy, hairs ? appressed, brown; lobes to 0.2
long, very obtuse and shallow. Petals red to purple; standard blade 6-8 mm high, 5-7
mm wide, claw 2-2.5 mm long; wing 4.5-6 mm long, 2-2.5 mm wide, claw 2-3 mm long,
lamellate sculpturing present but poorly developed; keel 4-5 mm long, 2-2.5 mm wide,
claw 2-3 mm long. Stamens 5-7 mm long, filaments united for the basal 2-4 mm, free for
mm
the distal 2-3.5 mm. Gynoecium

and lower half of the style hairy
1.5-2.5 mm long, ovary 2-3 mm
2-2.8 cm high, 2.1-2.9 cm wide,
5.5-8.5 mm
long, upper and lower surfaces of the ovary

sides
(ovary
glabrous), hairs brown, ? appressed; stipe
long, style 1.5-3 mm long; ovules 1. Fruit 2.5^4 cm long,
elongated, the apex somewhat pointed, weighing ca. 20 g
or less when dry, drying dark brown spotted with pale brown (probably green when fresh
though reported on herbarium labels as red and as brown), smooth; stylar remnant insignif
icant; mesocarp 2-4 mm thick, pale brown, quite soft, granular; endocarp 1-2 mm thick,
brown, woody, hard, not very fibrous (note: these notes of fruit structure are based upon a
= 11.
single, possibly immature fruit). Chromosome number: n
Phenology. Flowering April to August, concentrated in June, July.
Distribution
(Fig. 17). Cuba; forest, swamp, river bank, wood pasture,
ca.
to
1000 m.
pine savanna;
Vernacular
name.
savanna and
Yaba.
Examined.
Additional
Specimens
Cuba. Habana:
Isla de Pinos, Los Indios, N. L. Britton et al. 14246
Isla de Pinos, Morton
9983 (US);
(NY); near Puentes Grandes, Bro. Le?n 3347 (NY); 2 km N Nueva Gerona,
Cerro Nueva Gerona, Isla de Pinos, Morton
10304
10121,10155
(US); Isla de Pinos, Herradura Beach, Morton

56
VOLUME 64
Villas:
Santa Clara towards
(US); near Nueva Gerona, Isla de Pinos, A. H. Curtiss 525 (G, MO, NY, PR).?Las
14 km NW Trinidad on road to Tope de
N. L. Britton & Cowell 10259 (NY); Trinidad Mountains,
Manicaragua,
in Trop. Botany 197 (GH); Cienfuegos,
between Rancho Luna and central Soledad,
Collantes, Harvard Course
et al. 379 (NY); Belmonte,
s.n. (US); W of Rio San Juan, crossing of road to Trinidad, Howard
Soledad,
Hodge
Jack 5664 (A).?
4999 (US); Limones,
Jack 5390 (GH); Belmonte,
Soledad, Cienfuegos,
N.L. Britton & Wilson 474 (NY).?Oriente:
S Matanzas,
Bayate, edge of Sabana Miranda,
Ekman 1925 (US); Sierra de ?ipe, R?o Piloto, edge of river, Ekman 9687 (F); lower valley of R?o Miel, Schafer
DEL R?O: San Diego de los Ba?os, Bro. L?on 4628 (GH, NY); near Vinales, Bro. Alain 4404
4349 (NY).?Pinar
Jack
Cienfuegos,
Matanzas:
hills
(GH); road to San Vicente,
21 km N Pinar del Rio, Harvard
Course
in Trop. Botany
10 (GH, NY).
(1860) considered A. cubensis to be a doubtful species, but, upon close in

spection, many characters separate it from the similar A. inermis, which appears to be ab
sent from Cuba. Andira cubensis has smaller flowers, amore densely hairy gynoecium, a
single ovule, and a fruit with amore pointed apex. It also tends to have only 3^4- (-5) pairs
of leaflets compared to 2-9 inA. inermis, which have a rounded to cordate base, whereas
Bentham
the leaflets are abaxially hairy when young.

those in A. inermis are obtuse. Additionally,
This Cuban endemic could not be included in the molecular phylogenetic
analysis,
because no recently collected leaf material was available. It seems most likely that its
closest relative is A. inermis, because of their morphological
similarities, and therefore A.
cubensis is placed with the species of the basally divergent clade I (Fig. 8). It is intrigu
ing that A. inermis, which is present throughout the Caribbean islands, appears to be ab
sent from Cuba.
R. T. Pennington,
Colombia.
sp. nov.?Type:
Antioquia:
1
G.
Ca?as
de
Jul
G. Galeano, R.
Gordas,
Lozano,
1984,
Mpio.
Boquer?n
Toyo,
Londo?o, R. Bernai & D. Restrepo 3963 (holotype: COL!; isotype: MO!).
5. Andira
taurotesticulata
Species bene distincta, quae dum florens Andira macrothyrsa et in fructu vel A. cori
?cea vel A. micrantha fortasse confusa possit. Ab A. macrothyrsa differt ovario in paginis
superioribus inferioribusque tantum (haud uniformiter ubique) dense pubescenti. Ab A.
cori?cea et A. micrantha fructibus non laevibus sed de sutura ad suturam costatis, foliolis
inferne sparse pubescentibus, non glabris bene insignis.
Tree to 25 m tall; buttresses absent; bark smooth; slash pale red brown, oxidizing
darker, 7 mm thick; twigs brown to grey-brown, bark cracking on older twigs, red-brown,
with ? appressed hairs, glabrescent. Stipules to 12 mm long, to 1 mm wide, with red
brown appressed hairs; leaf axis 9-30 cm long; rhachis sparsely hairy, glabrescent, hairs
red-brown, ? erect; stipels 1-3 mm long, persistent; petiolules 3-8 mm long, indumentum
like that of rhachis; leaflets in 3-5 (-10) pairs, 4.5-15 cm long, 2-7.3 cm wide, subcoria
ceous, elliptic, narrowly elliptic, or lanceolate to narrowly obovate, base obtuse to
rounded, apex obtuse to rounded (rarely acute), generally with an acumen to 15mm long,
mucronate becoming slightly retuse, glabrous adaxially, sparsely hairy abaxially, glabres
cent, hairs pale to red-brown, ? erect to ? appressed, to 1.0 mm long; primary vein chan
nelled adaxially; secondary veins 8-11 (-14), ? plane to slightly sunken adaxially, ? plane
to slightly raised abaxially (occasionally prominently raised), pattern eucamptodromous
or ? completely brochidodromous,
becoming brochidodromous
tertiary veins plane adax
ially and abaxially. Panicles axillary and terminal, 6-25 cm long, with red-brown ? ap
pressed hairs at branch tips, glabrescent towards the base; bracts 1.5-2 mm long, with red
brown ? appressed hairs; pedicels 0.5 mm long or flowers sessile; bracteoles 1-1.2 mm
long, indumentum
like that of bracts. Flowers
7-10 mm
long. Calyx

3-4 mm
long, hairy
ANDIRA
2003
57
to very sparsely hairy, hairs red-brown, ? appressed; lobes 0.2-0.5 mm long, obtuse, shal
low. Petals white, the standard marked with red; standard blade 7-10 mm wide, 5.5-7.5
mm high, claw 1.5-3 mm long; wing 4-6 mm long, 2-3.5 mm wide, claw 3-4 mm long,
sculpturing absent; keel 4-5 mm long, 2.5-3 mm wide, claw 4 mm long. Stamens 8-8.5
mm
long, filaments united for the basal 3.5-4.5 mm, free for the distal 2.5-3.5 mm, vex
illary stamen 6 mm long. Gynoecium 7-8.2 mm long, ovary hairy on upper and lower sur
faces with the stipe and sides of ovary sparsely hairy to only the upper and lower surfaces
of the ovary hairy, hairs ? appressed; stipe 3-3.5 mm long, ovary 2-4 mm long, style
1.2-1.5 mm long; ovule 1. Fruits 5.4-8 cm long, 4.4-8 cm high, 4-7 cm wide, ? globose,
100-300 g when dry, pale brown or dark brown marked with pale brown (both
weighing
fresh and dry), the dark brown layer falling off as fruits mature, rough, distinctly ribbed
from suture to suture; stipe 5-6 mm long; suture raised adaxially with a central groove,
less obvious below; stylar remnant not apparent; mesocarp 2-8 mm thick, brown, hard,
finely granular; endocarp 2-7 mm thick, pale, woody, fibrous with distinct ridges extend
ing into the mesocarp which mark the ribs seen on the fruit surface. (S. Diaz 1279: "fresh
fruit with pale brown exocarp, pale green mesocarp, cinnamon endocarp.") Chromosome
number unknown. Figs. 3C, 4B(iii), 18.
Phenology. Flowering May to November
Distribution
(Fig. 19). Panama (Dari?n, San Bias), Colombia

(Antioquia, Boyac?,
Meta, Quindio, Santander, Valle), Venezuela
(T?chira), Ecuador (Pichin
cha, Sucumbios, Napo); in montane forest and its transition to rain forest (700-2000 m)
in Colombia and Venezuela, and in lowland rain forest in Ecuador.
Vernacular name. Cojones de toro (Antioquia).
Cundinamarca,
Additional
La Escalera,
Specimens
Examined.
E of Tres Bocas, Kirkbride
Dari?n:
Panama.
& Duke
1315
Cuasi-Cana
trail between
San Blas:
(MO, NY).?
Croat 26024 (MEXU, MO).
Cerro Campamento
and
road, Km 12, in
El Llano-Carti
Control Project,
vicinity of Gorgas Lao Mosquito
Colombia.
16 km E San Rafael along Guatap?-San
Rafael road, Brant &
Antioquia:
Mpio. San Rafael,
Roldan 1533 (E, MO); Mpio. San Carlos, Correg. Alto Samana, vereda Miraflores,
finca "El Despesero,"
Calle
San Luis, right bank Rio Samana, vereda el Port?n, road to the autopista
jas et al. 8584 (HUA, NY); Mpio.
C?rdenas 2501 (COL, JAUM); Mpio. San Luis, Autopista Medell?n-Bogot?,
vereda la Jose
Medell?n-Bogot?,
C?rdenas
fina, quebrada laMariola,
Sector R?o Samana-R?o
l?n-Bogot?,
MO); Mpio.
Josefina,
& J. G. Ram?rez
Claro,
San Luis, Autopista Medell?n-Bogot?,

& Estrada
270 (JAUM, MO);
Cogollo
2743 (JAUM); Mpio.

San Luis, Autopista Medel
la vereda Tulip?n, Cogollo & Estrada
199A (JAUM,
Sector R?o Samana-R?o
Claro, camino hacia la vereda La
12 km from Au
San Luis, carretera hacia Aquitania,
Mpio.
camino
2741,
hacia
et al 3762 (JAUM); Mpio.

vereda la
topista Medell?n-Bogot?,
Cogollo
et al. 4306 (JAUM); Parque Nacional Natural "Las Orqu?deas" Sector
quebrada laMariola,
Cogollo
4145 (COL, JAUM); Mpio.
Calles, right bank of R?o Calles, Cogollo
Josefina,
vereda
la Josefina, Hoyos & J. J. Hern?ndez

286 (JAUM, MO); Mpio.
de la vereda la Josefina a la vereda la Holanda, Hoyos & J. J. Hern?ndez
334 (JAUM, MO); Mpio. San
vereda la Josefina, camino al Tulip?n, ca?o laMariola, Hoyos & J.J. Hern?n
Luis, Autopista Medell?n-Bogot?,
camino
dez 917
Cerro road, 11 km above Frontino around el Aeropuerto,

(JAUM, MO); Frontino-El
Luteyn & Lebr?n
7196 (COL, MO, NY, US); Mpio. San Luis, Quebrada "La Cristalina," J. G. Ram?rez & L?pez 950 (HUA,
de Providencia,
valle del R?o Anor?,
JAUM, MO), 1161 (COL, HUA, JAUM, MO); Mpio. Anori, Corregimiento
entre Dos Bocas y Anor?, Buenos Aires, 4 km from Providencia,
Soejarto 3954 (GH); Mpio.
Ituango, Km 9 of
Luteyn
road to vereda La Hundida

6444 (HUA, MEXU, MO).?Boyac?:
(WSW of Ituango), Zarucchi & Betancur
near P?ez, Espinal & Montenegro
1698 (COL).?CUNDINAMARCA:
Cerro Quinini, Idrobo 5337 (COL).?Meta:
vereda El Caney, Ca?o Agua Dulce, Forero
et al. 10205 (COL).?QuiNDlO:
road from Pijao to
Restrepo,
Paramo de Chili, Gentry et al. 65327 (COL, E, MO).?SANTANDER:
de Virol?n,
Mpio. Charal?, Corregimiento
Vereda El Reloj, S. D?az 1279 (COL).?VALLE:
frente al vivero de la secretar?a
Sevilla, Hacienda La Esmeralda,
de Agricultura
y Fomento, Cuadros 643 (US); Mpio. Sevilla, via aMorroazul,
quebrada La Raquelita, Devia 941
Venezuela.
T?CHIRA: Parque Cazadero,
16 km NW San Crist?bal, Liesner &
(MO).
Quebrada Cazadero,

58
18. Andira
FIG.
face.
Fruit. M.
cence),
taurotesticulata.
A. Habit.
B. Broad
leaflet. C. Narrow
F. Calyx, opened to show inner surface. G. Standard

K. Gynoecium,
I. Keel petal, inner surface. J. Androecium.
E. Flower.
leaflet. D. Abaxial
leaflet surface.
inner surface. H. Wing petal, outer sur

section. L.
shown above in longitudinal
in longitudinal
section, showing wall structure. (Based on: A
et al. 334; C, H. Cuadros
G. Lozano et al. 3963; B, D, S. Hoyos
Fruit
E-K,
petal,
also
VOLUME 64
(leaf), W. Devia 941; A (inflores

643; L-M, W. Devia 941.)

ANDIRA
2003
59
A. taurotesticulata
FIG.
19. Distribution
o? Andira
taurotesticulata.
et
11706 (MO); Cerro Las Minas,
S of main road from Santa Ana, 17 km SE Santa Ana, Steyermark
Guariglia
120019
al 119886
18 km SE Santa Ana, Steyermark
(MO); Cerro Las Minas,
slopes leading to Cerro Azul,
van der Werff & Ortiz 5553f 5556, 5576 (E, MO).
Ecuador. NAPO: Can
(MO); Distr. Lobatera, La Cazadora,
ton Orellana, Reserva El Chuncho,
1 km W Rio Payamino, NW Coca, Baker 7025 (NY); Cant?n Orellana, Via
1 km W R?o
de Zorros, Pozo de Jaguar I, Palacios
3204 (QCNE); Cant?n Orellana, Reserva El Chuncho,
& M. Aulestia
524, 525 (E, K, QCA, QCNE).?PICHINCHA:
Quito,
Payamino, NW Coca, R. T. Pennington
reserva Faun?stica
Reserva
Juan Manuel
Palacios
12274
Durini,
(QCNE).?SUCUMBIOS:
Lago Agrio,
Field Station, Neill 10214 (QCNE).
Laguna Grande, near Catholic University
Cuyabeno,
is the only species that grows inmontane forest (to 2200 m),
Andira taurotesticulata
and extending along the
in Colombia,
occurring in all three cordilleras of the Andes
to Venezuela.
It is also found in the Caribbean coastal ranges in
Cordillera Occidental
Panama (in Dari?n and San Bias), but in Ecuador it is found at lower altitudes (to 200 m).
It is clearly distinct by virtue of its small flowers, large fruit, and leaflets that are sparsely
hairy abaxially. There is wide variation in leaflet size, shape, and indumentum, in flower
size, and the extent of the gynoecial indumentum, which reflect this species' wide eco
logical and geographical range. The variation appears continuous, and the fruit are uni
form. The Ecuadorian specimens are somewhat disjunct from the rest of the range, but are
similar; the only difference is an endocarp with more abundant stone
morphologically
cells (Gemeinholzer
1995).
Flowering specimens of A. taurotesticulata might be confused with A. macrothyrsa.
The wing petals of A. taurotesticulata lack sculpturing, and the ovary is hairy on its upper
and lower surfaces with the sides of the ovary sparsely hairy, or only the upper and lower
surfaces of the ovary are hairy. In contrast, A. macrothyrsa has wing petal sculpturing, and
its ovary is uniformly densely hairy. In fruit, A. taurotesticulata might be confused with
A. cori?cea and A. micrantha,
the other rain forest species o? Andira with large fruit. The
fruits of A. taurotesticulata are distinctly ribbed from suture to suture, whereas the fruit of

60
VOLUME 64
the other species are relatively smooth and lack the distinct ribs. Moreover,
these other
in
Guianas
and
central
Brazil
have
restricted
distributions
the
(A. cori?cea)
(A. mi
species
are
A.
and
from
taurotesticulata.
crantha),
widely disjunct
The name for this species comes from its local name in Colombia:
"cojones de toro."
Brazil.
Ducke, Bol. T?cn. Inst. Agron. N. 2: 31. 1944.?Type:
macrothyrsa
Amazonas: Esperan?a, 22 Oct 1942, A. Ducke 1036 (holotype: RB!; isotypes: K!
R?RB-2
sheets! US!).
PERU.
Andira multistipula var. peruana N. F. Mattos, Loefgrenia 53: 2.1971.?TYPE:
16 Feb 1924, J. G. Kuhlmann 1505 (holotype: RB!; isotypes: RB
R?o Huall?ga,
6. Andira
2 sheets!).
Tree to 40 m tall, often a canopy emergent; buttresses slight; bark smooth and grey;
dark brown; slash pale red-brown, slowly oxidizing red-brown; twigs dark brown,
often with the outer bark splitting to reveal paler bark beneath, densely hairy, glabrescent,
wood
hairs brown, short, appressed; older twigs with pale, raised, elongated lenticels. Stipules
early caducous (not seen); leaf axis 10-33 (-40) cm long; rhachis with appressed brown
hairs, glabrescent, hairs short; stipels to 2 mm long, caducous; petiolules 3-9 mm long,
indumentum like that of rhachis; leaflets in 4-8 pairs, 4.5-11.5 cm long, 1.8?4 cm wide,
narrowly ovate, elliptic, or narrowly obovate (rarely narrowly elliptic), subcoriaceous,
base obtuse (rarely acute); apex obtuse, often retuse or with an acumen to 10 mm long,
glabrous adaxially, hairy to sparsely hairy abaxially, hairs short, appressed, red-brown,
<0.2 mm
long; primary vein channelled adaxially; secondary veins 8-12, plane to very
becoming
slightly raised adaxially, slightly raised abaxially, pattern eucamptodromous
the veins generally curving uniformly, tertiary veins plane adaxially
brochidodromous,
and plane to slightly raised abaxially. Panicles terminal or axillary (axillary inflorescences
are smaller), 8-30 cm long, with dense red-brown appressed hairs at branch tips, glabres
cent towards the base; bracts 2 mm long, early caducous, with dense appressed red-brown
hairs; pedicels to 1mm long or flowers subsessile; bracteoles 1mm long, early caducous,
indumentum as bracts. Flowers 9-11 mm long, unpleasantly scented. Calyx 4-5 mm long,
with dense, red-brown, appressed hairs; lobes 0.5-1.2 mm long, acute or obtuse, the apex
to greenish, the standard with a central red-brown to
acuminate. Petals white-yellow
carmine marking; standard blade 5.5-8 mm wide, 4.5-5 mm high, claw 2-2.5 mm long;
wing 4-5 mm long, 1.5-2 mm wide, claw 3-3.5 mm long, lamellate sculpturing present;
keel 2.5^ mm long, 1-1.5 mm wide, claw 3.2^1 mm long. Stamens 6-7 mm long, fila
ments united for the basal 2-4.5 mm, free for the distal 2-3 mm, the vexillary stamen 4.5
mm
6.5-8 mm long, the ovary with dense red-brown appressed hairs,

long. Gynoecium
the indumentum extending to the top of the stipe and base of the style; stipe 2.5-3.5 mm
long, ovary 2.5-3 mm long, style 1.5 mm long; ovules 1-2. Fruit 5-5.5 cm long, 3^1 cm
high and wide, elongated, weighing 20-30 g when dry, green and strongly scented, dry
ing smooth, brown to red-brown; mesocarp 2-2.5 mm thick, reddish brown, granular; en
docarp 1.5 mm thick, woody with coarse, pale fibers. Chromosome number unknown.
Phenology. Flowering October to February.

Distribution
(Fig. 20). Colombia (Choc?, Valle), Ecuador (Carchi, Napo,
Peru (Loreto), Brazil (Acre, Amazonas);
in terra firme and seasonally flooded
also caatinga (low forest on white sand). Andira macrothyrsa
is disjunct across
all the collections from Colombia and some from Ecuador are from the Pacific

Los R?os),
forest and
the Andes;
coast.
2003
id
ANDIRA 61
- -
A. macrothyrsa
A a.
chigorodensis
FIG. 20. Distribution
Vernacular
names. Uxi de morcego
of Andira macrothyrsa
and A. chigorodensis.
(Brazil, Acre);
casub guesmoteig
(Aw?; Ecuador,
Carchi).
Additional
Specimens
Examined.
Colombia.
Choc?:
de Utr?a, NE de
Parque Nacional
alrededor de la quebrada la Chunga, F. Garcia C.
Utria, entre alManglar
y la falda de lamonta?a
near mouth of Quebrada
443 (MO).?Valle:
Bah?a de Malaga,
la Sierpe, Gentry et al. 40434
Ecuador.
CARCHI: Tulcan Cant?n, Parroquia Tobar Donoso,
Sector Sabalera, Reserva
MO).
la Ense?ada
de
& Agualimpia
(COL, JAUM,
Ind?gena Aw?,
Cerro Centinela, Mon
1345 (MEXU, QCNE).?Los

R?OS: Quevedo Cant?n,
Tipaz et al. 1225 (E,MO, QCNE),
ta?as de Da, 10 km E Patricia Pilar, Palacios
& Freir? 7444 (QCNE).??apo:
Carretera Holl?n-Loreto
Km 75,
cerca de R?o Guantaraco, Neill et al. 8056 (K,MO, QCA, QCNE); Cant?n Orellana, Est. Exp.
INIAP-Payamino,
5 km NW de Coca, Palacios
ca. 1 km after R?o Guataraco
4818 (MO); Cant?n Loreto, Sector R?o Guataraco,
on road Holl?n-Loreto,
left side road, R. T. Pennington
& M. Aulestia 523 (E, K, QCA, QCNE).?SUCUMBIOS:
Reserva
1 km N Laguna Grande, hectare plot 1, Valencia 67876 (QCA).

Peru. Loreto:
Cuyabeno,
tourist camp, halfway between
Indiana and mouth of Napo, Gentry et al. 60683 (MO);
above mouth of Rio Napo, Gentry et al. 37168
Explorama Tourist Camp, R?o Amazonas
(MEXU,
Faunistica
Yanamono,
Yanamono,
Explorama
T. D. Pennington
& Ruiz 13550 (K); Prov. Requena,
Iquitos, Quistacocha,
Sapuena, Bagaz?n-R?o
Estaci?n Experimental
V?squez & Jaramillo 8745 (MO); Maynas Province,
IIAP,
Iquitos, Allpahuayo,
Brazil. ACRE: Mpio. Rio Branco, estrada para Porto Acre Km 13, ramai a es
V?squez et al. 13802 (E, MO).
3057 (NY); Mpio. M?ncio
querda, 4 km da estrada Colonia Cinco Mil, Cid Ferreira & Nelson
Lima, estrada
et al. 5247 (NY); Mpio. M?ncio Lima, estrada do Isac, 4 km from
para o lugar Bar?o, Km 30-32, Cid Ferreira
et al. 10950 (MEXU, NY); Sub-base do Cruzeiro do Sol, Marinho
27 (RADAM),
vie. Serra
city, Cid Ferreira
MO,
NY);
Ucayali,
Prance et al. 12388 (US); Cruzeiro do Sol, pr?ximo ao Acampamento

do Projeto RADAM, Rosa 668
Alto Solim?es, Mpio.
Sao Paulo de Oliven?a,
estrada do Bon?im,
(NY).?Amazonas:
logo ap?s a saida da
cidade de Sao Paulo de Oliven?a, H. C. de Lima et al. 2783 (K).
da Moa,

VOLUME 64
62
most similar to A. chigorodensis

is morphologically
Andira macrothyrsa
(see note
under that species, no. 7), but may also be confused with A. inermis subsp. inermis. Andira
macrothyrsa differs from A. inermis in the short, appressed hairs on the abaxial leaflet sur
faces and in its whitish to greenish flowers with a red-brown to carmine spot in the cen
ter of the standard petal. Andira inermis has glabrous leaflets and pink flowers with a
white spot on the standard.
In some specimens from Brazil (Acre and Amazonas: Cid Ferreira 5247, Rosa 668,
H. C. de Lima 2783) the leaflets are blunt-ended, almost oblong, matt brown abaxially,
with the secondary veins curving as they approach the margin. These characters provide
no basis for recognition of subspecific taxa, because states that are intermediate between
those of this group and the remaining specimens of A. macrothyrsa are found in two spec
imens?L. Marinho 27 (Brazil, Acre) and V?squez & Jaramillo 8745 (Peru, Loreto).
7. Andira
R. T Pennington,
Colombia.
sp. no v.?Type:
Antioquia: 21
chigorodensis
10 Jul 1981, J. Brand & A. Cogollo 114 (holo
km de Apartado hacia Chigorod?,
type: HUA!; isotypes: COL! MO!).
Floribus et fructibus A. macrothyrsae Ducke similis, praecipue foliis majoribus et

floribus minoribus, foliolis angustis cum indumento subter tenue, pallido.
Tree 15-20 m tall; presence of buttresses unknown; bark unknown; slash with clear
ex?date; twigs thick and stout, brown to dark brown, hairy, glabrescent, hairs brown, fine,
tangled. Stipules 7-8 mm long, to 5 mm wide, hairy, hairs pale red-brown, appressed; leaf
axis 37-55 cm long; rhachis sparsely hairy, glabrescent, hairs pale, fine, ? erect; stipels 1
mm long, broad, hairy, caducous; petiolules 4-7 mm long, indumentum like that of
rhachis; leaflets in 7-9 pairs, 7-14 cm long, 3^4 (-4.5, non-terminal leaflets) cm wide,
to subcoriaceous,
base obtuse to
thick-chartaceous
narrowly elliptic to lanceolate,
rounded; apex acute to obtuse, often with an acumen to 7 mm long, glabrous adaxially,
hairy abaxially, hairs 0.1-0.2 mm long, appressed, fine, pale with red-brown at base, ca
ducous and leaving the red-brown bases (older leaflets appear dotted red-brown abaxially
at 30x magnification);
primary vein channelled adaxially; secondary veins 12-13, plane
adaxially, slightly raised abaxially, pattern brochidodromous,
tertiary veins plane adaxi
to
and
raised
Panicles
ally
plane
slightly
abaxially.
axillary (appearing terminal, but in the
a
cm
axil of
terminal bud), 20-25
long with dense, pale brown to pale red-brown ? ap
pressed hairs, glabrescent towards the base; bracts 1.5 mm long, with dense pale red
brown, ? appressed hairs; bracteoles 0.75-1 mm long, indumentum like that of bracts.
Flowers 5-6 (-7) mm long, subsessile. Calyx 3-3.5 mm long, with dense ? appressed
hairs; lobes 0.3-0.5 mm long, obtuse, slightly acuminate. Petals yellow marked with red;
standard blade 6 mm wide, 4 mm high, claw 2 mm long; wing 3 mm long, 1mm wide,
claw 2.5 mm long, sculpturing absent; keel 2 mm long, 0.75-1 mm wide, claw 3 mm long.
Stamens 5 mm long, filaments united for the basal 1.5-2.5 mm, free for the distal 1.75-3
4.7 mm long, ovary hairy, hairs ? ap
mm, vexillary stamen 3.5 mm long. Gynoecium
mm
pressed, pale red-brown; stipe 1.5
long, ovary 2 mm long, style 1.2 mm long; ovules
2. Fruits 6 cm long, 4 cm high, 4 cm wide, elongated, weighing ca. 20 g or less when dry,
very dark brown, appearing smooth but minutely tuberculate (best seen with lens or mi
croscope), stipe 5 mm long; suture a fine raised line adaxially, not obvious below; stylar
remnant tiny; mesocarp 3.5 mm thick, brown, fibrous, granular; endocarp 2-3 mm thick,
woody,
fibrous. Chromosome
number unknown. Figs. 3B, 4B(ii),
13B, 21.

ANDIRA
2003
FIG. 21. Andira
A. Habit. B. Abaxial
leaflet surface. C. Flower. D. Calyx, opened to show
chigorodensis.
E. Standard petal, inner surface. F. Wing
petal, outer surface. G. Keel petal, inner surface.
H. Androecium.
I. Gynoecium,
section. J. Fruit. (Based on: A-I, J. Brand &
et al. 3579.)
A. Cogollo
114; J, E. Renteria
inner
surface.

64
Phenology.
vember,
Flowering
VOLUME 64
July (only one record); fruiting records from September, No
February.
January,
(Fig. 20). Colombia
Distribution
(Antioquia);
in rain forest; to 300 m.
Examined.
Colombia.
Turbo, 8 km de Carepa hacia Chigorod?,
Antioquia:
de Lomas Aisladas, Renteria et al. 3579
(HUA, JAUM, MO); Mpio. Turbo, corregimiento
Renteria 4634 (JAUM, MO); Mpio.
camino Malag?n-Chigorod?,
(JAUM-2
sheets), MO); Mpio. Chigorod?,
13 km W of El Jardin, Finca El Amparo, Zarucchi & D. C?rdenas 4215 (COL, HUA, MO).
C?ceres,
Additional
Specimens
Brand & Cogollo
215
Andira chigorodensis most closely resembles

leaves, 37-55 cm long, with narrowly elliptic or
5-6 (-7) mm long. The leaves of A. macrothyrsa
rowly ovate, elliptic, or narrowly obovate leaflets,
Andira
8. Andira
chigorodensis
is endemic
A. macrothyrsa.
It differs in its larger
lanceolate leaflets, and smaller flowers,
are 10-35(-40)
cm long and have nar
and the flowers are 9-11 mm
to the Chigorod?
region of Antioquia,
long.
Colombia.
galeottiana
Standley, Contr. U.S. Nati. Herb. 20(6): 217. 1919.?Type: Mex
ico. Veracruz: Catemaco, 26 Apr 1894, E. W. Nelson 424 (holotype: US!; iso
type: NY!).
Tree to 30 m tall; buttresses absent; bark scaling; slash pinkish brown; sapwood yel
lowish, heartwood reddish brown; twigs densely hairy when young, glabrescent, hairs red
brown, tangled, erect. Stipules to 5 mm long, to 1 mm wide, early caducous, with red
brown ? appressed hairs, glabrescent; leaf axis 10-30 cm long; rhachis hairy, glabrescent,
hairs red-brown, tangled, erect; stipels 2-3 mm long, caducous; petiolules 3-6 (-7) mm
long, hairy, hairs red-brown, tangled, erect; leaflets in 4-5 pairs, 4-12.5 cm long, 2.3-6
cm wide, elliptic, narrowly obovate to broadly obovate, subcoriaceous to coriaceous, base
obtuse (rarely acute or rounded), apex obtuse to rounded, often with an acumen to 5 mm
long, or retuse, glabrous adaxially, hairy abaxially, hairs erect, red-brown, >0.2-1.0 mm
long; primary vein channelled adaxially; secondary veins (6-) 8-12, slightly sunken adax
ially, raised abaxially, pattern brochidodromous,
tertiary veins plane adaxially and raised
abaxially. Panicles terminal or axillary, 15-50 cm long, with dense, tangled, erect pale
red-brown hairs at branch tips, glabrescent towards the base; bracts 1-1.5 mm long, with
? erect, early caducous brown hairs; pedicels 2-3 mm long; bracteoles 1mm long, indu
mentum like that of bracts. Flowers 13-17 mm long. Calyx 7-7.5 mm long, with ? erect,
pale brown hairs; lobes 1-1.5 mm long, acute or right-angled. Petals pink to violet; stan
11-14 mm high, 9-12 mm wide, claw 4 mm; wing 8-10 mm long, 4^.5 mm
wide,
long, lamellate sculpturing present; keel 8-10.5 mm long, 4^.5 mm
mm
wide, claw 5.5-6
long. Stamens 13-13.5 mm long, filaments united for the basal 7-8
mm, free for the distal 4-5.5 mm, vexillary stamen 10-10.5 mm long. Gynoecium
13.5-15 mm long, glabrous, or with 1-3 scattered hairs on the lower ovary surface; stipe
5.5-6 mm long, ovary 4^.5 mm long, style 3.5-5 mm long; ovules (1-) 2. Fruit 6-10 cm
dard blade
claw 5 mm
long, 4.3-8.5 cm high, 5-8.0 cm wide, elongated, weighing 40-125 g when dry, green be
coming dark brown or black when mature, drying brown to dark brown, smooth, flattened
adaxially and below with broad ribs often running diagonally from base to apex, suture a
broad groove adaxially; stylar remnant minute or absent; mesocarp 5-10 mm thick, dry
ing pale brown, soft, with spongelike texture and fibers extending from the endocarp; en
docarp 3-5 mm thick, buff, woody, fibrous, thickened along both sutures. Chromosome
number
unknown.

ANDIRA
2003
110
105
100
95
90
85
80
75
o? Andira
65
70
60
65
galeottiana
55
50
45
40
and A. verm?fuga.
Phenology. Flowering April to July.

Distribution
(Veracruz, Oaxaca, Tabasco, Chiapas); wet
(Fig. 22). Mexico
banks, seasonally flooded zones, also in non-inundated forest.
Vernacular
Representative
names.
Macayo/a
(Veracruz,
Oaxaca,
35
Tabasco);
lombrisero
sites, river
(Oaxaca).
M. E. Beccara
Chiapas: Tierra Colorada, Villahermosa,
s.n. (MEXU);
1 km SW junction to Emiliano Zapata, road Villahermosa-Es
7606 (MEXU); Mpio. Reforma,
(MEXU); La Arena, Palenque, F. Miranda
Laiana (Chinantla), Galeota
19498 (MEXU).?OAXACA:
3464 (US); Dist. Ju
SPECIMENS EXAMINED. Mexico.
s.n. (MEXU); Palenque, Fuentes

c?rcega, Grether & Quero 1579
Petrolero Cactus, Tenorio

H. M. Hernandez
& Torres 335 (MO); Mpio.
Santa Maria
chit?n, 6.6 km W Mago??
por camino a Guichieovi,
418 (MO); Mpio. Tuxtepec, Tuxtepec, Mart?nez Calder?n
1369 (NY,
Santa Maria, H. M. Hern?ndez
Cnimalapa,
& Sarukh?n 9067 (NY); Acatl?n de P?rez
MEXU, MO); Mpio. Tuxtepec, Ejido Benito Ju?rez, T. D. Pennington
Campo
(MEXU, US); Dist. Choapain, San Juan Lalane, Schuttes & Reko 852 (F, GH); Mpio. Tux
Figueroa, Salazars.n.
9451 (A, F, U); Tolosita, L. Williams
9612 (A,
tepec, Las Pinas, Sousa 3667 (MEXU, US); Ubero, L. Williams
R?o Rosario,
carretera E
Savanna Area,
Barlow
30/186D
F, U).?TABASCO:
(MEXU);
Huimanguillo
de Huimangillo
Ch?velas & P?rez J. ES892
hacia Francisco
(NY); Km 35 de la deviaci?n
Zapata-Tenonique,
3125 (A, F, NY); Mpio. Centro, Laguna Estancia
Rueda, C. Cowan et al 2294 (MO, NY); Balanc?n, Matuda
Novelo & Ramos
Vieja, cerca de Luis Gil P?rez, S Villahermosa,
La Venta, Laguna Yucateco,
edge of lake, near Rio Chicozapote,
on road to Balanc?n, T?llez & E. Mart?nez
junction to Tenosique
1770
(MEXU); Mpio. Huimanguillo,

Novelo & Ramos 1786 (MEXU);
905 (MEXU); 900 m from junction Chatle-El

on road to Balanc?n, T?llez & E. Mart?nez 948 (MEXU).?VERACRUZ:
Mpio. San Juan Lalana, Santi
207 km S de Catemaco,
16846
Jalahui, carretera a San Juan del R?o, Calzada
(MEXU);
Hueyapac
Triunfo,
ago
10 km N
6 km from

66
de Ocampo,
Cedillo
& Calzada
near Huaxpala,
Playa Vicente,
Catemaco, Faden et al. 76/125
Hanson
17 (F); San Lorenzo Tenochtitl?n,

et al ES-4301
Ch?velas
(MEXU);
Ch?velas
VOLUME 64
et al. ES-2396
(MEXU); Mpio.
Island, Lake
Agaltepec
Mpio. Catemaco,
1.5 km N Tabanca,
10 km E Catemaco,
(F, K); along E edge Lago Catemaco,
121 (MO);
Men?ndez
Catemaco,
(R?o Coscoapan),
Laguna de Sontecomap?n
& Nee 7637 (F, MO);

171 (GH, MEXU-2
de Sontecomap?n,
& G?mez-Pompa
sheets); 2 km Adelante
Nevling
Panga, R?o
ca. de C?rdoba,
840 (GH, MEXU,
& G?mez-Pompa
Chocam?n,
Teschaacu, Nevling
US); P. Est. Chocam?n,
de los Tuxtlas, R?o Tecesapa,
al E de Seteapan, Sousa 3267 (MEXU); Mpio.
Salazar s.n. (MEXU); Region
Barra
Catemaco,
a 2 km de Coyame
Lago Catemaco,
13407
(MEXU); Fortuno,
grove, Wing 25 (GH).
Coatzacoalcos
sobre camino
river, Williams
8483
a Tebanca,
(F, US);
orilla de Lago Catemaco,

Sousa et al.
on
Tenochtitl?n
village
path to orange
and A. verm?fuga are

their wide geographical
separation, A. galeottiana
Despite
one
site.
and flo
differ
restriction
their
related;
by only
closely
plastomes
Vegetatively
are
A.
two
of
with
its
the
but
the
fruit
galeottiana,
large
species
remarkably similar,
rally,
spongy mesocarp, is unique in the genus. These fruits, especially when
is dry and filled with air gaps, appear well adapted to dispersal by water, as
suggested by Pennington and Sarukh?n (1968). Rovirosa (1890) reported these fruits as
floating in the Rio Grijalva and its tributaries, and Mario Sousa (pers. comm.) reports that
fruit are regularly washed up on beaches in the Gulf of Campeche. Yet, water is unlikely
to be the exclusive means of dispersal, because when fresh, the mesocarp is fleshy, indi
soft and somewhat
themesocarp
cating probable dispersal by vertebrates.

Andira galeottiana has hard timber, which is used in the construction of houses,
bridges, and railway sleepers. Schuhes (herb, label, Schuhes 852) reports use as a ver
mifuge and to treat fungal diseases of the skin, Salazar (herb, label, unnumbered collec
tion) reports an extract of the bark used as a febrifuge, and Antonio and Heinrich (herb,
label, Antonio & Heinrich GUI224) report "fruits" (presumably seeds) used as a rat poi
son when mixed with maize flour.
9. Andira
verm?fuga (Martius) Bentham, Comm. legum. gen. 44. 1837. Geoffroea ver
m?fuga Martius in Spix andMartius, Reise Bras. 2: 788. 1828. Vouacapoua ver
Brazil. Minas
m?fuga (Martius) Kuntze, Revis, gen. pi. 1: 212. 1891.?Type:
s.n.
Gerais: "Inter fr?tices ad Salgado, et in campis editis Taboleiro," Martius
(holotype: M!).
spinulosa Martius in Spix and Martius, Reise Bras. 2: 788. 1828. Andira
spinulosa (Martius) Bentham, Comm. legum. gen. 44. 1837. Vouacapoua spinu
losa (Martius) Lyons, PL nam. 396. 1900.?Type:
Brazil. Minas Gerais: "in
Geoffroea
prov. Minarum campis editis siccis ad Contendas," Martius s.n. (holotype: M!).
Andira paniculata Bentham, Comm. legum. gen. 45. 1837. Vouacapoua paniculata
Brazil.
"Barbacena et
(Bentham) Kuntze, Revis, gen. pl. 1: 212. 1891.?Type:
s.n.
Ponte d'Erva," Pohl
(holotype: K!; isotypes: K! M, PR!, photos of M iso
type: G! GH!).
Andira kuhlmannii N. F.Mattos,
S?o Paulo:
Loefgrenia 40: 2. 1970.?TYPE: Brazil.
Moji-Mirim, margem da rodovia para Campinas, pr?ximo ? entrada da Fazenda
Holambra, M. Kuhlmann 3945 (holotype: HB; isotypes: K! SP).
to 12 (-25) m tall with spreading crown, occasionally

a shrub, able to root
or
buttresses
bark
with
absent;
thick,
very slight
sprout;
rough
deep vertical fissures;
slash pale brown to red-brown to orange-brown,
with
red ex?date; wood
generally
to
often
the hairs ?
buff/cream;
swollen, densely
twigs
sparsely hairy, glabrescent,
Tree

2003
ANDIRA
67
appressed, red-brown; bark cracking on older twigs; lenticels absent. Stipules to 5 mm

long, to 1mm wide, caducous; leaf axis 6-30 cm long; rhachis brown to red-brown,
hairy to sparsely hairy, glabrescent, hairs pale brown, erect; stipels 1-2 mm long; peti
olules 2-5 mm long; leaflets (2-) 3-5 (-6) pairs, 4-11 cm long, 1.8-6.6 cm wide,
broadly elliptic, elliptic, broadly obovate (rarely ovate, broadly ovate, suborbiculate, or
to coriaceous, dark green, shiny adaxially,
biculate to narrowly obovate), subcoriaceous
matt abaxially with red-brown indumentum, base obtuse to rounded (rarely acute), apex
obtuse to retuse (rarely acute or emarginate), glabrous adaxially, hairy to sparsely hairy
abaxially, hairs erect, pale red-brown, >0.2-1.0 mm long; primary vein channelled
veins 5-11, slightly sunken adaxially, raised abaxially, pattern eu
brochidodromous
brochidodro
(occasionally
completely
camptodromous
becoming
Panicles
terminal
veins
and
raised
(more
mous),
tertiary
plane adaxially
abaxially.
to
rarely axillary), densely covered red-brown erect hairs at branch tips, glabrescent
secondary
adaxially;
wards
the
base;
bracts
narrow,
caducous,
ca.
1.5-2
mm
long,
with
red-brown,
ap
pressed to erect hairs; pedicels 2-A mm long; bracteoles narrow, caducous, 1mm long,
indumentum like that of bracts. Flowers 12.5-18 mm long. Calyx 6-7 mm long, brown
to deep purple-brown,
sparsely hairy, hairs red-brown to pale red-brown, ? appressed;
lobes 1-2 mm long, acute to obtuse. Petals pink to purple; standard blade 9-15 mm
9-12 mm high, claw 3.5-6 mm long; wing 8-10 mm long, 3.5-6 mm wide, claw
mm long, lamellate sculpturing present; keel 7-10 mm long, 3.5-5 mm wide,
claw 4-6.5 mm long. Stamens 12-16 mm long, filaments united for the basal 5-9.5 mm,
free for the distal 3-7 mm, vexillary stamen 8-11 mm long. Gynoecium
12-15 mm long,
ovary glabrous or the upper and lower surfaces sparsely hairy, hairs ? erect, pale with
wide,
4-6.5
bases; stipe 4-6.5 mm long, ovary 3-5.5 mm long, style 2.5-4.5 mm long;
(2-) 4-6. Fruits 2.4-4 cm long, 1.2-2.5 cm high, 1.7-2.5 cm wide, elongated,
ca. 20 g or less when dry, green, strongly scented, drying brown (more rarely
weighing
red-brown or dark brown), wrinkled, suture a fine raised line or indiscernible adaxially,
indiscernible abaxially; stylar remnant tiny; mesocarp 0.5-2 mm thick, fibrous, often
drying with oily granules; endocarp 1-3 mm thick, pale red-brown, woody, fibrous.
red-brown
ovules
Chromosome
number unknown.
Fig. 4A.
Phenology. Flowering April to October, but the great majority of records from Au
gust and September (occasional records throughout the year).
Distribution
(Fig. 22). Peru (Pasco), Bolivia (Pando, Santa Cruz), Brazil (Acre, Ama
zonas, Bahia, Cear?, Distrito Federal, Goi?s, Maranh?o, Mato Grosso, Minas
Piau?, R?ndonia, Sao Paulo); in cerrado and gallery forest.
Vernacular names. Angelim preto, mata barata, angelim branco (Brazil).
Representative
Hartshorn
et al. 2918
Specimens
Examined.
Peru.
Pasco:
PANDO: Manuripi,
Bolivia.
(MO).
Prov. Chiquitos,
Serran?a de Santiago,
Gerais,
Cerro de Pasco, central selva, Palcazu Valley,

35 km N Puerto America,
A. Jardim 1050 (K,
near Santiago de Chiquitos,
5 km ENE Santiago,
Cruz:
MO).?Santa
Serran?a de Santiago, Santiago de Chiquitos,
Daly et al. 6272 (NY); Prov. Chiquitos,
near mouth of Rio Macauhan
mani & A. Jardim 1291 (MEXU).
Brazil. Acre:
camino
a la cueva, Ma
of Rio Yaco),
150 km SE Hu
(A, F, U, US).?AMAZONAS:
Mpio. Humait?, Janssen 139 (K); Transamaz?nica,
Sanaiotti
287 (E, INPA).?Bah?a:
mait?, 22 km E Bodoc?,
Mpio. Gentio do Ouro, ca. 6 km after Santo Ina
cio on Xique-Xique-Gentio
do Ouro road, R. T. Pennington
& Brito 256, 257, 258, 259 (CEPEC, FHO, K);
& Brito 265 (CEPEC, FHO, K); Mpio. tt>oti
Mpio. Barreiras, ca. 2 km along road to airport, R. T. Pennington
Krukoff
(tributary
5483
rama, rd. Ibotirama-Seabra
& Brito 270, 271, 272, 273 (CEPEC, FHO, K);

(BR-242) Km 8, R. T. Pennington
et al. 20 (?B).?Cear?:
Fazenda
Serra de Ibiapaba, Campo Grande,
Correntina,
Jatob?, Rezenda
880 (F).?DISTRITO
FEDERAL: Bacia do Rio S?o Bartolomeu,
et al. 5450 (US); Chapada
Dahlgren
Heringer
da Contagem,
Irwin et al. 7954 (MO, US); Fazenda Agua Limpa, near Vargem Bonita,
18 km SSW Brasilia
Mpio.

68
et al 194 (UB); Chapada

Alvarenga
Fazenda Breij?o,
(NY); Chapada dos Veadeiros,
et al. 199 (E, UB); Mpio. Goi?s, S?o Joaquim, Gibbs et al. 2713 (K, NY); Mpio. Goi?s, Serra
Bridgewater
ca. 38 km NE Formosa,
Irwin et al. 15191 (F,
Dourada, Gibbs 2782 (K, NY, UB); near C?rrego Estrema,
W
12
km
of
rd
from
of Presidente
Presidente
MEXU,
NY); Mpio.
Kennedy,
village
BR-153-Itapor?,
TV
tower, Ratter
dos Veadeiros,
et al 3595 (E, F, K, UB).?Goi?s:

11 km E Cavalcante,
W. R. Anderson
VOLUME 64
Mpio.
7294
Vian?polis,
e? al. 8298
Fe?nho, Plowman
(GH, NY, US).?MARANH?O:
along Riber?o
602 (NY); Mpio. Lor?to, between
do Graja?, Riacho Grande, L. Duar?e & Castellanos
R?os Balsas and Parna?ba, 35 km S Lor?to, Eiten & Eiten 4513 (NY); Mpio. Carolina, Transamazonian
High
and BR-010,
Pedra Caida, 35 km N Carolina,
Serra da Baleia, E.L. Taylor et al. E1248
way, BR-230
(K, NY,
et al. 6416
Serra do Vi?va,
12 km da Cidade, Cid Ferreira
Grosso:
Santa Terezinha,
US).?Mato
Mpio.
Kennedy,
arredores
Fazenda
Primavera
de Bar?o
152 km NNE Xavantina,

155 (K); Mpio. Barra do Gar?as,
4.5
(NY, US); Brotas, 30 mi NE Cuiab?, Collenette
km S C?rrego Tangur?, Eiten & Ei?en 9849 (NY, US); watershed
between Amazonas
and Rio Paraguaya,
1-10
56241
10 km W S?o Felix, Richards 6482
km W Alto Araguaia,
Brasilia-Acre
(NY, UB);
highway, Maguire
transect 2,
Fazenda
road MT-170,
track west,
3 km N of junction with BR-364,
Itamarati,
(K, NY, UB);
Porto Buriti, margem
G?rais:
449, 460, 461 (E, INPA).?Minas
esquerda do Rio Paracatu, de F.
7120 (NY, UB); Cristalina,
140 (K); Patos de Minas, A. P. Duarte 3282 (NY); Felixlandia,
Heringer
17740 (US); Mpio. Unai, margem da rod. BR-251,
Paracatu, barranco do Rodo via, Heringer
perto da entrada
s.n. (RB).?ROND?NIA:
904 (SP).?PlAU?:
Alencar
Rd. Vilhena-Col
da Fazenda Paulista, B. A. S. Pereira
Sanaiotti
Almeida
orado,
Fazenda
20 km from Vilhena,
Joana, Felippe
to Piracicaba,
road S?o Manuel
221174
(UB); Mpio.
et al 2810 (GH, NY, US).?S?O

PAULO: Araraquara, Usina Tamoio,
14 km E de S?o Manuel,
18 km N Botucatu,
(US); Mpio. Botucatu,
along
near Est. "13 Maio" of the old railroad, /. S. Gottsberger
& C. J. Campos
12
Ja?, Leit?o Filho et al. 12927 (US).
Zarucchi
Santa
Brotas,
94
Andira verm?fuga ismost closely related toA. galeottiana

(no. 8; see notes under that
for
differential
characters).
species
Specimens of A. verm?fuga may be divided into two groups, one characterized by
thick, swollen twigs, the other by thin, leptocaul twigs, which are more usual in Andira.
two forms overlap somewhat geographically,
though the majority of the thin
are from the northeast of the range of A. verm?fuga (Bahia and
twigged collections
Maranh?o). No differences can be found to separate these groups in leaf, flower, or fruit
in which the twigs appear intermediate (e.g.,
characters, and there are some collections
R. T. Pennington
249, 258, C. A. Cid 6416). This indicates little basis for splitting A.
verm?fuga subspecifically. Other specimens show new growth from swollen twigs to be
These
slender and leptocaul, suggesting that the underlying basis for the differences may be
related, e.g., repeated burning may induce formation of thick twigs.
environmentally
This is corroborated by personal observations
of a population
of A. verm?fuga (at
a
Fazenda Agua Limpa, Distrito Federal, Brazil), which showed
gradation from tall
trees with slender twigs in gallery forest to stunted trees with thick twigs in open cer
rado only 50 m away. Andira kuhlmannii N. F. Mattos appears to be based upon a spec
imen from a large, gallery forest individual of A. verm?fuga, and the name is here placed
in synonymy.
Several interesting collections from the northwestern edge of the range are assigned
to A. verm?fuga. Zarucchi et al. 2810 from Rond?nia appears in all vegetative and fruit
characters to be A. verm?fuga, but the notes indicate that the twigs are hollow and house
ants. The information given is insufficient to determine whether this is only a casual as
et al. 8719 (Rond?nia, Brazil), Krukoff 5843 (Acre, Brazil) and
sociation. Prance
et al. 2918 (Pasco, Peru) all represent large trees to 25 m tall, but with flow
Hartshorn
ers 12.5-14 mm long, which is somewhat small for A. verm?fuga. They were collected
"transition forest" (sensu Ratter, 1992) on the edge of Amazonia,
extending
of
A. verm?fuga into this ecosystem. Prado and Gibbs (1993) and Pennington
the range
et al. (2000) give several examples of this type of distribution of a woody species, which
in seasonal

ANDIRA
2003
69
is principally found in dry vegetation but also on the fringes of Amazonia. One exam
ple is the legume Poeppigia procera Presl. It should be noted that A. verm?fuga has also
savanna of Humait?
been collected
in the isolated Amazonian
(A. Jans s en 139,
Sanaiotti 287).
Martius (1828) published the two names G. verm?fuga and G. spinulosa on the same
page of the same publication. Bentham (1860) cited G. spinulosa
verm?fuga, which therefore takes priority.
in the synonymy
of A.
ex Bentham, Comm. legum. gen. 45. 1837. Vouacapoua hu

Brazil. Minas
(Bentham) Kuntze, Revis, gen. pi. 1: 212. 1891.?Type:
Gerais: "ad Chapado do Par?nan," Martius s.n. (holotype: M!).
Andira laurifolia Bentham, Comm. legum. gen. 45. 1837. Vouacapoua
laurifolia
Brazil. Minas Gerais:
(Bentham) Kuntze, Revis, gen. pl. 1: 212. 1891.?Type:
Fa?ado, Pohl s.n. (holotype: K!; isotype: W, photo: G!).
10. Andira
humilis Martius
milis
Andira pauciflora Bentham, Comm. legum. gen. 45. 1837.?Type:

Gerais: "Ad Barbacena et Ponte d'Erva," Pohl s.n. (lectotype,
Brazil.
Minas
here designated:
K!).
Andira humilis var. cordata N. F. Mattos, Loefgrenia 40: 3. 1970.?TYPE:
S?o Paulo: Itahy, 10 Dec 1929, /. /. de Lima s.n. (holotype: RB!).
Brazil.
suffrutex forming mats up to 10 m in diameter, aerial shoots to 50 cm tall

Geoxylic
(occasionally shrublike, to 2 m tall); bark brown, Assuring vertically; slash pale brown to
pale red-brown with some clear ex?date that oxidizes red; twigs brown to buff, bark crack
ing on older twigs, with short, red-brown appressed hairs, glabrescent. Stipules to 7 mm
long, to 1 mm wide, with short, red-brown appressed hairs; leaf axis 9^45 cm long;
rhachis sparsely to very sparsely hairy, glabrescent, hairs short, red-brown, appressed;
stipels 1^1 mm long; petiolules 1-5 mm long, indumentum like that of rhachis; leaflets in
4-7 pairs, 4-12.5 cm long, 1.3-4 cm wide, elliptic, narrowly elliptic (rarely narrowly obo
to coriaceous, shiny, dark green
vate, oblanceolate, ovate to lanceolate), subcoriaceous
matt
and
the
venation
base
obtuse, rounded (occasionally
adaxially, paler
abaxially,
paler,
truncate or ? cordate), apex obtuse to rounded (occasionally
acute), generally retuse,
glabrous adaxially, glabrous to sparsely hairy abaxially, hairs minute (<0.2 mm long), pale
with a red-brown base, appressed; primary vein channelled adaxially; secondary veins
8-11, plane to slightly sunken adaxially, ? raised abaxially, pattern eucamptodromous be
coming brochidodromous,
tertiary veins plane adaxially and abaxially. Panicles terminal,
10-30 cm long, hairy to very sparsely hairy at branch tips, glabrescent towards the base,
hairs red-brown, short, ? appressed; bracts 1.5 mm long, with sparse, red-brown, short,
appressed hairs; pedicels 1^ mm long; bracteoles 0.75 mm long, indumentum like that
of bracts. Flowers 14-16 (-19) mm long. Calyx 6-8 mm long, deep purple, sparsely hairy
to glabrous except on margins of lobes, hairs red-brown to golden-brown,
short, ap
pressed; lobes 0.5-1.7 mm long, obtuse to acute. Petals violet to purple; standard blade
10-12 mm wide, 9-12 mm high, claw 3.5-4.5 (-6) mm long; wing 7.5-10 (-12) mm long,
4-5 mm wide, claw 4-6 mm long, lamellate sculpturing present; keel 7-9 (-12) mm long,
4-5 (-6) mm wide, claw 4-6.5 (-8) mm long. Stamens 10-13 (-16) mm long, filaments
united for the basal 5-9 mm, free for the distal 3-6 mm, vexillary stamen 8.5-10 (-12)
mm long. Gynoecium
11.5-16 mm long, glabrous; stipe 4.5-6.5
(-7.5) mm long, ovary
3.5-5.5 (-6.5) mm long, style 3-4 mm long; ovules 4-5 (-7). Fruits 2.8-5.3 cm long,
ca. 20 g or less when dry, green
2-2.8 cm high, 2-2.8 cm wide, elongated, weighing

70
of Andira
VOLUME 64
humilis.
turning yellowish when ripe,

long, but many fruits falling
groove, less obvious below;
soft when ripe, fibrous with
sweet smelling, drying wrinkled, dark brown; stipe 5-10 mm

off without a stipe; suture raised adaxially with a central
stylar scar tiny; mesocarp 0.5-2 mm thick, pale yellow and
oily granules; endocarp 0.5-1 mm thick, brown, woody, fi
brous.
unknown.
Chromosome
number
Phenology. Flowering May to November.

Distribution
(Fig. 23). Brazil (Bahia, Distrito Federal, Maranh?o, Mato Grosso, Mato
Grosso do Sul, Minas Gerais, Para, Paran?, Pernambuco, Rio Grande do Norte, Sao
Paulo), Paraguay (Amambay, Caaguaz?, San Pedro); in cerrado and other open, fire-prone
vegetation.
Vernacular names. Mata barata, angelim

h?o); manga do campo (S?o Paulo).
(Minas Gerais);
angelim
rasteiro
(Maran
ca. 100 km WSW

Brazil.
Specimens.
Bah?a: Espig?o Mestre,
Barreiras, W. R. An
26967 (K); Mpio. Morro do Chap?u,
(NY); Ibiquara, ca. 25 km N Barra da Estiva, Harley
Serra do Trombador,
6 km S Morro
do Chap?u, Irwin et al. 32461
Feira de Santana,
(MO, NY, US); Mpio.
& Brito 239, 240, 241, 242 (CEPEC, FHO, K); Mpio. Angical,
road Bar
Campus da UEFS, R. T. Pennington
Km 35, R. T. Pennington
& Brito 266, 267, 268, 269 (CEPEC, FHO, K).?
reiras-Ibotirama
(BR-242)
Representative
derson
et al. 36865

2003
71
ANDIRA
(SP); 30 km S Brasilia on road to Belo Horizonte,

ca. 20 km by road N Alto Para?so, W. R.
(NY).?Goi?S:
Chapada
Anderson
6231 (NY); Serra Geral da Paran?, 1 km E S?o Jo?o da Alian?a, W. R. Anderson
7895 (NY); Serra
on road to Jatai, Irwin & Soderstrom
do Caiap?, ca. 60 km S Caiaponia
7436B (NY); Serra dos Cristais,
3 km
W Cristalina,
Irwin et al. 9824 (NY); Chapada dos Veadeiros,
Rio das Almas, 47 km from Alto Para?so de
Distrito
Fazenda
Federal:
Irwin & Soderstrom
Goi?s
509.?Maranh?O:
Limpa, Heringer
Para?so
16202
dos Veadeiros,
on road to Teresina
5 km from Alto
Agua
5662
de Goi?s,
de Goi?s
Mpio. Lor?to,
& L. T. Eiten 5457
et al 499, 500, 501, 502, 503; Chapada dos Veadeiros,

R. T. Pennington
on road to the National
et al. 505, 506, 507, 508,
Park, R. T. Pennington
45-50 km S Lor?to on road between Santa Estev?o and Fazenda Santa Es
GROSSO: 15-120 km beyond

(NY); Grajah?, Lisboa s.n. (RB).?MATO
s.n.
to
et
Anna
al.
road
56291
Santa
das Chapadas, Malme
Cuiab?,
UB,
(NY,
US);
Araguaia,
Maguire
do Sul: road between Aquiduana
e? al 354 (E, UB); on road
and Miranda, Bridgewater
Grosso
(F).?Mato
tev?o, G. Eiten
Alto
to Porto Murtinho
from Bonito,
e? al
Gerais:
363, 364 (E, UB).?Minas
e?
do
Cavalcan?i
al s.n.
Serra
Cabrai,
Lagoa
e? al. 15968 (US); 27 km SW
Fazenda Geriza, Felippe 36 (US); Mpio. Lavras, Heringer
to Goveia,
Irwin e? al 22146a
(F,MO, NY, RB, UB, US); Morro das Pedras, ca. 40 km
e? al. 25699 (MO, NY, US); between Sete Lagoas and C?rvelo, R. S. Sanios & Casiellanos
Santa, B?rrelo
Serra do Cachimbo,
Dombrowski
9799
5491
Bridgewaler
(F); Joaquim
road Cuiab?-Santar?m,
(US); Rio
das Cinzas,
Felicio,
Km
Barra
856, Prance
do Perdizes,
e? al
Haischbach
25053
Mpio.
Santa Luzia,
(SPF); Mpio. Caet?,

on road
Diamantina
NE Patrocino,
(US).?Paran?:
24071
Irwin
(K).?Para:
Jaguariahyva,
7200
(US); Mpio. Campo Mour?o,

19955 (F, NY).?PERNAMBUCO:
Haischbach
15930
(F, US); Mpio. Arapoti, Haischbach
Campo Mour?o,
Sitio de Lima, /. L. S. de Lima 01 (RB).?Rio
GRANDE DO NORTE: road Cear? Mirim-Touros,
Km 25, Bamps
5078 (NY); Natal, Moacyr
8 (MO).?ROND?NIA:
4 km from Vilhena,
Vieira e? al 622 (NY).?
Alvarenga
S?O PAULO: Morro
Pellado, Edwall 5650 (SP); Mpio. Moji-Gua?u,

Campos das Sete Lagoas, Faz. Campin
G. Eiten & L.T. Eiten 2363 (F, US); Mpio.
estrada Pirassu
inha, just N Rio Moji-Gua?u,
Pirassununga,
Forero 8278 (SP); Mpio.
S?o Carlos, N of city of S?o Carlos, de Freitas Campos 31 (UB, US);
nunga-Emas,
18 km N Botacatu,
14 km E S?o Manuel,
road near old rail
Mpio. Botacatu,
along S?o Manuel-Piracicaba
road station, Gottsburger
et al
Bicuda
Hernandes
2131
? margem
do rodovia municipal,
5 km de Vitoriana,
(UB); Mpio. Botacatu,
rodovia Avar?-Sao
/. Mattos
14530a
Manoel,
(SP); 37 km de Avare,
(SP);
Amambay:
de Cerro Cora, Fern?ndez
Casas & Molero
Paraguay.
Parque Nacional
between Caaguaz?
and Yh?, Fern?ndez
Casas & Molero
6362 (MO); entre Yh? y
1538
Sarapui, Yano 05 (SP).

4079 (NY).?CAAGUAZ?:
San Blas, Fern?ndez
Casas
km N Arroyo Guaranungua,
al. 1497 (MO); Dist. Lima,
734
3851 (NY); 3 km S Arroyo Taruma, Zardini & Vel?squez 25331 (K); 2

& Vel?zquez 25676 (MO).?SAN
PEDRO: 4 km S de R?o Verde, P?rez et
Estancia Carumbe, Pedersen
8510 (A, K); Alto Paraguay, Primavera, Woolston
& Molero
Zardini
(K, U).
Andira
humilis is easy to recognize by its geoxylic suffrutex growth form, which is

in
the
genus and was described in detail by Warming (1892). It appears that occa
unique
A.
humilis
sionally
(e.g., Amorim et al. 1822, R. T. Pennington 239, 246, 247, 266) may
form a shrub or tree, as occurs in other species of geoxylic suffrutices, such as Kielmey
era rubriflora (Guttiferae; J. A. Ratter, pers. comm). In the case of A. humilis, this aber
rant habit may be the result of the prolonged absence of fire or possibly due to hybridiza
tion with shrub or tree species (see discussion under "Intraspecific cpDNA polymorphism
and potential hybridization in Andira").
There is a distinct geographic partitioning of variation in the indumentum of the
calyx. Specimens with glabrous calyces are restricted to western Minas Gerais, S?o
Paran?, and Distrito Federal (Fig. 11), whereas specimens with sparsely
are
found to the northeast of these areas. I have not recognized these spec
hairy calyces
imens as subspecif?c taxa, because there are intermediates (particularly in Paraguay and
Paulo, Goi?s,
Distrito
Federal) and other specimens that demonstrate that the same plant can produce
two inflorescences
in which the flowers have calyces with widely different amounts of
indumentum.
Bentham (1837) published the three names A. humilis, A. laurifolia, and A. pauciflora
on the same page of the same publication. He later (1860, 1862)
placed A. pauciflora in

72
the synonymy of A. humilis, but maintained A. laurifolia. Handro

one species, A. humilis, and therefore this name takes priority.
R. T. Pennington, sp. nov.?Type:
11. Andira macrocarpa
Reserva ?tnica Huaorani, Maxus road and pipeline
23 Jul 1994, N. Pitman, A. Dik & C. Aulestia 631
E! MO!).
VOLUME 64
(1969) recognized
Ecuador.
construction
only
Napo: Aguarico,
project, Km 98,
(holotype: QCNE!;
isotypes:
Species insignis propter foli?la parva pilis longiusculis patulis obtecta et fructus mag
nos gravesque laeviusculos endocarpio lignoso fibroso. Foliis A. vermifugae & A. suri
namensi similis, praecipue fructibus majoribus differt. Foliis A. galeottianae similis, prae
mollibus,
duris, solidis (in A. galeottiana mesocarpiis
spongiosis)
cipue mesocarpiis
recedit.
Emergent tree more than 30 m tall, 1m in diameter; presence of buttresses unknown;

bark and slash unknown; twigs dark brown, with sparse, red-brown appressed hairs when
young, glabrescent, bark cracking vertically and flaking. Stipules early caducous (not
seen, probably small); leaf axis 6-12 cm long; rhachis dark red-brown, hairy, hairs red
brown, erect, >0.2-1.0 mm long; stipels small (ca. 0.5 mm long, but caducous, and per
haps longer on younger leaves); petiolules 4-6 mm long, indumentum like that of rhachis;
leaflets in 2-3 pairs, 5.5-9 cm long, 2.5^ cm wide (the lower leaflets often the smallest),
narrowly obovate (rarely elliptic), subcoriaceous, base obtuse to rounded, apex rounded,
often slightly retuse, glabrous adaxially, hairy abaxially, hairs long, erect, red-brown; pri
mary vein channelled adaxially; secondary veins 7-10, slightly sunken adaxially, promi
becoming brochidodromous,
tertiary
nently raised abaxially, pattern eucamptodromous
veins plane to slightly impressed adaxially and raised abaxially. Inflorescences and flow
ers not seen. Fruits 10-11 cm long, 7.5-8 cm high, 8 cm wide, elongated, weighing ca.
300 g when dry, flattened adaxially, drying very dark brown and slightly ridged, the sur
face smooth to the touch; stipe 5-8 mm long; suture obvious, sunken adaxially, promi
nently raised below; stylar remnant tiny or not apparent; mesocarp 4-18 mm thick, finely
granular with small fibers, drying very hard, brown; endocarp 4-15 mm thick, brown,
woody, fibrous, thickened along the sutures. Chromosome number unknown. Fig. 24.
Vernacular name. Incgamuncga (Huaorani; Ecuador).
is known only from the type collection gathered in rain forest in
Andira macrocarpa
Napo, Ecuador (Fig. 25). Although this species has never been collected in flower, its dis
tinctiveness merits description. No other species have this combination of small leaflets
with long, erect hairs on their undersurfaces, and a large, relatively smooth fruit with a
woody, fibrous endocarp. The leaflets resemble those of A. verm?fuga and A. galeottiana
in their shape, venation, and indumentum. Neither of these species is sympatric; more
over, A. verm?fuga has small fruit, and the large fruit of A. galeottiana has a unique meso
that is soft and somewhat spongy with numerous air gaps when dry. The
carp morphology
leaflets might also be confused with those of A. surinamensis, but differ in their erect in
dumentum. Andira surinamensis also has a small fruit. The other sympatric species with
large fruit is A. taurotesticulata, but the fruit of this species are distinctly ridged, and have
an endocarp composed principally of parenchyma and stone cells, whereas the endocarp
of A. macrocarpa comprises mostly woody fibers. The leaflets of A. taurotesticulata gen
erally are acuminate, whereas those of A. macrocarpa are rounded or slightly retuse at the
is more
apex, and the venation on the undersurfaces of the leaflets of A. macrocarpa
prominently
raised.

2003 ANDIRA 73
FIG. 24. Andira

Fruit
in section,
A. Habit. B. Adaxial
leaflet surface. C. Abaxial
macrocarpa.
wall structure. (Based on Pitman et al. 631.)
leaflet surface. D. Fruit. E.
showing
1994. The tree from which the type

I attempted to re-collect this species inDecember,
a
was
cut down because of concerns
road
and
had
been
collection
made grew alongside
over
must
fruit
about the damage that falling
500 g when fresh) might cause
(which
weigh
to passing vehicles.
surinamensis
(Bondt) Splitgerber ex Amshoff, Meded. Bot. Mus. Herb. Rijks
Univ. Utrecht 52: 60. 1939. Geoffroea surinamensis Bondt, Dissertatio medica
surinamensis, 8, figs 1-8. 1788. Geoffrea ob
inauguralis de cortice Geoffraeae
4:
Bot.
Mat.
Med.
46. 1812, nom superfl. Andira retusa ? suri
tusifolia Stokes,
namensis (Bondt) DC, Prodr. 2: 476. 1825. Vouacapoua surinamensis
(Bondt)
12. Andira

74
VOLUME64
Kuntze,
Revis,
gen. pi. 1: 212.
1891?TYPE:
SURINAME. Bondt
s.n. (holotype:
L!).
pubescens L. C. Richard, Actes Soc. Hist. Nat. Paris 1(1): 111. 1792. Ge
offroea retusa Poiret in Lamarck, Encycl. 8: 182. 1808, nom. superfl. Andira re
tusa (Poiret) DC, Prodr. 2(2): 475. 1825. Vouacapoua retusa (Poiret) Lyons, PL
nam. 396. 1900. ?Type:
French Guiana. Cayenne, L. C. Richard s.n. (lecto
Geoffroea
type, here designated: P!, isolectoype: P!).

Andira retusa var. oblonga Bentham, J. Proc. Linn. Soc, Bot. 4, Suppl. ("A Synop
Brazil.
121.1860.?Type:
Para: Santarem, Spruce 914
sis of the Dalbergieae"):
K!
K!
here
RB!).
designated:
isolectotypes:
(lectotype,
Andira surinamensis var. ovatifoliolata N. F. Mattos, Loefgrenia 58: 3. 1973.?TYPE:
Brazil.
Para: Faro, 19 Aug
1907, A. Ducke
s.n. [RB 8395]
(holotype: RB!).
Tree to 40 m tall with spreading crown in open situations; large trees buttressed; bark
brown to dark brown, scaling; slash pinkish brown or reddish brown, oxidizing darker;
clear, gelatinous ex?date from cuts after several days; twigs dark brown, sparsely hairy,
glabrescent, hairs red-brown, appressed; lenticels not apparent. Stipules 2-3 mm long,
narrow, caducous; leaf axis 10-30 cm long; rhachis sparsely hairy, glabrescent, hairs
to red-brown, appressed; stipels 1-3 mm long; petiolules 3-5 mm
short, golden-brown
long, indumentum like that of rhachis; leaflets in 3-5 pairs, 3-15 cm long, 1.5-6 cm wide,
narrowly obovate, broadly obovate, narrowly elliptic, elliptic, broadly elliptic, oblong to
suborbiculate, subcoriaceous, base rounded, obtuse to acute and occasionally
slightly de
current
or
somewhat
cordate,
apex
retuse
to rounded,
occasionally
acute
and with
an acu
long (to 20 mm long on sapling leaflets), glabrous adaxially, sparsely hairy

to red-brown; primary
hairs
short (<0.2 mm long), appressed, golden-brown
abaxially,
vein channelled adaxially; secondary veins 7-13, sunken or slightly sunken adaxially,
men
to 7 mm
raised abaxially, pattern eucamptodromous

tertiary veins
or occasion
to
and
raised
Panicles
terminal
plane
slightly impressed adaxially
abaxially.
cm
to
at
branch
11-28
with
brown
hairs
the
brown
ally axillary,
long,
pale
tips, glabres
cent towards the base, hairs short, appressed; bracts 2 mm long, hairy to sparsely hairy,
to red-brown; pedicels 2-4 mm long; bracteoles 1mm long, indu
hairs golden-brown
mentum like that of bracts. Flowers 13-18 mm long. Calyx 6-7 mm long, with red-brown
hairs; lobes 0.5-1.5 mm long, acute, occasionally obtuse. Petals pink-purple to purple, the
standard with a central pale marking; standard blade 10-12 mm wide, 9-12 mm high,
claw 2.5-4.5 mm
long; wing 8-11 mm long, 3^4-mm wide, claw 4-5 mm long, lamellate
sculpturing present; keel 7-9 mm long, 3-5 mm wide, claw 5-6 mm long. Stamens (9-)
12-13.5 (-17.5) mm long, the filaments united for the basal 7-11 mm, free for the distal
3-6.5 mm, vexillary stamen 8-12.5 mm long. Gynoecium
13-17 mm long, very sparsely
hairy on the upper and/or lower surface of the ovary, hairs red-brown, erect; stipe 3-8 mm
long, ovary 4.5-6 mm long, style 3-4 mm long; ovules 3-4. Fruits 4-6 cm long, 2-3.8 cm
high, 2.1-4 cm wide, elongated, weighing ca. 20 g or less when dry, green, slightly glau
cous, smelling very sweet, drying dark brown and distinctly wrinkled; stipe 12mm long;
suture raised adaxially and below, but hard to discern because of wrinkles; stylar remnant
minute; mesocarp 4-7 mm thick when fresh, thinner when dry, green, fibrous, drying with
abundant air spaces; endocarp 2-8 mm thick, woody,
mosome number unknown. Figs. IB, 2B, E.
Phenology. Flowering year-round.
Distribution
(Fig. 25). Trinidad; Colombia
fibrous, brown, drying paler. Chro
(Amazonas, Meta,
Vichada),

Venezuela
75
ANDIRA
2003
of Andira
macrocarpa
and A. surinamensis.
(Amazonas, Anzoategui, Apure, Bol?var), Guyana, Suriname, French Guiana, Ecuador

(Napo), Peru (Loreto, Madre de Dios), Bolivia (Pando, Santa Cruz), Brazil (Amazonas,
in forest, especially along rivers and
Para, Roraima, Maranh?o, Cear?, Mato Grosso);
as
an
savanna (pers. obs. in Guyana).
tree
in
in
also
isolated
forest
savannas,
gallery
arisoru (Warao), canelito negro,
Vernacular names. Yatuinay (Guahibo; Colombia);
blanco
sob?
wonka
winka,
(Venezuela); koraro, bat seed,
(Yekuana),
(Panare), palo
pilon,
maha, wild mango
(Guyana); rode kabbes, koeraroe ibiberoe (Suriname); mangarana,
manga brava, sucupira da v?rzea, angelim (Brazil); ituayuro (Peru); visguero del barbe
cho, visguero (Bolivia); large jumbie bean (Trinidad).
Representative
serve, Gill
12621
Specimens.
Trinidad.
Toco Road,
Valencia,
N. L. Britton
et al. 1776
(NY); Arena
Re
(NY).
Amazonas:
478 (COL, COAH).?

bocas del Rio Yari en el R?o Caquet?, Pab?n & Mahecha
Colombia.
cerca al terminal de transportes, Qui?ones
Los Llanos, R?o
2764 (COL).?Vichada:
META: Villa Vecencio,
r?o
4079 (US).
Venezuela.
AMAZONAS: Depto. Atabapo, Ca?o Negro,
Orinoco, Puerto Carre?o, Cuatrecasas
con el R?o Cuncunuma,
R?o Mapire,
arriba desde la confluencia
Steyermark et al. 126212 (NY).?ANZOATEGUI:
afl. norte
Camejo
estaci?n
medio,
galer?as del Cinaruco
del R?o Orinoco

y Achaguas,
Rosales
& Valles 79 (NY).?Apure:
Musinacio,
entrando por la Laguna de Calceta, R. G?mez

Distritos
et al. 588
Pedro
(NY).?
VOLUME 64
76
toward Caicura, Boom & Grillo

vie. Panare, village of Corozal, 6 km from Maniapure
Cede?o,
of
Moku-moku
Mts.
in
NW
of
Kanuku
Creek, A. C. Smith 3447
(COL, NY).
drainage
Guyana.
slopes
Suriname.
Schulz 7321 (COL); Sectie O, Sta
433,462,463.
(A, MO, U); District 8, Armai, R. T. Pennington
s.n. (US).
s.n. (US); Cayenne, Martin
hel 57 (MEXU).
French
Guiana.
Crique
Jacques, Wachenheim
BOL?VAR: Dist.
6460
Ecuador.
Napo:
et al. 7747 (E, MO, QCNE).

Peru.
Cuyabeno,
Laguna Cocha, Palacios
a orillas del R?o Nanay, 25 km SW Iquitos, Torres 160 (MO, NY).?Madre
58003
(MO).?
junction of R?os La Torre and Tambopata, Gentry & Jaramillo
Faunistica
Reserva
Prov. Maynas,
Loreto:
Nina Rimi,
DE Dios: Tambopata Reserve,

von Humboldt,
UCAYALl: Bosque Nacional
a Puerto
Pando:
camino
Bolivia.
Cobija,
Mar?a Km 88, Gentry et al. 36390 (MO, NY).

road Pucallpa-Tingo
& Hartshorn
2093 (NY).?SANTA
San Francisco, Meneces
Rico,
Brazil. Amap?: Rio Araguar?,

El Refugio, Killeen & P?rez 6682 (K, MO).
Ecol?gica
Rio Negro, Santa Isabel, Ducke 145 (A)?
Froes & Black 27684 (INPA).?AMAZONAS:
Island of S?o Luis, Froes 11805 (A,
Cear?:
Fortaleza, estrada do aeroporto, Ducke 2432 (US).?Maranh?o:
GROSSO: Aripua?a, Km 245 da BR-174, M. G. Silva & A. Pinheiro 4327 (F).?
GH, K, NY, U, US).?MATO
do Igarap? Curupira, M. Silva & R. Sousa 2284
Para: Santarem, Km 35 da estrada do Palh?o, acampamento
Cruz:
Velasco,
cachoeira
Reserva
do Pared?o,
(COL, US).?Rond?NIA:
Km
Manaus-Caracari,
et al. 296
F. E. Miranda
Ji-Paran?, Rio Machado,
Mpio.
515, bank Igarap? Dias, Steward et al. 135 (GH).
(INPA).?RORAMA:
estrada
surinamensis might be confused with A. verm?fuga, A. humilis, A. galeottiana,

A.
and A. galeottiana differ in their much larger
and
macrocarpa. Andira macrocarpa
fruits and, like A. verm?fuga, by the longer, erect indumetum on the abaxial leaflet surface.
Andira humilis has short, appressed hairs on its leaflet undersurfaces, like those of A. suri
Andira
namensis, but it is a geoxylic suffrutex rather than a tree.

Most previous workers (e.g., Mattos 1979) have cited
(Bondt) Splitgerber ex Pulle; however, J. C. Lindeman
Pulle (1906) made this combination in synonymy, which
the ICBN (Art 34.1; Greuter et al. 2000). The correct
this species as A. surinamensis

(pers. comm.) pointed out that
is inadmissible under terms of
attribution
is A. surinamensis
ex Amshoff.
(Bondt) Splitgerber
The timber of A. surinamensis is useful in construction. It is also resistant in water
and is used for cattle water troughs in Venezuela. Milliken
(1997) reported that in Ro
as
a
a
tree
is taken
decoction of the bark of this
raima, Brazil,
vermifuge, as reported by
as
a malaria remedy.
Bondt (1788) in the first description of the species, and also
anthelmia
(Vellozo) Bentham, Comm.
legum. gen. 44. 1837 [as "an
thelmintica"', the combination also proposed by Macbride,
1940, and Toledo,
1946]. Lumbricidia anthelmia Vellozo, Fl. flumin. 306. 1829. Andira anthelmia
var. acuminata Bentham, Fl. bras. 15(1): 294. 1862, nom. superfl. Vouacapoua
13. Andira
anthelmia
(Vellozo) Kuntze, Revis, gen. pi. 1: 212. 1891.?LECTOTYPE, here

designated: Vellozo's figure (Fl. flumin., icon. 7: 104. 1831).
Andira stipulacea Bentham, Comm. legum. gen. 43. 1837.?TYPE: BRAZIL. Bahia:
Pohl s.n. (holotype: K!).
stipulacea var. bahiensis Bentham, J. Proc. Linn. Soc, Bot. 4, Suppl. ("A
119. 1860. Andira legalis var. bahiensis (Ben
Synopsis of the Dalbergieae"):
Brazil.
Bahia: Blanchet
tham) N. F. Mattos, Loefgrenia 40: 3. 1970.?Type:
Andira
607 (holotype: BM!).

Andira frondosa var. longifoliolata N. F. Mattos, Loefgrenia
58: 3. 1973.?TYPE:
BRAZIL. Rio de Janeiro: Restinga de Jacarepagu?, 23 Sep 1958, E. Pereira,
Liene, Sucre & Duarte 4311 (holotype: HB; isotypes: RB-2 sheets!).
Tree to 25 m tall with large, broad, spreading crown in open situations; buttresses ab
to brown, Assuring vertically and scaling on larger trees; slash pale
sent; bark grey-brown

2003
77
ANDIRA
salmon pinkish, oxidizing red-brown, with clear ex?date and small amount of red ex?date;
buff to yellowish,
streaked; twigs often with crowded persistent stipules, stipule
scars, and leaf scars, with sparse red-brown appressed hairs, glabrescent; bark pale brown
to buff, Assuring vertically; lenticels few or absent. Stipules 1.5-6 cm long, generally >0.5
wood
cm wide, persistent, ovate, pale brown, with sparse, red-brown appressed hairs, glabres
cent and generally glabrous at maturity; leaf axis 24-55 cm long; rhachis dark to pale
brown, sparsely hairy when young, glabrescent, hairs red-brown, erect; stipels 1-9 mm
1.5^4.5 mm
long, indumentum like that of rhachis; leaflets in (3-) 4-6

4.1-12.5
(-7.5) cm wide, elliptic, nar
(-7) pairs, (2.2-)
(-20.5) cm long, (1-) 2.2-6.5
obovate
rowly
(rarely narrowly ovate, narrowly elliptic, broadly obovate to oblanceolate),
thick-chartaceous
(rarely chartaceous or subcoriaceous),
shiny dark green adaxially, matt
long; petiolules
abaxially, base obtuse or rounded, often slightly decurrent, apex obtuse or rounded (rarely
acute), often slightly retuse or with a short acumen to 10mm long, glabrous adaxially ex
the hairs erect,
cept the groove of the primary vein very sparsely hairy, glabrescent,
?
?
to
to
hairs
red-brown to
erect,
very sparsely hairy abaxially,
sparsely
appressed
mm
>0.2-1.0
veins
vein
channelled
whitish,
8-13, ?
long; primary
adaxially; secondary
to
sunken
plane
slightly
adaxially, raised abaxially, pattern eucamptodromous
becoming
brochidodromous,
tertiary veins plane to slightly impressed adaxially and raised abaxially.
Panicles
11-35 cm long, terminal and axillary, hairy to sparsely hairy at branch tips,
glabrescent towards the base, hairs red-brown, ? appressed; bracts 2-6 mm long, narrow,
caducous, sparsely hairy, hairs red-brown, appressed; pedicels 2-6 mm long; bracteoles
1.5-2.5 mm
long, indumentum like that of bracts. Flowers 19-24 mm long. Calyx 8-10
mm long, purple-brown, with sparse, red-brown, appressed hairs, indumentum most dense
on lobes; lobes 0.3-2 mm long, obtuse to acute. Petals rose-violet to purple; standard
blade 14-16 mm wide, 13-15 mm high, claw 6-7 mm long; wing 10-13.5 mm long,
4.5-6.5 mm wide, claw 6-10 mm long, lamellate sculpturing present; keel 9-12 mm long,
5-6.5 mm wide, claw 7-10 mm long. Stamens white, 15-20 mm long, filaments united for
the basal 8.5-14 mm, free for the distal 4-7 mm, vexillary stamen 12.5-16.5 mm long.
15.5-21 mm long, ovary hairy, stipe and style sparsely hairy to upper and
Gynoecium
lower surface of ovary hairy, stipe and style with scattered hairs, hairs red-brown, ap
pressed; stipe 5-6 (-9) mm long, ovary 4.5-7 mm long, style 4.5-6 mm long; ovules 4-5.
cm long, 2.5-4
cm high, 2.2-4 cm wide, elongated, weighing ca. 20 g or less

when dry, strong smelling, dark brown, this surface layer thin, green beneath, drying dark
brown with surface appearing smooth but minutely wrinkled (best seen with lens or mi
croscope); stipe to 12mm long, but fruit usually breaking off without a stipe; suture raised
Fruits 3-6.2
adaxially, ? undetectable abaxially; stylar remnant slight, but often obvious at very apex
of fruit; mesocarp
1.5-3 mm thick, green to pale lime-green, drying pale brown, fibrous
and granular, somewhat air-filled and soft; endocarp 1-2.5 mm thick, brown to pale
brown, woody, fibrous. Chromosome number: n = 10, 11. Figs. 1A, 26.
Phenology. Flowering September to December.
Distribution
(Fig. 27). Brazil (Alagoas, Bahia, Espirito Santo, Rio de Janeiro); in

forest
and
rain forest. I have also seen this species growing along river banks in
restinga
a
in
and
Bahia,
seasonally inundated area in Rio de Janeiro; it can clearly tolerate water
seems
and
it
logging,
likely that its fruits may be secondarily dispersed by water.
Vernacular
names. Angelim
coco,
angelim
preto
(Bahia);
angelim
Janeiro).

coco
(Rio de
78
VOLUME 64
FIG. 26. Andira
anthelmia
and A. legalis. A. A. anthelmia,
habit. B. A. anthelmia,
abaxial leaflet surface.
leaflet. D. A. legalis, abaxial leaflet surface. E. A. legalis, stipule. F. A. legalis, flower. G. A. legalis,
calyx, opened to show inner surface. H. A. legalis, standard petal. I. A. legalis, outer surface of wing petal with
lamellate sculpturing. J. A. legalis, keel petal, inner surface. K. A. legalis, androecium. L. A. legalis, gynoecium,
C. A. legalis,
also
shown
above
in longitudinal
section. M.
fruit. O. A. legalis, fruit,
281; C-F, R. T. Pennington
307;
Pennington
nington, 183; O, G. P Lewis & H. C. de Lima
tion. N. A. anthelmia,
sec
A. anthelmia,
gynoecium,
partially sectioned to show wall structure. (Based on: A-B, R. T.
F-L,
A. Lima
51-944;
M, E. Pereira
et al
1196.)

4156; N, R. T. Pen
ANDIRA
2003
50
55
79
45
35
of Andira
40
anthelmia.
Examined.
Brazil.
Alagoas:
without
27 (RB).?Bah?a:
Additional
Specimens
locality, Uch?a
Estrada Mara?-Ubaitaba,
Bel?m 1879 (NY, UB); Coaraci, cocoa plantation, Bel?m & R. S. Pinheiro 2949 (US);
120 (CEPEC); Mpio.
II
Ilh?us, Fazenda Serra Grande, W of CEPEC cacao plantation, Cabruca, Hummel
Mpio.
h?us, estrada Ilh?us-Itabuna,
pr?x. a CEPLAC, H. C. de Lima et al 3866 (CEPEC); Mpio. Dh?us, CEPLAC-I1
et al. 183 (CEPEC, FHO, K); Mpio.
h?us road, 2 km from CEPLAC, R. T. Pennington
Jussari, Fazenda Santa
to Palmira, then 1 km on right hand turn, R. T. Pennington
Antonio de Baixo, Km 16 on road from BR-101
205,
et al. 212 (CEPEC, FHO,
286 (CEPEC, FHO, K); Mpio. Marau, road Mara?-Ubaitaba
et al 227, 238 (CEPEC,
R. T. Pennington
K); Mpio. Ubaitaba, banks of Rio de Contas, by road Ubaitaba-Marau,
R. T. Pennington
& Brito 281, 282
5 kms before CEPLAC,
Ilh?us, road Ilh?us-Itabuna,
K); Mpio.
& F. A. de Car
and road to Jussari, R. T. Pennington
Jussari, Junction of BR-101
(CEPEC, FHO, K); Mpio.
bank of Rio dos Frades, R. T. Pennington
valho 294 (CEPEC, FHO, K); Mpio. Porto Seguro, BR-101,
<feF. A.
de Carvalho
297 (CEPEC, FHO, K); Mpio. Prado, 4 km along track on right side road from Prado to Alcoba?a,
R. T. Pennington
? F. A. de Carvalho 312 (CEPEC, FHO, K); Mpio. Una, Fazenda S?o Fafael, margem do Rio
1996 (US).?ESPIRITO
Alian?a, R. S. Pinheiro 250 (US); estrada Santa Ines-Rio Bahia, Km 7-9, R. S. Pinheiro
FHO,
360 (RB).?MINAS
/. G. Kuhlmann
GERAIS: Est. Exp. de Agua Limpa,
Collatina,
DE JANEIRO: Praia de Grumari, D. Araujo 1843 (NY); Mpio. Campos, Lagoa de
(RB, UB).?R?o
et al 2593 (RB); estrada dos Ban
4044 (GUA); Silva Jardim, Poco d'Anta, Carauta
Cim, D. Ara?jo & Maciel
da Trjuca, Machado
77 (RB); Guanabara,
deirantes, Morro do Calemb?, Lanna Sobrinho 649 (US); Restinga
Santo:
Gomes
Restinga
estrada do Pan?as,
2864
de Jacarepagu?, Pereira
et al 291 (US).
et al. 4156
(NY); Goitacazes,
Lagoa
Feia, Ponta Grossa
Segadas-Vianna

dos Fildagos
borough,
80
VOLUME 64
closely related to A. legalis (Fig. 9), which has similar

The
large, persistent stipules.
species are clearly distinguished by their fruits: A. anthelmia
has small, bat-dispersed fruits, which are never longer than 6.2 cm, whereas A. legalis has
large, rodent-dispersed fruits 5.6-12 cm long. Vegetatively
they may be separated by the
Andira
anthelmia
is most
denser, more deeply red-brown indumentum on the stipules, leaflet undersurface, inflo
rescence axis, and calyx of A. legalis. Andira anthelmia might also be confused with A.
ormosioides, which has similar, large flowers, but lacks large, persistent stipules.
14. Andira
legalis (Vellozo) Toledo, Arq. Bot. Estado S?o Paulo 2(2): 29. 1946. Lumbri
cidia legalis Vellozo,
Fl. flumin. 306. 1829. Vouacapoua
legalis (Vellozo)
Kuntze, Revis, gen. pi. 1: 212. 1891.?LECTOTYPE, here designated: Vellozo's
figure (Fl. flumin., icon. 7: 105. 1831.)
Andira frondosa Martius ex Bentham, Comm. legum. gen. 44. 1837. Vouacapoua
(Martius ex Bentham) Kuntze, Revis, gen. pi. 1: 212. 1891.?TYPE:
frondosa
Brazil.
Brasiliae
Rio de Janeiro: "In arenosis
sylvarum ad Capo Frio et alibi circa oram

orientalis maritimam," Martius s.n. (holotype: BR!; isotype: M).
Tree to 20 m tall, simply branched; buttresses absent; bark pale brown to grey-brown,
rough, Assuring vertically; trunks of saplings marked with leaf and stipule scars, stipules
often persistent; slash buff to pale red-brown, oxidizing darker red-brown, a little red ex
?date; wood buff to yellowish buff; twigs swollen, with crowded persistent stipules, leaf
scars and stipule scars, with red-brown, erect hairs, glabrescent; lenticels not apparent.
Stipules 1.5-4.5 cm long, >0.5 cm wide, persistent, ovate, with deep red-brown, appressed
leaf axis 15-30 (-60) cm long; rhachis with erect, red-brown hairs,
glabrescent;
stipels to 9 mm long; petiolules 3-7 mm long, indumentum like that of
in 4-7 (-9) pairs, 5.5-15 (-30) cm long, 2.3-7 (-11) cm wide, elliptic,
leaflets
rhachis;
narrowly obovate, ovate (rarely narrowly elliptic to broadly elliptic), coriaceous, base ob
tuse, rounded, ? truncate or very slightly cordate, apex obtuse or rounded, often slightly
retuse or with an acumen to 7 mm long, glabrous adaxially except scattered hairs in the
groove of the primary vein, hairy to sparsely hairy abaxially, indumentum most dense on
veins, hairs red-brown, erect, >0.2-1.0 mm long; primary vein channelled adaxially; sec
hairs, glabrescent;
ondary veins 8-13, slightly sunken adaxially, raised abaxially, pattern eucamptodromous
tertiary veins plane to impressed adaxially and raised abax
ially. Panicles 10-50 cm long, terminal (more rarely axillary), densely covered with erect,
red-brown hairs; bracts 5.5-10 mm long, ? ovate, densely covered with long, ? appressed,
red-brown to buff hairs; pedicels 2-3 mm long; bracteoles 7-10 mm long, ? ovate, indu
mentum like that of bracts. Flowers 19-23 mm long. Calyx (7-) 9-10.5 mm long, brown
purple, hairy to densely hairy, hairs red-brown, often long; lobes 1-3 mm long, obtuse to
acute. Petals purple, standard with a pale marking at center; standard blade (12-) 16-17
mm wide, (12-) 15mm high, claw 5-7 mm long; wing 13-14 mm
long, (3.7-) 6-7.2 mm
wide, claw 6.5-8 mm long, lamellate sculpturing present; keel (8.2-) 11-13 mm long,
(4-) 5-6 mm wide, claw 6-8.4 mm long. Stamens (12.5-) 15 mm long, filaments united
for the basal 8-11 mm, free for the distal 3-6.4 mm, vexillary stamen 12.5-15 mm long.
(14.5-) 17.2-18 mm long, ovary sparsely hairy, stipe and style very sparsely
Gynoecium
to
upper (basal half only) and lower surface of ovary hairy with a few hairs extend
hairy
down
the stipe, hairs red-brown, ? appressed; stipe 5-6 mm long, ovary (5.5-) 7 mm
ing
long, style (4-) 5.2-6 mm long; ovules 2-A. Fruits 5.6-12 cm long, 4.5-8.7 cm high,
4.5-9 cm wide, elongated, weighing
100-300 g when dry, rough, pale brown speckled

2003
81
ANDIRA
50
55
35
45
of Andira
40
legalis.
(both fresh and dry); stipe to 5 mm long, sutures not obvious, sunken adaxi
ally, slightly raised abaxially; stylar remnant not obvious; mesocarp 4-10 mm thick, pale
brown, hard, finely granular; endocarp 3-18 mm thick, brown, woody, fibrous. Chromo
some number unknown. Figs. ID, 26.
Phenology. Flowering August to November.
dark brown
Distribution
(Fig. 28). Brazil (Bahia, Espirito Santo, Minas Gerais, Pernambuco, Rio
de Janeiro); in restinga and rain forest.
Vernacular names. Angelim
preto, angelim roxo (Bahia); angelim coco (Rio de
Janeiro).
Additional
Specimens
Examined.
Brazil. Bah?a: Mara?, Bel?m 1830 (CEPEC, MEXU, NY, UB, US);
7.3 km na estrada Serra Grande-Itacar?,
Fazenda Lagoa do conjunto
Mpio. Uru?uca, Dist. Serra Grande,
et al 3508, 3509 (CEPEC, NY); Mpio. Porto Seguro, junction on right
Fazenda Santa Cruz, A. M. de Carvalho
at Km 16 on road Povoada Santa Cruz, 1 km up this road, Mattos
Silva et al. 340 (CEPEC, K); Area Controle
da Caraiba Metais, Noblick 2299 (K, UEFS); Mpio. Porto Seguro, junction on right at Km 16 on road Povoada
1 km up this road, R. T. Pennington
Santa Cruz,
& Lewis
191 (CEPEC, FHO, K); Mpio. Mara?,
road
et al 214, 215 (CEPEC, FHO, K); Mpio. Alcoba?a,
Mara?-Ubaitaba
1 km N do centro

82
VOLUME 64
da cidade, R. T. Pennington
& F. A. de Carvalho 305,307,308,310
(CEPEC, FHO, K); Teixeira de Freitas, Vale
do Rio Alcoba?a,
7.3 km N Serra Grande on road to Itacar?, Fazenda
dos Santos 2108 (US); Mpio. Uru?uca,
SANTO: Reserva
Lagoa do Conjunto e Fazenda Santa Cruz, W. W. Thomas et al 8533 (CEPEC, NY).?ESPIRITO
J. G. Kuhlmann
478 (NY).?Per
da CVRD, Linhares, DA.F.
128 (RB); Vitoria, Fazenda Maruhype,
DE JANEIRO:
NAMBUCO: Mata de Dois limaos, margem estrada para o acuda do Prata, Lima 51-944
(K).?R?o
near Brejo de Bezerra, D. Araujo 4612
Almeida de Jesus 2044 (NY); Mpio. Maca?,
Guanabara, Marambaia,
near Sambaqui da Beirada, D. Araujo 8079 (GUA); Mpio. Cabo Fri?, dune system
(GUA); Mpio.
Saguarema,
Florestal
loteamento da Praia de Itauna, D. Araujo et

(GUA); Mpio. Saguarema,
Comoros da Lagoa Vermelha, D. Araujo & Mauro 8614 (GUA); Mpio. Ar
W.
rail do Cabo, restinga de Massambaba,
8700 (GUA); Recreio dos Bandeirantes,
Zacara, D. Araujo & Maciel
06529 (RB); Mpio. Marica, Barra
Rio Doce, J. G. Kuhlmann
Hoehne
5718 (SP), 5895 (COL, SP); Goitacazes,
at Dama
Branca, D. Araujo
al. 8597
(GUA); Mpio.
et al. 8326
Saguarema,
de Marica, pr?x. a Lagoa de Marica,

Cabo, L. B. Smith 6554 (US).
Lewis
& H. C de Lima
1196 (K); Mpio.
Cabo Fri?, Cabo Fr?o, Arraial
do
Andira legalis is mostly closely related to A. anthelmia and A. ormosioides. For ac

counts of how to distinguish these species, see the notes under A. anthelmia (no. 13) and
A. ormosioides
(no. 15). The collection Duarte 4263 from Minas Gerais listed under A.
ormosioides may be A. legalis (see discussion of A. ormosioides).
legum. gen. 44. 1837. Andira anthelmia var.
J.
Proc. Linn. Soc, Bot. 4, Suppl. ("A Synop
(Bentham) Bentham,
Brazil.
120. I860.?Type:
sis of the Dalbergieae"):
Tingua, Schott s.n. (holo
type: K!; isotype: F!).
15. Andira
ormosioides
Bentham,
Comm.
ormosioides
Tree (though can flower when very small) to 30 m tall, long-boled with small crown
(even in open situations), with small buttresses; bark pale brown to grey-brown, Assuring
vertically and flaking slightly; slash pale red-brown; wood buff/cream; twigs brown, with
erect, red-brown hairs, becoming paler, glabrescent. Stipules to 16 mm long, to 1 mm
wide, moderately persistent, with red-brown, ? appressed hairs;
long; rhachis with red-brown erect hairs, glabrescent; stipels 2-3
2-5 mm long, indumentum like that of rhachis; leaflets in (3-)
on sterile shoots) cm long, 1.8-7 cm wide, elliptic to narrowly
leaf axis 9-30 (-35) cm

(-6) mm long; petiolules
4-5 pairs, 4.7-15.5
(-19
obovate, subcoriaceous,
shiny, dark green adaxially, base obtuse to rounded, often very slightly decurrent, apex ob
tuse to rounded, occasionally with an acumen to 7 mm long, glabrous adaxially except
scattered hairs in the groove of the primary vein, hairy to sparsely hairy abaxially, indu
dense on veins, hairs red-brown, erect, >0.2-1.0 mm long; primary vein
adaxially; secondary veins 8-11, slightly sunken adaxially, raised abaxially,
mentum most
channelled
pattern eucamptodromous becoming brochidodromous,

tertiary veins plane to slightly im
cm
and
raised
Panicles
10-30
pressed adaxially
long, terminal (more rarely ax
abaxially.
mm
erect
with
bracts
3-7
red-brown,
hairs;
illary),
long, with red-brown, ? appressed
mm
mm
3-7
bracteoles
1-2.5
hairs; pedicels
long;
long, indumentum like that of bracts.
18-23 mm long. Calyx 9-10 mm long, with red-brown, ? appressed to ? erect
lobes
0.8-2 mm long, obtuse to acute. Petals pink to purple; standard blade 17-20
hairs;
mm wide, 15-19 mm high, claw 6-7 mm long; wing 12-15 mm long, 5-6.5 mm wide,
claw 8-9 mm long, lamellate sculpturing present; keel 10-12 mm long, 4-7 mm wide,
claw 8-10 mm long. Stamens 15-20 mm long, filaments united for the basal 8.5-11 mm,
Flowers
free for the distal 5-10 mm, vexillary stamen 11.5-15 mm long. Gynoecium
18.5-20 mm
long, ovary sparsely hairy, stipe and style sparsely hairy, hairs red-brown, ? appressed;
stipe 6-7 mm long, ovary 7 mm long, style 4.5-6 mm long; ovules 5-8. Fruit 4.7-5.6 cm
long, 3.4-4.1 cm high, 3.4-4.1 cm wide, elongated, weighing ca. 20 g or less when dry,

83
ANDIRA
2003
50

mosioides
(see discussion);
of Andira
45
55
ormosioides.
if so, the locality
35
The collection
is a considerable
40
A. P. Duarte
range extension
4263
probably
for the species.
represents A. or
smooth, very dark brown with green beneath when fresh, drying very dark brown, ap
tuberculate (best seen with lens or microscope);
pearing smooth but minutely
stipe 6-8
mm long; suture slightly raised adaxially, flanked by two shallow grooves, very slightly
raised abaxially; stylar remnant raised at apex of fruit; mesocarp
3^1 mm thick, pale
mm
2-3
thick, dark brown, woody,
greenish white, drying pale brown, granular; endocarp
fibrous. Chromosome
number unknown. Fig. 3H.

Phenology. Flowering June to December.
Distribution
(Fig. 29). Brazil (Bahia, Espirito Santo, Minas Gerais, Rio de Janeiro,
S?o Paulo); in restinga, rain forest, and semi-deciduous
forest.
Vernacular names. Angelim pedra (Espirito Santo); angelim tento, angelim rosa, an
gelim amargoso (Rio de Janeiro); Jacaranda rajada (S?o Paulo).
Additional
& F. A. de Carvalho
164/79
(RB).?Minas
Examined.
Specimens
306, 309
Gerais:
Brazil. Bahia: Mpio. Alcoba?a,

1 km N do cidade, R. T. Pennington
Santo: Linhares, Res. Florestal da CVRD, Foli
(CEPEC, FHO, K).?Espirito
Patos de Minas, A. P. Duarte 4263 (NY; see discussion);
5 km de Grama Juiz

84
VOLUME 64
de Fora, Heringer
2700 (RB); Carangola, Rio Carangola, Leoni & Leoni 856 (K); 400 km E Belo Horizonte,
Caralinga, Fazenda Montes Claros, Lopes &
vicinity of Rio Manhuac?,
Valley of Rio Doce, Estac?o Biol?gica
DE Janeiro:
Serra da Estrela, de Almeida s.n. (RB); Barreira de Soberbo, Dias s.n.
Andrade 850 (K, RB).?Rio
s.n. (RB 81810) (RB);
s.n. (RB 54881) (RB); Rio de Janeiro, Horto Florestal
(R); Rio de Janeiro, Horto Florestal
528 (F, NY, RB, UB); Mpio. Mage, Cachoeiras
29 (RB); Guanabara, /. G. Kuhlmann
Avellar, /. G. Kuhlmann
de Macacu,
Para?so, near CPRJ, banks of Rio Para?so, H. C. de Lima et al. 4253 (RB); estrada do Recreo dos
s.n. (RB 76106), Machado
s.n. (RB 75296)
da Trjuca, Machado
Lutz 1493 (R); Restinga
Bandeirantes,
(RB);
10503 (RB); Mpio. Mag?,
& Pessoa
do RJ, Martinelli
(IIIo), Dist. Para?so, Centro de Primatologia
Mag?
s.n. (R); Ubatuba,
PAULO: Aeto da Serra, de Andrade
Para?so, Sampaio 6 (RB); Gavea, E. Ule LVII (R).?SAO
da Cruz 13 (NY, SP).
pr?ximo a Base Norte do Instituto Ocean?grafico,
Mpio.
is
Field observation in Bahia of growth form and fruit indicates that A. ormosioides
distinct from the three species to which it is most similar: A. fraxinifolia, A. legalis, and
A. anthelmia. The fruit of A. ormosioides are small (4.7-5.6 cm long, vs 5.6-12 cm long
in A. legalis) and have a pale greenish white mesocarp (in comparison to the green meso
and A. anthelmia). Andira ormosioides has a long bole and small
carp of A. fraxinifolia
and A. anthelmia have a short bole
crown, even in open situations, whereas A. fraxinifolia
and a broad, spreading crown.
Herbarium specimens of A. ormosioides can be difficult to separate from large-flow
ered specimens o? A. fraxinifolia
and particularly from specimens of A. legalis and A. an
thelmia that have lost their characteristic large stipules or were collected without stipules.
An example is A. P. Duarte 4263, which has the dense red-brown indumentum on the
leaflet undersurface, inflorescence axis, and calyx characteristic of both A. ormosioides
and A. legalis. In the absence of fruit or stipules, it is not possible to determine this spec
imen with certainty. This specimen was collected in Patos de Minas, western Minas
Gerais, ca. 600 km from the Atlantic coastal forests, where both A. ormosioides and A. le
galis grow. Forests in Patos de Minas are semideciduous, with a few patches of dry de
ciduous
forests
on
calcareous
outcrops
(A. T. Oliveira-Filho,
pers.
comm.).
Andira
ormo
forests of the Rio Doce valley of eastern Minas

in the semi-deciduous
are
to
the
rain forests, whereas A. legalis grows strictly
which
coastal
transitional
Gerais,
in rain forests and restingas. The Duarte specimen from the dry area of Patos de Minas is
sioides
more
occurs
likely to be A. ormosioides
and is therefore included here.
fraxinifolia Bentham, Comm. legum. gen. 44. 1837. Vouacapoua fraxinifolia

Brazil. Minas Gerais:
(Bentham) Kuntze, Revis, gen. pi. 1: 212. 1891.?Type:
prope Itamb?, Pohl s.n. (lectotype, here designated: K!; isolectotype: K!).
Andira parvifolia Martius ex Bentham, Comm. legum. gen. 44. 1837. Vouacapoua
(Martius ex Bentham) Kuntze, Revis, gen. pi. 1: 212. 1891.?TYPE:
parvifolia
Brazil. Minas Gerais: "in campis altis Serro Frio," Martius s.n. (holotype: M!).
16. Andira
Andira pisonis Martius ex Bentham, Comm. legum. gen. 44. 1837. Vouacapoua piso
nis (Martius ex Bentham) Kuntze, Revis, gen. pi. 1: 212. 1891.?Type:
Brazil.
Bahia: "Inter virgulta in arenosis adMucuri," Martius s.n. (holotype: M!).
Andira rosea Martius ex Bentham, Comm. legum. gen. 44. 1837. Andira fraxinifolia
var. rosea (Martius ex Bentham) Bentham, Fl. bras. 15(1): 294. 1862.?TYPE:
Brazil.
S?o Paulo: "In sylvis aetemis supra Serra do Mar, provinciae Sancti
Pauli ad fazienda dos Negros," Martius s.n. (holotype: M!).
ex Glaziou, Bull. Soc. Bot. France 52, M?moire
3: 151.
Rio de Janeiro: Nova Friburgo, Glaziou 20274 (holotype:
P!; isotypes: BR, K!; photo of B isotype: G!).
Andira micans
1906.?Type:
Taubert
Brazil.

ANDIRA
2003
85
BRAZIL. Bahia: be
Loefgrenia 40: 1. 1970.?Type:
tween Len?ois and Itaberaba, 15 Sep 1956, E. Pereira 2064 (holotype: R!; iso
type: RB!).
var. latifoliolata N. F. Mattos, Loefgrenia 40: 2. 1970.?TYPE:
Andira fraxinifolia
s.n.
13 Aug 1949, O. Machado
Brazil.
Rio de Janeiro: Restinga da Gavea,
(holotype: RB!).
Brazil.
Andira anthelmia var. gracilis N. F. Mattos, Loefgrenia 40: 3. 1970.?Type:
S?o Paulo: Limeira, Navarro de Andrade 167 (holotype: R; isotype: RB!).
Brazil.
Bahia: Km
Andira bahiensis N. F. Mattos, Loefgrenia 45: 2. 1970.?Type:
Andira handroana N. F. Mattos,
80 between Betanha
and Canavieiras,
13 Jul 1964, N. T. Silva 58414
(holotype:
UB; isotype: CEPEC!).
N. F. Mattos, Loefgrenia 53: 1. 1970.?TYPE: BRAZIL. Per
Andira pernambucensis
nambuco: Rio Formoso, Engenho S?o Manoel, 3 Sep 1954, J. Falc?o, Engler &
E. Pereira 944 (holotype: RB!; isotypes: RB-2 sheets!).
var. lanceata N. F. Mattos, Loefgrenia 58: 3. 1973. Brazil.
S?o
Andira fraxinifolia
24 Aug 1963, J. Mattos & N. F. Mattos 8383 (holo
type: SP!).
58: 2. 1973.?Type:
Andira pisonis var. emarginata N. F. Mattos, Loefgrenia
Brazil.
Bahia: Igua?u, 30 Dec 1922, P. Campos Porto s.n. (lectotype, here des
Paulo: 2 km N of Atibaia,
ignated: RB!).
Brazil.
Andira pisonis war. puberula N. F. Mattos, Loefgrenia 58: 2. 1973.?Type:
/.
G.
Kuhlmann
do
Rio
Santo:
1934,
Dur?o,
Doce,
Linhares,
Apr
Lagoa
Espirito
163 (holotype: RB!).
Shrub or small tree to 12 m tall, with broad spreading crown in open situations; but
tresses absent; bark grey-brown, fissuring vertically and flaking (larger trees); slash pale
brown to pale red-brown, oxidizing darker, occasionally with slight red ex?date; wood
to dark brown, older twigs paler, often buff/whitish, hairy to
sparsely hairy, glabrescent, hairs erect. Stipules 2-9 mm long, to 1mm wide, caducous,
with sparse, red-brown ? appressed hairs; leaf axis 6-21.5 (-25) cm long; rhachis sparsely
hairy (occasionally more densely hairy), glabrescent, hairs red-brown, erect; stipels 1-5
mm long; petiolules 2-3 (-5) mm long, indumentum like that of rhachis; leaflets in (2-)
3-7 pairs, 2-12 cm long, 0.7^1.2 cm wide, elliptic, narrowly elliptic, narrowly obovate,
cream/buff;
twigs brown
to subcoria
oblanceolate
(more rarely lanceolate to broadly obovate), thick-chartaceous
ceous (rarely chartaceous), dark green, shiny adaxially, matt abaxially, base obtuse to
rounded (rarely acute), often very slightly decurrent, apex acute, obtuse to rounded, gener
ally with an acumen to 7 mm long, glabrous adaxially except scattered hairs in the groove
of the primary vein, hairy to very sparsely hairy abaxially, indumentum most dense on
.0mm long; primary vein channelled adaxially;
sunken adaxially, raised abaxially, pattern eu
camptodromous becoming brochidodromous,
tertiary veins plane adaxially and slighty
raised or raised abaxially. Panicles 4-30 cm long, terminal and axillary, with red-brown,
erect hairs, glabrescent towards the base; bracts 2-3.5 mm long, with appressed, red-brown
hairs; pedicels 2.5-4 mm long; bracteoles 1.5-2 mm long, indumentum like that of bracts.
veins, hairs pale to red-brown, erect, >0.2-l
secondary veins 6-10, ? plane to slightly
Flowers 13-17 mm long. Calyx 6-7 mm long, brown to purplish, hairy to sparsely hairy,
hairs ? appressed, most dense on lobes; lobes 0.25-1.75 mm long, obtuse to acute. Petals
pink to purple, the standard with a white central marking; standard blade 10-14 mm wide,
10-13 mm high, claw 4-5 mm long; wing 8.5-11.5 mm long, 5-7 mm wide, claw 5-7 mm

86
VOLUME 64
long, lamellate sculpturing present; keel 8-11 mm long, 4-6 mm wide, claw 5-7 mm long.
Stamens 9-15.5 mm long, filaments united for the basal 5-9.5 mm, free for the distal 4-8
13.5-18 mm long, ovary hairy, stipe and
mm, vexillary stamen 9-12 mm long. Gynoecium
to
of
and
lower
surfaces
ovary sparsely hairy with scattered hairs
upper
style sparsely hairy
on sides of ovary and stipe and style, hairs red-brown, ? appressed; stipe 3-6.5 mm long,
ovary 5-8 mm long, style 4.5-6 mm long; ovules (3-) 4-6 (-7). Fruits 2.5-6 cm long,
1.8-4 cm high, 1.6-3.8 cm wide, elongated, weighing ca. 20 g or less when dry, green,
appearing smooth but irregularly ridged (best seen with lens or micro
scope), drying dark brown to brown; stipe 4-10 mm long; suture raised adaxially, obscure
abaxially; stylar scar raised at apex of fruit; mesocarp 1-3 mm thick, green when fresh, dry
ing brown, hard, granular, with air spaces; endocarp 1-7 mm thick, brown, woody, fibrous.
Chromosome number: n = 11. Figs. 2C, 3G, 4A(i), 5A, 30.
sweet-smelling,
Phenology. Flowering year-round.

Distribution (Fig. 31). Brazil (Alagoas, Bahia, Cear?, Distrito Federal, Espirito Santo,
Goi?s, Minas Gerais, Paran?, Pernambuco, Rio de Janeiro, Santa Catarina, S?o Paulo); in
restinga, rain forest, campo rupestre, and secondary vegetation, often seen isolated in pas
tures.
Vernacular names. Angelim branco, angelim preto (Bahia); angelim coco (Bahia, Es
pirito Santo); angelim amargoso (Minas Gerais); angelim pedra (Minas Gerais, Rio de
Janeiro); angelim da folha grande, pau de morcego,
quaiseara, Jacaranda de morcego,
fruta de c?valo (S?o Paulo); pau angelim (Santa Catarina).
e? al
Specimens.
Brazil. Alagoas:
Fazenda S?o Louren?o, Andrade-Lima
Fleixeiras,
Mpio. Sa?de, Cachoeira do Paulista, 6 km along road starting at Km 10 of road Sa?de-Jacobina,
1055 (CEPEC, E); Serra Jacobina, Blanchet
2723 (E, BM, F, K, NY); Mpio.
Jacobina, Serra do
na estrada para Morro do Chap?u, A. M. de Carvalho
et al 6141
Tombador, ca. 8 km SW da sede do municipio,
/. G.
(E, NY); Mpio. Saude, Cachoeira do Paulista, 6 km along road starting at Km 10 of road Sa?de-Jacobina,
Jardim 71 (CEPEC, FHO); Mpio.
100 m up hill from Club Tomoromba,
R. T. Pennington
194
Ilh?us, Oliven?a,
Representative
7 (F).?Bah?a:
Amorim et al
road to Pontal, junction on left, about 300 m up this

Ilh?us, ca. 3 km N Oliven?a,
200 (CEPEC, FHO, K); Mpio.
track, R. T. Pennington
Ilh?us, Agua da Oliven?a, R. T. Pennington
201, 202
& A. M. de Carvalho 204, 228 (CEPEC,
(CEPEC, FHO, K); Mpio. Jussari, Fazenda Teimoso, R. T. Pennington
to Palmiras, ca. Km 15, R. T. Pennington
& A. M. de Carvalho
207
FHO, K); Mpio.
Jussari, road from BR-101
et al. 211, 213 (CEPEC, FHO,
road Mara?-Ubaitaba
(CEPEC, FHO, K); Mpio. Mara?,
Km 4 (off road to Pedras from main road Oliven?a-Una),
R. T. Pen
Una, road to Independencia
Jacobina, Km 20 along road Jacobina toMorro do Chap?u
nington e? al 234, 236 (CEPEC, FHO, K); Mpio.
& Brito 248,249,250;
18 km E Morro do Chap?u on BR
(BR-324), R. T. Penningion
Mpio. Morro do Chap?u,
K); Mpio.
& Brito 251 (CEPEC, FHO, K); Mpio. Len?ois,

5 km from Len?ois on road to BR-242,
052, R. T. Penningion
ca. 2 km N Alcoba?a,
R. T. Pennington
& Brito 274, 275, 276, 278, 279; Mpio. Alcoba?a,
R. T. Pennington
&
F. A. de Carvalho
Queiroz et al.
Itua?u, arredores do Morro da Mangabeiras,
1610 (K, NY).?Cear?:
Federal:
Santana, Poranga, S. J. Filho 63 (RADAM).?Distrito
Campus of Uni
10064 (UB); Fazenda Agua Limpa, near Vargem Bonita, ca. 18 km SSW Brasilia
versity of Brasilia, Heringer
TV Tower, Ratter
163
(NY); Mpio.
et al 3791
Linhares,
SANTO: Lagoa do Dur?o, Linha Res., Rio Doce, J. G. Kuhlmann

da CVRD, H. C de Lima 1711 (RB).?GOI?S:
Mpio. Santo An
Cabteiro do Para?so, Elias de Paula 3147 (UB).?MINAS
GERAIS: San
(UB).?ESPIRITO
Reserva Florestal
tonio Descoberto,
Corrego da Fazenda
tana do Riacho, Serra do Cipo, pr?x. do Rio Cipo, de Barros 1356 (SP); Mpio. Jaboticatubas, Km 123 ao longo
de rodovia Lagoa Santo-Concei?ao
do Mato Dentro, Joly & Samir s.n. (SP); Vi?osa, Mexia 4932 (A, F, GH, U,
de Mata Verde, estrada para Lamban, E. Pereira 5788 (RB); Rio Vermelho,
Pedra Menina,
US); Concei?ao
Guaratuba,
Vargem da Ang?lica, Morro da Virada do Mato Virgem, Pirani et al s.n. (SP).?Paran?:
da entrada da Baia, Braga 2364 (RB); Itacar?, opp. Marungaia,
Dus?n 16511 (A); Mpio. Trjucas do Sol,
15117 (NY, US); Mpio. Antonica,
Rib. Lagoa Vermelha Haischbach
18102 (NY, US);
Matul?o, Haischbach
Fazenda
Morro
Mpio.
Morretes,
(NY).?Pernambuco:
do Cabo, Graziela
Col.
Florestal,
Recife,
Hatschbach
Mata
20206
27144
(RB); Mpio.
Senges, Rio do Funil, Hatschbach
de Janeiro: Cabo Fri?, Arraial
Irm?os, Lima 48-126
(RB).?Rio
Reserva Biol?gica
de Jacarepagu?, Lanna 124 (RB); Mpio. Maca?,
de Dois
7 (NY); Guanabara,

ANDIRA
2003
87
leaflet surface. D.
A. Habit. B. Node of rhachis with stipels. C. Abaxial
30. Andira fraxinifolia.
E. Calyx, opened to show inner surface. F. Standard petal, inner surface. G. Outer surface of wing petal
K. Gy
with lamellate sculpturing. H. Detail of lamellate sculpturing. I. Keel petal, inner surface. J. Androecium.
section. L. Fruit. M. Fruit in section, showing wall structure. O. Seed.
noecium, also shown above in longitudinal
FIG.
Flower.
(Based on: A-K,
R. T. Pennington
228; L-M,
R. T. Pennington
202)

88
60757065
55
VOLUME 64
403550
45
o? Andira fraxinifolia.
Restinga da Praia de Carapebus, H. C. de Lima 656 (RB); Mpio. Nova Igua?a, pr?x. a Lixeira, H. C. de Lima
N Joinvile, J. Mattos
1173 (RB); Mpio. Parati, Parati-Mirim, H. C. de Lima 3673 (RB).?Santa
Catarina:
12530 (SP); Campo Maseaiamb?,
Palho?a, Reitz & Burkart 5617 (US); Mpio.
Itaja?, Itaja?, Praia Braba, L. B.
Smith & Klein 7288 (NY, US); Brusque, Mata do Hoffman,
Veloso 98 (RB).?SAO
PAULO: Canan?ia, Una do
s.n. (SPF); near Moji-Mirim,
M.
Cardoso, Morro do Pereirinha, de Barros 2238 (SP); S?o Vicente, W. Hoehne
1477 (NY); Juqui?, Capueira, na margem do Rio Juqui?, M. Kuhlmann 4694 (SP); Ubatuba, Praia de
Kuhlmann
J. Mattos
13802 (SP); Mpio.
de Jureia, Pirani 818 (SPF); Mpio.
Perequer?-Ass?,
Iguape, esta?ao ecol?gica
Jundinhy, C. Smith 14 (SP); Sao Paulo, Instituto de Bot?nica, Sugiyama 888, 889 (SP).
Andira fraxinifolia
is a widespread and variable species, which is most likely to be
confused with A. anthelmia, A. ormosioides, and A. legalis in the rain forests and restinga
forests of Atlantic coastal Brazil, and with A. verm?fuga, in the cerrado woodlands of Cen
tral Brazil. Andira ormosioides, A. legalis, and A. anthelmia have flowers 18-24 mm long,
are 13-17 mm long. Andira fraxinifolia
whereas those o? A. fraxinifolia
also lacks the
large, persistent stipules that characterize A. legalis and A. anthelmia. The gynoecium of
A. verm?fuga is either glabrous or at most sparsely hairy on the upper and lower surfaces
it is uniformly hairy. The fruits of A. verm?fuga dry with
only, whereas in A. fraxinifolia
a wrinkled surface, whereas inA. fraxinifolia
they dry smooth. Furthermore, A. fraxinifo
lia does not grow in the cerrado woodlands, which is the main habitat for A. verm?fuga.
In the cerrado biome, A. fraxinifolia
is only occasionally
found in gallery forests.

2003
ANDIRA
89
Vegetative characters, particularly leaf and leaflet size and the amount and color of
the indumentum, are variable inA. fraxinifolia. Careful study of over 300 specimens from
the entire range of the species demonstrates that the variation in these features is contin
uous and therefore not a basis for maintaining
the species or varieties described by N. F.
Mattos and others (see synonymy above). My own field observations revealed consider
able variation in leaf and leaflet size and indumentum in very limited geographical areas
(e.g., specimens R.T. Pennington 211, 213 from Mara?, Bahia).
Several specimens from Jacobina, Bahia, are here placed in A. fraxinifolia but may
represent a new species (Pennington 1996). These specimens have fruit and flowers with
the characters of A. fraxinifolia,
but show an unusual growth form; they are small trees
with flattened rather than rounded crowns and with widely spreading branches.
Bentham (1837) published four names on the same date and same page; here their
types are included in one species for the first time. The epithet fraxinifolia was chosen be
cause it has been most widely
used.
ex Bentham, Comm. legum. gen. 45. 1837.?Type:

s.n. (holotype: BR!;
Rio
Santo:
Doce,
1827, Wied-Neuwied
Espirito
BM! K! M, NY!; photo of M isotype: F!).
17. Andira
n?tida Martius
Brazil.
isotypes:
shrub to tree 25 m tall; buttresses absent or very small; bark brown,

and
Assuring vertically
scaling (large trees), or pale brown, often marked with patches of
lichen (shrubs and small trees in restinga); slash pale red-brown to buff, oxidizing darker,
often with slight red ex?date; wood buff/cream; twigs red-brown to dark brown, or cov
Multi-stemmed
ered with pale buff bark (which may be scraped away to reveal dark bark beneath), hairy,
hairs red-brown, appressed, or glabrous; lenticels occasionally numerous, elongated, pale.
Stipules to 10mm long, to 1mm wide, caducous, hairy to sparsely hairy, glabrescent; leaf
axis 4-15 (-25) cm long; rhachis pale brown to brown, the pulvinus often darker, sparsely
hairy or glabrous, hairs short, red- brown, ? appressed; stipels 1 (-2) mm long, caducous;
(-5) pairs, 2.4-11.5
(-14) cm long, 1.7-4.5 (-6) cm wide, elliptic, broadly
ovate
ovate,
elliptic, narrowly
(rarely narrowly elliptic, suborbiculate to broadly ovate),
coriaceous to thick-chartaceous, base obtuse, rounded to slightly cordate, often slightly
decurrent, apex acute, obtuse, rounded, often slightly retuse or with a short acumen to 10
mm long, glabrous adaxially, very sparsely hairy abaxially or glabrous, hairs red-brown,
short (<0.2 mm long), appressed; primary vein channelled adaxially; secondary veins
6-13, ? plane to slightly sunken adaxially, ? plane to slightly raised abaxially, pattern eu
camptodromous becoming brochidodromous,
tertiary
leaflets in 2-4
veins plane adaxially and abaxially. Panicles 6-30 cm long, terminal and axillary, with
sparse to very sparse, short, appressed, red-brown hairs at branch tips, becoming less hairy
towards the base; bracts 1.5-2 mm long, narrow, caducous, sparsely to very sparsely hairy,
hairs red-brown, ? appressed; pedicels to 2 mm long or flowers sessile; bracteoles 0.5-1
mm long, indumentum like that of bracts. Flowers 10-13 mm long. Calyx 4-5.5 mm long,
red-brown to reddish to ? black, with sparse red-brown, appressed hairs or glabrous ex
cept around the margins of the lobes; lobes 0.2-0.5 mm, obtuse. Petals pinkish white to
purple, the standard pale with violet markings or pinkish with a central white marking;
standard blade 8-8.5 mm wide, 7-8 mm high, claw 3-3.5 mm long; wing 5.6-8 mm long,
3-3.5 mm wide, claw 3-4 mm long, lamellate sculpturing present; keel 5-7.5 mm long,
2.6-3.5 mm wide, claw 3-4.5 mm long. Stamens 8.5-10 mm long, united for the basal
4.7-8 mm, free for the distal 2-4.5 mm, vexillary stamen 6-7.5 mm long. Gynoecium

90
VOLUME 64
9-13 mm
long, glabrous or stipe, lower portion of style, upper and lower surfaces of the
ovary very sparsely hairy, hairs red-brown, appressed; stipe 2.5-5.5 mm long, ovary 3-5
mm long, style 2.2-4.5 mm long; ovules (1-) 2-4. Fruits 3-7 cm long, 2.5-5 cm high,
2.4-5.5 cm wide, elongated, weighing ca. 20 g or less when dry, green, often glaucous,
smooth or ridged from upper to lower suture, drying ? smooth or wrinkled, brown to very
dark brown with or without a waxy bloom; stipe to 8 mm long (many fruits breaking off
without a stipe); suture prominently raised adaxially, raised but obscure abaxially; stylar
remnant raised and obvious or imperceptible; mesocarp 4-7 mm thick when fresh, 1-2.5
mm
dry, pale green to green, fibrous, sweet-smelling,

drying brown, hard,
slightly granular; endocarp 2-5.5 mm thick, brown to dark brown, drying paler, fibrous.
Chromosome number unknown. Figs. 3C, 4A(iii), 5B, 13B, 32.
Phenology. Flowering September to January (toMay in Espirito Santo).
thick when
Distribution
(Fig. 33). Brazil (Bahia, Espirito Santo, Pernambuco, Rio de Janeiro); in

on white sand, closed restinga forest, and rain forest.
scrub
open restinga
Vernacular names. Angelim branco, angelim da praia (Bahia); angelim coco (Espir
ito Santo); angelim (Pernambuco).
REPRESENTATIVE Specimens.
Brazil. BAHIA: Santa Cruz da Cabr?lia, Almeida & dos Santos 103 (CEPEC,
Sa?da de Itaju do Colonia
356 (US); Mpio. Uru?uca, Dist. Serra Grande,
para Itap?, Km 20, Almeida
estrada Serra Grande-Ilh?us,
3 km from Distrito, Amorim et al. 347, 348 (CEPEC); Itacar?, Bel?m & R. S. Pin
heiro 2988 (MEXU, UB, US); Mpio.
et al.
Ilh?us, ca. 7 km along road Oliven?a-Vila
Brasil, A. M. de Carvalho
US);
3309 (CEPEC); Mpio. Porto Seguro, Esta??o Ecol?gica

do Pau Brasil, Euponino
167 (US); Mpio. Nova Vi?osa,
Km 9 along road Nova Vi?osa-Mucuri,
Farney et al. 2615 (RB); Mpio. Mucuri, Rio Mucuri, Farney et al. 2653
on road to
16 km S Cumuruxatiba
Salvador, pr?ximo ao aeroporto, M. L. Guedes 972 (RADAM);
(RB); Mpio.
et al. 18086 (K); Mpio.
Prado, Harley
Ilh?us, grounds of CEPLAC,
G. P. Lewis & A. M.
Prado, 12 km S Prado, estrada para Alcoba?a,
Copuva,
G. Hatschbach
Ubaitaba,
G. P. Lewis
& J. M.
756
791
(CEPEC, E, NY); Mpio.

Nova Vi?osa,
(K); Mpio.
1 km S Porto de Campinhos,
estrada para
(K); Mpio. Mara?,
1029 (K, MEXU); Mpio. Valen?a, Km 13 da estrada Valen?a para
1050 (K); Mpio. Belmonte,
estrada Belmonte-Itapebi,
Km 26, Mat
Guaibim, G. P. Lewis & A. M. de Carvalho
tos Silva & Hage 585 (CEPEC, K, UB); Mpio.
186 (CEPEC, FHO,
Ilh?us, just S of Oliven?a, R. T. Pennington
Santa Cruz da Cabr?lia,
10 km N Porto Seguro on road to Santa Cruz Cabr?lia, R. T. Pennington
&
K); Mpio.
G. P. Lewis 188 (CEPEC, FHO, K); Mpio.
Ilh?us, ca. 3 km W Oliven?a, R. T. Pennington
192,198
(CEPEC,
& A. M.
Silva
J. G. Jardim
de Carvalho
48748
de Carvalho
ca. 15 km S Itabuna on road BR-101, R. T. Pennington

et al. 209 (CEPEC, FHO,
Buerarema,
road Porto de Campinhos-Ubaitaba,
et al. 221 (CEPEC, FHO, K);
Marau,
et al. 235 (CEPEC, FHO, K); Mpio.
R. T. Pennington
Mpio. Una, near Pedras, Km 5 on road to Independencia,
et al. 288, 290, 291, 292 (CEPEC, FHO, K); Mpio. Alcoba?a,
Km
Salvador, dunas de Itapua, R. T. Pennington
FHO, K); Mpio.
K); Mpio.
4 on road Alcoba?a-Teixeira
de Freitas, R. T. Pennington
& F. A. de Carvalho
ca. 2 km N Alcoba?a,
1 km from the sea, R. T. Pennington
& F. A. de Carvalho 301 (CEPEC, FHO,
Alcoba?a,
K); Mpio. Mata de S?o Jo?o, estrada do Coco, em dire?ao a Sau?pe, G.C.P. Pinto 429/81
(RADAM); Mpio.
14 km N Serra Grande, off road to Itacar?, W. W. Thomas et al 9469 (CEPEC);
Itacar?, "Campo Cheiroso,"
Barrol?ndia, Est. Experimental
Mpio. Belmonte,
"Gregorio Bondar," 48 km E BR-101 on road to Belmonte, W.
W. Thomas et al. 9881 (CEPEC).?ESPIRITO
SANTO: Mpio. Vila Velha, restinga de Lagoa do Milho, D. Araujo
& Peixoto 324 (RB); Itaunas, G. P. Lewis et al. 1636 (K); Mpio. Linhares, Vale do Rio Doce, Reserva Florestal
da Cia., H. C. de Lima 1668 (RB); Mea?pe,
estrada Rodovia do Sol, 10 km depois de Guarapari, H. C. de Lima
2922 (K, MEXU); Mpio. Concei?ao
do Barra, Ita?nas, ap?s a ponte para a cidade velha, H. C. de Lima 2966
(MEXU, RB); Mpio. Linhares, Reserva Florestal de Linhares, Sucre 8656 (K, NY).?Pernambuco:
Recife,
Mata
de Dois
limaos, Andrade
Lima 55-1988
(K); Cabo, ?rea-projecto
Suape, Mata do Zumbi, Andrade-Lima
105 (F); Ilha Itamaraca, G. A. Ramage
s.n. (BM).?Rio
DE JANEIRO:Mpio. Quissam?, Mata
do Caio, Farney et al. 3425 (E, RB).?SERGIPE:
Mpio. Santo Amaro de Brotas, 8 km after "Terminal Portuario"
of Aracaju, Farney 2880 (RB); Mpio. Santo Amaro de Brotas, 5 km after "Terminal Portuario" of
Aracaju, Far
& Medeiros-Costa
ney 2930
Gomes
Santo Amaro de Brotas, Sap?-Fejeunde

(RB); Mpio.
Arauari, near Torre de Embratel, Farney
2994 (RB); Mpio. Estancia, Rod. Abais-BRIOl,
1 km W Abais, Mattos Silva et al. 3027 (CEPEC).

&
2003
ANDIRA 91
leaflet surface. C. Flower. D. Calyx, opened to show inner

surface of wing petal with lamellate sculpturing and detail of
sec
I. Gynoecium,
also shown below in longitudinal
sculpturing. G. Keel petal, inner surface. H. Androecium.
dried fruit. L. Fruit in section, showing wall structure. M. Seed. (Based on: A,
tion. J. Fresh fruit. K. Wrinkled
288 (leaves), H C. de Lima 1668 (inflorescence);
B, R. T. Pennington
C-I, A. Ribeiro 62; J, L, R. T. Pennington
292; K, R. T. Pennington
291.)
FIG.
32. Andira
surface. E. Standard
nitida. A. Habit.
petal,
B. Abaxial
inner surface. F. Outer

92
50
FIG.
55
45
33. Distribution
35
o? Andira
VOLUME64
40
nitida.
Andira nitida ismost likely to be confused with A. humilis, A. marauensis, and A. car
valhoi. Andira humilis is distinguished by its unique geoxylic suffrutex growth form, but if
this is not noted on herbarium specimens, the two species may be distinguished by A. ni
tida's smaller flowers (10-13 mm as compared to 14-16 (-19) mm inA. humilis). Further
more, A. humilis is a cerrado species that is not sympatric with A. nitida, which is restricted
to the rain forests and restingas of coastal Brazil. Andira nitida does grow alongside A. car
valhoi in Bahian restingas, but the latter is clearly distinct because of itsmuch larger (5-10
cm long, 4.8-8 cm high, 4.5-9 cm wide), brown fruits. Additional differential characters
are provided by the larger flowers (14-15 mm) and leaf axis (8.5-25 cm) of A. carvalhoi.
Andira nitida is also sympatric with A. marauensis
in Bahian coastal rain forest. The dif
ferences between these two species are outlined under A. marauensis
(no. 18).
Field observations in Bahia revealed two sympatric variants. One is a rain forest tree
(also growing in damp areas in restinga) with generally glabrous, thick-chartaceous leaflets
and a smaller, smooth fruit, which dries wrinkled. The other is a shrub or small tree, which
can tolerate dry restinga habitats; it has coriaceous, abaxially sparsely hairy leaflets and a
larger fruit, which
is ridged from the lower to the upper suture and dries smooth. Imade

ANDIRA
2003
93
(mostly sterile) of both growth forms in Bahia and found them

I initially considered them as possibly distinct species (A. n?tida
distinct morphologically.
and "A. sp. nov. 3" in Pennington 1995, 1996), and they were coded as separate terminal
taxa in cladistic analyses. Subsequent examination of more herbarium specimens showed
extensive
collections
that differences between the putative taxa are not consistent, especially at the northern and
southern ends of the range. It is not possible to ascribe either putative taxon a unique
combination of character states, and they are therefore not formally recognized.
18. Andira
Brazil.
N. F. Mattos, Loefgrenia 45: 1. 1970.?Type:
R.
P.
R.
12 Jan 1967,
Belem &
S. Pinheiro 3089 (holotype: UB!;
marauensis
Mara?,
Bahia:
isotype:
CEPEC!).
Tree to 35 m tall (though flowering from 3 m tall); buttresses absent; bark grey
brown, Assuring vertically and flaking in small plates; slash pale brown, oxidizing darker;
twigs brown to very dark brown, with very sparse appressed hairs, glabrescent; bark
thicker and cracking on older twigs; lenticels on older twigs, elongated. Stipules ca. 2.5
mm long, to 1mm wide, caducous, sparsely hairy; leaf axis 3-10 cm long; rhachis very
sparsely hairy, glabrescent, hairs short, red-brown, ? appressed; petiolules 3-7 mm long;
stipels not seen (either absent or early caducous); leaflets (1-) 2 (-3) pairs, 2.5-8.5 cm
terminal
long, 1.1-3.7 cm wide, elliptic, narrowly obovate (rarely broadly obovate),
leaflets often the most distinctly obovate, subcoriaceous, base acute to obtuse, slightly de
rounded (rarely slightly retuse or with an acumen to 2 mm long),
very
glabrous adaxially,
sparsely hairy abaxially, hairs red-brown, short (<0.2 mm long),
vein
channelled adaxially; secondary veins 8-10, ? plane adaxially,
appressed; primary
raised
slightly
abaxially, pattern brochidodromous,
tertiary veins plane adaxially and
or
cm
raised
Panicles
5-11
plane
slightly
abaxially.
long, terminal and axillary, with
at
red-brown
hairs
branch
sparse, short, appressed
tips, glabrescent towards the base;
bracts narrow, caducous, ca. 2 mm long, sparsely hairy, hairs red-brown, ? appressed;
1-2 mm long; bracteoles caducous, not seen. Flowers 9-11 mm long. Calyx
pedicels
4^.5 mm long, deep purple, glabrous except around the margins of the lobes; lobes
0.2-0.3 mm, obtuse, shallow. Petals pink to purple; standard blade 7.5-10 mm wide, 6-8
mm high, claw 2.5-3.5 mm long; wing 5-7 mm long, 2.5-3.5 mm wide, claw 4-4.5 mm
long, lamellate sculpturing present; keel 5-7 mm long, 3-3.5 mm wide, claw 3.5-4.5 mm
current, apex obtuse,
long. Stamens 8-9 mm long, united for the basal 4-6.5 mm, free for the distal 2-3 mm,
vexillary stamen 5.5-6 mm long. Gynoecium 9.5-12.5 mm long, glabrous or with 1, 2, or
3 hairs; stipe 3.5-5 mm long, ovary 3-4 mm long, style 3-3.5 mm long; ovules 1-2. Fruit
weighing ca. 20 g or less when dry, drying smooth (only seen as rotted fragments on the
forest floor; pers. obs.). Chromosome
number unknown.
Phenology. Flowering January (3 records) and May (1 record).

Distribution
(Fig. 34). Brazil (Bahia); in Atlantic coastal rain forest.
Vernacular
Additional
names.
Specimens
no Km 46 da Rod. BA-001
R. P. Bel?m & Pinheiro
Angelim.
Examined.
Brazil.
(Ilh?us-Una),
Amorim
Bah?a: Mpio. Una, Reserva Biolog?a do Mico Le?o, estrada

et al. 1411,1238,1908
(CEPEC, NY); Mpio. Mara?, Mara?,
(MEXU, UB, US); Mpio. Uru?uca, Dist. de Serra Grande, 7.3 km
3091 (NY, UB), 3142

na estrada Serra Grande-Itacor?,
et al. 3510 (CEPEC, NY); Mpio. Una, Reserva Biolog?a do
A. M. de Carvalho
Mico Le?o, estrada no Km 46 da Rod. BA001
/. G. Jardim et al. 90 (CEPEC, FHO); Mpio. Una,
(Ilh?us-Una),
border of Fazendas Maruim
on road Oliven?a-Buerarema,
and Dois de Julho, 33 km SW Oliven?a
Maruim,

94
50
FIG.
Mori
et al
13800
102 (CEPEC, K-2
(NY); Mpio.
34. Distribution
Ilh?us, Fazenda
35
45
55
of Andira
VOLUME 64
40
marauensis.
Barra do Manguinho,
Km
11 da Rod.
Ilh?us-Oliven?a,
Voeks
sheets).
ismorphologically
Andira marauensis
similar to rain forest trees of A. nitida, but it
differs in its smaller leaves, its obovate, blunt-ended leaflets with an acute to obtuse base,
smaller flowers, and fewer ovules. Itmust be considered endangered given the tiny area
of suitable rain forest habitat remaining in southern Bahia.
19. Andira carvalhoi R. T. Pennington & H. C. Lima, Kew Bull. 50: 559. 1995.?TYPE:
Brazil.
Bahia: Mpio. Ilh?us, estrada Oliven?a-Vila
Brasil, a 7 km de Oliven?a,
13 Jan 1981, A. M. de Carvalho et al. 491 (holotype: CEPEC!; isotype: RB!).
with creeping rhizomes, or occasionally a small tree to
Shrub, often multi-stemmed
10m tall with spreading branches; buttresses absent; bark grey-brown to brown, Assuring
vertically and flaking slightly; slash 2-4 mm thick, pale brown to red-brown, some red ex

2003
ANDIRA
95
udate; wood hard, dense, pale brown to cream, streaked (in section: with concentric rings
of brown and buff and a dark ring at center); twigs brown to dark brown with whitish
waxy bloom, older twigs grey-brown, bark splitting vertically; very sparsely hairy, hairs
short, appressed, red-brown, glabrescent. Stipules to 14 mm long, to 1 mm wide, with
sparse, red-brown appressed hairs, glabrescent; leaf axis 8.5-25 cm long (^10 cm long,
sterile branches); rhachis with very sparse, red-brown, appressed hairs, glabrescent, cov
ered with waxy bloom; stipels to 1mm long; petiolules 4-10 mm long, indumentum like
that of rhachis; leaflets in 2-3 (-4) pairs, 6.7-18 cm long (proximal leaflets sometimes 4.5
mm long), 2.8-7 cm wide (proximal leaflets sometimes 2.3 mm wide), elliptic to narrowly
ovate, coriaceous, dark green, shiny, the venation paler, older leaflets tending to become
b?llate with themargins inrolled, base obtuse, rounded (rarely ? truncate), apex obtuse to
rounded (rarely acute), often slightly retuse or with an acumen 3^1 mm long, glabrous
adaxially, glabrous abaxially except the primary vein with very sparse, appressed, red
brown hairs; primary vein channelled adaxially; secondary veins 8-13, plane adaxially,
slightly raised abaxially, pattern brochidodromous,
divergence angle wide, tertiary veins
cm
Panicles
and
5-30
abaxially.
plane adaxially
long, terminal, with red-brown, appressed
hairs at branch tips, glabrescent at base, the axes dark, almost black, glaucous at base;
bracts 4-5 mm
long, narrow, moderately

persistent, with short, appressed, red-brown
mm
mm long, narrow, moderately persistent,
bracteoles
1.5-3
1.5-2
hairs; pedicels
long;
mm
indumentum like that of bracts. Flowers 14-15
long. Calyx 6-7 mm long, with very
sparse, appressed, pale red-brown, short hairs; lobes 0.5-1 mm long, acute to obtuse.
Petals violet, the standard with a central white marking; standard blade 10 mm high, 12
mm wide, claw 5 mm long; wing 9 mm long, 4-4.5 mm wide, claw 6 mm long, lamellate
sculpturing present; keel 8 mm long, 3.5-4 mm wide, claw 6 mm long. Stamens 11mm
long, filaments united for the basal 5.5-6.5 mm, free for the distal 4.5 mm, vexillary sta
men 9-9.5 mm long. Gynoecium
13-13.5 mm long, with sparse short appressed hairs on
the lower surface of the ovary; stipe 4.5 mm long, ovary 4.5 mm long, style 4-4.5 mm
long; ovules 3. Fruits 5-10 cm long, 4.8-8 cm high, 4.5-9 cm wide, elongated, weighing
100-300 g when dry, flattened adaxially, rough, pale brown with dark brown specks (both
fresh and dry); stipe 5-13 mm long; suture raised adaxially and abaxially; stylar remnant
not visible; mesocarp 6-15 mm thick (fresh), pale greenish white to pale green, slowly ox
idizing red-brown when cut, drying pale brown, hard, dry, finely granular; mesocarp 1.5-2
mm thick; endocarp 3-9 mm thick (fresh), brown, woody, fibrous. Chromosome number
unknown. Figs. 3F, 4A(ii), 35.
Phenology. Flowering in October.
Distribution
(Fig. 36). Brazil (Bahia); in open scrubby restinga on white
sand.
Additional
Specimens
Examined.
Brazil.
Bah?a:
rod. BR-030,
Porto
de
Mara?,
Mpio.
et al. 178 (CEPEC, K); Mpio.
Km II, A. M. de Carvalho
Ilh?us, estrada Ilh?us-Canavieiras,
Campinhos-Mara?,
Km 33, A. M. de Carvalho 622 (CEPEC); Mpio.
Km 9, A. M. de Carvalho &
Ilh?us, estrada Oliven?a-Maruim,
Faria 2547 (CEPEC, UEFS); Mpio. Dh?us, 7 km on road Oliven?a-Vila
et al. 3308
Brasil, A. M. de Carvalho
Km 10, Gentry & Zardini 50011
Ilh?us, estrada Pontal-Oliven?a,
(CEPEC); Mpio.
Ilh?us,
(CEPEC); Mpio.
estrada Oliven?a-Maruim,
Km 5-8, M. P. M. de Lima et al. 20 (CEPEC, RB-2 sheets); Mpio.
Ilh?us, estrada
Km 7-10, Martinelli
et al. 11102 (CEPEC, RB); Mpio.
Km 5,
Ilh?us, estrada Oliven?a-Una,
Oliven?a-Maruim,
Fazenda Jairi, Mattos Silva et al. 1199 (CEPEC, K); Mpio.
ramai com en
Ilh?us, Fazenda Barra do Manguinho,
trada no Km
10 da estrada
Km 10, Mattos
Silva et al. 1393 (CEPEC, K); Mpio.
Ilh?us,
Pontal-Oliven?a,
?
Fazenda
Barra
do
ramai com entrada no Km
10 da estrada
(junto
Manguinho),
Mattos
Silva et al 1877 (CEPEC, K); Mpio.
Km 6, R. T.
Ilh?us, dirt road Oliven?a-Maruim,
Ilh?us-Oliven?a,
et al 181 (CEPEC, FHO, K); Mpio.
R. T. Pennington
Pennington
Ilh?us, just S Oliven?a,
184, 185, 229
197 (CEPEC, FHO, K); Mpio. Mara?,
Ilh?us, ca. 3 km N Oliven?a, R. T. Pennington
Fazenda
Guanabara

96
FIG.
opened
35. Andira
to show
carvalhoi.
inner surface.
A. Habit.
B. Abaxial
F. Standard
233; C, A. M.
surface.
C.
de Carvalho
D. Flower. E. Calyx,
of wing petal with lamellate
also shown
J. Gynoecium,
Inflorescence.
petal, inner surface. G. Outer surface

H. Keel petal, inner surface. I. Androecium.
sculpturing and detail of sculpturing.

section. K. Fruit. L. Fruit
below in longitudinal
T Pennington
leaflet
VOLUME 64
in section, showing wall structure. M.

491; D-J, M. P. M. de Lima et al. 20; K-L,

Seed. (Based on: A, B, R.

R. T. Pennington
197.)
2003
ANDIRA
50
55
45
o?Andira
97
35
40
carvalhoi.
road Mara?-Porto
de Campinhos Km 20, R. T. Pennington

et al 216, 217 (CEPEC, FHO, K); Mpio. Una, road
Km 42, ca. 0.5 km down road to Pedras, R. T. Pennington
et al 232, 233 (CEPEC, FHO, K);
8.9 km from Oliven?a, W. W. Thomas et al 8969 (CEPEC, NY); Mpio.
Ilh?us, Road Oliven?a-Maruim,
Mpio.
then 3 km W, W. W. Thomas et al 9726 (CEPEC, NY); Mpio.
Ilh?us, 10 km S Hh?us airport on road to Oliven?a,
on road toMaruim, W. W. Thomas et al. 10950 (E, NY).
Ilh?us, 8.9 km SW Oliven?a
Oliven?a-Una,
Andira carvalhoi is only likely to be confused with A. n?tida from which it differs by
itsmuch larger, brown fruits. It also has larger flowers and a longer leaf axis; the flowers
of A. n?tida are 10-13 mm long and the leaf axis rarely exceeds 15 cm.
Although A. carvalhoi is abundant in some sites (e.g., at Oliven?a, Bahia), itmust be
considered endangered because of its restricted range. Its habitat of sandy coastal restinga
is under great pressure for development
and is highly prone to soil erosion if cleared.
the fruits of A. carvalhoi are dispersed by large rodents (Pennington & de Lima
Moreover,
1995) and may not be distributed adequately in the absence of these animals, as has
been demonstrated for Hymenaea
courbaril (Leguminosae-Caesalpinioideae;
Asquith et
al. 1999). It seems likely that many areas of restinga where A. carvalhoi grows do not

98
support healthy populations of large rodents, given

vegetation and proximity to centers of population.
VOLUME 64
the high levels of disturbance
of the
Brazil.
parviflora Ducke in Arq. Inst. Biol. Veg. 2(1): 47. 1935.?Type:
Amazonas: Manaus, estrada do Aleixo, 27 Apr 1932, A. Ducke RB 23865 (lecto
type, here designated: RB!; isolectotypes: F! G! K! RB-2 sheets! U!).
20. Andira
Tree to 20 m tall, buttresses absent; bark smooth, pale brown, cracking vertically, not
flaking, some horizontal markings; slash bright orange-brown; wood cream; twigs pale
brown (or dark brown), with red-brown, erect, tangled hairs, glabrescent; bark Assuring
vertically on older twigs. Stipules to 6 mm long, up to 5 mm wide, densely covered with
erect, tangled red-brown hairs; leaf axis 2.5-17.5 cm long; rhachis with red-brown erect,
tangled hairs; stipels 0.5-1.5 mm long; petiolules 0.5-1.5 mm long, indumentum like that
of rhachis; leaflets in (2-) 3^1 (-5) pairs, 2-10 cm long, 1-4 cm wide, narrowly obovate,
elliptic (rarely oblanceolate), coriaceous, base obtuse or rounded (rarely slightly cordate),
apex obtuse, often with a short acumen to 5 mm long, glabrous adaxially, abaxially with
sparse red-brown, erect hairs, >0.2-1.0 mm long, the veins more densely hairy; primary
vein channelled adaxially; secondary veins 10-15, slightly sunken adaxially, raised abaxi
the brochidodromous
veins
ally, pattern eucamptodromous
anastamosing very close to themargin, tertiary veins impressed to slightly impressed adax
ially and raised abaxially. Panicles 3-18 cm long, axillary and terminal, densely covered
with red-brown, erect, tangled hairs; bracts 2.5-3 mm long, 1.5-2.5 mm wide, densely cov
ered with pale brown, ? appressed hairs; bracteoles 1.5-2 mm long, 0.75-1 mm wide, in
dumentum like that of bracts. Flowers 6-7.5 mm long. Calyx 3-4 mm long, lilac to purple,
with
sparse, appressed, pale brown hairs. Petals sessile, whitish, the standard with purple
marking; standard blade 6.2-7.5 mm wide, 5-5.5 mm high, claw 1.5mm long; wing 4.5-5
mm long, 2.8-3.2 mm wide, claw 2-2.5 mm long, sculpturing absent; keel 4-5 mm long,
2-2.5 mm wide, claw 2.5-3 mm long. Stamens 5 mm long, filaments united for the basal
1.5-2.5 mm, free for the distal 2-3 mm, vexillary stamen 3 mm long. Gynoecium 4.7-5.2
mm long, the ovary with pale brown, ? appressed hairs on upper and lower surfaces, indu
mentum extending to the top of the stipe and base of the style with scattered hairs at the
sides of the ovary, or ovary sparsely hairy on upper and lower surfaces only; stipe 1-1.2
mm long, ovary 2 mm long, style 1.5-2 mm long; ovules 1-2. Fruits 3-4.1 cm long,
2.4-3.2 cm high, 2.4-3.2 cm wide, ? globose to elongated, weighing ca. 20 g or less when
dry, green, drying red-brown to dark brown to almost black, appearing smooth but some
what tuberculate (best seen with lens or microscope);
stipe insignificant (fruit ? sessile); su
ture indistinct adaxially and abaxially; stylar remnant tiny; mesocarp
1.5 mm thick, pale
mm
1.5-4
thick
brown, granular, hard; endocarp
(thickened along upper side), cream to
non-fibrous.
Chromosome
number
unknown. Fig. 37.
pale brown, hard,
to
Phenology. Flowering February
July.
Distribution
(Fig. 38). Brazil (Amazonas, Para); in terra firme forest on sandy and
clay soil, and also in low forest on white sand.
names.
Vernacular
Sucupira
vermelha,
acupu
rana,
sucupira
chorona
(Amazonas);
andira uchi (Para).

Additional
Rod.
BR-174,
INPA/WWF
Specimens
Km
64,
1302.4500.2
Examined.
Brazil.
then 34 km E on ZF3,
Amazonas:
Fazenda
(K); estrada Manaus-ltacoatiara,
Distrito Agropecuario
da Suframa,
1302 of DBFF project, Ackerly et al
Km 26, Reserva Florestal Ducke, Adair s.n. (INPA);
Esteio,
Manaus,
Reserve

ANDIRA
2003
99
V?rSeUAAM yC?ltf_s
leaflet surface. E. Flower.

leaflet surface. D. Adaxial
FIG. 37. Andira parviflora. A, B. Habit. C. Abaxial
F. Calyx, opened to show inner surface. G. Standard petal, outer surface. H. Wing petal, outer surface. I. Keel
in longitudinal
K. Gynoecium,
also shown below
section. L. Entire fruit
petal, inner surface. J. Androecium.
(above) and in section (below), showing wall structure. (Based on: A, E-K, E. Soares 106; B-D, W. Rodrigues
11179; L, G. T. Prance
et al 9074.)

100
FIG.
38. Distribution
of Andira
parviflora,
A. cujabensis,
VOLUME 64
and A. cordata.
Manaus, Reserva Florestal Ducke, Alu?sio 115 (INRA); Manaus, Reserva Florestal Ducke, L. Co?lho s.n. (INPA);
near Rio Taruma, near Cachoeira Alta, Ducke 2229 (COL, INPA); Manaus, Ducke 21348 (F); Manaus,
estrada
90 km NNE Manaus, Distrito Agropecuario
da S?frama, re
do Taruma, Ducke RB 23866 (RB); Mpio. Manaus,
serve 1501 of DBFFproject,
"Km 41," Kukle 23 (K), Kukle & Boom 44 (K, MEXU);
Proj. Radam, Rio Cauabari,
afl. do Rio Negro, Marinho
526 (NY); Manaus,
track from Km 63, road Manaus-Itacoatiara,
Prance et al 9074
Km 60, Prance & Ramos 23563 (US); estrada Manaus-Itacoat
(COL, F, GH, K); estrada Manaus-Caracara?,
do inventario florestal, W. Rodrigues
7990 (INPA); estrada Manaus-Itacoatiara,
iara, Km 130, ?rvore XIV-104
reserve Km 51, W. Rodrigues
et al 11179
8484 (INPA); Manaus, WWF/INPA
DBFF
85, W. Rodrigues
Km 30, M. F. Silva et al. 91 (F); Rio Uaup?s,
island above rapids
(FHO, INPA, K); estrada Manaus-Caracara?,
at Ipanor?, D. W. Stevenson et al 958 (K).?Para:
do aeroporto, L. S. Co?lho et al. 270
C.PT ?as proximidades
estrada da barragem do Caran?, pr?ximo ao igarap?, E. Soares 104 (INPA).
(INPA); Porto Trombetas,
Km
This distinctive species is the only species o? Andira from the Amazonian region with
red-brown hairs on the leaflet undersurfaces. The timber of A. parviflora is used for con
struction and general carpentry (Loureiro & da Silva 1968).
21. Andira
cujabensis Bentham, J. Proc. Linn. Soc, Bot. 4, Suppl. ("A Synopsis of the
tribe Dalbergieae):
120. 1860. Vouacapoua cujabensis (Bentham) Kuntze, Revis,
gen. pi. 1: 212. 1891.?TYPE: BRAZIL. Goi?s: between Arragas and Navidade,
Gardner 3654 (holotype: K!; isotypes: BM! E-2 sheets! F! G! GH-2 sheets! K!).

2003
ANDIRA
101
lanei N. F. Mattos, Loefgrenia 40: 1. 1970.?TYPE: BRAZIL. Mato Grosso:

Tres Lagoas, Fazenda Canaan, Feb 1969, F. Lane s.n. (holotype: HB; isotype:
K!).
Andira
a shrub) to 12 m; buttresses absent; bark thick, fissured; slash

(occasionally
wood
honey-brown;
pale straw-colored; twigs dark brown to pale brown, densely hairy,
hairs pale brown to pale red-brown, erect, tangled, bark splitting on older twigs to reveal
paler bark beneath; lenticels often numerous, pale, elongated. Stipules to 5 mm long, 3
Tree
mm wide
or narrower, early caducous, densely covered with pale brown to red-brown

hairs; leaf axis 5-23 cm long; rhachis densely hairy, glabrescent, hairs buff to pale brown
to red-brown, erect, tangled; stipels 1-2 mm long, caducous; petiolules 1-5 mm long, in
dumentum like that of rhachis; leaflets in (2-) 3^4 (-5) pairs, (3-) 4-10.5 cm long, 2-5.5
cm wide,
broadly elliptic, elliptic, ovate, narrowly ovate (rarely broadly obovate to nar
coriaceous, base rounded to cordate, apex obtuse (rarely acute or
obovate),
rowly
a short blunt acumen to 5 mm long or slightly retuse, glabrous adax
often
with
rounded),
ially, densely hairy abaxially and the epidermis often not visible, glabrescent, hairs gen
erally buff to pale brown, occasionally red-brown, erect, tangled, >0.2-1.0 mm long; pri
mary vein channelled adaxially; secondary veins 8-10, impressed to slightly impressed
or ?
adaxially, raised abaxially, pattern eucamptodromous
becoming brochidodromous
completely brochidodromous,
tertiary veins plane adaxially and raised abaxially. Panicles
(4.5-) 10-32 cm long, axillary and terminal, densely covered with pale red-brown, long,
tangled hairs; bracts 1.5-2 mm long, 1mm wide, caducous, with long, pale red-brown
hairs; bracteoles 1mm long, 0.5 mm wide, indumentum like that of bracts. Flowers 5.5-7
mm long, sessile. Calyx 3^4- mm long, black to dark purple, sparsely to very sparsely
hairy, the margins of lobes hairy, hairs long, red-brown; lobes 0.2-0.5 mm long, obtuse,
sometimes with the apex
markings; standard blade
long, 1.5-2.2 mm wide,
2.5-3 mm long. Stamens
acuminate. Petals dirty white, the standard with mauve-purple

5-7 mm wide, 4-5 mm high, claw 2 mm long; wing 3.5-4.5 mm
sculpturing absent; keel 2.5-4 mm long, 1-2 mm wide, claw
4.5-5.5 mm long, filaments united for the basal (1.8-) 2.8-3.5
free for the distal 1.5-2 mm; vexillary stamen 4 mm long. Gynoecium
5-5.75 mm
on
or
on
its
lower
surface
both
the
and
lower
surfaces, or the en
upper
long, ovary hairy
tire distal half of the ovary hairy, hairs long, pale; stipe 1.5-2 mm long, ovary 1.5-2 mm
long, style 1.75-2 mm long; ovules 1-2. Fruit 2.6-4.2 cm long, 2-3.2 cm high, 2-3.3 cm
mm,
wide, ? globose, weighing ca. 20 g or less when dry, green, drying brown to dark brown,
smooth, appearing smooth but minutely tuberculate (best seen with lens or microscope);
stipe \-4 mm long; suture a slight depression adaxially, not visible abaxially; stylar rem
nant absent; mesocarp
1.5-2 mm thick, hard, granular, tinged slightly greenish; endocarp
1-2.5 mm thick, pale brown, woody, very hard, non-fibrous. Chromosome
number un
known. Fig. 2D.
Phenology. Flowering January toMarch (occasional records as early as November or
as late asMay).
Distribution (Fig. 38). Brazil (Mato Grosso, Mato Grosso do Sul, Goi?s, Para); in cer
rado and gallery forest. Oliveira Filho (1992) demonstrated
that near Cuiab? (Mato
Grosso), A. cujabensis can tolerate seasonal water-logging, which may be the explanation
for its ability to grow in gallery forest.
Vernacular names. Angelim branco, angelim do cerrado, cascudinho,
Grosso); angelim amargoso (Goi?s); andira (Para).

sucupira (Mato
VOLUME64
102
Additional
W. R. Anderson
38339
Examined.
7150 (NY);
8 km N Terezina by road, W. R. Anderson
Brazil. Goi?s:
ca. 15 km S of Goi?s Velho,
7293 (NY, QCA); Serra Dourada,
by road, W. R. Anderson
& Kummrow
(NY); Urua?u, Elias de Paula 3277 (UB); arredores de Guarai, Hatschbach
Specimens
11 km E of Cavalcante
10051
(K); Serra do Caiap?,

ca. 8 km S Cavalcante,
eiros,
UB, US); Mpio. Piren?polis,

Hidrel?trica
da Serra da Mesa,
50 km S Caiap?nia,
road to Jata?, Irwin et al 17962 (INPA); Chapada dos Vead
Irwin et al. 23984 (F, G, UB); 8 km S Niquel?ndia,
Irwin et al. 34872 (MO, NY,
Mac?do
4352 (US); margem direita do Rio Tocantins,
canteiro de obras da Usina
B. A. S. Pereira et al 1552 (NY); Rio Java?s, /. M. Pires & M. R. Santos 16257

(INPA); estrada para Formoso, a 6 km W de Cariri, J. M. Pires & M. R. Santos 16637 (US); Serra Dourada, Rizzo
4142 (RB); Mpio. Santa Isabel, Una do Bananal, Parque Nacional
do Araguaia,
Fazenda Bareira Branca, cam
inho para Riozinho,
F. C. da Silva et al 257 (SP, UB); Mpio. Goi?s, Souza Lima 208 (RADAM, RB).?Mato
GROSSO: 35 km ENE Barra do Gar?as, W. R. Anderson

at western
11293 (SPF); Mpio. Chapada dos Guimar?es,
9691
(NY); Mpio. Coxim, Rio Taquar?, W. R. Anderson

11361
edge of Chapada dos Guimar?es, W. R. Anderson
Geographical
Society Base Camp, de Castro R2102 (E, K, U,
22 km S of Royal
(NY); Corrego de Maribondo,
a 25 km da BR-163,
estrada
Fazenda Mission?ria,
Rio Teles Pires,
UB); Mpio.
para Porto dos Gauchos
Sinop,
et al. 6259 (F); Mpio. Cuiab?, Rio Caxipozinho,
Cachoeira Cachoeir?o,
Cid Ferreira
pr?ximo a Cachoeira V?u
et al. 6541 (INPA); Mpio. Nova Andradina,
de Noiva, Cid Ferreira
Casa Branca, Hatschbach
31876
(NY);
34085 (MEXU, US); 1 km NE Garap?, Irwin & Soder str?m
Mpio. Cuiab?, Parque Aguas Quentes, Hatschbach
6551 (F, US); Serra do Roncador, Rio Turvo, 210 km N Xavantina,
Irwin et al 16179 (F, G); 15 km S Xavan
of Serra do Agaupei,
J. H. Kirkbride & Lieras
tina, Irwin et al. 16873 (F, GH); Km 197 on road from Caceres-S
3024 (F, US); 8 km S Xavantina, M. C. G. Kirkbride
near old road,
1597 (UB); Livramento,
Fazenda Rozalina,
M. Mac?do
et al. 56394 (FHO,
& Assump??o
2265 (UB); 100 km N Cuiab? on road to Diamantino,
Maguire
NY, UB, US);
Luciara,
Santa Terezinha,
(SP); vicinity of Veu de Noiva,

ena, Prance & Schaller 26266
Chapada
Santa Terezinha, perto do Lago Portugu?s, /. Mattos

15539
Prance et al 18965 (K, U, US); Fazenda Santa Filom
270 km N Xavantina, Royal Geographical
Society Base Camp, Ramos
Fazenda
dos Guimar?es,
(GH, NY);
& R. Souza 156 (K); Vale de Sonhos, 80 km N Barra do Gar?as on road to Xavantina,
Ratter et al 2323 (E, K,
U, UB); Pantanal, Rio Negro, Fazenda Santa Teresa, Schaller 160 (NY); Km 165 da rodovia Cuiab?-Santar?m
entroncamento, M. G. Silva & Rosario 4939 (NY); Mpio. Luciara, lake 2 km NW Luciara, W. W. Thomas et al
4312
51
do Sul: Mpio. Nova Andradina, MS-134

a 30 km de Nova Andradina, Leite & Klein
a 22 km de Nova Andradina,
Nova Andradina,
MS-134
Pastore
& Klein
77
do Rio Araguaia,
? Santana do Araguaia,
Lemes & Mileski
158
margems
pr?ximo
Grosso
(K).?Mato
(RADAM,
RB); Mpio.
(RADAM).?Para:
Fazenda
(RADAM, RB); near Reden??o,
Inaj?, N.T. Silva 4808 (NY, US).
Prof. Getulino,
Ratter
et al. 6869V
(E); Rio Araguaia,
Fazenda
do Rio
In the Brazilian cerrados, sterile specimens of A. cujabensis might be confused with

A. verm?fuga-, however, the smaller (5.5-7 mm long), white flowers of A. cujabensis are
completely distinct from the larger (12.5-18 mm long), pink to purple flowers of A. ver
m?fuga. For a separation from A. cordata see that species (no. 22).
Although Bentham (1860) spelled the specific epithet "cujabensis" in the protologue,
two orthographic variants have been widely used, "cuiabensis" (Bentham 1862; I have
used this incorrect spelling on many determination slips) and "cuyabensis" (Mattos 1979).
22. Andira
Arroyo ex R. T. Pennington & H. C. Lima, Kew Bull. 50: 562.

Brazil.
Bahia: 30 km W Barreiras, 12 Jan 1977, G. Hatschbach
MBM!;
(holotype:
isotypes: G! K! MEXU!).
cordata
1995.?Type:
39477
a shrub; buttresses absent; bark thick (to 1 cm),

Tree 6 (-15) m tall, occasionally
to grey-brown, Assuring vertically; slash pale red-brown, 5-20 mm thick, a little
red ex?date; wood buff streaked with cream; twigs dark brown to brown, older twigs
paler, glabrous (or rarely sparsely hairy, glabrescent); lenticels numerous, elongated, pale
brown. Stipules early caducous, not seen; leaf axis 4.5-15 cm long; rhachis glabrous (or
rarely with sparse, pale brown, erect hairs), dark brown peeling to reveal red-brown or
pale brown beneath; stipels 0.5 mm long, caducous; petiolules 1.5-4 mm long, glabrous
brown
(or sparsely hairy); leaflets in 2-4 pairs, 2-7 cm long, 1.4^- cm wide

(the lower leaflets
2003
103
ANDIRA
to 5 cm wide),
broadly elliptic to ovate, (rarely suborbiculate, elliptic, narrowly ovate to

ovate),
broadly
thinly coriaceous, base truncate, cordate (rarely rounded); apex obtuse to
rounded (the terminal leaflets sometimes truncate), often slightly retuse, occasionally with
an acumen to 4 mm long, glabrous (rarely very sparsely hairy abaxially, hairs erect); pri
mary vein channelled adaxially; secondary veins 8-10, plane to slightly impressed adaxi
ter
ally, slightly raised abaxially, pattern eucamptodromous becoming brochidodromous,
cm
to
Panicles
5-25
veins
raised
and
abaxially.
long,
slightly
tiary
plane
plane adaxially
axillary and terminal, sparsely covered with pale-brown hairs at branch tips, glabrescent
at base; bracts and bracteoles caducous, not seen. Flowers 6.5-7 mm long, sessile. Calyx
3-3.5 mm long, purple to black, glabrous (or with a few scattered pale appressed hairs),
except themargins of the lobes sparsely hairy, hairs pale, ? erect; lobes 0.2-0.4 mm long,
obtuse. Petals whitish to pale lilac, the standard with purplish markings; standard blade
5-6.5 mm wide, 4.5-5 mm high, claw 2-2.5 mm long; wing 4-4.5 mm long, 1.5-2.5 mm
wide, claw 2.5-3 mm long, sculpturing absent; keel 3-3.5 mm long, 1.5-2 mm wide, claw
3 mm long. Stamens 5.5-6 mm long, filaments united for the basal 2.5-4 mm, free for the
5.5-6.5 mm long,
distal 1.5-2.5 mm; vexillary stamen 3.5-4 mm long. Gynoecium
mm
mm
mm
2-2.5
1.75-2
long; ovules 2.
long, style
long, ovary
glabrous, stipe 1.75-2.2
cm
cm
?
cm
Fruit 3-3.6
wide,
high, 2.5-2.9
globose, weighing ca. 20 g or
long, 2.5-2.9
less when dry, green with pale yellowish specks when young, drying dark brown, almost
tuberculate (best seen with lens or microscope);
appearing smooth but minutely
mm
remnant
not apparent; mesocarp 1-2 mm thick, gran
sutures
and
3.5
stylar
stipe
long;
black,
ular, hard; endocarp 2^- mm
thick. Chromosome
January to April.
number unknown. Figs. 3A, 4B(i),
Phenology. Flowering
Distribution
(Fig. 38). Brazil (Maranh?o, Bahia, Goi?s, Tocantins);
Vernacular name. Gr?o de galo (Bahia).
Additional
Anderson
in cerrado.
ca. 100 km WSW Barreiras, W. R.

Examined.
Brazil. Bahia: Espig?o Mestre,
(NY), 36797 (NY, UB); Mpio. Correntina, Faz. Jatob?, margem da estrada que da acesso
da Silva 1352 (UB); Mpio. Barreiras, Elias de Paula 3122 (UB); Mpio. Correntina, Fazenda
& Collares
Rio Formoso, M. M. Fern?ndez
17 (RADAM, RB); drainage of Rio Corrente,
Specimens
et al 36607
a guarita, Aparecida
Formoso do Guara,
Rio Piau, ca. 150 km WSW
al. 31521
39.
(UB); Mpio.
Barreiras, Irwin et al 14855 (F, G); Mpio. Barreiras, 5 km NW Barreiras, Irwin et

ca. 2 km up road to airport, R. T. Pennington
& Brito 261, 262, 263, 264
Cocos, Lagoa do Pratud?o, S. B. da Silva & M. G. de Lima 372 (NY); Mpio. Formoso
Barreiras,

do Rio Preto, proximo ao rio Riach?o, pr?ximo a vereda Olhas d'Agua, Walter et al. 228 (US).?GOI?S:
Serra
G?rai de Goi?s, Rio da Prata, 6 km S de Posse, Irwin et al. 14420 (F, G, NY); Rio Corda, regi?o de Xamboi?,
E. Oliveira
1446 (UB); 10?45'S, 47?15'W, Orlandi 91 (RADAM, RB).?MARANH?O:
7?12'S, 47?25'W, J. A.
Ferreira
& C. A. Miranda
304 (RADAM, RB); Mpio. Graja?, J. A. Ferreira & C. A. Miranda
322 (RADAM,
& G. Gottsberger
15-9482
S?o
Codo, Fazenda Canto do Ro?a, /. Go?tsberger
RB); Mpio.
(NY); Mpio.
em dire?ao ? Balsas, C. A. Miranda
& J. A. Ferreira
Raimundo
das Mangabeiras,
355 (RADAM, RB); duas
Black 1630a (NY); near Carolina on road N to Estreito,
transect area 2,
leguas de Carolina, J. M. Pires &G.A.
Ratter & V. P. de Lima 6716
olina,
Ponto
Taguatinga,
and BR-010,
35 km N of Car
(E); Mpio. Carolina, Transamazonian
hwy, BR-230
towards Serra de Baleia, E. L. Taylor et al E1259
(NY).?TOCANTTNS:
Mpio.
estrada Taguatinga-Mimoso
do Oeste, Km 14, B. A. S. Pereira
1604 (NY).
Tur?stico W
Andira cordata is the only tree species of Andira in the Brazilian cerrados with en
tirely glabrous foliage, or with only very poorly developed indumentum on the leaflet un
dersurfaces. Andira verm?fuga and A. cujabensis both have well-developed
indumentum
on the leaflet undersurfaces. Although many specimens of A. humilis are glabrous, this
species is completely distinct in its geoxylic suffrutex growth form.
This
species
closely
resembles A. cujabensis
and is parapatric

(sensu Mayr,
1982,
104
VOLUME 64
cordata. A. Bark of trunk. B. Portion of twig with lenticels twig. C. Habit. D. Node of
stipels. E. Flower. F Calyx, opened to show inner surface. G. Standard petal, inner surface. H. Wing
K. Gynoecium,
also shown at right in longitu
petal, outer surface. I. Keel petal, inner surface. J. Androecium.
of C and
dinal section. L. Fruit. (Based on: A, R. T. Pennington
264; B-D, R. T. Pennington
262, inflorescence
FIG.
39. Andira
rhachis with
E^K, G. Hatschbach
39477;
L, J. M. Pires
1630a.)

2003
ANDIRA
105
p. 275) with it. The combined phylogenetic

analysis indicates that these taxa are closely
from A. cujabensis by its lack of indu
related (Fig. 8). Andira cordata is distinguished
mentum (on foliage, gynoecium, and calyx). Andira cordata also has smaller leaflets that
generally are not acuminate and secondary veins that are plane adaxially and less raised
abaxially. Some specimens of A. cordata (e.g., Gottsberger 15-9482 and Taylor E-1259)
have sparsely hairy foliage, but the indumentum is very poorly developed in comparison
to that of A. cujabensis, and the gynoecium remains glabrous.
Brazil. Ama
23. Andira micrantha Ducke, Arq. Inst. Biol. Veg. 2(1): 48.1935.?Type:
zonas: Manaus, estrada do Aleixo,
10May 1932, A. Ducke KB 23864 (lectotype,
here designated: RB!; isolectotypes K! U! US!).
Tree to 35 m
tall; presence of buttresses unknown; bark brown, smooth; slash red

clear
brown with
ex?date; twigs reddish brown (or very dark brown), glabrous; lenticels
absent. Stipules early caducous (not seen). Leaf axis 3-14 cm long; rhachis glabrous;
stipels 0.5-2 (-6) mm long, caducous; petiolules 3-7 mm long, glabrous; leaflets in 2
pairs, 3.3-13 cm long, 1.1-5.1 cm wide, narrowly ovate, elliptic (rarely lanceolate,
broadly elliptic to narrowly obovate), thickly chartaceous to subcoriaceous, base obtuse to
rounded, often slightly decurrent, apex obtuse with an acumen to 15 mm long, glabrous
(seedling leaflets with sparse, short, red-brown indumentum abaxially); primary vein
channelled adaxially, the groove flattened, particularly at the base; secondary veins 6-8,
plane adaxially, very slightly raised abaxially, pattern eucamptodromous
becoming
brochidodromous,
tertiary veins plane adaxially and abaxially. Panicles 3.5-15 cm long,
axillary and terminal, hairy to sparsely hairy with red-brown, ? appressed hairs at branch
tips, glabrescent below; bracts and bracteoles early caducous, not seen; pedicels 0.3-0.5
mm long. Flowers 6.5-7 mm long. Calyx 3-4 mm long, glabrous or with a few scattered
appressed red-brown hairs, the lobes with sparse red-brown hairs; lobes 0.2-0.4 mm long,
obtuse (or rarely 90?). Petals white to cream with the standard marked with lilac; standard
blade 6-6.5 mm wide, 4.5-5 mm high, claw 1-2 mm long; wing 3.7-5 mm long, 2-2.2
mm wide, claw 2-2.8 mm long, sculpturing absent; keel 2.7-3.5 mm long, 1.6-1.9 mm
wide, claw 2.2-3 mm long. Stamens 5 mm long, fllaments united for the basal 1.4-2.6
mm, free for the distal 0.9-2.5 mm, vexillary stamen 3 mm long (all 10 stamens united in
from two specimens). Gynoecium 4.6-5.5 mm long, with sparse to very
sparse red-brown appressed hairs that become less frequent towards the stipe; stipe 1.4-2
mm long, ovary 1.7-2 mm long, style 1.3-1.8 mm long; ovules 1 (-2). Fruits 9-10 cm
100-300 g when dry, green ripen
long, 6.5 cm high, 6-6.3 cm wide, elongated, weighing
to
on outer edge), drying very dark
with
least
mesocarp purple (at
ing grey
grey-brown,
almost
wrinkled
and
brown,
black, shallowly
tuberculate, with pale spots
minutely
(lenticels?; best seen with lens or microscope);
stipe 2 mm long, 6-8 mm wide, short and
stout; suture a thin raised line adaxially, not visible abaxially (dried fruits tend to crack
flowers dissected
along the upper suture); stylar remnant absent; exocarp thin, black in section; mesocarp
4-8 mm thick, pale brown to brown, granular, very hard; endocarp 5-9 mm thick, pale
brown,
very
hard,
non-fibrous.
Chromosome
number
unknown.
Phenology. Flowering March to June.

Distribution
(Fig. 40). Brazil (Amazonas); in terra firme forest, on both sandy and
clay soils, though some records are from river banks.
Vernacular
names.
Acapu-rana,
sucupira
vermelha,
sucupira
chorona.

106
50
55
60 70 65
of Andira
micrantha,
A. cori?cea,
VOLUME 64
4035 45
and A. grandistipula.
Examined.
Reserva Florestal Ducke, Adair s.n.
Specimens
Additional
Brazil. Amazonas:
Manaus,
estrada do Taruma, Ducke RB 23401 (G, K, RB, U); Manaus,
(INPA); Manaus, Ducke RB 21365 (F); Manaus,
terreno da SIDERAMA,
Loureiro et al. s.n. (INPA);
estrada do Aleixo, Ducke RB 35077 (G, INPA); Manaus,
reserve 1501 ("Km 41") of DBFF project, A. A.
90 km NNE Manaus, Distrito Agropecuario,
Mpio. Manaus,
Oliveira
ter Egler,
(MEXU,
et al. 420
(K); Manaus, Reserva Florestal Ducke, quadra 23, O. Pires 29 (INPA); Manaus, Reserva Wal
estrada Mau?, Prance el al. 11635
Km 64, Prance ei al 9049 (U); Manaus,
road Manaus-Itacoatiara,
ei al. 15011 (U);
Prance
NY, U); Basin Rio Negro, Rio Cuieras just below mouth Rio Brancinho,
1534 (US); Manaus, Reserva Florestal Ducke, ao lado da

& Chagas
Igarap? da Cachoeira Alta, W. Rodrigues
Km 84, ?rvore no.
7929 (COL, INPA); estrada Manaus-Itacoatiara,
?rvore no. 1578, W. Rodrigues & Osmarino
7992 (INPA); Manaus, Cachoeira Baixa do Tarum?, W. Rodrigues &
V-13 do inventario florestal, W. Rodrigues
L. Co?lho 8557 (INPA).

is a rain forest species and ismost likely to be confused with A. co
ri?cea and A. taurotesticulata, which also grow in this habitat and possess large fruit.
These two species are not, however, sympatric with A. micrantha, which grows in central
Amazonia
in the vicinity of Manaus. Andira micrantha may be distinguished from A. co
Andira micrantha
ri?cea by the persistent stipules

whereas stipules of A. micrantha
and presumably tiny; no stipule
fruit that are ribbed from suture
15 mm long and 5 mm wide,

on any specimens examined)
Andira taurotesticulata has
sparse hairs on the undersur
face, whereas
leaflets glabrous.
of the latter. These reach

are caducous (not present
scars have been observed.
to suture and leaflets with
the fruit of A. micrantha are smooth and the
Pulle, Rec. Trav. Bot. N?erl. 6: 267. 1909.?Type:

Sectie O, tree number 61, 15May 1907, Boschbeheer
(lectotype, here designated: U!).
24. Andira
cori?cea
Boschreserve

Suriname.
(B. W.) 61
2003
ANDIRA
107
R. Benoist, Bull. Mus. Hist. Nat. (Paris) 25(4): 296. 1919.?

Guiana. Maroni, Wachenheim 79 (lectotype, here designated: P!;
P!).
Andira wachenheimii
Type: French
isolectotype:
Tree 25 (^10) m tall, with small buttresses; bark grey-brown, Assuring vertically,
twigs glabrous, very dark brown, almost black, with nu
flaking; slash orange-brown;
merous pale lenticels. Stipules to 15 mm long, to 5 mm wide, quite persistent, often
sparsely hairy at the tips; leaf axis 7-25 cm long; rhachis glabrous, very dark brown,
this layer splitting to reveal red-brown beneath; stipels absent; petiolules 5-6 mm long,
glabrous; leaflets in 3 pairs (4 pairs seen on a sapling leaf), 5.2-15 cm long, 2.5-5.5 cm
wide, elliptic, narrowly obovate to narrowly ovate, base rounded (rarely obtuse), apex
obtuse (rarely rounded) with an acumen to 15 mm long, glabrous; primary vein chan
nelled adaxially; secondary veins 7-10 (-13), ? plane adaxially, ? plane to slightly
raised abaxially, pattern eucamptodromous
tertiary veins
becoming brochidodromous;
cm
and
branch
Panicles
10-22
and
terminal,
axillary
long,
abaxially.
adaxially
plane
tips hairy to sparsely hairy, glabrescent below, hairs red-brown, ? appressed; bracts and
bracteoles early caducous (seen on only one specimen), 2 mm long, 2 mm wide, dark
brown, hairy atmargins. Flowers 6-7 mm long, ? sessile. Calyx 3-4 mm long, glabrous
except around the margins of the lobes; lobes 0.5-0.75 mm long, obtuse to 90?. Petal
color unknown; standard blade 6-7 mm wide, 5-5.5 mm high, claw 1-2.5 mm long;
wing 4.5-5.5 mm long, 2.3 mm wide, claw 3 mm long, sculpturing absent; keel 3-4.5
mm long, 1.5-2.5 mm wide, claw 3-3.5 mm long. Stamens 4.5-6 mm long; filaments
united for the basal 2.5-4 mm, free for the distal 2-2.5 mm, vexillary stamen 2.5-4.5
mm long. Gynoecium
6-6.5 mm long, glabrous; stipe 1.75-2 mm long, ovary 2-3 mm
long, style 1.5-2 mm long; ovules 1. Fruits 7-7.5 cm long, 6-6.5 cm high, 6.5 cm wide,
? globose, weighing
100-300 g when dry, green to grey-green, drying dark brown (al
most black) with slight waxy bloom, surface slightly ridged and cratered; stipe to 1 cm
long; suture not visible; stylar remnant absent; mesocarp 5-8 mm thick, pale red-brown,
hard, granular; endocarp 5-8 mm thick, very hard, pale brown, non-fibrous. Chromo
some
number
unknown.
Phenology.
Guiana).
Distribution
Flowering
May
to October
(May, Suriname; August-October,
French
(Fig. 40). French Guiana and Suriname; in rain forest.

names. Kobon, roode kabbes, reddie kabbes, koeraroe (Arawak), akoeli
lebi (Par?maca; French Guiana).
tjerere (Suriname); St. Martin rouge, lebi kiabici/kiabici
Vernacular
Specimens
Examined.
Sectie O, B. W. 1909 (COL, U, US); Sectie O, tree 61,
Suriname.
I, B.W. 5450 (U); Brownsberg,
1, tree 5, B. W. 4092 (U); Zanderij
(U); Forest Reserve Zanderij
tree 1301, B. W. 6870 (A); Brownsberg,
B. W. 6870 (U); Emmaketen,
1.5 km from main camp in direction of
small Hendrik peak, Daniels
in bosje op te savanna ten O. van de Brinck
& Jonker 868 (U); Sabanpassie
Additional
B. W. 3933
158 (U); via secta ab Moengo

heuvel, Heyligeri
tapoe ad Grote Zwiebelzwamp,
bij Km 3 in boss, Lanjouw &
Lindeman 441 (U); Poika, Schulz 7625 (U); Upper Coesewijne
River, ca. 20 km SW Poika, Schulz 7953 (U);
Naturreservaat
Teunissen
& Wildschut
11830
Brinkheuvel,
(U); Dist. Broko
Sabanpasi-savannecomplex,
van Donselaar
1743 (U).
St. Laurent, BAFOG 5IN (U);
French Guiana.
pondo, 8 km ESE of Brownsberg,
1201 (U); Cayenne, BAFOG
1264 (U); route de l'Est, near the bridge on the Comte River,
Cayenne, BAFOG
ca. 52 km S Cayenne,
56 (NY); piste de St. Elie-interfluve
Km 15.7,
Champagne
Sinnamary/Counamama
Orstom
3046 (B, CAY, K); route de Saint Laurent ? Cayenne,
Km
12, 50M (U); route de
camp, Pr?vost
? l'Acarouany,
Km 4.500,
c?t? nord et ? 10 m de la route, 231M (U); route de l'Acarouany,
Km 4.300, c?t? gauche
4.300, c?t? gauche et en bordure de la route, 7699 (U); route de l'Acarouany,
bordure de la route, 7733 (U).
Charvein

Km
et en
108
VOLUME 64
ismost likely to be confused with A. taurotesticulata (no. 5) and A.

the
other rain forest species with large fruit; see those species for a
(no. 23),
discussion of their differences.
CTFT (1989) reported that the timber of A. cori?cea is used in the construction of
of items such as
wooden houses and has commercial potential for the manufacture
snooker cues and umbrella handles.
Andira
cori?cea
micrantha
Pulle
specimen,
(1909) cited two collections with his description of A. cori?cea. The flowering
collected on 15May 1906, is selected here as the lectotype.
Amshoff, Bull. Torrey Bot. Club 75(4): 394. 1948.?Type:

grandistipula
Guyana. Kaieteur Plateau, Kaieteur Savannas, 6May 1944, B. Maguire & D. B.
Fanshawe 23273 (holotype: NY!; isotypes: A! F! K! U!).
25. Andira
Small tree 2.5-6 m tall; presence of buttresses unknown; bark and slash unknown;
twigs glabrous, very dark brown, almost black, this outer layer splitting and peeling to
reveal red-brown beneath; lenticels pale. Stipules to 7.5 cm long, to 6 cm wide, broadly
obovate, pale green drying pale brown, glabrous, persistent, crowded at shoot tips and
around inflorescence bases; leaf axis 21-30 cm long; rhachis very dark brown, almost
this layer thin and peeling to reveal red-brown beneath, glabrous; stipels absent
(perhaps present on young leaves and caducous); petiolules 7-13 mm long, dark brown,
almost black, this layer thin, peeling to reveal red-brown beneath, glabrous; leaflets in
(2-) 3-4 pairs, 11-18 cm long, 4.5-10 cm wide, elliptic, broadly elliptic (rarely nar
black,
rowly obovate, some lower leaflets occasionally

broadly ovate to ovate), coriaceous,
base rounded, apex rounded (rarely obtuse), often with a short acumen to 3 mm long,
glabrous adaxially, sparsely hairy abaxially, hairs minute (<0.1 mm long), appressed,
red-brown; primary vein channelled adaxially; secondary veins 8-11, ? plane to slightly
sunken adaxially, raised abaxially, pattern eucamptodromous
becoming brochidodro
tertiary veins plane adaxially and plane to slightly raised abaxially. Panicles
cm long, terminal, dark brown, almost black, sparsely covered with short, ap
pressed, red-brown hairs at branch tips, glabrescent at base; bracts not seen; pedicels
2-4 mm long, very dark brown, almost black, glabrous; bracteoles not seen. Flowers ca.
mous,
17-32
15mm
long. Calyx dark brown, almost black, ca. 6 mm long, glabrous except the mar
gins of the lobes sparsely covered with red-brown hairs; lobes 0.5-1 mm long, obtuse.
Petal color unknown (only old flowers seen); standard blade ca. 12 mm wide, 10 mm
high, claw ca. 3.5 mm long; keel and wings not seen. Stamens ca. 10mm long, filaments
united for the basal ca. 6 mm, free for the distal ca. 4 mm. Gynoecium
ca. 13-15 mm
long, very sparsely hairy on lower surface of the ovary, hairs red-brown, ? appressed;
stipe ca. 4 mm long, ovary ca. 6 mm long, style ca. 4-5 mm long; ovules 1 (single ovary
dissected). Fruit 4-7.8 cm long, 3.4-6 cm high, 3.6-6.5 cm wide, elongated, broad, flat
tened adaxially, weighing
100-300 g when dry, without odor, greenish yellow when
brown
and
dark brown, some areas somewhat yellow, appearing
ripe, drying pale
smooth but minutely wrinkled and tuberculate (best seen with lens or microscope);
su
ture a fine line adaxially, raised abaxially; exocarp thin, almost papery (fresh fruit);
5-12 mm thick, greyish brown, hard, finely granular, often splitting along
mesocarp
suture
when dry; stylar remnant minute; endocarp 3-13 mm thick, cream-buff,
upper
non-fibrous,
very
Phenology.
hard.
Chromosome
Flowering
in May
number
(single
unknown.
record);
two fruiting
records
November.

in June and
ANDIRA
2003
Distribution
areas
of
(Fig. 40). Guyana,
restricted
109
to the Pakaraima Mountains;
in scrub in
savanna.
Specimens
Examined.
Region:
Additional
Pakaraima Mts, Imbaimadai,
Guyana. Mazaruni-Potaro
7972 (U, US);
1992 (COL, K, US); Pakaraima Mts,
Imbaimadai Creek W of Imbaimadai, Pipoly
Hoffman
Pakaraima Mts, Mt. Membaru, Maas & Westra 4299 (K, U); Upper Mazaruni River Basin, Partang Rapids, near
et al 43878 (NY-2 sheets).
mouth Partang River, Maguire
is distinguished by its large, persistent stipules from the other

grandistipula
II
of
Andira (defined by an endocarp of stone cells). None of the
clade
of
eight species
collections examined had fully mature flowers, and the petal color is unknown.
Andira
Brazil.
Arroyo ex R. T. Pennington, sp. nov.?Type:
Amap?: Rio
Oiapoque, 3 km E mouth Rio Mutura, 22 Sep 1960, H. S. Irwin, J. M. Pires & L.
Y. Th. Westra 48442 (holotype: NY!; isotypes: F! U! ?B!).
26. Andira
praecox
tervequinata floribus majoribus et statura altiore recedit (arbor grandis

m
ad
30
alta, non frutex vel arbuscula usque ad 7 m alta).
usque
m
Tree to 30
tall; presence of buttresses unknown; bark and slash unknown; twigs
brown, densely covered with red-brown appressed hairs, becoming grey-brown, glabres
cent. Stipules to 10 mm long, to 1mm wide, caducous, with appressed, red-brown hairs;
Ab Andira
leaf axis 3-10 (-18.5) cm long; rhachis with sparse, red-brown, appressed hairs; stipels
1-2 mm long; petiolules 2-5 mm long, sparsely to densely hairy, hairs red-brown, ? ap
pressed, short (<0.2 mm long); leaflets in (1-) 2 (-3) pairs, 3.5-9.5 (-14) cm long, 1.9?4.3
(-6) cm wide, narrowly obovate (rarely elliptic), terminal leaflets distinctly obovate,
to subcoriaceous,
(rarely acute or rounded), slightly decur

retuse (or rarely with an acumen to 4 mm
long), glabrous adaxially, sparsely hairy abaxially, hairs short, appressed, red-brown or
pale with a red-brown base; primary vein channelled adaxially; secondary veins 8-12,
thick-chartaceous
rent, rounded
(rarely obtuse),
base obtuse
apex slightly
plane adaxially, raised abaxially, pattern eucamptodromous

tertiary veins plane adaxially. Panicles 9-15 cm long,
axillary and terminal, with red-brown hairs at branch tips, hairs ? appressed, glabrescent
at base; bracts caducous, not seen; pedicels 1-2 mm long, glabrous; bracteoles caducous,
not seen. Rowers
6.5-7 mm long. Calyx 3-3.5 mm long, glabrous except the margins of

the lobes sparsely covered with red-brown hairs; lobes 0.5 mm, obtuse. Petal color un
known (probably white); standard blade 5.5 mm wide, 4.5 mm high, claw 1.5 mm long;
wing 3.5-5 mm long, 2-2.5 mm wide, claw 2.2 mm long, sculpturing absent; keel 3-4 mm
long, 2-2.5 mm wide, claw 2.2 mm long. Stamens 5-5.5 mm long, filaments united for
the basal 2-3 mm, free for the distal 1.5-2 mm, vexillary stamen 3-4 mm long. All gy
noecia seen developing into fruit; ovaries swollen, sparsely hairy on upper surface; stipe
ca. 3mm long, style 2.5-3 mm long; ovules 1-2. Fruits 3.1-4.1 cm
long, 2.9-3.6 cm high,
2.8-3.6 cm wide, ? globose, weighing ca. 20 g or less when dry, very dark brown, almost
(with brown pock-marks where the surface is broken) and
slightly wrinkled
tuberculate with a slight waxy bloom (best seen with lens or microscope);
stipe
long; sutures raised adaxially and abaxially; stylar remnant insignificant; meso
carp 1-2 mm thick, pale brown, hard, granular; endocarp 2-5 mm thick, buff, woody, non
black,
minutely
to 9 mm
fibrous,
very
hard.
Chromosome
number
unknown.
Fig.
41.

110
VOLUME 64
FIG. 41. Andira praecox. A. Habit with developing

young fruit. B. Habit with mature fruit. C. Abaxial
leaflet surface. D. Calyx, opened to show inner surface. E. Standard petal, inner surface. F. Wing petal, outer sur
face. G. Keel petal, inner surface. H. Androecium.
I. Gynoecium,
also shown at right in longitudinal
section. J.
Old flower with developing
fruit. K. Entire fruit (at left) and in section (at right), showing wall structure. (Based
on: A, C-J, H. S. Irwin et al. 48442;
B, K, D. C. Daly
et al. 3923.)

Phenology.
ber
111
ANDIRA
2003
and
of Andira
praecox,
in September
Flowering
A. tervequinata,
A. trifoliolaia,
and A. unifoliolata.
(single record); two fruiting
records in Decem
January.
Distribution
Vernacular
Additional
(Fig. 42). Brazil (Amap?); in terra firme forest.

names. Andiroba jaruba, sucupira vermelha.
Brazil. Amap?: Mpio. Mazag?o,
road toward
BR-156,
incomplete
3923 (K, NY); Mpio.
5-10 km SW Rio Preto, Daly & Cardoso
Macap?,
Vila Nova,
18 km SSW Cupixi, Rabelo et al 3199 (ULM, US); Serra do Navio, Ro
Specimens
Monte
75-80
Dourado,
road
Cupixi-Rio
Macap?,
Examined.
km WSW
drigues 2917 (TNPA).

Andira praecox ismost similar toA. tervequinata, especially in leaf morphology, but
differs in several characteristics, especially in its smaller flowers (6.5-7 mm long, whereas
those of A. tervequinata are 9 mm long). Andira praecox grows in rain forest and is a tree
15-30 m in height. In contrast, A. tervequinata is a shrub or small tree to only 7 m high,
which grows in savanna and associated gallery forest habitats. The twigs of A. praecox are
grey-brown, whereas those of A. tervequinata are very dark brown, generally with almost
black outer bark, which peels to reveal red-brown bark beneath. Andira praecox leaves
generally have five leaflets (occasionally trifoliolate, or with seven leaflets), but A. terve
quinata always has five leaflets or trifoliolate leaves. Leaflet shape differs subtly. In A.
leaflets are narrowly obovate (rarely elliptic), but in A. tervequinata they are
praecox,
broadly elliptic to broadly obovate. Further diagnostic characters may be found in the
fruit, but fruiting collections of both species are few and poor. Andira praecox generally
has smaller fruits (to 4.1 cm long), and its fruit surface (when dry) is dark brown to almost

112
VOLUME64
black, whereas the fruit of A. tervequinata

brown with pale, raised specks.
is larger (5.5 cm long) and generally more
red
R. T. Pennington, G. Aymard & N. Cuello, Novon 7: 72.

tervequinata
1997.?Type:
Venezuela.
Bol?var: Distrito Heres, west bank of R?o Trueno
km
of
35
W
Alto,
Chiguao, high plateau, ecotone between gallery forest and sa
27. Andira
vanna, 23 Mar
1985, Huber 10345 (holotype: NY!;

INPA, K, MO, MY, MYF! P, PORT, RB, US, VEN).
isotypes AAU,
E! HBG,
tree or shrub to 7 m tall; presence of buttresses unknown; bark and slash un

twigs very dark brown with raised elongated lenticels, outer bark very dark brown
Small
known;
(almost black), this layer often flaking to reveal red-brown beneath, with sparse, red
brown, ? appressed hairs, glabrescent. Stipules to 7 mm long, to 1mm wide, caducous,
with red-brown, appressed hairs; leaf axis 4-16 cm long; rhachis longitudinally ridged,
dark brown flaking to reveal red-brown beneath, glabrous; stipels tiny; petiolules 3-8 mm
long, sparsely covered with appressed hairs; leaflets in 1-2 pairs, 3.5-11 cm long, 2-5 cm
wide, broadly elliptic to broadly obovate, coriaceous, base obtuse, apex obtuse to rounded,
generally retuse, occasionally emarginate, glabrous adaxially, with short (<0.2 mm long),
appressed, pale hairs abaxially; primary vein channelled adaxially; secondary veins 6-7,
plane adaxially, very slightly raised abaxially, pattern eucamptodromous
becoming
brochidodromous,
tertiary veins plane adaxially and abaxially. Panicles 10-12 cm long,
axillary and terminal, sparsely covered with red-brown, appressed hairs; bracts 2 mm
long, caducous; pedicels 1-1.5 mm long; bracteoles not seen (presumably small and early
caducous). Flowers 9 mm long. Calyx 4 mm long, glabrous except a few scattered hairs
on the lobes; lobes 1mm long, obtuse, the apex with a pointed acumen. Petals pale pur
plish white; standard blade 8 mm wide, 6 mm high, claw 3 mm long; wing 6 mm long, 3
mm wide, claw 4 mm long, sculpturing absent; keel 5 mm long, 3 mm wide, claw 4 mm
long. Stamens 6-7 mm long, the filaments united for the basal 4-5 mm, free for the dis
tal 1-2 mm, vexillary stamen 4.5 mm long. Gynoecium 9-9.5 mm long, the upper surface
and distal end of the lower surface of the ovary with sparse hairs; stipe 3.5-4 mm long,
ovary 3 mm long, style 2.5 mm long; ovules 1. Fruits 5.5 cm long, 3.8 cm wide, 3.8 cm
ca. 20 g or less when dry, green or grey-green, drying dark
high, ? globose, weighing
brown to red-brown, the surface with pale, raised specks; suture raised below, slightly
raised adaxially; stylar remnant minute or absent; mesocarp 3 mm thick, pale, amorphous;
endocarp 4-5 mm thick, pale, very hard. Chromosome number unknown. Fig. 43.
Phenology. Flowering inMarch and with young and mature fruits inMay.
Distribution
(Bol?var: Distrito Heres and Distrito Piar); in
(Fig. 42). Venezuela
in
tree
savanna dominated by Trachypogon plumosus
shrub
and
islands
in
forest,
gallery
was
on conglomerated
this
(G. Aymard, pers. comm;
vegetation
sandstones), and in the
ecotone between savanna and gallery forest; 350 to 500 m.
Vernacular
name. Pil?n rebalsero.
Additional
Specimens
Examined.
Venezuela.
Bol?var:
Distrito Heres, a lo largo del R?o El Trueno al
N de la base del Guaiquinima
6147
Distrito Piar, R?o Aparam?n
at Kambay
NY,
(MO,
PORT);
Aymard
Tepui,
mer? rapids, SE base of Amaruay-tepui,
W of Aparam?n-tepui,
E of Auyan-tepui,
Holst & Liesner 2798 (E,
MO); Distrito Piar, R?o Aparam?n, Kambay-mer?
rapids, ca. 3 km SE of SSE corner of Amaruay
tepui, Liesner
& Hoist
20674
(E, MO).

ANDIRA
2003
113
D. Flower. E. Calyx,
leaflet surface. C. Inflorescence.
tervequinaia. A. Habit. B. Abaxial
inner surface. F. Standard petal, inner surface. G. Keel petal, inner surface (above), and wing
K. Gynoecium
in longitudinal
section. L. Fruit. M. Fruit in section,
petal, outer surface (below). H. Androecium.
showing wall structure. (Based on: A-K, O. Huber 10345; L-M, B. Holsi & R. Liesner 2798.)
FIG. 43. Andira
opened
to show

114
VOLUME64
Andira tervequinata might be confused with A. trifoliolata (no. 28) and A. praecox
from A. trifoliolata were discussed in de
(no. 26); see notes under the latter. Differences
tail by Pennington et al. (1997) and are summarized here. Andira trifoliolata has trifolio
late leaves and leaflets that are glabrous, with a plane primary vein on the upper surface
and an acuminate
apex. Andira tervequinata has leaves with 3-5 leaflets, which have
on the undersurface, a channelled primary vein on the upper sur
hairs
short, appressed
face,
a retuse
and
or
emarginate
apex.
Brazil.
Ducke, Arq. Inst. Biol. Veg. 4(1): 22. 1938.?Type:
Rio Curicuriare, 20 Feb 1936, A. Ducke RB 35079 (lectotype, here
designated: RB!; isolectotypes: G! K! U!).
28. Andira
trifoliolata
Amazonas:
to 20 (-30) m tall; presence of buttresses unknown; bark dark brown, slash

hard, yellow, fibrous; twigs dark brown (or almost black), sparsely hairy,
occasional
glabrescent;
pale, elongated lenticels. Stipules to 9 mm long, to 1mm wide,
with
brown, caducous,
appressed, red-brown hairs at tips, glabrous near base; leaves
leaf
axis 2.5-9 (-12) cm long. Rhachis dark brown, this layer
trifoliolate,
pinnately
Tree
yellow; wood
peeling to reveal pale brown beneath, glabrous; stipels 1mm long, caducous; petiolules
3-7 (-8) mm long, glabrous; leaflets 1 pair, 3.7-11.5 cm long, 1.9-5 cm wide, broadly
elliptic, elliptic (rarely suborbiculate and some terminal leaflets narrowly obovate), sub
coriaceous, base obtuse to rounded and often slightly decurrent, apex obtuse with a
short acumen to 5 mm long, glabrous; primary vein plane adaxially; secondary veins
5-8, plane adaxially, slightly raised abaxially, curving uniformly, pattern ? completely
or ? eucamptodromous
or mixed,
brochidodromous
tertiary veins plane adaxially and
or
cm
Panicles
6.5-17.5
terminal,
abaxially.
axillary
long, with sparse red-brown hairs
at branch tips, glabrescent below; bracts early caducous (seen only on young inflores
cence of a single specimen), 5 mm long, with sparse red-brown hairs; pedicels to 0.3
mm long or absent; bracteoles 2-2.5 mm long, early caducous,
indumentum as on
bracts. Flowers 8-10 mm long. Calyx 4-5 mm long, black, glabrous or with a few scat
tered appressed red-brown hairs, the lobes very sparsely hairy becoming more hairy at
the margins, 90? to obtuse, 0.5-1 mm deep. Petals white or dull flesh-colored,
the stan
dard probably marked with red or purple; standard blade 7.5-9 mm wide, 6.8-8 mm
high, claw 2-2.2 mm long; wing 6-7 mm long, 3-3.5 mm wide, claw 3-3.5 mm long,
sculpturing absent; keel 4-5.7 mm long, 2.2-3 mm wide, claw 3.5-4 mm long. Stamens
6-7 mm long, filaments united for the basal 3-4.7 mm, free for the distal 1.2-3 mm,
7.2-8.5 mm long, the ovary sparsely hairy
vexillary stamen 4.5-5 mm long. Gynoecium
on upper and lower surfaces and often on the sides; stipe 2.5-3.5 mm long, ovary
2.5-3.5 mm long, style 1.5-1.7 mm long; ovules 1-2. Fruit 2.3-3.5 cm long, 2-2.7 cm
high, 2-2.7 cm wide, ?
brown or almost black,
with pale
microscope),
ture a thin raised line
1.5-2 mm
ca. 20 g or less when dry, green, drying dark

globose, weighing
smooth
but
appearing
minutely wrinkled (best seen with lens or
where
surface
is broken; stipe 15-25 mm long; su
pock-marks
adaxially, not visible abaxially; stylar remnant tiny; mesocarp
to dark brown (or greenish), hard, granular; endocarp 2-3 mm
thick, pale
number unknown.
thick, pale buff, very hard, non-fibrous. Chromosome
Phenology. Flowers recorded in February, June, July, and October.
Distribution
(Guainia); Venezuela
(Fig. 42). Colombia
(Amazonas);
zonas); along rivers, but also in terra firme and secondary forest.

Fig. 4B.
Brazil
(Ama
ANDIRA
2003
115
Colombia.
Guain?a: R?o Atabapo, Region Cacaual, R. Rodrigues &
Specimens
Examined.
forest near mu
Venezuela.
Amazonas:
San Carlos de R?o Negro,
(COL, INPA, TJDBC-2 sheets).
et al 1432 (US); Depto. Casiquiare, R?o Casiquiare,
Laguna del Pucibe and
nicipal playing field, Christenson
5 km N village, near savanna "morichal,"
area, Colella et al 2020 (FHO, NY); Depto. Atabapo, La Esmeralda,
Additional
Acero
221
small laja along lower part

81 (K); Depto. Rio Negro, Mucuriapi,
551 (FHO); road from San Fernando
(NY); Depto. Atabapo, L. Delgado
km from San Fernando, Gentry & Tillett 10889 (MO); Depto. Atabapo,
et al. 125856
nidades de Culebra y Huachamacari,
(NY); Rio
Steyermark
43368
Brazil. AMAZONAS: Barcelos, Ducke
(F, NY, U, US).
Adderley
Coomes
of R?o Baria, Davidse & Miller 26752

to Santa Barbara,
12^0
de Atabapo
entre las Comu
R?o Cunucunuma,
Casiquiare,
RB 17293
near Solano, Wurdack
&
(RB, K); Rio Negro, Rio

764 (NY); estrada Manaus-Porto
trecho entre os Rios Castanho e Tupana,
Velho,
Curicuari, O.C. Nascimento
M. F. Silva et al. 924 (INRA); estrada Manaus-Porto
Velho, Km 40-30, M. F. Silva 979 (INPA).
Andira trifoliolata is the only species in the genus that has all leaves trifoliolate. It is
likely to be confused only with A. praecox, which rarely has a few trifoliolate leaves, and
A. tervequinata, which has 1-2 pairs of leaflets; neither of these species is uniformly tri
foliolate like A. trifoliolata. For other diagnostic characters, refer to the discussion of A.
praecox (no. 26) and A. tervequinata (no. 27).
The collection Gentry & Tillett 10889 may represent a new species. It is said to have
been taken from a shrub, whereas all other records of A. trifoliolata are from trees. More
over, the leaflets have sparse, appressed hairs on their lower surfaces, whereas all speci
mens
of A. trifoliolata are glabrous. Yet, the specimen has immature fruit and lacks flow
ers; more complete material is necessary to determine its true status.
The fruit and bark of A. trifoliolata may be used as a contraceptive (herbarium label:
Christenson et al. 1432, US).
Brazil.
Ducke, Arq. Inst. Biol. Veg. 4(1): 22. 1938.?Type:
estrada
do
3
Mar
A.
Ducke
RB
35078
Manaus,
Aleixo,
1937,
(holo
type: RB!; isotypes: G! K! INPA! RB-5 sheets! U!).
29. Andira
unifoliolata
Amazonas:
Tree to 35 m tall with small buttresses; bark rough, flaking; slash unknown; twigs
dark brown, with very sparse, red-brown hairs when young, rapidly glabrescent, bark of
older twigs splitting to reveal paler bark beneath; lenticels raised, pale, elongated. Stipules
to 4 mm long, very early caducous (seen only on a seedling), with sparse, red-brown hairs;
petiole 5-16 (-18) cm long, glabrous, dark brown, this layer peeling to reveal pale brown
stipels to 2 mm long (seen only on seedlings); petiolule 3.5-6 mm long,
unifoliolate
(1-2 pairs leaflets in seedlings), leaflet 4.7-13 cm long, 1.3-5 cm
glabrous;
wide, elliptic (to narrowly ovate), dark green, shiny adaxially, much paler and matt abax
ially, the venation pale, thick-chartaceous (seedlings) to coriaceous, base obtuse (or rarely
acute), often slightly decurrent, apex acute to obtuse with an acumen to 10 mm long,
beneath;
glabrous adaxially, very sparsely hairy abaxially, glabrescent, hairs short, appressed; pri
mary vein plane adaxially; secondary veins 6-7, plane adaxially, raised or only slightly
raised abaxially, pattern disorganized,
eucamptodromous with occasional brochidodro
mous linkages (particularly at the leaflet tip), the veins curving uniformly, tertiary veins
plane adaxially and abaxially. Panicles 2^4.5 cm long, axillary (and terminal), with sparse,
pale red-brown hairs at branch tips, glabrescent proximally; bracts and bracteoles early ca
ducous (not seen); pedicels 0.8-1 mm long. Flowers 6-7 mm long. Calyx 3-4 mm long,
dark brown or ? black, glabrous or with a few scattered appressed pale red-brown hairs,
the margins of the lobes sparsely to very sparsely hairy; lobes obtuse with small pointed
tips, 0.3-0.8 mm. Petals white, the standard marked with red; standard blade 6 mm wide,

116
VOLUME 64
i\??M?^
L?\r\^
A. Habit. B. Abaxial
leaflet surface. C. Flower. D. Calyx, opened to show
FIG. 44. Andira unifoliolata.
inner surface. E. Standard, inner surface. F. Wing petal, outer surface. G. Keel petal, inner surface. H. Androe
also shown below in longitudinal
section. J. Fruit. K. Endocarp.
cium. I. Gynoecium,
(Based on: A-I, A. Ducke
61 A; J. A. Ducke 30578.)

andira
2003
117
5 mm high, claw 1.5 mm long; wing 4.5-4.8 mm long, 2-2.2 mm wide, claw 3 mm long,
sculpturing absent; keel 3.3-4 mm long, 2-2.2 mm wide, claw 3 mm long. Stamens 6 mm
long; filaments united for the basal 1.5-2.5 mm, free for the distal 2-3.8 mm; vexillary
stamen 4 mm long. Gynoecium
7 mm long, very few scattered hairs on ovary upper sur
face; stipe 3 mm long, ovary 2 mm long, style 2 mm long; ovules 1-2. Fruit 2.5-3.7 cm
long, 2-2.8 cm high, 2-2.8 cm wide, ? globose, weighing ca. 20 g or less when dry, green,
very dark brown, almost black, with a slight waxy bloom,
turning yellow, sweet-smelling,
mm long; suture a thin line adaxially, not detectable
3-4
very slightly wrinkled; stipe
remnant
abaxially; stylar
tiny; mesocarp pale reddish brown, hard, granular; endocarp un
known. Chromosome
number unknown. Figs. IC, 2F, 44.

Phenology. Flowering February toMarch.
Distribution (Fig. 42). Brazil (Amazonas, Para, Roraima);
sandy and clay soils.
Vernacular
names.
zonas); andira-uchi
Sucupira
amarela,
sucupira
chorona,
in terra firme rain forest on
sucupira
vermelha
(Ama
(Para).
Additional
Specimens
Examined.
Brazil. Amazonas:
Reserva Florestal Ducke, Adair s.n.
Manaus,
Reserva Florestal Ducke,
?rvore do fenel?gico
(INPA); Manaus,
20, Alu?sio s.n. (INPA); Mpio. Mau?s, Rio
?cima do lugar S?o Sebasti?o, Cid Ferreira 4268 (INPA); Manaus,
estrada do Aleixo, Ducke 674
Apoquitaua
s.n. (INPA), Mac?do
s.n. (INPA), F. C. Mello
(F, GH, K, NY, US); Manaus, Reserva Florestal Ducke, Lourival
et al. s.n. (INPA); Manaus,
Reserva Florestal Ducke;
estrada Manaus-Itacoatiara,
Km 133, Monteiro
& F.
s.n.
Mello
Reserva
Florestal Ducke, W. Rodrigues
& Loureiro
5953
(INPA); Manaus,
(INPA); Manaus,
Florestal Ducke,
?rvore do fenel?gico
& D. Co?lho 7554 (INPA); estrada Man
13, W. Rodrigues
Km 190, ?rvore XXVI-37
do inventario florestal, W. Rodrigues
7994 (INPA); estrada Man
aus-Itacoatiara,
Km 170, picada XXII, ?rvore 58 do inventario florestal, W. Rodrigues
e? al. 8502 (INPA);
aus-Itacoatiara,
Reserva
estrada Manaus-Porto
Velho, Rio Castanho, margem

direita, estrada para
Reserva
Florestal Ducke,
estudo de associa?ao,
J. A. C.
(INPA); Manaus,
entre as cachoeiras
e Jandi?, L. S. Co?lho 103
de Tremalhetinha
Trombetas,
do Caran?, entrada para a Barragem Velha, E. Soares 5 (INPA).?Roraima:
reservatorio,
Maciel
o Careiro, M. F. Silva e? al. 373

Rio
Souza s.n. (INPA).?Para:
estrada
(INPA); Porto Trombetas,
UHE de Samuel, are? do futuro
104 (INPA, RB, UB).
leaves are unique to A. unifoliolata, and this species cannot be confused

it is most similar to A. trifoliolata. The two
any other of Andira. Morphologically
are
is uniformly
species
easily distinguished
trifoliolate,
vegetatively: A. trifoliolata
whereas A. unifoliolata
is uniformly unifoliolate. Andira unifoliolata also has flowers
6-7 mm long compared to 8-10 mm long in A. trifoliolata. Andira unifoliolata has useful
Unifoliolate
with
timber.
Doubtful
Andira
sect. Aristobulia
Andira
amazonum
and Excluded
Names
Bentham, Comm. legum. gen. 43. 1837.?LECTOTYPE, here des

Andira
amazonum
Martius ex Bentham. = Watairea guianensis Aublet, Hist,
ignated:
t.
Guiane
2:
302.
1775.
755,
pi.
Martius
ex Bentham,
Comm.
legum.
gen.
43.
1837.?TYPE:
Rio Negro, Martius s.n. (holotype: BR!; isotype: M). = Vatairea guianensis
Hist. pi. Guiane 2: 755, t. 302. 1775; see also de Lima (1982).
Andira
BRAZIL.
Aublet,
araroba Aguiar, Gazeta M?dica

da Bahia 10: 353. 1878.?Type:
Est. 1-4.
J. M. Aguiar, Memoria
sobre a araroba, Ed. Imprensa Econ?mica, Bahia. 1879. =

VOLUME 64
118
Bentham, Comm. legum. gen. 44. 1837, nom

americana Aublet, Hist. pi. Guiane, suppl. 9. 1775.
Andira
8: 26. 1936; see
(Aguair) Ducke, Ann. Acad. Brasil. Ciencias
Vataireopsis araroba
also de Lima (1980).
aubletii
superfl.
Vouacapoua
Brazil.
Andira bracteosa Bentham, Comm. legum. gen. 43. 1837.?Type:
Para, Sieber
s.n. (holotype: BR!; isotypes: B-W, HAL). = Vatairea guianensis Aublet, Hist. pi.
Guiane 2: 755, t. 302. 1775; see also de Lima (1982).
Andira
cinerascens Martius,
in sched.; see also de Lima
(1982).
Andira gabonica B?illon, Adansonia 6: 219. 1866.?TYPE: Duparquet

=
Aganope gabonica (B?illon) Polhill, Kew Bull. 25: 269. 1971.
32 (holotype: P!).
Andira horsfieldii Leschenault, Ann. Mus. Nati. Hist. (Paris) 16: 481, t.12. 1810.?TYPE:
= Euchresta
Leschenault, Ann. Mus. Nati. Hist. (Paris) 16: 481, t.12.1810.
horsfieldii
rar.
PL
31.
1838.
148, pi.
(Leschenault) Bennett,
jav.
Andira
ex J. St. Hilaire, Diet. Sei. Nat. 2: 137. 1804, nom. superfl. =

americana Aublet, Hist. pi. Guiane, suppl. 9. 1775.
racemosa Lamarck
Vouacapoua
Suriname. Am
Andira villosa Kleinhoonte, Rec. Trav. Bot. Ne?rl. 22: 404.1925.?Type:
bei
dem
Pulle
319
1920,
Coppenameflusse
Raleighstrom, Aug
(holotype: U!).?The
not
it does not clearly
does
and
the
floral
reveals
that
Andira,
type
belong
morphology
Itmatches a specimen from Peru (Loreto: Iquitos,
fit any genus of Papilionoideae.
San Juan, 19 Sep 1945, A. Ducke 1823), which Ducke described as Hymenolobium
velutinum; however, it is not a species o? Hymenolobium
(pers. obs.; H. C. de Lima,
pers. comm.). Both specimens constituting the type collection have flowers but lack
fruits; thus, it is impossible
to determine
their affinities.
Andira
Brazil.
zehntneri Harms, Repert. Spec. Nov. Regni Veg. 17: 443. 1921.?Type:
Bahia, Barra, Oct 1912, Zehntner 2097 (holotype: M; isotype: RB!). = Lonchocarpus
sp.
Skolemora pernambucensis
Arruda in H. Koster, Travels Brazil 498. 1816.?TYPE: un
known.?It
is not known whether Arruda's types are extant (Stafleu & Cowan 1976).
The description of the anthelminthic properties of this species suggests that this name
may apply to a species o? Andira.
ACKNOWLEDGMENTS
This work was financially
Research Council
(SERC)
supported initially by the Science and Engineering
and the Royal Botanic Gardens, Kew, and completed with the support of the Royal Botanic Garden Edinburgh.
The Gatsby Charitable Foundation
funded research on intraspecific cpDNA polymorphism,
carried out by Liz
in Guyana was supported by The Commonwealth
and Government
of Guyana
Iwokrama
I am grateful to my Ph.D. advisers Roger Polhill, Robert Scotland, Chris Leaver, and
Programme.
the guidance of Brian Styles,
Bennett, who supervised the start of this research. I particularly acknowledge
Caddick.
Fieldwork
Rain Forest
Mike
who
very sadly died before my
thesis work was
complete.
I also thank: Maureen
Warwick

(artwork), Robert Mill
ANDIRA
2003
119
work and cladistic analysis), Tony Cox (molecular work),

(Latin diagnoses), Mark Chase (advice on molecular
David Harris (numer
Peter Gasson
(fruit anatomy), Bente Klitgaard
(pollen data), Iain Prance (biogeography),
ous discussions
and Barbara
Richard Brummitt, Mark Watson,
and reading an entire draft), David Mabberley,
comments on in
MacKinder
(constructive
(nomenclatural
issues), Denis Filer (databasing), Peter Hollingsworth
of Leucaena was an inspiration for this
Colin Hughes
(whose monograph
traspecific cpDNA polymorphism),
Serena Marner, Anne Sing, Alison Strugnell (help with ad
and who collected Andira inermis inMexico),
for invaluable help on my arrival
ministrative
(advice on all topics and particularly
problems), Gwilym Lewis
leaves and fruit of Mexican
in Brazil), Lourdes Rico (who collected
(who
species), Clive Foster, Phil Griffiths
work,
of Brazilian
(discussion
species), Eimear Nie Lughadha,
expertly grew species of Andira), Ary Oliveira-Filho
of Mex
inMexico,
discussion
for fieldwork
in Guyana), Mario Sousa (hospitality
Charlie Stirton (arrangements
ican species), Martin Cheek (who collected Andira inermis in Africa), Jeff Doyle
(molecular systematics),
Larry
Jane Doyle, Paul Manos,
(cladistics and critically reading portions of this manuscript),
Kelly, Anne Bruneau
(cladis
(molecular systematics), Kevin Nixon
Soreng, Stephan Maumont
I particularly
thank Andr? Carvalho for help and
and Dada). For my fieldwork,
enormous hospitality,
Jim Ratter, Sam Bridgewater,
and
de Lima, Alvaro Cogollo,
and also Haroldo Cavalcante
I
also
Taiman
dos
For
Fernando
Herminio
with
fieldwork
thank
Ribeiro.
Carvalho,
Brito,
("Guga")
help
Felipe
Beth Dickson,
tic advice
David
of Clados
Jo?o
Isolde Ferraz, W. Wayt Thomas, Bruce Nelson, William Rodrigues, Niro Higuchi,
Lino Veloz, Terry Pennington,
Milton Aulestia,
Plana, Fernando Montenegro,
Tamara N??ez, Efrain Frere, Mar?a Margarida Fiuza de Mel?, M. Sugiyama, Nelson Zamora, Ma
da Silva, Jes?s Idrobo, Jos? Luis Fern?ndez, Bruce Hoffman
(especially for collecting Andira gran
Santos, Lenise
Guedes,
Alex Monro,
Alu?sio,
Robert
Karen Hanson,
and copies
Neill,
noel Claudio
Vanessa
drawings were provided by Rosemary Wise. The following herbaria

for this study: A, BM, BR, CEPEC, COL, CR, E, F, FHO, G, GH, HUEFS,
and Isaac Johnson. All botanical
disiipula),
provided
material
and/or
facilities
INBIO,
INPA, JAUM, K, M, MEXU,
MO,
NY, OXF,
P, PR, PRC, QCA,
QCNE,
R, RB, RADAM,
SP, U, UB,
US, USP.
LITERATUREcited
1990. Official methods of analysis,
15th ed. Arlington:
1985. Methods
of coding variable morphological
Zool. 34: 236-345.
AOAC.
Association
J.W.
Archie,
features
of Official
Analytical
taxonomic
for numerical
Chemists.
1999. The fruits the agouti ate: Hymenaea

Asquith, N. M., J. Terborgh, A. E. Arnold, and C. Mail?n.
seed fate when its disperser is absent. J. Trop. Ecol. 15: 229-235.
evidence of evolutionary
radiations.
and morphological
Bateman, R. M. 1999. Integrating molecular
ular systematics
London: Taylor
Bennett, M. D.
1984. Nuclear
J. P. Gustafson,
G.
Bentham,
and plant evolution,

and Francis.
ed. P. M. Hollingsworth,
and its manipulation.

York: Plenum Press.
architecture
469-502.
New
R. M. Bateman,
and R. J. Gornall,
In Gene manipulation
inplant
courbaril
InMolec
432-471.
improvement,
ed.
de leguminosarum
generibus. Vienna: J. P. Sollinger.
commentationes.
Ann. Wiener Mus. Naturgesch.
2(1): 107-109.
generibus
leguminosarum
a tribe of the Leguminosae.
A synopsis of the Dalbergieae,
Proc. J. Linn. Soc, Bot. 4, Supplement:
118-123.
1837. Commentationes
De
_1839.
_1860.
24-25;
_1862.
Andira.
Bondt, N.
In Flora
1788. Dissertatio
brasiliensis,
medica
ed. C. F. P. von Martius,
inauguralis de cortice Geoffraeae

la flore de Centre Afrique ?cologie
15(1): 291-299.
surinamensis.
Munich:
R. E. 1985. Zoogeographie
geogr. 12: 445^61.
J. A. 1989. Quantitative
Chappill,
Buskirk,
patterns
and tectonic history
of Jamaica
E. J. H.
(Centre Technique
du Cirad.
Forestier
Tropical).
1989. Bois des Dom-Tom.
Tome
3: Strate
International
and the northern Caribbean.
characters in phylogenetic
5: 291-296.
analysis. Cladistics
1: 8-9. Cambridge: Cambridge University
1976. The seeds of the dicotyledons,
et rejicienda spermatophytorum,
Cowan, R. S. 1959. Andira. In Nomina gen?rica conservanda
ett and F. A. Stafleu. Taxon 8: 295.
Corner,
F. Fleischer.
Leiden.
Y. 1997. Catalogue
de
sommaire, distribution.
ligneuse de savanes. Central African Republic: Centre Orstom de Bangui.
internationales
de la nomenclature
Briquet, J. 1906. R?gles
botanique adopt?es par le Congr?s
de Vienne 1905. Jena.
Botanique
Boulvert,
CTFT
Syst.
analysis.
Tomel, Guyane.
partment

de
J. Bio
Press.
ed. H. W. Rick
Nugent-sur-Marne,
De
120
VOLUME 64
In Zoogeography
T. W. 1988. Geological
constraints on Caribbean biogeography.
of Caribbean
Donnelly,
Press.
15-37. Ithaca, New York: Cornell University
serts, ed. J. K. Liebherr,
as one character taxonomy. Syst. Bot.
J. J. 1992. Gene trees and species trees: molecular
systematics
Doyle,
144-163.
1990. A chloroplast-DNA
J. J., J. A. Doyle,
and A. H. D. Brown.
phylogeny
tives of soybean (Glycine subgenus Glycine)', congruence with morphological
lution 44: 371-389.
Doyle,
T. Kajita, and H. Ohashi.

E. E. Dickson,
J. J., J. L. Doyle,
J. A. Ballenger,
in
taxonomic
correlations and insights
rbcL
the
gene
chloroplast
Leguminosae:
lation. Amer. J. Bot. 84: 541-554.
Doyle,
of the wild
perennial
and crossing
in
17:
rela
groups. Evo
of the
1997. A phylogeny
into the evolution of nodu
A. 1944. New or noteworthy Leguminosae

of the Brazilian Amazon. Bol. Inst. Agron. Norte 2: 30-31.
Press.
Emmons, L. H. 1990. Neolropical
rainforesi mammals:
afield guide. Chicago: Chicago University
to elucidate the history,
1999. Using organelle markers
Ennos, R. A., W. T. Sinclair, X-S. Hu, and A. Langdon.
and plant evolution,
ed. P. M.
In Molecular
and evolution
of plant populations.
sysiemalics
ecology
Ducke,
R. M. Bateman, and R. J. Gornall,

1-19. London: Taylor
Hollingsworth,
and J. I. Sprent.
Faria, S. M. de, H. C. de Lima, A. A. Franco, E. S. F. Mucci,
from south east Brazil. PL <&Soil 99: 347-356.
and Francis.
1987. Nodulation
of legume
trees
J. S. 1988. Hennig86,
version 1.5. New York: Port Jefferson.
in camouflage. Cladistics
Phenetics
6: 91-100.
I. K., and J. J. Skvarla. 1981. The pollen morphology
of the subfamily Papilionoideae
Ferguson,
(Leguminosae).
in legume systematics,
In Advances
Part 2, ed. R. M. Polhill and P. H. Raven,
859-896.
Kew: Royal
Farris,
_1990.
Botanic
Ferguson,
Gardens.
I. K., B. D. Schrire,
tionships
and R. Shepperson.
of the tribe Sophoreae

1994. Pollen morphology
and rela
In Advances
and Papilionoideae
in
Caesalpinioideae
(Leguminosae).
Part 6, Struciural botany, ed. I. K. Ferguson
and S. C. Tucker, 58-96. Kew: Royal
between
subfamilies
legume systematics,
Botanic Gardens.
versus specialization:
the evolution of feeding strategies in frugivorous bats.
Fleming, T. H. 1986. Opportunism
In Frugivores
and seed dispersal,
ed. A. Estrada and T. H. Fleming, 285-384. Dordrecht: Dr W. Junk Pub
lishers.
Fox, D. P. 1969. Some
of the cold hydrolysis

technique for staining plant tissues by the Feulgen
17: 226.
Cytochem.
1976. Foraging behaviour of solitary bees: implications
for out
Frankie, G. W., P. A. Opler, and K. S. Bawa.
forest tree species. J. Ecol. 64: 1049-1057.
crossing of a Neotropical
von Pflanzen der Insel Kuba I. Kulturpflanze
18: 189-197.
Fritsch, R. 1970. Chromosomenzahlen
reaction.
characteristics
J. Histochem.
Fucsk?, M.
1914. Studien ?ber den Bau der Fruchtwand

der Papilionaceen
und die hygroskopische
der H?lsenklappen.
106: 160-215.
Bora
B. 1995. A survey of fruit anatomy in the genus Andira (Leguminosae).
Gemeinholzer,
Unpublished
of Edinburgh and Royal Botanic Garden Edinburgh.
sis, University
Bewegung
MSc
the
and P. F. Stevens.
ex
in the delimitation
1997. Vagaries
of character states in quantitative variation?an
perimental
study. Syst. Biol. 46(1): 112-125.
In Advances
and the phylogeny
of the Leguminosae.
in legume systematics, Part 2,
Goldblatt, P. 1981. Cytology
ed. R. M. Polhill and P. H. Raven, 439. Kew: Royal Botanic Gardens.
F. R. Barrie, H. M. B?rdet, V. Demoulin,
T. S. Filgueiras, D. H. Nicolson,
P. C. Silva,
Greuter, W., J.McNeill,
Gift, N.,
J. E. Skog, P. Trehane, N. J. Turland, and D. L. Hawksworth.

2000. International code of botanical nomen
clature (St. Louis Code). Regnum Veg. 138: 1-474.
1976. World guide to tropical drift seeds and fruits. New York: Quadrangle/New
Gunn, C. R., and J. V. Dennis.
York Times Book Co.
Hallwachs,
W.
1986. Agoutis
nosae). In Frugivores
Junk Publishers.
the inheritors of guapinol

(Dasyprocta punclala):
(Hymenaea
and seed dispersal,
ed. A. Estrada and T. H. Fleming,
285-384.
coubaril:
Dordrecht:
Legumi
Dr W.
ao estudo da unidade de dispers?o

1969. Contribui??o
ex
da pl?ntula
de Andira humilis Mart,
Sao Paulo: Universidade
de Sao Paulo Faculdade de Filosof?a, Ciencias
(Leguminosae-Lotoideae).
e Letras, Bol. 349 (Bot?nica 27).
aux lois de la nomenclature
de compl?ment
Harms, H. 1904. Proposition
botanique de 1867. Notizbl. K?nigl.
Bot. Gart. Berlin, Appendix
13: 23.
J. A., C. E. Hughes,
and R. W. Scotland.
1997. Primary homology
characters and charac
Hawkins,
assessment,
Handro, W.
Benth.
ter states. Cladistics
13: 275-283.

2003
ANDIRA 121
P. S., W. L. Crepet, and D. L. Dilcher.

1992. The fossil history of the Leguminosae:
phylogenetic
and biogeographic
In Advances
Part 4, The fossil
record, ed. P. S.
implications.
Herendeen
and D. L. Dilcher,
85-160. Kew: Royal Botanic Gardens.
Jodrell labora
dissertation,
Herridge, D. 1992. The wood anatomy of the genus Andira. Unpubl. undergraduate
tory, RBG Kew, and Bristol Polytechnic.
Herendeen,
L. J. 1979. A revised
Hickey,
classification
of the architecture
2d ed., ed. C. R. Metcalfe

and L. Chalk,
cotyledons,
Hopkins, H. C. 1983. The taxonomy, reproductive biology
in Africa and Madagascar.
J. Linn. Soc,
mosoideae)
of dicotyledonous
leaves. In Anatomy
Oxford: Clarendon Press.
of the di
25-39.
and economic
Bot.
potential
87: 135-167.
of Parkia
Mi
(Leguminosae:
H. C, and M. J. G. Hopkins.
1983. Fruit and seed biology of the Neotropical
In
species of Parkia.
no.
2
of
the
British
Tropical rainforest:
ecology and management,
publication
Ecological
Society,
special
ed. S. C. Sutton, T. C. Whitmore,
and A. C. Chadwick,
197-209. Oxford: Blackwell
Scientific.
Janzen, D. H. 1982. Simulation of Andira fruit pulp removal by bats reduces seed pr?dation by Cleogonus wee
Hopkins,
vils. Brenesia
19-20:
165-170.
Janzen, D. H., G. A. Miller,
J. Hackforth-Jones,
C. M. Pond, K. Hooper, and D. P. Janos. 1976. Two Costa Rican
seed shadows of Andira
inermis (Leguminosae).
57: 1068-1075.
Ecology
J. H. 1804. Andira. In Dictionnaire
des sciences naturelles dans lequel on traite m?thodique
ment des diff?rents ?tres de la nature, ed. F. Cuvier, 2: 137. Paris: Levrault, Schoell and Cie.
bat-generated
Jaume St. Hilaire,
Jussieu, A. L. de. 1789. Genera plantarum,

Lamarck, J. B. A. P. M. 1783. Encyclop?die
o?Coursetia
Lavin, M. 1988. Systematics
_1993.
Biogeography
and A. Delgado
373. Paris: Herrisant.

m?thodique.
1: 171. Paris: Panckoucke.
Botanique,
21: 1-167.
Syst. Bot. Monogr.
37: 1-87.
Syst. Bot. Monogr.
variation
(Leguminosae):
morphological
(Leguminosae-Papilionoideae).
and systematics o?Poitea
(Leguminosae).
S. 1990. Pollen brush of Papilionoideae

systematic utility. Amer. J. Bot. 77: 1294-1312.
1993. Origins and relationships
of tropical North
Lavin, M., and M. Luckow.
boreotropics
hypothesis. Amer. J. Bot. 80: 1-14.
Lavin, M.,
Lavin, M., R. T. Pennington,
B. B. Klitgaard,
legumes (Fabaceae): Delimitation

and M. Sousa. 1995. Phylogenetic
gioid
Lavin, M.,
in the context
America
and
of the
J. I. Sprent, H. C. de Lima, and P. E. Gasson. 2001. The dalber

of a pantropical monophyletic
clade. Amer. J. Bot. 88: 503-533.
and biogeography
of the Tribe Robinieae
systematics
(Legumi
45: 1-165.
nosae). Syst. Bot. Monogr.
2000. Africa,
the odd man out: molecular
Lavin, M., M. Thulin, J.-N. Labat, and R. T. Pennington.
biogeogra
phy of dalbergioid
legumes (Fabaceae) suggests otherwise.
Syst. Bot. 25: 449-467.
1993. Late Quaternary
environmental
and climatic changes in central Brazil. Quaternary Res. 39:
Ledru, M.-P.
90-98.
Leishman,
M. R.,
dence.
and M. Westoby.
8: 205-214.
1994. The
role of large seed size in shaded
conditions:
Experimental
evi
Funct. Ecol.
Lewis,
G. P. 1987. The
Lima,
H. C.
legumes ofBahia. Kew: Royal Botanic Gardens.

1980. Revis?o
taxon?mica
do g?nero Vataireopsis
Ducke
32: 21-40.
de.
drigu?sia
-1982.
Revis?o
taxon?mica
do g?nero Vatairea Aublet
Ro
(Leguminosae-Faboideae).
(Leguminosae-Faboideae).
Arch.
Jard. Bot. Rio de
Janeiro 26: 173-214.

-1991.
Tribo Dalbergieae
dos frutos, sementes e pl?ntulas e
(Leguminosae
Papilionoideae)?morfolog?a
sua aplica??o na sistem?tica. Arch. Jard. Bot. Rio de Janeiro 30: 1^2.
Tulasne
Lima, H. C. de, A. M. de Carvalho, and C. G. Costa. 1990. Estudo taxon?mica do g?nero Diptychandra
Anais XXXV Cong. Nac. Bot., Brasil: 175-181.
(Leguminosae-Caesalpinioideae).
A. A., and M.
do Interior.
Loureiro,
da Silva.
1968. Cat?logo
das madeiras
da Amazonia,
vol. 2. Bel?m: Ministerio
M.
1995. Species concepts: assumptions, methods

and applications.
Syst. Bot. 20: 589-605.
or renamed spermatophytes
8: 26. 1940.
mostly Peruvian. Candollea
C. F. P. von. 1828. Geoffroya.
In Reise in Brasilien
by J. B. von Spix and C. F. P. von Martius,
Luckow,
J. F. New
Macbride,
Martius,
Munich:
Marx,
Freitas
G. A.
2: 788.
M. Lindauer.
1987. A suite of mutants
that modify
pattern formation
in pea leaves. PI. Molec.
Biol. Reporter
5(3):
311-335.
N. F. 1979. O g?nero Andira Lam. (Leguminosae
1982. The growth of biological
thought. Boston:
Mattos,
Mayr,
E.
McDade,
L.
no Brasil. Acta Amazon.

Papilionoideae)
Harvard University
Press.
1990. Hybrids and phylogenetic

I. Patterns
systematics
for cladistic analysis. Evolution
44: 1685-1700.
of character
expression
implications

9: 241-266.
in hybrids
and their
Milliken,
W.
1997. Traditional
Milliken,
W.,
R. P. Miller,
VOLUME 64
122
in Roraima,
medicine
anti-malarial
Brazil.
Econ.
Bot.
51: 212-237.
of Brazil, 79. Kew: Royal Botanic Gardens.

S. A., B. M. Boom, and G. T. Prance. 1981. Distribution
coastal forest tree species. Brittonia 33: 233-245.
Atroari
of the Waimiri
1992. The ethnobotany
and E. L. Wandelli.
S. R. Pollard,
In
dians
Mori,
patterns
of eastern Brazilian
and conservation
Illinois: C. Thomas.
of medicinal pla??s of Middle America.
Springfield,
law. Syst. Zool. 27: 324-345.
and the biogenetic
1978. Ontogeny,
phylogeny,
paleontology,
K. C. 1992. Clados, version 1.2. New York: Trumansberg.
Dada, version 0.86 beta test. New York: Trumansberg.
J. F. 1981. Alias
Morton,
G.
Nelson,
Nixon,
_1993.
K. C,
Nixon,
and J. I. Davis.
1991. Polymorphic
values
taxa, missing
and cladistic
analysis.
Cladistics
7:
concept.
Cladistics
6:
233-241.
K. C,
Nixon,
and Q. D. Wheeler.
1990. An
of the phylogenetic
amplification
species
211-223.
'murundus'?the
of Brazilian
Filho, A. T. de. 1992. The vegetation
nity. J. Trop. Ecol. 8: 465^86.
Palmer, J. D., G. P. Singh, and D. T. N. Pillay. 1983. Structure and sequence
190: 13-19.
plast DNAs. Mol. Gen. Genet.
on the plant commu
island-effect
Oliveira
evolution
of three legume
chloro
and phytochemical
1987. Ecological
C. M., and M. J. Koziol.
diversity of arillate seeds in Aglaia
a study of vertebrate dispersal in tropical trees. Philos. Trans., Ser. B. 316: 303-333.
Papilionoideae,
systematics o? Andira (Leguminosae,
Pennington, R. T. 1994. The taxonomy and molecular
Oxford University:
Dalbergieae).
unpublished DPhil thesis.
Pannell,
(Meli
aceae):
_1995.
Cladistic
Amer.
bergieae).
and morphological
Molecular
_1996.
DNA
analysis of chloroplast
J. Bot. 82: 526-534.
restriction
data provide
site characters
resolution
in Andira
at different
tribe
Dal
(Leguminosae:
hierarchical
levels
in Andira.
45: 496-515.
Syst.Biol.
_2002.
Proposal
to conserve
the name Andira,
nom. cons.
(Fabaceae)
with
a conserved
type. Taxon
51:
385-386.
R. T., G. Aymard, and N. Cuello.
1997. A new
Pennington,
from the Venezuelan
Guayana. Novon 7:12-1 A.
species
of Andira
(Leguminosae,
Papilionoideae)
and biology of Andira

R. T., and B. Gemeinholzer.
2000. Cryptic clades, fruit wall morphology
Bot. J. Linn. Soc. 134: 267-286.
(Leguminosae-Papilionoideae).
R. T., M. Lavin, H. Ireland, B. Klitgaard,
and J. Preston. 2001. Phylogenetic
of prim
Pennington,
relationships
itive papilionoid
intron irnL. Syst. Bot. 26: 537-556.
legumes based upon sequences of the chloroplast
Pennington,
Pennington,
R. T., and H. C. de Lima.

their distributions.
1995. Two new
Kew Bull.
species
of Andira
and the influence
of dispersal
in deter
50: 557-566.
mining
R. T., D. E. Prado, and C. A. Pendry. 2000. Neotropical
Pennington,
seasonally dry forests and Quaternary veg
etation changes. J. Biogeog.
27: 261-273.
T. D., and J. Sarukh?n.
1968. Arboles
206-207. Mexico:
Instituto Nacional
tropicales de Mexico,
Pennington,
de Investigaciones
Forestales.
R. A., and R. Riggins.
1987. The nature of cladistic data. Cladistics
3: 201-209.
L. Elzevir.
81. Leiden: F. Hack; Amsterdam:
Piso, W. G. 1648. De medicina
brasiliensi,
and J. A. Coddington.
1991. On missing
entries in cladistic
Platnick, N. L, C. E. Griswold,
7: 337-343.
Pimentel,
1969. Notes
on east African
Dalbergieae.
Kew: Royal
In Advances
R. M.
Polhill,
-1981.
233-242.
Botanic
Prado, D. E., and P. E. Gibbs.

ica. Ann. Missouri
Prance,
G. T. 1985. The
Pergamon
-1987.
changing
of species
distributions
Polhill
in the dry seasonal
Cladistics
and P. H. Raven,
forests of South Amer
80: 902-927.
forests.
In Amazonia,
ed. G. T. Prance
and T. E. Lovejoy,
146-165.
Oxford:
Press.
of Neotropical
and Qualernary
plants. In Biogeography
hisiory
and G. T. Prance, 46-65. Oxford: Oxford University
Press.
in Iropical Amer
1906. An
enumeration
of the vascular plants known from Surinam. Leiden: E. J. Brill.
1909.
Beitr?ge zur Flora Surinams. Rec. Trav. Bot. Ne?rl. 6: 251-293.
1992. Transitions
between cerrado and forest vegetation
in Brazil. In Nalure and dynamics
Neue
_1909.
Ratter,
1993. Patterns
Bot. Gard.
Biogeography
ica, ed. T. C. Whitmore
Pulle, A. A.
Kew Bull. 23: 488^*90.

Dalbergieae.
Part 1, ed. R. M.
Gardens.
analysis.
J. A.
esl-savanna
boundaries,
ed. P. A. Furley,
J. Proctor,
and J. A. Ratter. London:
Chapman

and Hall.
offor
2003 ANDIRA 123
Ratter,
J. A., S. Bridgewater,
and J. F. Ribeiro.
R. Atkinson,
1996. Analysis
of the floristic composition
of the
Brazilian
cerrado vegetation
H: comparison
of the woody
of 98 areas. Edinb. J. Bot. 53:
vegetation
153-180.
Richards, A. J. Plant breeding systems, 2d ed. London: Chapman and Hall.

L. H., and S. J. Brunsfeld.
1992. Molecular
InMolecular
evidence and plant introgression.
Rieseberg,
systemat
ics of plants, ed. P. S. Solus, D. E. Soltis, and J. J. Doyle,
151-176. New York: Chapman and Hall.
M. G. M. van. 1985. Fruits of the Guianan flora. Utrecht: Utrecht University,
Institute of System
Roosmalen,
atic Botany.
J. N.
Rovirosa,
408^114.
1890. Productos
P. 2000. Cryptic
Rudall,
tematics:
London:
Stafleu,
coding
vegetales
characters
characters
en las palizadas
in monocotyledons:
for phylogenetic
Stevens, P. F. 1991. Character

553-583.
y sus afluentes.
Naturaleza,
and coding revisited. In Homology

homology
ed. R. W. Scotland and R. T. Pennington,
analysis,
Taylor and Francis.

and R. S. Cowan.
1976. Taxonomic
F. A.,
del Grijalva
states, morphological
Literature,
variation,
ser. 2,
and sys
114-123.
ed. 2, vol. 1:A-G. Regnum Veg. 94: 1-1136.

and phylogenetic
analysis: a review. Syst. Bot.
in papilionoid
Stirton, C. H. 1981. Petal sculpturing
legumes, la Advances
M. Polhill and P. H. Raven, 771-788.
Kew: Royal Botanic Gardens.
in legume
systematics,
1:
16:
Part 2, ed. R.
E. K. James, and T. Naisbitt.

structure with special reference to
1994. Nodule
Sutherland, J.M., S. G. Mclnroy,
the tribes Sophoreae, Genistieae
In Advances
and Thermopsideae.
Part 5, The ni
and J. I. Sprent, 41-56. Kew: Royal Botanic Gardens.
trogen equation, ed. D. McKey
Swofford, D. L. 1999. PAUP*. Phylogenetic
(*and other methods). Version 4, beta 2.
analysis using parsimony
Thiele,
Sinauer Associates.
Sunderland, Massachusetts:
K. 1993. The holy grail of the perfect character:
9: 275-304.
Toledo, J. F. Observa??es
28-31.
1946.
criticas
sobre nomes
the cladistic
de algunas
treatment of morphometric
plantas brasileiras.
Arq. Bot. Estado
data. Cladistics
S?o Paulo
2(2):
W. G.
1843. Repertorium
botanices
1: 803-804. Leipzig: F. Hofmeister.
systematicae,
1892. Lagoa Santa: Et bidrag til den biologiska
Selsk.
plantegeografi.
Kongel. Danske Vidensk.
Math. Afd., ser. 6, 6: 153-488.
Skr., Naturvidensk.
In Silvics of North America,
vol. 2, Hard
Weaver, P. L. 1989. Andira inermis (W.Wright) DC (SO-ITF-SM-20).
1-7 (USDA Forest Service Agriculture Handbook
DC: US Department
of Agri
woods,
654). Washington,
Walpers,
E.
Warming
culture.
P. 1988. Indirect and direct methods
in systematics.
In Ontogeny and systematics,
ed. C. J. Humphries,
Press.
University
forests of Africa: a preliminary
review. Gard. Bull. Singapore 29: 57-71.
White, F. 1976. The underground
T. C, and J.M. Williams.
1967. Radio tracking of homing bats. Science
155: 1435-1436.
Williams,
M. F, M. J. Sanderson, B. G. Baldwin,
and M. J. Donoghue.
1993. Monophyly
of aneuploid As
Wojciechowski,
internal transcribed spacer sequences. Amer.
tragalus (Fabaceae): evidence from nuclear ribosomal DNA
J. Bot. 80:711-722.
Weston,
27-56.
Wright,
-1787.
W.
New
York: Columbia
1777. Description
and use of the cabbage-bark
An account of the medicinal
plants growing
tree of Jamaica.
Philos.
in Jamaica. Lond. Med.
Trans.
67: 507-512.
J. 8: 217-295.

124
VOLUME64
APPENDIX 1
Restriction
Restriction
site characters
study by Pennington
Character2
3
used
to assess
phylogenetic
Sites
of species
position
not included
of Andira
Enzyme
Probe
Asel
regionb
Site Diagnostic
Mutation0
MB8+9
3.6=1.8+1.7
1.8 = 1.05+
For
Clade H (site absence)
Asel
MB8+9
Asel
MB11
2.0=1.7
Asel
MB11
3.7 = 2.0+1.7
MB 13
1.0 = 0.6 + 0.4
n.d.
+ n.d.
Clade
III (site presence)
Clade
I (site presence)
Clade m
(site absence)
I (site absence)
Clade
Asel
Clal
MB2
1.6=1.4
+ n.d.
Clade m
(site absence)
10
EcoRI
MB2
1.5 = 1.2 + n.d.
Clade n
(site absence)
11
EcoRI
MB2
2.6 = 2.35 + n.d.
Plastome Group I and

Clade H (site presence)
12
EcoRI
MB3
3.85 = 3.5 + n.d.
Clade U (site absence)d
5.2 = 4.0+1.2
Clade H (site presence)
MB8+9
3.6 = 3.2 + n.d.
Clade
19
HindHI
MB2
20
Hindm
34
XmnI
MB3
1.8 = 1.2 + 0.6
35
XmnI
MB11
0.96 = 0.6 + n.d.
I (site presence)
A. verm?fuga; A. humilis
RTP 269 (site presence)
Clade
I and Clade H
(homoplasy;
37
in the
(1995).
MB13
XmnI
3.5 = 3.4 + n.d.
Clade m
site absence)
(site presence)
numbers of those of Pennington

(1995, Table 2).
bProbe regions refer to the Vigna chloroplast genome
(Palmer et al. 1983).
(kb). Fragments not detected, either because of small size (probable mi
cFragment sizes are given in kilobases
as "n.d." (not detected).
problems, are designated
gration off the end of the gel) or possible probe homology
for an accession
of A. inermis (T. D. Penningion
dAbsence of this restriction site was also an autapomorphy
of A. inermis included in this study had this re
13558) included in Pennington
(1995); however, all accessions
Character
striction
site, and its absence
is diagnostic
for clade H.

2003
125
ANDIRA
APPENDIX 2
Inferred
Plastome
Types
of Andira
Accessions
and the accessions

of each species ordered numerically;
collectors are
species are listed alphabetically
AMC = A. M. de Carvalho; CEH = C. E. Hughes; HCL = Haroldo Cavalcante
de Lima; JR = J. A.
l Claudio da Silva; MS = M.
Ratter; LR = L. Rico; MC = M. Cheek; MCh = Mark Chase; MdS = Man
= R. T.
TDP = T. D. Pennington.
Accessions
marked with an asterisk are those
Sugiyama; RTP
Pennington;
screened in this study; the remainder are those screened previously
in the col
(1995). Notations
by Pennington
umn marked "Clade": I = restriction site mutations
characteristic
of clade I; II = restriction site mutations
char
The
abbreviated:
= restriction site mutations

characteristic of clade III; PI = restriction site mutations
char
of plastome
site mutations
characteristic
of the monophyletic
group I; PITH restriction
group of
(1995), but no muta
plastome group I and clade II [these accessions were screened previously
by Pennington
tions diagnostic
for clade II were screened].
acteristic
of clade II;m
acteristic
Species
Voucher
Herbarium
A. anthelmia
RTP 227
CEPEC,
FHO, K
Population
Clade
Locality
ffl
Brazil:
Bahia;
14?19'S,
A. anthelmia
RTP 281
CEPEC,
FHO, K
Brazil:
Bahia;
14?51'S,
A. anthelmia
RTP 282
CEPEC,
FHO, K
39?21'W
Brazil:
39?04'W
Bahia;
14?51'S,
39?04'W
A. anthelmia
RTP 208
CEPEC,
FHO, K
Brazil: Bahia;
A. anthelmia
RTP 286
CEPEC,
FHO, K
Brazil: Bahia;
A. anthelmia
RTP 294
CEPEC,
FHO, K
15?06'S,
15?05'S,
in
RTP 217
A. carvalhoi
RTP 229
CEPEC,
FHO, K
pi/m
CEPEC,
FHO, K
pi
Brazil:
CEPEC,
FHO, K
Brazil:
CEPEC
AMC*
pi
Brazil:
39?06'W
Bahia;
14?57'S,
A. cordata
RTP 262
CEPEC,
FHO, K
pi/n
Brazil:
A. cordata
RTP 263
CEPEC,
FHO, K
ii
Brazil:
RTP 264
CEPEC,
FHO, K
39?01'W
Bahia;
12?07'S, 45?06'W
Bahia;
12?07'S,
A. cordata
39?01'W
Bahia;
15?10'S,
A. carvalhoi
39?01,W
Bahia;
14?57'S,
RTP 233
39?24'W
Brazil: Bahia;
13?58'S,
A. carvalhoi
39?33'W
Brazil: Bahia;
15?06'S,
A. carvalhoi
39?33'W
45?06'W
Brazil: Bahia;
12?0rS,
45?06'W
A. cujabensis
RTP 467*
E,UB
A. cujabensis
RTP 474*
E,UB
ii
Brazil: Goi?s;
A. cujabensis
RTP 478*
E,UB
Brazil: Goi?s;
A. cujabensis
RTP 485*
E,UB
ii
Brazil: Goi?s;
Brazil:Goi?s;
16?44'S, 52^7^
16?52'S,
16?30'S,
16?44'S,

52^0^
52^3^
52?37'W
Species
VOLUME 64
126
Herbarium
Voucher
Population
Clade
Locality
A. cujabensis
RTP 486*
E,UB
Brazil: Goi?s;
A. cujabensis
RTP 493*
E,UB
Brazil: Goi?s;
A. cujabensis
RTP 497*
E,UB
Brazil: Goi?s;
52?37'W
16?44,S,
16?44'S,
47*23^
13?55'S,
A. cujabensis
RTP 498*
E,UB
ii
Brazil: Goi?s;
13?55'S,
A. cujabensis
RTP 503*
E,UB
ii
RTP 213
FHO, K
CEPEC,
in
Brazil:
RTP 236
FHO, K
CEPEC,
in
Brazil:
RTP 274
FHO, K
CEPEC,
39?03'W
Bahia;
15?10'S,
A. fraxinifolia
47?23'W
Bahia;
14?12'S,
A. fraxinifolia
47?23'W
Brazil: Goi?s;
13?55'S,
A. fraxinifolia
52?37'W
Brazil:
39?06'W
Bahia;
12?35'S,
41?16'W
A. fraxinifolia
RTP 280
CEPEC,
FHO, K
Brazil:
A. fraxinifolia
RTP 283
CEPEC,
FHO, K
Brazil:
A. fraxinifolia
RTP 318
CEPEC,
FHO, K
A. fraxinifolia
RTP 285
CEPEC,
FHO, K
Brazil: Bahia;
A. fraxinifolia
RTP 287
CEPEC,
FHO, K
Brazil: Bahia;
Bahia;
12?32'S,
14?56'S,
ni
39?01'W
15?05'S,
15?05'S,
RTP 298
FHO, K
CEPEC,
in
Brazil:
RTP 302
FHO, K
CEPEC,
Brazil:
RTP 303
FHO, K
CEPEC,
Brazil:
RTP 249*
A. fraxinifolia
RTP 250
39?14'W
39?!^
Bahia;
17?30'S,
A. fraxinifolia
39?33,W
Bahia;
17?30'S,
A. fraxinifolia
39?33'W
Bahia;
17?13'S,
A. fraxinifolia
39?01'W
Brazil: Bahia;
14?55'S,
A. fraxinifolia
41?14'W
Bahia;
39?!!^
CEPEC,
FHO, K
Brazl: Bahia;
CEPEC,
FHO, K
Brazil:
llo02'S,40?43'W
Bahia;
11?02'S,40?43,W
A. fraxinifolia
MS 889
K,SP
ni
Brazil:
23?S,
A. galeottiana
A. galeottiana
A. humilis
LR s.n.
MCh
s.n.*
RTP 239
CEPEC,
pi
Mexico:
pi
Mexico:
in
Brazil:
FHO, K
S?o Paulo;
38?W
Oaxaca;
17?N, 95?W
Chiapas;
17?N, 91?W
Bahia;
38^7^
12?15'S,
A. humilis
RTP 246
CEPEC,
FHO, K
Brazil:
Bahia;
11?09'S,40?32,W
A. humilis
RTP 247
CEPEC,
FHO, K
Brazil:
Bahia;
llo09,S,40?32/W

2003
ANDIRA
Voucher
Species
Herbarium
127
Clade
Population
Ahumilis
RTP 267*
CEPEC,FHO,K
PI
Ahumilis
RTP 268*
CEPEC,
FHO, K
PMI
Locality
Brazil:Bahia;
12?10'S, 44^3^
Brazil:
Bahia;
44?43'W
12?10'S,
CEPEC, FHO,K
RTP 269
Ahumilis
PI
Brazil: Bahia;
12?10'S, 44?43'W
RTP 499*
Ahumilis
PI
3
E, UB
Brazil:
Goi?s;
47?23/W
13?55'S,
RTP
A. humilis
PI 3
E, UB
500*
Brazil:
Goi?s;
47^3^
13?55'S,
RTP
Ahumilis
PI3
E, UB
501*
Brazil:
Goi?s;
47?23'W
13?55'S,
RTP 502*
Ahumilis
PI
3
E, UB
Brazil:
Goi?s;
47?23'W
13?55'S,
Ahumilis
RTP 506*
E, UB
Ahumilis
RTP 507*
E, UB
PI
4
Brazil:
Goi?s;
14?07,S,47?31/W
PI4
Brazil:
Goi?s;
14?07,S,47?31'W
A. humilis
RTP 508*
E, UB
PI 4
Brazil:
Goi?s;
14?07'S, 47?31'W
RTP 509*
A. humilis
PI 4
E, UB
Brazil:
Goi?s;
47?31'W
14?07'S,
A.humilis
MdS
1*
PI 5 Brazil:
D. F.;
47?50'W
15?55'S,
Ahumilis
2*
MdS
PI 5
Brazil:
D. F;
47?5(TW
15?55'S,
MdS Ahumilis
3*
PI
5
D. F. ;
Brazil:
47?5(TW
15?55'S,
4*
MdS Ahumilis
MdS
Ahumilis
PI
5
5*
D. F.;
15?55'S, 47?5(W
Brazil:
5 PI
D. F.;
Brazil:
47?50/W
15?55'S,
inermis A.
TDP13558
1 K
Costa Rica: Puntarenas;

83?19/W
09?07'N,
A.m^w
CEH1673
FHO, K
3579
K
3
I
Mexico:
Oaxaca;
16?22/N,94?11'W
miroi?A.
MC
A.
inermis
merm?5
RTP 512*
A.
RTP 580*
I4
E, QCNE
E, INBIO
Cameroon,
SW Province;
03?58'N,
09?16'E
00?33'S,
Napo;
77?50^
Ecuador:
Rica: Puntarenas;
83^0^
Costa
08?42,N,
A.
?n*/row
RTP
589*
E, INBIO
5 I
Rica: Puntarenas;
83?3(TW
Costa
08?42'N,
A. te^a/w
RTP
305
CEPEC,
FHO, K
HI
Brazil:
Bahia;
17?30'S,
Alegalis
RTP 307
CEPEC,FHO,K
ffl
39?!^
Brazil:Bahia;
17?30'S,

39?!!^
128
VOLUME64
Species
Voucher
A.
legalis
RTP 308
legalis
HCL
Herbarium
FHO, K
CEPEC,
Population
Clade
Locality
ffl
Brazil: Bahia;
Brazil: Rio de Janeiro;

22?55'S, 43?10'W
17?30,S,39?11'W
A.
s.n.
A. macrothyrsa
TDP 13550
PI
Peru: Loreto;
00?S, 73?W
A. macroihyrsa
RTP 523*
E,QCNE
PI
Ecuador:
00?40'S,
A. marauensis
AMC
s.n.
CEPEC
ffl
Brazil:
Bahia;
15?09'S,
A. nilida
RTP 300
CEPEC,
FHO, K
ffl
Brazil:
A. niiida
RTP 304
CEPEC,
FHO, K
ffl
Brazil:
A. niiida
RTP 313
CEPEC,
FHO, K
CEPEC,
FHO, K
ffl
Brazil:
RTP 288
CEPEC,
FHO, K
ffl
Brazil:
39?14'W
Bahia;
13?02'S,
A. nitida
RTP 289
CEPEC,
FHO, K
ffl
Brazil:
RTP 292
CEPEC,
FHO, K
ffl
Brazil:
RTP 301
CEPEC,
FHO, K
ffl
Brazil:
A. nitida
RTP 316
CEPEC,
FHO, K
ffl
Brazil:
A. nitida
RTP 237
CEPEC,
FHO, K
ffl
Brazil:
CEPEC
ffl
Bahia;
Brazil:
RTP 433
A. surinamensis
RTP 458
FHO, K, U, US
ffl
A. surinamensis
RTP 462
FHO, K, U, US
ffl
A. surinamensis
RTP 463
FHO, K, U, US
ffl
FHO, K, U, US
ffl
39?06'W
Bahia;
15?18'S,
A. surinamensis
39?13/W
Bahia;
15?06'S, 30?\1^
15?10'S,
AMC 3309
38?23"W
Bahia;
17?30'S,
A. nitida
38?23'W
Bahia;
13?02'S,
A. nitida
38^3^
Bahia;
13?02'S,
A. nitida
39?!^
Bahia;
17?13'S,
A. nitida
39?!^
Brazil: Bahia;
17?30'S,
III
39?13'W
Bahia;
17?30'S,
RTP 311
39?05'W
Bahia;
17?30'S,
A. niiida
Napo;
77?10'W
39?04'W
Guyana:
0A?\3^, 58^8^
Guyana:
04?13'N,
58*58^
Guyana:
0A?\3f"H, 58^8^
Guyana:
04?09,N,
59^2^
A. verm?fuga
RTP 256*
CEPEC,
FHO, K
PI
Brazil: Bahia;
A. verm?fuga
RTP 257*
CEPEC,
FHO, K
PI
Brazil:
11?15/S,42?52,W
Bahia;
11?15,S,42?52,W
A. verm?fuga
RTP 258*
CEPEC,
FHO, K
PI
Brazil:
ll015'S,
A. verm?fuga
RTP 259
CEPEC,
FHO, K
PI
Bahia;
42^2^
Brazil: Bahia;
11?15'S, 42^2^

ANDIRA
2003
Herbarium
129
Species
Voucher
A. verm?fuga
RTP 265
CEPEC, FHO,K
PI
A. verm?fuga
RTP 270
CEPEC,FHO,K
PI/H
Population
Clade
Locality
Brazil: Bahia;
12?07'S, 45?06'W
Brazil:
12?07'S,
RTP 271
verm?fuga
FHO, K
CEPEC,
PI
Brazil:
Bahia;
43?15'W
Bahia;
43?15'W
12?07'S,
A. verm?fuga
RTP 272
CEPEC, FHO,K
PI/II
Brazil: Bahia;
12?07'S, 43?15'W
A. verm?fuga
RTP 273
CEPEC,FHO, K
PITH
Brazil:
Bahia;
12?07'S,
RTP 465*
A. verm?fuga
E,UB
PI
43?15'W
Brazil: Goi?s;
16?52'S,
52?20'W
E,UB
PI
Brazil: Goi?s;
E,UB
PI
Brazil: Goi?s;
verm?fuga
RTP 471*
E,UB
PI
Brazil: Goi?s;
16?52'S, 52?20'W
A. verm?fuga
RTP 504*
E,UB
PI
Brazil: Goi?s;
14?07'S, 4713^
A. verm?fuga
JR 7232V*
PI
Brazil: Goi?s;
A. verm?fuga
RTP 466*
A. verm?fuga
RTP 469*
16?52'S,
16?52'S,
14?07'S,
NUMERICALLIST OF SPECIES
1. A.
inermis
la. A.
inermis
subsp.
not determined)
(subspecies
inermis
14. A. legalis
15. A. ormosioides
lb. A.
inermis
lc. A.
inermis
subsp. glabricalyx
subsp. rooseveltii
17. A. nitida
2.
3.
A. jaliscensis
A. multistipula
4.
A. cubensis
5.
A.
16. A. fraxinifolia
18. A. marauensis
19. A. carvalhoi
6.
20. A. parviflora
21. A. cujabensis
22. A. cordata
7.
23. A. micrantha
9.
A. verm?fuga
10. A. humilis
25. A. grandistipula
26. A. praecox
12. A.
11. A. macrocarpa
surinamensis
28. A.
13. A. anthelmia
29. A. unifoliolata
taurotesticulata
A. macrothyrsa
A. chigorodensis
8. A. galeottiana
24. A. cori?cea
27. A.
tervequinata
trifoliolata

52^0^
52^0^
47?13'W
130
VOLUME 64
INDEXTONUMBERED COLLECTIONSEXAMINED
The numbers
Species
presented
in parentheses
above.
refer to the corresponding
3775 (la).
1070 (lb).
P., & J. A. Cede?o 7430 (3).
P., et al. 8127 (28); 8357 (28).
species
A., & W. Rodrigues

F., et al. 3102(3).
Acevedo,
P., & A. Sinca
Aubreville,
Acevedo,
R., & J. Sosa
Ayala,
Acevedo-Rodr?guez,
Acevedo-Rodr?guez,
Ackerly, D., et al. INPA/WWFl302.4500.2
Adams, C. D. 7843 (la).

Aguilar, R. 228 (la).
Alain 4404 (4).
Albert de Escobar, L., & J. I. Santa 3527
Albert
de Escobar,
H. A.
(la);
13885
(12).
8233
?feL. Delgado
133 (la).
C, et al. 700 (la).
G.,
(12).
B.
Azuara,
(20).
Azurdia,
BAFOG 51N (24); 1183 (12); 1201 (24); 1264 (24).

<fcE. Z. Bailey
L. H.,
Bailey,
149 (la);
52 (la);
321
(la); 1352 (12); 1657 (12).
(la).
L., et al. 1848 (5).
13367
681
List of
Aymard,G. 6147 (27); 7991 (12); 9263 (28).

Aymard,
7751 (16).
J. B., & J. R. Guilamon
Baker, M. A. 7025 (5).
Balslev, H. et al., 84821 (la); 97505 (3).
Bamps, P. 5078 (10).
Baitello,
Alencar (RB 54889) (12); (RB 61016) (9).

Allard,
in the text and in the Numerical
(la).
Allen, P.H. 873 (la); 938 (la); 1770 (la); 2955 (la);
4435 (la); 4470 (la); 4482 (la); 5220 (la); 5458
(la); 5609 (la); 7190 (la).
Almeida, J. 12 (13); 356 (17); 2044 (14).
Alu?sio, J. 115 (20); 224 (29); 226 (29).
393 (la); 432

1090 (la).
Barbosa, C, et al. 1856 (la).
Barkley, F. A., & M. Hern?ndez
Barlow, F. D., 30/186D
(8).
D., et al. 194 (9).

D. L. 145 (21).
A. M. A., et al. 347
Amorim,
Barreto,M. 937 (16); 1515 (16); 1782 (16); 5485 (10);

5491 (10); 5590 (16); 5592 (16); 5608 (16);
5764 (16); 10096 (16).
J., & T. S. dos Santos
Almeida,
103 (17);
130 (17).
Alvarenga,
Amaral,
(17); 348
(17);
1055
(16); 1238 (18); 1411 (18); 1896 (16); 1908 (18).

C. W.
Anderson,
332
W. R., et al. 7159
(10); 36024
(16); 36607
(22); 36865 (10).

Andrade-Lima
& Medeiros-Costa
105 (17);
154 (17).
et al. 7 (16).
F. W. 464 (le).
Andrade-Lima
Andrews,
Bartlett, H. H.,
(9);
3441 (9).
Arana, A. 27 (la).
Araujo, D. 1843 (13); 3553 (16); 4612 (14); 5289

(16); 6176 (16); 6598 (16); 7390 (16); 8079
(14); 8295 (16); 9687 (16); 9763 (16).
4874 (16).
D., & R. Henriques
3683 (16); 4044
D., & N. C. Maciel
Beard,
Araujo,
(13);
6031 (16).
D., & A. Magnanini
D,. & J.Mauro 8614 (14).
D., & A. L. Peixoto 324 (17).
Araujo, D., et al. 3461 (16); 8326 (14); 8597

Archer, W. A. 2050 (la); 2563 (la).
L. 2805 (la).
Aristeguieta,
Atchinson,
E. 276
(4).
16319
(la).
(la).
(la).
S. G.
18686 (la).
S. G., et al. 19556
Becker, R. M. 42 (21).
Beck,
(3).
Belem, R. P. 1830 (14); 1879 (13).

R. P., ?feR. S. Pinheiro
Belshaw,
C. M.
Bena,P.
1167(12).
2445
3370
(16); 2550
(16);
(la).
A. 3658 (lb); 3796

Benitez-Paredes,
Bernai, H. Y. 158 (la).
Black, G. A. 54-17372
(la).
Black,
Black,
(lb).
G. A., & Klein 54-17197

(la); 54-17372
(la).
G. A., & P. Ledoux 50-10366
(la); 50-10386
(12).
Black,
4259 (16);4465 (16); 7514 (16); 8700 (14).

Araujo,
40602
2591 (16); 2807 (17); 2949 (13); 2988 (17);

3089 (18); 3091 (18); 3142 (18).
L. M. 3-7 (la).
N., & M. Heinrich GUI224
(8).
Antonio, T. 3616 (la).
da Silva, M., et al. 1352 (22); 2738
Aparecida
Araujo,
(la).
J. S. 157 (la).
P. 1336 (la).
Beck,
Andrews,
Araujo,
?feT. Lasser
J. H. 6332
Beaman,
Beard,
Belem,
Antonio,
Araujo,
W. N.
C.
Bautista,H. P. 533 (17); 1145 (17); 1368 (17).
(12).
Anderson,W. R. 6231 (10); 6456 (10); 6771 (21);

7150 (21); 7293 (21); 7294 (9); 7895 (10); 9560
(21); 9691 (21); 10051 (21); 11292 (la); 11293
(21); 11361 (21); 36797 (22).
Anderson,
Bangham,
Barbosa,
G.A.,
et al. 54-16454
(12)
Blanchet, J. S. 607 (13); 2723 (16); 3089 (16); 3137

(10); 3672 (16).
Boege, W. 1244 (8).
Bohrer, C. 47 (16).
Bonder, G. 2465 (16).
(14).
Bonpland 1599 (la); 3901 (la).

Boom,
B. M.
Boom,
B. M.,
B. M.,
Boom,
6738
(la); 6791 (la); 7198 (la).

?feA. L. Weitzman
5233 (la).
<&
M. Grillo
6460

(12).
2003 ANDIRA 131
Boschbeheer, (B.W.)45 (12); 57 (12); 61 (24); 1909

(24); 1944 (12); 2216 (12);2458 (12);2579 (12);
3547 (12); 3933 (24); 4092 (24); 4210 (24);
5245 (24); 5252 (24); 5450 (24); 6870 (24).
?feCunningham
1385 (la).
Bowie,
N0.3
(16).
114 (7); 178 (la); 215
J., ?feA. Cogollo
M. Gonz?lez
988 (6).
Brand, J., ?fe
1533 (5).
Brant, A. E., <&J. Roldan
Bravo, H. 8 (8); 142 (8).
D.E.
Breedlove,
A. M.
Brenes,
257
37711
Cedillo,
(16); S447
S243
Bridgewater,
S.,
?feJ. Fonseca
Filho
S., et al. S241
Bridgewater,
(21); S246
Britton,
1127(3) (la).
E. G., & D. W. Marble
Britton,
N. L. 2906
Britton, N. L.,
Britton, N. L.,
(10); S363
2328
Bunting,
Christenson,
(la).
10259
(4).
(la); 6944
(la).
(la).
?feH. de Cabrera
Cabrera
Salge, N. 13 (la).
S. 317 (la).
R., et al. 8584 (5).
Calder?n,
Callejas,
(la).
J. L, et al. 11500
Campos,
A.
Campos,
1352 (la);
9 (16).
7028
(8); 7378
(la).
(la); 18688
1381 (la).
(lb).
Capucho
J. P. P., et al. 2593
M., et al. 2020 (28).

J. E. R. 91 (17).
C. L. 155 (9).
Collenette,
Cooper,
(la).
133 (16).
E. 7083 (la); 7600
A.
D. 81 (28).
G. P., & G. M.
(la);
10139
Slater 9 (la);
(la).
147 (la);
215
(la); 265 (la).

Coradin,
(14).
?feJ. G. Ram?rez
Cordeiro,
2741
(5); 2743
M.
L. 256 (la).
Carrasquilla,
Carvajal, A. 263 (la).
G. 953
Colonnello,
Contreras,
C?rdenas,D. 1300 (la); 1485 (la); 2501 (5).

1346 (la).
Carlson, M. C. 511 (la);
(20).
(5); 199A (5).
Collares,
Coomes,
Carauta,
C?rdenas,
270
Cogollo, A., et al. 6036 (5); 6346 (5); 6353 (5); 6390
(5); 6400 (5); 6486 (5);6521 (5); 6583 (5); 6907
(5); 6958 (5); 7281 (5).
Constantino,
J.M.
D.,
A., & C. C. Estrada
Cogollo,
Candida, M.,
C?rdenas,
L., et al. 394 (3).

L. S., et al. 103 (29); 270
Co?lho,
Co?lho,
Colella,
Porto, P. 2650 (16).

et al. 168 (16).
481 (la).
Campos
(9); 6541 (21); 7250 (3); 9948 (la); 10824 (3);

10896 (la); 10950 (6).
Co?lho,L. 115 (20); 394 (23).
Calzada, J. I. 19 (8); 988 (8); 14929 (8); 16846 (8);

16880 (8); 19293 (la).
Calzada,
C. A., & J. Lima 3611 (3).

Cid Ferreira, C. A., & B. W. Nelson
3057 (6).
Cid Ferreira, C. A., et al. 6259 (21); 6270 (la); 6416
P. 12 (10); 62 (10); 236 (10).

A. J., et al. 4339 (la).
Cobra, L. A., & J. Oliveira 69 (9).
Co?lho, D., & C. Mota 621 (3).
G. S., et al. 4125 (12).

Burchell, W. J. 8491 (21); 8544-2 (21).
?feG. Matta 4767 (la).
Burger, W.,
E.,
Cid Ferreira,
Clewell,
Bunting,
Cabrera,
Cid Ferreira,C. A. 3380 (la); 3382 (la); 4268 (29);

5247 (6).
Claussen,
(la).
2243 (la).
S. H. 910 (2); 1313 (2).
G. S. 7707 (la); 7795 (la); 7976
C. A. C. 2728
(1).
(la).
F. 1583 (la).
G. M., et al. 1432 (28).
Choussy,
Britton, N. L., et al. 1776 (12); 14246 (4).

Broadway, W. E. 611 (la); 3170 (la); 6667
R. J. 12630
F., et al. 765
Chiang,
(21).
(la); 4848 (4).

?feJ. F. Cowell
1018 (la);
?feP. Wilson
474 (4).
(la).
Brooke, W. M. A. 5825
D. 412 (la).
Ch?zaro, M., et al. 5474 (lb).
Chevalier, A. 7773 (le); 24917
(8).
Ch?vez,
(9).
S354
(10); S364 (10).
Bullock,
Ch?velas,
Ch?velas,
?feE. Cardoso
(la).
(le).
U. 581 (la).
J., & L. A. P?rez 80 (8); ES-892
(8).
J., & C. Zamora ES-4742
(la); ES4890
J., et al. ES-2396
(8); ES-4301
(8).
Ch?velas,
(la);
(la).
Ib??ez-Garc?a S435
S.,
Brunt, M.
J. D. 4203
Chavarria,
(la).
(16).
Brooks,
(3).
T. B., et al., 1738 (9).
17 (8).
T., & J. I. Calzada
Cavalcanti,
Chapman,
Bridgewater,
Bristan,N.
G.
H. 56 (24).
Champagne,
Chan, F, & R. Lira 4707
15595
?feA.
S.,
(7).
(9).
12619
Bridgewater,
(la); 6136
Castillo,
P. I. S. 2364
J. E. M.
5218
(la);
A. 25336 (9).
Castellanos,
Castillo, A. 2353 (12).
Castillo,
Brand,
Braz?o,
103 (la); 584
(la); 6430 (la).
1115 (la).
J. 830 (la).
P. 238 (la); 3164
Cavalcante,
28 (16).
Box, H. E.
Braga,
R. R.
Casta?eda,
Cordovil,
(5).
Correa,
L., et al. 6510 (16).

L, et al. 825 (16).
S., et al. 119(9).
J. A. 22 (16).
& M. A. Magana
Cowan, C,
1136 (la);
1300 (la).
3046
(la).
Cowan, C, et al. 2294 (8).

Cowan, R. S. 38846 (la).
Croat, T. B.
17753
(3); 17759

(la);
19235
(la);
19235
132
(la); 19313 (la); 20388 (la); 20467 (la); 24471

(la); 24861 (la); 26024 (5); 30320 (3); 32787
(la); 61182 (la),
da Cruz, N. D.
Cuadros,
H. 643
13 (15).
(5); 2965
(la).
Cuatrecasas, J. 4079 (12); 16224 (la); 16857 (la);

17410 (la); 21441 (la); 21461 (la).
J., & L. Willard
Cuatrecasas,
26061
1265 (la).
Curran, H. M. 23 (la); 71 (la);
Curtiss, A. H. 525 (3).
(la).
Cumings
da Silva,
S. B. 67 (21).
M. G. de Lima
S. B., ?fe
da Silva,
B. E. 880 (9).
Dahlgren,
?feR. Cardoso 3923
Daly, D. C,
Daly, D. C, et al. 6272 (9).
168 (le).
L. 10345
Damazio,
Daniels,
Davidse,
Davidse,
Delgado,
Delgado,
372
89 (5).
(5).
A., et al. 480 (2).
E. 311 (la).
(la).
D?az,C, et al. 670 (3); 1295 (la); 1315 (3).
(26).
D?az, M. G. 544
(la).
S. 1279 (5).
L. Th. 9799
Dombrowski,
D?az,
(10).
(10).
47 (12); 57 (la).
R., & E. Gonzales
J. 2389 (la); 2389 (la); 2823 (la).
Donnell-Smith,
Dorantes,
J., et al. 874 (la).
(12).
A. G. H., <&F. P. Jonker 868 (24).
G. 18661 (la); 18831 (la).
<&A. C. Gonzalez
12316
Dom?nguez,
(12);
13913
G.,
Davidse,
G., & J. S. Miller 26752 (28).

G., et al. 16784 (12); 18661 (la);
?feG. Herrera
<&
G. Donahue
34185
31242
(la).
(la).
8513
dos Passos, B. C, & L. A. Dambios

dos S. Schettini, C. M. 4 (16).
dos Santos, T. S., & L. A. Mattos

18831
(la).
(la),
Duarte,
A. P., & E. Pereira
de Bruijn, J. 1358 (12).

de Carvalho, A. M. 2544
Duarte,
L., & A. Castellanos
de Carvalho,
de Carvalho,
Dubs,
(19).
A. M., & S. Faria 2544 (17); 2547 (19).
A. M., et al. 178 (19); 491 (19); 622 (19);
764 (16); 2421 (16); 3180 (16); 3308 (19); 3309

(17); 3508 (14); 3509 (14); 3510 (18); 3891
(10); 6141 (16); 6475 (18).
de Castro, R. A. R2102
(21).
de F. Almeida,
E. 140 (9).
de Freitas Campos,
J.M. 31 (10); 72 (10).
de Goes 24 (16).
<&E. A. L?pez
(16).
de Haas, Sr., et al. 122 (9)
de Lima, A. 49-313
(17); 49.169
?feJ. Caruzo
3059
?feJ. Guedes
2731
et al. 2741
Duarte,
3261
4724
(16).
602
(9).
108 (16); 146 (9).
Ducke, A. 599 (la); 674 (29); 1035 (3); 1036 (6);

1822 (3); 2229 (20); 8395 (12); 11509 (la);
17234 (12); 17293 (28);21348 (20);21365 (23);
23401 (23); 23864 (23); 23865 (20);23866 (20);
35073 (12); 35076 (la); 35077 (23); 35078 (29);
35079 (28).
Duke,
(17).
A.
841 (la).
J. A., & J. D. Kirkbride
14070
(la).
Dus?n, A. 8756 (16); 10673 (10); 16493 (16); 16511

(16); 17253 (16).
1089 (la); 3231
Duss,A.
(la).
Dwyer, J. D. 2338 (la); 11419 (la); 1463A (la);
(16).
11130A(la).
Edwall 1621 (16);4004 (16); 5650 (10).

(3); 2783
(16).
Duke, J.A. 7514 (la); 9781 (la); 11120 (la); 11122

(la); 12004 (la); 13500 (la); 14357 (la); 8412
(3), 11017(3) (la).
327 (16).
(3).
(la); 2767
L., & R. S. Santos

B. 475 (la).
Dugand,
de Lima, H.C. 506 (16); 656 (16); 1173 (16); 1668

(17); 1711 (16); 1712 (14); 2922 (17); 2951
(16); 2966 (17); 3866 (13).
de Lima, H. C,
de Lima, H. C,
de Lima, H. C,
Silva
Duarte, A. P. 2856 (10); 3282 (9); 4263 (15); 9724

(16); 10555 (16).
de Barros,F. 1356 (16); 2238 (16).
de Grande, D. A.,
de Haas, Sr. 2950
(21).
dos Santos,T. S. 954 (16); 2108 (14); 2182 (16); 3062

(14); 4432 (13).
G., & T. Hawkins
Davidse,
C,
(21).
J. I. A-5
D?via, W. 941 (5).

Dias, A. C. 29 (16).
D?az, C, & J. Ruiz 953
(22).
Davidse,
Davidson,
(21).
Delgado, L. 44 (12); 551 (28).
(20).
(12); 21714 (12).
Davidse,
de Santos, R. R., et al. R1242

de Silva, M. F. 373 (29).
de Sousa Silva, et al. 65 (la).
del Valle,
C. 2890
G.,
de Queiroz, L. P., et al. 2957 (16).

de Saint-Hilaire, A. 1755 (10).
de Sant'Ana, S. C, et al. 113 (17).
del Llano, M.
(la),
Dalziel
Dami?o,
de Mello
Dean
et al. INPAAVWF2303.4846
J.M.,
P. M., et al. 20 (19).

Filho, L. E. 3008 (17).
de Queiroz, L. P. 1610 (16).
de Lima, M.
de Souza Lima, J. P. 208

15552 (la).
198 (la).
D'Arcy, W. G. 4727 (la).

D'Arcy, W. G., ?feK. Sytsma 14583
da Silva Araujo, J., et al. 108 (17).
da Silva, F. C, et al. 257 (21).
da Silva,
VOLUME 64
(6);
2950 (16); 3673 (16); 3845 (16); 4363 (16);

4366 (16).
J. B. 429P
Edwards,
(la).
124 (la), 1421 (12).
Egler, W. A. 625 (la).
Eggers,
B.

2003 ANDIRA 133
Eiten,G. 1724 (10); 4513 (9).

L. T. Eiten 2262 (10); 2363 (10); 2381
Eiten, G., ?fe
(10); 3331 (16); 5457 (10); 8440 (19);9849 (9).
Ekman,E. L. 1925 (4); 1975 (4);4326 (la); 6232 (4);
9687 (4); 12587 (la);H12587 (la).
Elias de Paula, J. 3122 (22); 3147 (16); 3277 (21).
Elias
de Paula, J., ?feC. A. Concei?ao

C. O. 418 (la).
1681 (la).
Erlanson,
91 (la).
<&B. I. A. Giacometto
Espina
Espina,
2994
(17).
Fern?ndez,
(la); 342 (la).

A. 6694 (12).
Fern?ndez,
M. M.,
?feJ. E. R. Collares
Fern?ndez-Casas,
J. <&
Carvalho
Fern?ndez-Casas,
J. F,
2823
?feJ. Molero
17 (22).
(la)
3790 (10); 3851
J. F, et al. 7452 (10).

Fern?ndez-Casas,
J. L., et al. 7477 (la); 12976 (la).
Fern?ndez,
304 (22); 322 (22).
Ferreira, J. A., ?feC. A. Miranda
<&
A. P. Araujo 61 (la).
Ferreira, M. C,
1907 (16).
(10).
(2).
Foli, D. A. 011 (16); 128 (14); 336 (15); 164/79 (15).

Fonnegra,
J. P. 3078
R.,
(la); 10047 (la).

et al. 1709 (la); 3022
(5); 3053
(5);
3092 (5).
(la).
Forero, E., et al. 1595 (la); 4004
(la); 4127
10889
A., & E. Zardini
Fonseca,
P. E., et al. 2713
Gibbs,
L. J., & H. Persaud
Godfrey,
1217 (la).
G?mez,
G?mez,
G?mez,
(la); 4695
R. K. 66999
(la).
V. 2864
(14).
L.D.
18517 (la).
L. D., et al. 22935
R., et al. 588 (12).
et al. 11333
F,
Gonz?lez,
J., & J. Casta?eda
Gonz?lez,
Forment, W. L. 737 (la).

Fosberg, F. R. 57497 (la).
13259
Foster, R. B., ?feE. Vivar
Gonzalez,
J., & A. P?rez

L., & V. Garza
Fredholm, A. 3176 (la).

Freitas 63 (3).
1042 (le).
Friis, L, ?feK. Vollesen
(16).
Glaziou, A. 6193 (16); 7600 (16); 10571 (13); 13684

(16); 19051 (16); 20274 (16).
Gonzalez,
(la).
(19).
(9); 2782 (9); 3262

R656 (9).
Gifford, D., & J. Ramos

Gill, J. R. 12621 (12).
(la); 8278 (10); 10205 (5).

ForestDept. 5655 (12); 7068 (la); 7692 (la).
Foster, R. B., et al. 11027
(28).
50011
Gentry,A., et al. 15668 (3); 30215 (3?); 36390 (12);

37168 (6); 40432 (la); 40434 (6); 42145 (la);
42184 (la); 43181 (5?);60863 (6); 65327 (5).
Gomes,
R. M. 1607 (la).
Fonzar, L. P. M., et al. 79 (la).
Forero, E., ?feR. Jaramillo 4434
(la).
L. J., et al. 1227 (la).

Gillespie,
Gillis, W. T. 9417 (la).
Giordano, L. C, et al. 08 (13).
Filho,
Folsom,
Se S. Tillett
Gentry, A.,
Gentry,
Gillespie,
S. J. 63 (16).
Fishlock, W. C. 111 (la).
Flores, G., et al. 26 (la); 1841 (lb); 4795
(la).
Gentry, A., & E. Renter?a 24163 (la).

Gentry, A., & D. Smith 36062 (la).
(10); 4079 (10); 6362 (10).
(16);
(la).
Gentry, A., & N. Jaramillo 58003 (12).

Gentry, A., & P. N??ez 69571 (la); 69572
317
K. 6206
A. 4179
Gentle, P.H. 15 (la); 1346 (la); 3345 (la); 3443 (la);

3902 (la); 4032 (la); 4794 (la); 4979 (la); 5325
(la); 9104 (la).
Gentry,A. 6373 (la); 6433 (la); 7887 (la); 7968 (la);
8041 (la); 8231 (la); 9267 (la); 9426 (la);
13455 (la); 43349 (la).
Gentry, A., & H. Cuadros 47545 (la); 55445
Gentry, A., & M. Fallen 17165 (la).
Gentry, A., & M. Horna 29375 (la).
1938 (la).
Ferreira, V F. 418
(6).
A. P. 71 (la).
Gamier,
Felippe, G. M. 36 (10); 78 (10); 94 (9).
Fiebrig,
(la).
Galusser,
Gardner,G. 1538 (9); 2330 (21); 2552 (9); 2816 (10);

3654 (21).
Fawcett, W. 8017 (la).

Fay, J.M. 4321 (la).
Fendler, A.
C, et al. 283
C. 22 (la).
A. 18 (la).
Gamon,J.
Garbier, A. P. 71 (la).
Gallardo,
Gardier,
Farney,C, et al. 2610 (16); 2615 (17); 2653 (17);

2930 (17); 3425 (17).
C.
(12).
A., & R. Torres 1695 (la).

443
F., & E. D. Agualimpia
H. 5122-A (la).
Garc?a-Barriga,
Faden, R. B., et al. 76/125

(8).
Falc?o, J. I. A., et al. 944 (16).
Farney, C. 2880 (17).
Feddema,
27684
Garcia,
M. Mosquina
2126 (la).
J., <&
Espinal, S. 195 (5); 1194 (la).
96 (5); 1698 (5).
Espinal, S., ?feE. Montenegro
Espinosa, G. 24 (la).
Euponino, A. 167 (17); 414 (17).
?feJ. C. Gomes
Froes, R., & G. A. Black
Galeotti,H. 3428 (8); 3464 (8).
Garcia,
(la).
Espina,
Farney, C,
Froes, R. 3 (la); 1836 (la); 20198 (16); 25660 (12);

12727/99 (13).
(la).
(la).
205
(la).
190 (la); 239 (la).
7009 (la); 7252 (la).
Goodland,R. 155 (9); 739 (10); 905 (10); 933 (10).

(la);
13259
(3).
Gottsberger,
Gottsberger,
Gottsberger,
Gottsberger,
G., et al. 23-31083

(9).
I. 2131 (10); 29R-29176
(9).
I., & C. J. Campos 420 (10).
15-9482
L, & G. Gottsberger
40169 (10); 12-221174 (9).

(22);
15
134
L, et al. 30582
Gottsberger,
E. 22/88 (16).
Grandez, C. 1433 (3).
(12).
Gouvea,
M.
(le).
(3); 2298
1916 (la).
M. L. 972
Guedes,
(17).
B. 766
Guerrero,
(2).
Guerrero, B., et al. 1262 (2).
Guillen, R. 2721 (la).
R. Chore 2913
Guillen, R., <&
Guillen,
R.,
<&S. Coria
Guillen,
R.,
?feT. Killeen
A. A.
2127
Heringer, E. P. 92 (10); 271 (16); 1970 (16); 2700

(15); 3555 (10); 3559 (10); 3706 (16); 3742
(16); 5411 (10); 6044 (16); 7120 (9); 7129 (9);
7200 (16); 7341 (9); 9029 (10); 9263 (9); 9814
(10); 9839 (10); 9844 (10); 10064 (16); 11175
(9); 11329 (16); 12228 (la); 12877 (9); 16202
(10); 17135 (10); 17440 (9).
E. P., & Paula 13927
E. P., & C. T. Rizzini
Heringer,
Heringer,
(la); 3194
(la).
Hernandes
Bicudo,
(la).
Hern?ndez,
Carvajal, L., <&J. D. L?pez

D. Castro 138 (la).
Guzm?n, M., <&
Gu?zar, E. 726 (la).
Guizar, E., <&F. Ramos 2050 (la).
A. 4840 (8).
G?mez-Pompa,
Hage, J. L., ?feE. B. dos Santos
233
Hern?ndez,
(la).
Hern?ndez,
Hern?ndez,
H., & L. Arosemena
Herrera,
1436 (13).
L. R., et al. 1538 (10).
H. M., & R. Torres 335 (8).

J. J., & S. E. Hoyos 450 (5).
R., & E. Matuda 2364 (la).
Hern?ndez,
351 (16);
(9).
E., & P. Segalen 343 (8).

H. 1127 (8); 2447 (8).
H. M. 418 (8).
Hern?ndez,
Gurgel 1648 (16).

Guti?rrez
(9).
17526
Heringer, E. P., et al. 674 (10); 5380 (9); 5388 (9);

5450 (9); 15968 (10).
(la).
2791
(la).
158 (24).
231 (la).
T. W. 2815 (12).
et al. 2645 (la).
Henkel, T. W,
Herb. Bernhardi
1513 (12)
Herb. Sagot 126 (12)
A. Brown 2756 (12).

F., <&
E., et al. 2114 (la); 2125 (la); 2145
(la).
Guedes, M.
P. C.
Henkel,
Gr?ger,A. 637 (12); 707 (12); 807 (12).

Gu?nchez,
E. R. 530
Heligers,
Heller,
4077
(la).
Grayum,
Graziela, Ed. App. 7 (16).
7252
P. J., ?feJ. Eggeling
Greenway,
Grether, R., ?feH. Quero 1579 (8).
Grijalva, A., et al. 322 (la).
Gudi?o,
Heithaus,
VOLUME 64
H. H.
1514 (la);
1777 (la).
1648 (la).
Hage,
Hespenheide,
Heudelot
380
Hahn, W.
Hinton, G. B. 3623 (la); 3805 (la); 4153 (la); 5877

(la); 9978 (la).
J. L., et al. 1335 (13).

Hahn, M. 216 (la); 1231 (la).
841 (10).
F. 1046 (la).
Hammel, B., et al. 16850
Handro,
Hansen,
Hanson,
Harley,
(la).
(la).
<&
M. N?e 7637
B. F,
R. M. 26967
(10).
et al. 10899
Hoehne, F. C. 729 (16); 5895 (14); 28294 (16).

Hoehne,W. 3177 (16); 5718 (14); 13765 (16).
(8).
(9); 18086
(17); 19158
(9);
22571 (16); SPF 35059 (16)

Harmon, W. E., <&J. D. Dwyer
Hart, J. 103 (la).
Hartshorn, G. S. 888 (la).
2658
Hartshorn, G. S., et al. 2918 (9).

Harvard Course in Tropical Botany,
Hassler, E. 7222 (10).
(la); 3983
(la).
10 (4); 197 (4).
Hatschbach,
G., & H. Hass 15795 (la).

?feKasper 41576 (16).
E. Moreira
9506 (16).
G., <&
Hatschbach,
G.,
?feP. Pelanda
Hatschbach,
G.,
?feJ.M.
Hatschbach,
G.,
<&F. J. Zelma
Hatschbach,
G.,
et al. 53973
Hatschbach,
G.,
27898
Silva 48748
49481
Hawkins,
T. 1260 (la);
(16).
(17).
(16).
(21); 58028 (13); 59549
(21); 61856 (16).

1264 (la).
Hoffman, B. 1992 (25).

Hoffman, B., & D. Gopaul 371 (12).
Holm, R. W, & H. H. Iltis 419 (la).
Holst, B., & R. Liesner 2798 (27).
R. A. 5497
Howard,
Howard,
(4).
R. A., & E. S. Howard
Howard,
R. A., & L.
19656
I. Nevling
5191
R. A., et al. 379 (4).

S. E., & J. J. Hern?ndez
286
Hoyos,
(la).
15951
(la).
Howard, R. A., & G. R. Proctor
Howard,
Hatschbach, G. 6513 (16); 7200 (10); 15117 (16);

15930 (10); 18102 (16); 18175 (16); 19955 (10);
20206 (16); 27144 (16); 31876 (21); 34085 (21);
38339 (21); 39477 (22).
Hatschbach,
(la).
162 (3).
V, & J. Gorham
Hodges,
O. 631, (16).
B., et al. 9399
R. M.,
Harley,
A. S. 16796
Hitchcock,
Halle,
(la).
(la);
(la);
13500
17006
(la).
(5); 334 (5); 376
(5); 917 (5).

Huashikat,
Huber,
O.
V. 49 (3); 925
10345 (27).
(3).
Huber, O., & C. Alarcon 6550 (12).

1389 (12).
Huber, O., & L. C?rdenas
Hughes, C. E. 452 (la); 461 (la); 574 (la); 788 (la);

853 (la); 963 (la); 1673 (la).
13927 (9).
Hummel, M. 120(13).
Hut, M. J., & E. Cabrera
I. A. S. 015 (16).
Huingo
Ib??ez-Garc?a,
A. A65
2422
(la).

(la).
2003 ANDIRA 135
Ib??ez-Garc?a,
?feA. Howe
A.,
Idrobo, J.M. 5337 (5).

Idrobo, J.M., ?feR. Jaramillo
Idrobo, J.M.,
A93
(la).
1277 (la).
Irwin, H. S., ?feT. R. Soderstrom
5662
Kuhlmann
(10); 5804
(9);
6140 (10); 6551 (21); 7544 (10); 7436B (10).

(9); 8093
(10); 9824
(10);
Kukle,
Jahn, A.
Laurent,
(12); 2244
(12); 3880
(12); 5589 (12).

Janssen, A.
399
44 (20).
180 (la).
E. 929 (la).
Lanjouw, J., & J. C. Lindeman
Lanna 124 (16).
387 (16).
Lanna & Castellano
Lanna Sobr., J. P. 649 (13).
Langlass?,
129 (la).
L., & J. Pierre 77 (la).
Leite, P. F, & R. M. Klein 518 (21).
Leit?o Filho, H. F., & F. R. Martins
5929
et al. 3211
Filho, H. F,
(16).
185 (4).
Letouzey,R. 6768 (1); 12438 (1); 14743 (la).

C. B. 2128 (la).
G. P. 1050 (17).
G. P., & A. M. de Carvalho
Lewis,
Lewis,
Lewis,
Lewis,
Le?n, B. 3347 (4);4628 (4); 6249 (4).
(la).
1061 (la).
L. A. 669 (la).
?feJ. A. Duke
55-1988
Lindeman,
1315 (5).
3024 (21).
Klug, G. 2763 (la); 2843 (la); 3108 (la); 4309 (la).

Knapp,
S.,
739
(la).
(5).
(17).
(12).
J. C,
Linden,
J. 2160
Liogier,
A. H.
Liogier,
<&
A. Rodriguez
C,
?feR. Schmalzel
3629
11706
(12).
&
J. H.
de Haas
2742
(10); 3109
(10);4547 (10); 5728 (16).
Liogier,
R. 98 (16).
Knab-Vispo,
R., & M. Guariglia
11459
Lindeman, J.C. 5351 (12); 6348 (12); 6890 (12).
J. H., ?feE. Lieras

Kirkbride, M. C. G. 1597 (21).
Kitson, A. 758 (1).
Klein,
Liesner,
S. M.
Lima,A. 07 (16); 105 (17);48-126 (16); 51-944 (14);
Killeen,
Kirkbride,
Liesner,
Liesner,
7413 (la).
T. J., <&
A. P?rez 6682 (12).
Killip, E. P. 40032 (la).
King, R. M. 713 (la); 1961 (la).
M. Goes 286 (16).
?fe
Kirizawa, M.,
4543
(4).
106 (8).
5566 (12);
R., & A. Gonz?lez
R., & B. Holst 20674 (27).
Le?n Cazares,
(la).
J. H.,
B., & Hioram
Le?n,
(le).
T. J., <&
R. Guillen
Kirkbride,
(17);
G. P., & C. E. Hughes

1737 (la).
G. P., & H. C. de Lima 1196 (14).
G. P., et al. 1636 (17).
Lewis,
554
Killeen,
745 (16); 791
796 (17); 812 (14); 1029 (17).
1521 (la).
Ju?rez, O., et al. 233
C. 354
12927
(16);
J.M., & E. Mileste

158 (21).
K. 64091402
(la).
Leonard, E. C. 3596 (la); 7237 (la).
Leoni, L. S., & A. M. Leoni 856 (15).
Lewis, W. H., et al. 1646 (la).

Le?o Costa, A. 1373 (17).
Kersting
(10).
Lems,
Jorgensen, P., 4595 (10).

Jung, S. L. 402 (16).
Kernan,
(16); 7347
(9).
Lewis,
Kenoyer,
1460 (la).
Lao, E. A.
Johnston, J. R. 872 (la).

Joly, C. A. 5992 (10).
R.
(24);
Lemes,
Jim?nez, A. 3088 (la); 3819 (la).

Johnson, H. 1140 (la).
Johnston, I.M. 45 (la); 559 (la);
Johnston, J. A 297 (la).
Kayap,
441
Lely, A. V 832 (le); P192 (le).
1050 (9).
Jardim, J. G. 756 (17).
Jardim, J. G., et al. 71 (16); 90 (18); 585
Jenman 3945 (la).
O. E.
& R. Gieschen
Lane, C,
(9).
Jardim, A.
Jennings,
P. 23 (20).
P., & B. Boom
Kukle,
Leit?o
139 (9).
Janssen, A., <&I. Gemtchujnicor
Janzen, D. H. 10457 (la).
Jaramillo, N. 575 (3).
29 (15).
Kuhlmann, J. G. 163 (16); 172 (16); 230 (10); 360

(13); 478 (14); 528 (15); 1505 (6); 2468 (16);
06529 (14); 13374 (13); 40598 (13); 106410
(13).
Kuhlmann,M. 282 (16); 1477 (16); 1571 (16); 3945
(9); 3950 (16);4694 (16); 10448 (16).
10272 (10); 14420 (22); 14855 (22); 15191 (9);

15873 (16); 16179 (21); 16713 (9); 16873 (21);
17126 (21); 17362 (9); 17962 (21); 18050 (16);
19631 (16); 20191 (16);20266 (16);20781 (16);
20981 (16);22094 (16);22836 (16);23984 (21);
25699 (10); 31521 (22); 32461 (10); 34872 (21);
48442 (26).
Jack, J. G. 4999 (4); 5351 (4); 5390 (4); 5664 (4);
7365 (4).
1239 (la).
Jansen-Jacobs, M. J., et al. 2037
(la).
et al. 24452 (la).
Krukoff,B. A. 695 (la); 5190 (la); 5483 (9).
2269
R. E. Schultes
<&
S., et al. 7954
A.,
Krapovickas,
Irm?oEdesio 6411 (16); 6416 (10).

Irwin,H. S. 10867 (10); 23384 (10); 74363 (10);
22146a (10).
Irwin, H.
et al. 79240
S. D.,
Koch,
(la).
(4).
12338
(la).
A. H., & P. Liogier
A. H., et al. 28353
28663 (la).
(la); 32872 (la).
Little, E. L. 6272 (la); 6341 (la); 13112 (la); 13235

(la); 13745 (la); 16329 (la); 26073 (la).
Loefgren,
A.,
508
(16); 3132

(16); 3727
(16).
Long,
L. E. 229
Lopes, M. A.,
(la).
?feP. M. Andrade
850
7196
(5).
S. 4005
P. J.M.,
Maas,
P. J.M.,
3293
(12).
Mattos
4299
(25).
Mattos
?feCustodio Filho 38 (16).

I. C. C,
<&S. Assump?ao
2265 (21).
M., et al. 407 (la); 637 (21); 1212 (la);
Mac?do,
1402
(la).
W.
61 (9).
G. 324 (16).
Machadhum,
Machado
de Campos, S. 69 (10).
Mac?do,
U.
Macqueen,
104 (29).
D. J. 462
Macqueen,
D.
(la).
J. R., & N. F. Mattos
(10); 8383
8259
(16).
Silva, L. A., & J. L. Hage 585 (17).

Silva, L. A., et al. 340 (14); 1199 (19);
1354
167 (16).
S., & W. J. Clark 14327 (la).
T., et al. 2480 (12).
McDowell,
G. 15014 (la); 15401 (la).
McPherson,
3004 (le).
Mearns
McDaniel,
et al. 49 (3?).
M., & J. Pimentel 23644
Meijer, W,
Machado, O. 77 (13); 452 (9); 452 (10); RB76106

(15).
Maciel,
(la); 6068
6035
(la).
Mavarro de Andrada
M.,
Mac?do,
13 (la).
M. E., & D. Taylor
(16); 1393 (19); 1810 (17); 1877 (19);3027 (17).

Matuda, E. 2109 (la); 3125 (8); 3569 (la); 38597
(la).
Mac?do, A. 2559 (10); 2587 (9); 4352 (21).

Mac?do,
FV.
Mattos,
(la).
?feJ. A. Tawjoeran
?feL.Y. Th. Westra
T. 3311
MacDougall,
(la).
Mattos Silva, L. A. 76 (9); 599 (16); 1393 (16); 2102

(16).
Lutzelburg 1548 (10); 1813 (10).

L?pez,
455
3388
Mattos, J. R. 10593 (16); 11542 (10); 12530 (16);

13563 (16); 13802 (16); 13818 (16); 14079 (16);
14530 (10); 15059 (10); 15118 (10); 15430 (16);
15539 (21).
Lundell, C. L. 4416 (la); 6098 (la); 6986 (la).
Maas,
E. M.
Mathias,
19 (16).
V. 16090 (13).
M. Lebron-Luteyn.
Luteyn, J. L., ?fe
Lutz, B. 1493 (15).
E., & R. Torres
Mart?nez,
Mason,
H.
Luetzelburg,
Mart?nez,
(la).
Mart?nez, E. M., & B. M. de Jes?s 3486 (la).
Martius, C. F. P. von 1157 (la).
1477 (12).
O., & S. M?rquez
Marulanda,
(15).
S., et al. 4480 (12).

L?pez-Palacios,
Lott, E. J. 2544 (2).
Loubry, D. 2426 (24).
Lozano, G., ?feO. Rangel 5245 (la).
Lozano, G., et al. 3963 (5).
Ludenwalt,
VOLUME 64
136
Mejia,
Mejia,
M., & T Zanoni

F., & L. Co?lho
6590
(la).
(la).
3463 (la).
et
al.
1225 (10);
Mendon?a,
E., & G. Hartshorn 2093
Meneces,
F. 121 (8).
Men?ndez,
Mello,
R. C,
(la).
A. Macqueen
J., <&
602
(la).
1610 (9).
(12).
Magallanes, J.A. S. 421 (2); 585 (2); 1008 (2); 1044

(2); 1383 (2); 1669 (2); 1710 (2); 2273 (2); 3556 Mexia, Y. 4932 (16); 5315 (16); 5448 (16).
Mildbraed 9422 (le).
(2);4197 (2);4404 (2).
Maga?a,
M. A. 2404
Maga?a,
M. A.,
Maguire,
B.,
(la).
?feS. Zamudio
<&
D. B. Fanshawe
291
(la).
23273 (25).
E. 070 (21).
Mileste,
C. W. 22 (la).
Miller,
J.S., & C. D. Sherman
Mill,
6369
(la).
Maguire, B., et al. 43878 (25); 56125 (9); 56193 (9); Miller, J. S., & C. M. Taylor 6028 (la).
56241 (9); 56291 (10); 56394 (21); 56403 (9).
Miranda, C. A., & J. A. Ferreira 355 (22).
Mahecha, G. 1084 (la); 5813 (la); (UDBC10203) Miranda, F. 4075 (la); 4150 (8); 5997 (la); 6609 (la);
7120 (la); 7606 (8).
(la); (UDBC 10210) (la).
6415 (la).
1064 (la).
A. Jardim 1291 (9).
Mamani,
F., <&
R. Andreata 90 (16).
Mamede, M. C. H., <&
Mantosani, W. 10/88 (10).
Miranda,
Mantovani,W. 980 (10); 1005 (10); 1036 (10); 1109

(10); 1232 (10).
Molina,
Mahecha,
G.,
Maitland,
T. W.
Marinho,
Martinelli,
Martinelli,
Martins,
<&
G. Jim?nez
L. R. 27 (6); 526
G. 5501 (16).
G., et al. 11102
H. F. 335 (16).
Moacyr
Molina, A. 1897 (la); 6790 (la); 6940 (la); 7225 (la);

8063 (la); 8633 (la); 10930 (la); 21925 (la);
22430 (la).
26025 (la).
A., & A. Molina
21549 (la).
A., & E. Montalvo
Molina,
Monteiro,
(20).
(16).
O. P. 147 (5?).
1724 (12); 2210
R. A.
Montes,
(19); 11365
F. E., et al. 296 (12).

T. 8 (10).
Alvarenga,
Morales,
J. 683
(12).
(la).
Mori, S. A. 6996 (la); 7074 (la); 10626 (16).
T. 14 (21).
P. 177 (16); 190 (16).
Martuscelli,
Mart?nez, C. 57 (la); 1308 (la).
Martins,
G. 1369 (8).
Mart?nez-Calder?n,
Mart?nez, E., ?feL. Rico 6106 (8).
170 (la).
Mart?nez, E., ?feO. T?llez
Mori,
Mori,
7900 (la).
S. A., & R. Dressler
S. A., et al. 11457 (17); 13800 (18).
Morton, C. V. 4686 (la); 5599 (la); 9983 (4); 10121

(4); 10155 (4); 10304 (4).
Murphy,
Narvaez,
H., & G. Pacca 723 (la).

L. H., & J. R. Olmos 69 (la).

2003 ANDIRA 137
et al. INPA/WWF1032
J. R. M.,
Nascimento,
(20);
4500 (20).
O. C. 764 (28).
Nascimento,
G. V. 913 (la).
?feN. Taylor 1618 (la).
Nash, G. V,
167 (16).
Navarro de Andrade
(16);278 (16);279 (16);280 (16); 281 (13);282

(13).
Pennington,
Pennington,
9255
?feJ. Dwyer
Nee, M., et al. 46951
Neill, D. 10214 (5).
Neill,
(la).
(la).
Pennington,
?feE. Gudi?o
D.,
10115
D., et al. 8056 (6).

?feE. Romero 4519
Nelson, C,
(la).
Pennington,
(la).
Pennington,
C, et al. 6166 (la).

E. W. 424 (8); 2638
Pennington,
(la); 3840 (la).

171 (8); 840 (8).
L., ?feA. G?mez-Pompa
Nevling,
E. F. 125 (9).
Nieustedt,
Nelson,
Noblick, L. R. 2921 (10); 3415 (10); 3534 (10).

Noblick,
Noblick,
L. R., ?feI. C. Britto, 4430

L. R., et al. 2299 (14).
A. 449
Nongonierma,
C. 249 (10).
(17).
(la).
Novaes,
Novelo,
A. 243
Novelo,
R. A.,
P. 6851
Nunez,
(8).
?feV. L. Ramos
1770 (8); 1786 (8).
(la).
P., ?feL. Qui?ones

A. J. 99 (la).
Oldeman, R. A. A. B718
Nunez,
6913
(la).
Oakes,
Oliveira,
A. A.,
et al. 420
203
(16); 205
R. T., & F. A. de Carvalho
294
(13); 297
R. T., & I. Johnson 433 (12).

R. T., & G. P. Lewis
188 (17);
189 (17);
190 (17); 191 (14).
Neill,
Nelson,
de Carvalho
(13); 300 (17); 301 (17); 302 (16); 303 (16); 304
(17); 305 (14); 306 (15); 307 (14); 308 (14); 309
(15); 310 (14); 311 (17); 312 (13); 313 (17); 316
(17); 317 (13); 318 (16).
Nee, M. 10785 (la); 31530 (3); 33333 (la).

Nee, M.,
R. T., & A. M.
(13); 207 (16); 208 (13).
Nash,
(17);
211 (16); 212 (13); 213 (16);214 (14);215 (14);

216 (19);217 (19);221 (17);222 (17);223 (17);
224 (17); 227 (13);231 (16); 232 (19);233 (19);
234 (16); 235 (17);236 (16);237 (17);288 (17);
290 (17); 291 (17);292 (17);458 (12);462 (12);
465 (9); 466 (9);467 (21);469 (9);470 (9);471
(9); 474 (9); 478 (21); 479 (21); 480 (9); 481
(21); 485 (21);486 (21); 491 (9);492 (21);493
(21);496 (21);497 (21);498 (21);499 (10); 500
(10); 501 (10); 502 (10); 503 (10); 504 (9); 506
(10); 507 (10);508 (10); 509 (10); 512 (la); 580
(la); 589 (la); 682 (5); 683 (5).
Pennington,
(la).
Pennington,
(23).
Pennington,
Oliveira, E. 330 (12); 1446 (22); 2032 (12).
R. T., & N. Zamora 612 (la).

R. T., et al. 181 (19); 183 (13); 209
T. D.
13558 (la).
T. D., & E. Freir? 13777 (la).
T. D., & J. Sarukh?n 9067 (8); 9093
(8);
9480 (la); 9622 (8).
P. I. 87 (10); 680 (16).

11 (9).
E., ?feS. de Fonseca
Opara, N. 609 (le).
Orlandi, R. P. 91 (22); 189 (17).
Ortega, F. 775 (3).
T. D.,
et al. 14044
(la);
14939
(la).
Oliveira,
Pennington,
Onishi,
Pereira,B. A. S. 67 (9); 76 (9); 768 (9);904 (9); 1134

(9).
Pereira, B. A. S., & D. Alvarenga
(la).
478 (12); 482 (3).
?feG. Mahecha
Pab?n, M.,
Palacios, W. 3204 (la); 4818 (la); 12274 (5).
?feE. Freir? 7444 (6).
Palacios, W.,
Pereira, E., & Egler 3355 (la).

Pereira, E., et al. 4156 (13); 4399
Pereira Neto, M. 86(10).
Pereira, R., ?tal. 517(16).
et al. 7747
Pennington,R. T. 184 (19); 185 (19); 186 (17); 187

(17); 192 (17); 194 (16); 197 (19); 198 (17); 199
(17); 200 (16);201 (16);202 (16); 228 (16); 229
(19);283 (16);284 (16);285 (16); 286 (13); 287
(16);464 (12).
R. T.,
M. Aulestia
?fe
523 (6); 524 (5); 525
(5); 537 (3).

Pennington,
R. T.,
?feH. S. Brito
238
(9);
Pereira,E. 1673 (16);2064 (16);4934 (la); 5788 (16);

7148 (9); 7154 (10); 8835 (21).
(12).
Pastore, U., ?feR. M. Klein 77 (21).
Pedersen, T. M. 8510(10).
Pena, B. S. 608 (3).
Pennington,
(9); 2823
Pereira, B. A. S., et al. 1552 (21).
Ortiz, G., et al. 2035
Palacios, W.,
2234
2900 (la); 3426 (la).
(13); 239
(10);
240 (10); 241 (10); 242 (10); 246 (10?); 247

(10?); 248 (16); 249 (16); 250 (16); 251 (16);
256 (9); 257 (9); 258 (9); 259 (9); 261 (22); 262
(22); 263 (22); 264 (22); 265 (9); 266 (10); 267
(10); 268 (10); 269 (10); 270 (9); 271 (9); 272
(9); 273 (9); 274 (16); 275 (16); 276 (16); 277
(16); 7132
(16).
P?rez, L., et al. 1497 (10).

P?rez, L. A. 182 (2).
P?rez, L. A., & M. Hern?ndez
Perrottet 31 (la).
Pickel, B. 5448 (16).
848 (2).
G. 13387 (16).
Pic?n, G., et al. 1576 (12),
Pickolt,
Pinheiro, R. S. 250 (13); 332 (13); 449 (16); 1302

(16); 1579 (16); 1596 (16); 1700 (16); 1996 (13);
2183 (16); 2251 (16).
Pinto, G. C. P. 56/85 (17); 189/82 (10); 418/81 (17);
429/81 (17).
11/85 (17).
Pinto, G. C. P., & H. P. Bautista
Piper, C.V. 5117 (la).
Pipoly, J. J. 7972 (25); 11265 (la).

J. J., et al. 17002
Pipoly,
(5); 17085
3582 (la); 4222 (la); 4273 (la); 4729 (la); 4794
(10).
Pires, J.M. 3394 (16); 4064 (12); 6295 (10); 16257

(21); 16637 (21); 51939 (12).
Pires,
J.M.,
Pires,
J.M.,
?feG. A. Black
et al. 16726
1630a
(22).
(12).
Reyes-Garc?a,
(9).
R. M., et al. 5214 (la).

291 (8).
F, <&C. Alvarez
Prance, G. T. 59051 (9); 59362 (la).
Prance, G. T., ?feJ. F. Ramos 23563 (20).
Prance, G. T., ?feG. B. Schaller 26266 (21).
G. T., et al. 4160
(12); 6944
(16); 7968
17136
19754
(la);
21472
(la);
(3);
(la);
26576 (la); 30248 (la).

Pr?vost, M. F. 648 (la); 1192 (la); 3046
Purpus, C. A. 10304 (la).
L. 2764
(12).
et al. 2238
B. V,
Rabelo,
(la); 2789
(la);
3199
(26);
3759
(la).
G. A. 221
(la).
R. S., & G. Rodrigues
T. P., et al. 2469
Ramamoorthy,
Ramage,
1108 (16).
Ramalho,
(la).
369 (la).
Ram?rez, N. 3083 (la); 3327 (12).
Ram?rez, J. G., ?feD. C. L?pez 950
Ram?rez, J. G., et al. 4284 (5).
Ramcharan
R.,
?feG. Flores
Ram?rez, V,
?feF. Morales
Ram?rez,
674
Ratter,
J. A.,
?feJ. Ramos
J. A.,
et al.
Ratter,
1043
?feDaniels
3414
Rebo,
B. P. 3671
(la);
(16).
3676
(21); 2323
(la).
Regnell, A. T. 468 (9);469 (10); 1572 (16).

Reitz,
Reitz,
Reitz,
P. R.,
Renteria,
?feKlein
E. 3765
(la).
Filho
3514
(la).
Rizzo, A. 4142 (21);4323 (21);4424 (9).

17062
G.
R.
(la).
Rodrigues,
1915 (la).
E. 221 (12).
Rodrigues,
G., & R. Ramalho
Rodrigues,
R., Se E. Acero
822 (16).
221
(28).
Rodrigues,W. 2917 (26); 7990 (20); 7991 (23); 7992

(23); 7994 (29); 8484 (20); 11179 (20).
Rodrigues,
W,
& A. Aubreville
Rodrigues,
W,
& J. Chagas
Rodrigues,
W,
& D. Co?lho
681 (12).
1534 (23).
7554 (29); 7863
Rodrigues,
W,
& L. Co?lho
8557
Rodrigues,
W,
& A. Loureiro
Rodrigues,
W,
& F. Mello
5953
4975
(29).
(23).
(29).
(la).
(21);
Rosa, N. A. 668 (6); 3645 (12).

J., & J. L. Valles 79 (12).
Rosales,
J., et al. 505 (12).
Rosales,
Rose,
J. N.
Rose,
J. N.,
1639 (la); 1782 (la); 3556 (la).

et al. 3385 (la).
Rossi, L., et al. 656 (16); 789 (16); 808 (16).
J. N. 186 (la); 472 (8).
Rovirosa,
Rudd, V E. 902
(8).
Rusby,H. H. 826 (la); 1161 (la); 1270 (la); 2353 (la).
P. R. 2270
(16).
P. R., & A. Burkart
Rizzini,
3246
18 (la).
S., & Mattos
Romero, R. 584 (la); 3142 (la); 5218 (la); 5444 (5);

6136 (la).
3595 (9); 3599 (10); 3791 (16);4123 (la); 6891

(9); 7151 (21); 6729V (9); 6869V (21); 7021V
(21); 7022V (21); 7044V (21); 7045V (21);
7691V (21).
Read,
G.
4273 (la).
W, & B. Wilson
et al. 8477 (29); 8502 (29).
Rodrigues, W,
Rojas, S. 230 (la).
Rojas, S., & R. Zuniga 232 (la).
343
(la).
(21); 2318
Rivas,
Rodrigues,
(lb).
334
Ratter,
Ratter,
(16).
Rodrigues,W, & Osmarino 6888 (29); 7929 (23);

8472 (29).
(5); 1161 (5).
(la).
J., ?feR. Sousa 156 (21).
J. A. 531 (la); 4123 (21); 5192 (la).
J. A., ?feV. P. de Lima 6716 (22).
Ramos,
J. J. 397 (la).
M.Y.
Rimachi,
Robles,
(16).
113 (la); 230 (la).
Quesada,F.
Questel, A. 2173 (la).
(9).
(la).
Riedel, L. 40 (16); 118 (16); 123 (la); 190 (16); 224

(16); 402 (la); 469 (21); 470 (9); 503 (21); 555
(10); 645 (10); 685 (10); 708 (la); 827 (21);
2207 (16); 2935 (10).
Roberty,
(24).
L. P., et al. 2957
Qui?ones,
A. E. 451
et al. 255
Ridley,
8719 (9); 9049 (23); 9074 (20); 11182 (12);

11635 (23); 12388 (6); 15011 (23); 18965 (21);
25053 (10).
Queiroz,
A.V. 20 (9).
A. J. 62 (17).
Richards, P. 6482 (9); 6760
Rezende,
Ricksecker,
Ponce,
G. R.
(la);
Ribeiro,
Ricksecker,
Polhill,
Proctor,
(la).
(3).
A. 1724 (la); 2366 (la).
A., & M. Sousa 2010 (la); 2074
2312 (la)
Pohl 1262 (9)
Prance,
D. 954
J. 275
Reyes-Garc?a,
Pittier,H. 3372 (la); 3870 (la); 3910 (la); 3999 (la);

4764 (la); 4769 (la); 8273 (la); 12079 (la).
T. 8298
(la).
Restrepo,
Revilla,
Pires, O. 29 (23).
Pitman, N., et al. 631(11).
Plowman,
E., & D. C?rdenas 4315 (la); 4373 (la).

E., et al. 1887 (la); 2322 (la); 3579 (7);
Renteria,
(5).
Renteria,
Pirani,J.R.818(16);1178(9).
Pirani, J. R., et al. 2189
VOLUME 64
138
5617
3907
(la); 4634
(16).
(7).
(16).
Saito Kunigoshi,
Salgado, O. 081
Sampaio,
Y. 4013
(16).
(9); 170 (9).
T. 6 (15).

2003 ANDIRA 139
Sanaiotti,T.M. 222 (12); 287 (9); 449 (9); 460 (9);

461 (9).
D. 1958 (5).
S?nchez, M., & G. Mahecha
Santos, M. M. 85 (22); 210
Santos, R. R. 14965 (la).
S?nchez,
9 (la); 9a (la).
Sastre, C. 7731 (la).

Saunders, J. 216 (la); 543
Schaller, G. 160 (21).
Schinini,
A., & R. Vanni
(5).
(la); 323 (la).
A. 2148 (2); 2950 (2).
Solis-Magallanes,
Sothers, C. A., ?feE. Pereira 397 (20).
(9).
Santos, R. S., & A. Castellanos
24071
(10).
(la).
25958
Schipp, W. A., 484 (la).

Schmeda, G. 1082 (10).
Schultes, R. E., & B. P. Reko
(la).
852
(8).
Schulz, J. P. 7166 (12); 7273 (12); 7321 (12); 7513

(12); 7625 (24); 7953 (24).
Schunke, J. 1958 (la); 3859 (3); 5725 (3); 8286 (3).
Schunke,
J.M.
181 (12).
Schwacke,C. A. W. 4239 (13);4527 (10); 5033 (16);

10753 (10).
Schweinfurth,
G.
1875 (le).
F,
Solano,
E. 298
Sothers,
C. A.,
et al. 291
et al. 136 (20).

(la); 7612
Soto Nunez,
J. C. 6237
Soto Nunez,
J. C,
?feB. Boom
2097
Soto Nunez,
J. C,
?feL. Cortes
2297
Soto Nunez,
J.C,
?feE. Mart?nez
Soto Nunez,
Soto Nunez,
Soto Nunez,
J.C,
J.C,
?feF. Sol?rzano
J. C,
et al. 40
?feS. Zarate
(la).
(la).
(la).
4043 (la).
12631 (la).
1292 (la); 1321 (la).

93 (la); 609 (la);
(la);
6238 (la); 8016 (la); 8187 (2); 9581 (la).

Sousa,M. 3267 (8); 3667 (8); 3697 (la); 9926 (la);
11964 (8); 12162 (la).
Sousa, M.,
Sousa, M.,
Sousa, M.,
S. E. F. 8533 (3); 9064 (la); 9283 (la).

Segadas-Vianna,
S. C. C-41 (la).
Soares, A. 104 (20).
Soares, E. 5 (29).
Soejarto, D. D., et al. 3954
Snedaker,
?feA. S. Magallanes
7382 (la).
?feS. Zarate 9945 (la).
et al. 4088 (la); 4371 (la); 5212
(la); 5503
(la); 5535 (la); 7369 (la); 7656 (la); 7960 (8);

10152 (la); 10207 (la); 10262 (8); 10777 (la);
11926 (la): 12083 (la); 12204 (la); 13207 (la);
13369 (la); 13407 (8).
(13).
Seler&Selerl841(la).
Seligson, D. 160 (la).
Sellow 39 (14).
Souza, A.
Sentenio, P. 3356 (la).

Sess? et al. 2012 (4).
Shafer, J. A. 2593 (la); 3196
Spruce,R. 914 (12);4115 (la); 17235 (12).
Shannon, W. C. 5025
Stahel, G. 57 (12);
(la); 4349
(16).
Sillman, M. S. 047 (21).

Silva, I. A. 260 (17).
Silva, M., & R. Sousa 2284
(12).
(28).
Silva, M. F, et al. 91 (20); 373 (29); 924
Silva, M. G. 3337 (12); 5933 (la).
Silva, M. G., & R. Bahia 3103 (12).
(28).
(12); 4415
Steinbach,
(21).
Silva,N. T. 4808 (21); 58315 (16); 58414 (16).

Simpson,
D. R. 675
Sintenis,
P. 3356
Stehl?,H. 166 (la); 483 (la); 1983 (la); 5381 (la);

6033 (la).
Stehl?, M.,
Silva, M. G., & J.Maria 3337 (9).

Silva, M. G., & A. Pinheiro 4327 (12).
Silva, M. G., & R. S. Pinheiro 4415 (21).
3627
(la);
(la);
(la);
(la);
(la);
(la);
Stearn, W. T. 1004 (la).
F. 979
Silva, M. G., & C. Rosario
109a (la).
Standley,P. C. 11189 (la); 14219 (la); 20383

20861 (la); 20991 (la); 21227 (la); 21796
21943 (la); 22245 (la); 23206 (la); 23409
28041 (la); 28954 (la); 29110 (la); 29342
29834 (la); 31106 (la); 31890 (la); 39992
40135 (la); 53570 (la); 60533 (la); 74130
79182 (la); 89347 (la).
(4).
(la).
Shapiro, G. 335 (la).

Sharp, A. J. 46110 (la).
Shepherd, G. J., et al. 5878
Silva, M.
1846 (14).
?feH. Stehl?
5181
(la).
J. 6710
(la).
Stergios, B., et al. 3461 (12); 13195 (12).
Stern, W. L. 10 (la).
Stern, W. L., et al. 1699 (la).
17150 (la).
Stevens, W. D., ?feO. M. Monteil
Stevenson
5622 (la).
Stevenson, D. W. 958 (20).
J. A. 2194 (la).
Stevenson,
Steward, W. C, et al. 135 (12).
J. A. 37865 (la); 39608
Steyermark,
(12).
(la).
P. 520 (10).
Slane, V 803 (la).
Smith, A. C. 3447 (12).
Sladen,
(la); 44779
(la);
46202 (la); 47462 (la); 62930 (la); 87679 (la).
14 (16).
Smith, D. N. 3923 (la).
Smith, D. N., et al. 1172 (la).
Smith, H. H. 18 (la).
Smith, L. B. 6554 (14).
Smith, C.
Smith, L. B., & R. Klein 7288 (16).

Smith, L. B., et al. 14853 (16).
Steyermark,
Steyermark,
Steyermark,
J. A.,
J. A.,
J. A.,
<&
G. Davidse
116474
<&
A. Gonzales
et al.
(la).
113652
119886
(5);
125856 (28); 26212 (12).

Stimson, W. R. 2013 (la).
J. J., et al. 4463
Strudwick,
Sucre, D. 3637
(16); 8443
(12).
(16); 8656

(17).
(la).
120019
(5);
140
et al. 6583
Sucre, D.,
10228
(16);
811 (16).
van derWerff, H., & R. Ortiz 5553 (5); 5556 (5); 5576
(16).
Sugiyama,M.
Tarto, L. 2 (16).
Taylor, C. M. 6325
(5).
van Donselaar,
(la).
Taylor, C. M., & H. J. Scott 7863 (la).
Taylor, E. L., et al. E1248
(9); E1259
(22).
Taylor, N. 238 (4).
L. O. A.
Teixeira,
L. O. A.,
et al. 1182 (23).
O., & E. Mart?nez
T?llez,
O., & J.Miller
905
10478
(8); 948
(8); 8079
(la).
(lb).
P. 19498
Tenorio,
(8).
P., & R. Torres 212 (la).
P., et al. 395 (la); 3108
Tenorio,
Tenorio,
16869
(la);
(lb);
19415 (la).
G. 3437
(la); 3836 (3).

R A., & J. T. Wildschut
Tessmann,
Teunissen,
D. W.
Thomas,
11830
(24).
K. 594
(la); 607
A. S. 54 (le).
Thornewill,
S. S. 756-393
Tillet,
0125
(3); 2910
(6).
(3).
V?squez,
R.,
V?squez,
R.,
?feF. Ayala 13397 (3).

?feC. Gr?ndez 2661 (3).
V?squez,
R.,
?fe
N.
1142 (3); 2708
Jaramillo
(6); 2718
V?squez,
R., et al. 5919
Velazco,
J. 1285 (12).
12814
(la);
13802
(la);
(6).
H. P. 98 (16).
Veloso,
?feE. L?pez
Ventura,
E.,
Ventura,
F. 12803
1140 (la); 3365
(la).
(8).
C. 287 (16).
Vieira, M. G., et al. 622 (10).
Voeks, R. 102 (18)
Viana-Freise,
R. J. 280
Wagner,
1644 (la);
(la);
B. M. T., et al. 228
1804 (la).
1481 (21); 2733
(22);
M. G. L., et al. 105 (16).

D. C,
?feF. Encarnaci?n
Wasshausen,
(21).
Wanderley,
Thomas,W.W, et al. 3962 (21);4312 (21); 8533 (14);

8969 (19); 9469 (17); 9726 (19); 9881 (17);
10055 (16); 10950 (19); 11196 (13).
Thomsen,
R. 2680
Walter,
313
(la).
J., et al. 1546 (la).
Thomas,
V?squez,
(la); 4728 (12); 8745 (6).
T?llez, O. 4733 (8); 5477 (la); 5521 (la); 9076 (lb).

T?llez,
J. 1743 (24).
?feJ.Morales
E.,
Varon,
B. C. 72 (16).
36 (12).
Teixeira,
Teixeira,
VOLUME 64
H. von
Wedel,
Weddell,
(la).
(la).
Tipaz, G., et al. 1225 (6); 1345 (6); 2215

A. 124 (la).
(la).
?feK. I.Miller
G. L.,
Webster,
1905 (la); 2017
M. H. 2898
?feA. Villalobos
T.,
Wendt,
T., et al. 3416

E., & Arnold
?fe
W.
1057 (la).
(la).
(la).
(la); 3420
Wendt,
West,
8718
(9).
4463
(la).
(la).
L. 877
(la).
S. Alverson
459
(la); 470
Toepffer,
White,
Tonduz,A. 6511 (la); 6874 (la); 9771 (la).
Wilhams, L. O. 2421 (la); 4727 (la); 6457 (la); 8483

(8); 9451 (8); 9484 (8); 9612 (8).
Torres, J. 160 (12).

Torres, R., & R. Cedillo
2783 (la).
Torres, R., & C. Mart?nez 6005 (8).
Torres, R., & M. A. Mart?nez 6606 (la).
Torres, R., & P. Tenorio 210 (la); 220 (la).
Torres, R., et al. 1279 (la); 2618
(la); 5361
(8); 5362
(la); 9729 (la).

J., & J.B. Alcorn
Treacy,
477
(la).
Trigos,R. C. 556 (la); 737 (la); 1070 (la); 1495 (la).

R. C,
Trigos,
& R. Torres
H. von 7834
Tuerckheim,
Tuez, & Cochran 29269

22 (16).
Tupinamba
Williams,
L. O.,
<&
V. Assis
Williams,
L. O.,
?feA. Molina
Williams,
L. O.,
?feT. P. Williams
Williams,
R. S. 419
1483 (la).
(la).
(la); 420
L. M., & R. F. Griggs

I. 6178 (la).
F. 7269
Woytkowski,
J. J., ?feL. S. Adderley

J. J., ?feJ. V. Monachino
T. G. 18268 (la).
H. 50 (12).
M. Castrillo
Zamora, N., ?fe
711 (la).
?feR. Garcia
Zanoni,
T.,
Zanoni,
T.,
Zanoni,
?fe
M. Mejia
T., et al. 16093
Usteri,
P. A.
Zardini,
E.,
P. F. J. 125-5/B
Valerio, M. 522 (la).

Valeur, E. J. 665 (la).
(la).
(6).
(la).
43368
(28).
39345
(la).
Zabelo,
L. 6650
R., et al. 67876
(la); 35016
Wright, C. 1160 (4); 1187 (4); 1188 (4); 2356 (4);

2358 (4).
Uribe,
Valencia,
(la).
(la).
Yuncker,
Ule, E. 7627 (12);LV11(15).
Valcarcel,
24614
Yano, T. 05 (10).
108 (1).
Tyson, E., & M. Kuns 1003 (la).
Tyson, E., et al. 4849 (la).
(la).
104 (16).
(10).
(la).
E. S. 25 (8).
R. H. 189 (la).
A. L. 734 (10).
Woolston,
Wurdack,
Turner, L.
Urban,
(10): 7453
14628
Wing,
Wurdack,
(16).
Underwood,
7430
(la).
Woodworm,
1131 (la).
Triana,J.
S.,
17748
(la);
(la); 21028
Zarucchi,
Zarucchi,
J. L., & J. Betancur
Zarucchi,
J. L.,
376
(la).
(la).
?feU. Velazquez
J. L. 2810 (9).
Zentner
2031
26927
25531
?feD. C?rdenas
(9).

6444
4215
17899
(la).
(la); 28788
(10); 25676
(5).
(7).
(la).
(10).
2003
ANDIRA 141
INDEXTO SCIENTIFICNAMES
are in roman
names
Accepted
type; the main
(Baill.) Polhill
118
frondosa Mart, ex Benth. 80

var. longifoliolata
N. F. Mattos
gabonica Baill. 118
Agouti
paca 14
Amerimnum
affine Spreng.
Juss. 3
44
Lam.
117
Benth.
sect. Euandira
sect. Lumbricidia
grandistipula
(Veil.) Benth. 3, 22, 31, 36

N. F. Mattos
3, 22, 31, 36
3, 31, 36
(Veil.) N. F. Mattos
subsect. Glabratae
N. F. Mattos
acuminata
Benth.
amazonum
Mart,
anthelmia
(Veil.) Benth.
3, 31, 36
117
4, 5, 6, 11, 12, 13, 15, 16,
(Veil.) J. F. Macbr.
(Veil.) Toledo 76
38, 42, 43, 69-72,76,

var. cordata N. F. Mattos
inermis
N. F. Mattos
Benth.
carvalhoi
82
85
118
4, 9, 10, 12,
49, 50,
Perr.
?feA. Rich.)
Polhill
4, 21,
33,
43, 44,45,
49-50,129
var. riedelii Benth. 44
jaliscensis
R. T. Penn.
5, 9, 10, 12, 21, 22, 23, 24,
25, 27, 28, 32, 33, 37, 38, 39, 42, 49, 50-52,
53, 129
R. T. Penn.
5, 8, 9, 10, 21, 23, 29,
laurifolia
ex R. T. Penn. & H. C. Lima 4, 5,
Pulle 4, 5, 13, 16, 21, 27, 29, 35, 37, 40,
41, 56, 59, 60,10?-108,129

cubensis Benth. 8, 16, 21, 23, 33, 34, 37, 40, 42,
129
53, 55-56,
Benth.
4, 5, 7, 12, 13, 17, 18, 21, 22,
23, 24, 25, 27, 28, 32, 33, 34, 35, 37, 39, 41,
2, 4, 5, 7, 8, 9, 10, 11, 12, 13,
14, 15, 16, 17, 18, 20, 21, 24, 25, 26, 27, 28,
30, 32, 33, 37, 39, 42, 84-S9,126,
legalis
Urban
42
N. F. Mattos
129
Benth.
66, 68
101
69, 71, 72
(Veil.) Toledo
4, 5, 6, 12,13,
14, 18, 21,
24, 25, 26, 27, 28, 29, 30, 32, 33, 36, 37, 38,
41, 78, 80-82,
84, 88, 127, 128, 129
var. bahiensis
76
(Benth.) N. F. Mattos
R. T. Penn. 14, 21, 29, 33, 37, 39, 41,
72-73, 75, 76, 129
Ducke 4, 5, 8, 10, 12, 13, 21, 24, 25,
macrothyrsa
macrocarpa
26, 28, 29, 30, 32, 37, 40, 41, 56, 59, 60-62,
64, 128, 129
marauensis
100-102,103,105,125,126,129
Kunth 44
Benth.
jamaicensis
kuhlmannii
lanei N. F. Mattos
9, 10, 18, 20, 21, 24, 25, 26, 28, 30, 32, 33,
129
34, 35, 37, 40, 41, 100,102-105,125,
fraxinifolia
51, 53, 56, 62,
(Guillem.,
subsp. grandiflora
Gillett ex Polhill 44,48
(de Wild.)
subsp. rooseveltii
37, 38, 40, 41, 61, 62-64,129

chiricana Pittier 44, 48
cinerascens Mart.
118
excelsa
12,
4*49,129
125, 129
cujabensis
1, 2, 3, 4, 5, 7, 8,10,
R. T. Penn. 21, 33,43,44,45,
subsp. glabricalyx
27, 28, 30, 32, 33, 34, 37, 40, 41, 92, 94-98,
cori?cea
127, 129
103,126,
62,129
76
R. T. Perm. & H. C. Lima
cordata Arroyo
92,
69
inermis 4, 21, 29, 33, 43, 44-48,
subsp.
13, 14, 15, 16, 17, 18, 19, 20, 21, 24, 25, 26,
chigorodensis
DC.
(W.Wright)
76
117
bracteosa
4, 5, 8, 12, 14, 16, 17, 18,
33, 34, 36, 37, 40, 42-50,

124, 127, 129
118
bahiensis
44
13, 14, 15, 16, 17, 24, 25, 26, 27, 28, 30, 32,
var. gracilis N. F. Mattos

85
var. ormosioides
(Benth.) Benth.
Aguiar
Benth.
?feA. Rich.
5, 15, 21, 24, 25, 26, 28,
19, 21, 23, 24, 25, 26, 28, 30, 32, 33, 35, 37,
ex Benth.
Benth.
Perr.
Amshoff
118
horsfieldii Lesch.
humilis Mart, ex Benth.
44
18, 20, 21, 22, 24, 25, 26, 27, 28, 29, 30, 32,
araroba
68, 72, 76,
29, 30, 32, 34, 37, 38, 41,106,108-109,129

handroana N. F. Mattos
85
33,34,37,38,41,76-30,82,84,88,125,129
var. acuminata Benth. 76
aubletii
12, 15, 21, 23, 24, 25, 26, 28,
Guillem.,
grandiflora
3, 31, 36
sect. Paucifoliolatae
subsect. Lumbricidia
anthelmintica
76
126, 129
sect. Aristobulia
anthelmia
84
30, 32, 33, 34, 37, 39, 41, 64-66,
36-37
anthelmia
Standl.
galeottiana
Andira
Andira
85
var. latifoliolata N. F. Mattos

85
var. rosea (Mart, ex Benth.) Benth.
Aganope
gabonica
are in italics.
Synonyms
var. lanceata N. F. Mattos
15
Aglaia
is in boldface.
entry for each
N. F Mattos
5, 21, 23, 24, 25, 27, 28,

129
32, 37, 40, 42, 92, 93-94,128,

micans Taub, ex Glaz. 64
micrantha
Ducke
5,15,
16, 21, 25, 27, 29, 35, 37,

108, 129
40, 41, 56, 59, 60,105-106,

142
microcarpa
Griseb.
55
Dalbergieae
multistipula
Ducke
5, 8, 21, 23, 24, 25, 27, 28, 29,
Dasyprocta
32, 37, 38, 41, 43, 51,5^-54,129

var. peruana N. F. Mattos 60
nitida Mart, ex Benth. 4, 5, 9,10,
11, 12,13,
Benth.
subsp. aurantica
18,
12, 13, 15, 20,
4, 5, 8, 9,11,
1, 2, 33
14
Diptychandra
aurantica Tul. 36
21, 24, 25, 26, 27, 28, 30, 32, 33, 37, 38, 40,
42, 89-93, 94, 97, 128, 129
ormosioides
Krug & Urb.

Euchresta
80, 82-S4, 88, 129

Benth. 66
Geoffroea
Ducke
4, 5, 21, 24, 25, 26, 28, 30, 32,
34, 37, 39, 41, 98-100,129

parvifolia Mart, ex Benth. 84
Benth. 69, 71
N. F. Mattos
pernambucensis
pisonis Mart, ex Benth. 84
pauciflora
ex R. T. Penn.
Arroyo
114, 115, 129

37, 40, 42,109-112,
racemosa Lam. ex J. St. Hil. 2, 3, 118
retusa (Poir.) DC. 74
riparia Kunth 44
rosea Mart, ex Benth.
sapindoides
spinulosa
surinamensis
(Bondt)
73
25, 27, 28, 29, 32, 37, 39, 40, 41, 56-60,
72,
?feN. Cuello
5,
109, 111,112-114,
Ducke
3, 5, 10, 21, 22, 25, 27, 29, 31,

35, 37, 38, 40, 111, 114-115,
117, 129
unifoliolata Ducke 3, 5, 6, 7, 13, 21, 22, 24, 25,
111,
(Mart.) Benth.
4, 5, 10, 12, 13, 17, 18,
19, 21, 24, 25, 26, 28, 30, 32, 33, 37, 39, 42,
76, 88, 102, 103, 128, 129
118
65, 66-69,72,
villosa Kleinhoonte
wachenheimii
Benoist
Harms
Caesalpinoideae
12
Cleogonus
118
15, 97
Coursetia
caribaea
Seem.
(Jacq.) Lavin
var. caribaea 34
107
49
var. novogaliciensis
var. viridiflora 49
Lonchocarpus
staudtii Harms
44
Lumbricidia
Veil.
36
Veil.
76
legalis Veil.
Meliaceae
15
Lavin & Sousa 49
36, 80
Millettia
rooseveltii
de Wild.
Mimosoideae
115-117,129
zehntneri
viridiflora
anthelmia
129
verm?fuga
118
15, 36, 49, 97
Leguminosae
Lennea
106, 108, 129
26, 28, 30, 31, 32, 34, 36, 37, 38, 40,
Ducke
J. R. Forst. & G. Forst.
2, 4, 5,
var. ovatifoliolata
N. F. Mattos
74
taurotesticulata R. T. Penn. 9, 10, 16, 21, 23, 24,
trifoliolata
15, 97
Kielmeyera
rubriflora71
76
21, 27, 29, 37, 40, 42,
44
21, 24, 25, 26, 28, 30, 32
velutinum
Inocarpus
R. T. Penn., Aymard
Ruiz & Pav. ex G. Don
nitidum
ex Amshoff
66, 69
Benth. 1, 2, 20, 23, 33, 118

Hymenolobium
flavum 21, 24, 25, 26, 28, 30, 32
84
Splitg.
74
71
Hymenaea
courbaril L.
6, 7, 13, 15, 16, 18, 21, 24, 25, 26, 28, 30,
32, 34, 37, 39, 42, 72, 73-76,
128, 129
tervequinata
Glycyrrhiza
undulata
Guttiferae
(DC.) Benth. 44, 48

(Mart.) Benth. 66
stipulacea Benth. 76
var. bahiensis Benth.
118
acutifolia Stokes 42
inermis W. Wright 42, 44
Stokes 72
obtusifolia
verm?fuga Mart.
5, 21, 27, 29, 35,
var. oblonga Benth. 74

var. surinamensis
(Bondt) DC.
(Lesen.) Benn.
Jacq. 2, 4
spinulosa Mart. 66, 69

surinamensis
Bondt 73
var. emarginata N. F. Mattos

85
var. puberula N. F. Mattos
85
praecox
Carvalho
ex Taub. 4
Rich.
pubescens
retusa Poir. 74
85
(Tul.) Lima,
Dussia
horsfieldii
paniculata
36
subsp. epunctata
36
21, 24, 25, 26, 27, 28, 30, 32, 33, 37, 39, 42,
parviflora
VOLUME 64
Myoprocta
14
Ostryoderris
brownii Hoyle
49
1, 36, 49,
Papilioniodeae
Parkia
biglobosa
multijuga
Pisum
49
15
118
50
Benth.
15
sativum L. 29
Poeppigia
procera C. Presl
69

& Costa
2003 ANDIRA 143
Jacq. 4
sapindoides DC. 44
anthelmia
(Veil.) Kuntze 76
(Benth.) Kuntze 55
100
(Benth.) Kuntze
cujabensis
(Benth.) Kuntze 84
fraxinifolia
Pterocarpus
cubensis
Pterodon
emarginatus
Robineae
49
35, 36
Vogel
frondosa
humilis
Skolemora
pernambucensis
33
Arruda
118
laurifolia
Trachypogon
guianensis
80
(Benth.) Kuntze 69
(Veil.) Kuntze 80
(Benth.) Kuntze 66
paniculata
Sophoreae
plumosus
Vatairea
inermis
(Mart, ex Benth.) Kuntze

(Benth.) Kuntze 69
(W.Wright) Lyons 42
legalis
112
Aubl.
117, 118
Vataireopsis
araroba (Aguair) Ducke
Vigna 124
Aubl. 2
Vouacapoua
americana Aubl. 2
118
(Mart, ex Benth.) Kuntze

(Poir.) Kuntze 74
(DC.) Kuntze 44
sapindoides
pisonis
retusa
spinulosa
(Mart.) Lyons 66
surinamensis
(Bondt) Kuntze
verm?fuga
(Mart.) Kuntze

66
73
84

Monograph of Andira (Legimnosae-Papilionoideae)

Uploaded by

Document Information

Original Description:

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Monograph of Andira (Legimnosae-Papilionoideae)

Uploaded by

Copyright:

Available Formats

Monograph of Andira (Leguminosae-Papilionoideae)

Author(s): R. Toby Pennington

This content downloaded from 158.39.37.3 on Wed, 3 Sep 2014 14:45:31 PM

of A. humilis with plastomes

characters and have no morphological

informative characters and

ation of species of Andira

are fully described and their ranges are mapped,

in eastern Brazil. All

and one new

share a root nodule morphology

This content downloaded from 158.39.37.3 on Wed, 3 Sep 2014 14:45:31 PM

is based upon examination of ca. 3,000 herbarium specimens

and field studies conducted

grow around the fringes of Amazonia,

frameworks for Andira based upon both chloroplast DNA (cpDNA) re

(1783) description of Andira racemosa provided no generic diagno

This content downloaded from 158.39.37.3 on Wed, 3 Sep 2014 14:45:31 PM

1839, 1860, 1862) features prominently in the taxonomic history of

sections. Section Paucifoli

This content downloaded from 158.39.37.3 on Wed, 3 Sep 2014 14:45:31 PM

The fibers are non-septate

and usually very thick-walled.

This content downloaded from 158.39.37.3 on Wed, 3 Sep 2014 14:45:31 PM

Stipules. Four species (A. multistipula, A. grandistipula, A. anthelmia, and A. legalis)

trifoliolata is the only species with uniformly trifoliolate leaves; A. tervequinata

pattern varies continuously

from entirely eucamptodromous

This content downloaded from 158.39.37.3 on Wed, 3 Sep 2014 14:45:31 PM

FIG. 1. Stipules and leaves

This content downloaded from 158.39.37.3 on Wed, 3 Sep 2014 14:45:31 PM

This content downloaded from 158.39.37.3 on Wed, 3 Sep 2014 14:45:31 PM

Inflorescences are terminal and axillary panicles, which vary in size,

ducous. There is little variation in theirmorphology

throughout Andira. The bracteoles

length varies continuously from 5 to 23 mm (Fig. 3).Within species,

geographical correlation, which is the basis for the de

wing petal sculpturing (Fig. 4A; character 9 in cladistic analysis) is present in

This content downloaded from 158.39.37.3 on Wed, 3 Sep 2014 14:45:31 PM

This content downloaded from 158.39.37.3 on Wed, 3 Sep 2014 14:45:31 PM

in Andira. Species with wing petal sculpturing: A, wing petal of A. ver

Pollen. The pollen morphology

This content downloaded from 158.39.37.3 on Wed, 3 Sep 2014 14:45:31 PM

inifolia contrasts with the pale green-white mesocarp of

A. ormosioides, and confirms the

and structure, and preservation of

species discovered by cpDNA data (Pennington 1995). Pennington and

This content downloaded from 158.39.37.3 on Wed, 3 Sep 2014 14:45:31 PM

(1914), is characteristic of legume fruits and consists of thin-walled, juicy

state of the endocarp dominated

formed by parenchyma and/or col

in this species. Field obser

These data suggest

that greater seed protection may be provided by these harder, thicker

Seeds of species of Andira can be considered "overgrown" (sensu Comer,

This content downloaded from 158.39.37.3 on Wed, 3 Sep 2014 14:45:31 PM

(de Lima 1991; pers. obs. for A. carvalhoi,

studies of this and other species are

have superficially similar morphology

valhoi, and A. verm?fuga, where

mosioides, have flowers of a correspondingly

that they, too, have distinct pollina

This content downloaded from 158.39.37.3 on Wed, 3 Sep 2014 14:45:31 PM