Pennington MonographAndiraLeguminosaePapilionoideae 2003

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

Source: Systematic Botany Monographs , Mar. 31, 2003, Vol. 64, Monograph of Andira
(Leguminosae-Papilionoideae) (Mar. 31, 2003), pp. 1-143
Published by: American Society of Plant Taxonomists
Stable URL: https://www.jstor.org/stable/25027903
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MONOGRAPH OF ANDIRA
(LEGUMINOSAE-PAPILIONOIDEAE)
R. Toby Pennington
Royal Botanic Garden Edinburgh
20a Inverleith Row
Edinburgh EH3 5LR, United Kingdom
Abstract. The principally Neotropical genus Andira Lam. (Leguminosae) comprises 29 species; one
species, A. inermis, includes three subspecies, two of which occur in Africa. Chloroplast DNA restriction site
analysis allows phylogenetic placement of 21 species o? Andira, including five species that were not previously
studied. These data are also used to investigate intraspecific chloroplast DNA polymorphism and to identify ac
cessions of A. humilis with plastomes derived from probable introgressive hybridization. A cladistic analysis of
12 morphological characters provides little phylogenetic resolution in comparison with a simultaneous analysis
of cpDNA restriction site data and morphology. Although some of the clades discovered by this analysis are
"cryptic," i.e., they are supported only by molecular characters and have no morphological synapomorphies, the
results provide a reasonable estimate of the phylogeny of Andira and are used as the basis of discussions of char
acter evolution, infrageneric classification, and biogeography. Previous infrageneric classifications of Andira
were based on the nature of the indumentum of the gynoecium, but glabrous ovaries have evolved in parallel at
least three times. Detailed analysis of fruit wall anatomy provides phylogenetically informative characters and
insights into dispersal biology. There have been at least five instances of independent evolution of large, puta
tively rodent-dispersed fruits and three of large, persistent stipules. The frequent changes in stipule morphology
are perhaps due to mutation in a single gene. Evidence points to a relatively recent (possibly Pleistocene) radi
ation of species of Andira in eastern Brazil. All species are fully described and their ranges are mapped, and 15
are illustrated. Five new species and one new subspecies are proposed: A. chigorodensis, A. jaliscensis, A.
macrocarpa, A. praecox, A. taurotesticulata, and A. inermis subsp. glabricalyx.
INTRODUCTION
Andira 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
closely related to Hymenolobium Benth. (Bentham 1860; Polhill 1981), a genus of 10-12
species confined to South America. They are similar florally and vegetatively, and they
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
type; Hymenolobium has samaras and Andira drupes. The two genera also differ in their
mode of germination: Andira is cryptohypogeal, whereas Hymenolobium is phanerohy
pogeal. Thus, Andira is considered a monophyletic group with synapomorphies of a dru
paceous fruit and cryptohypogeal germination (Pennington 1995, 1996).
The close relationship of Andira and Hymenolobium is confirmed by recent chloro
plast trnL intron nucleotide sequence data (Pennington et al. 2001). The two genera form
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

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2 SYSTEMATIC BOTANY MONOGRAPHS VOLUME 64
molecular systematic studies (Doyle et al. 1997; Lavin et al. 2001) indicate that Dal
bergieae is not monophyletic. This confirms earlier suggestions, based upon consideration
of fruit, seed, and seedling morphology, that Dalbergieae are not a natural group (de Lima
1991).
This taxonomic revision is based upon examination of ca. 3,000 herbarium specimens
and field studies conducted in five Neotropical countries from 1991 to 1997. Species de
limitation in Andira has been regarded as problematic, and the only recent revision (Mat
tos 1979), covering the Brazilian species, is inadequate for species identification, particu
larly for taxa from the Atlantic coastal rain forest and coastal restinga forest of
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 grow around the fringes of Amazonia, which is an
area of diversification for Andira.
Phylogenetic frameworks for Andira based upon both chloroplast DNA (cpDNA) re
striction site data and morphology have been presented by Pennington (1995, 1996). The
phylogenetic analyses presented in this monograph are drawn from these studies, but
modifications in coding of some characters are proposed based upon new observations of
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,
1996). The refined phylogenetic hypotheses presented here are used to re-evaluate infra
generic classification in Andira, to present the first biogeographic hypotheses to explain
the evolution and diversification of the genus, and to investigate the evolution of mor
phological features, such as stipules, flowers, and fruits.
TAXONOMIC HISTORY
Andira first came 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.
Piso (1648) described the use of the seeds of A. fraxinifolia as an anthelminthic in Per
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 in many floral and vegetative characters.
Determining the first valid publication of the name Andira has been problematic. It
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
generico-specifica" to validate simultaneously the generic and specific names could not

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2003 ANDIRA 3
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
mosa 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.
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, the maintenance 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, it was 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, it would displace A. inermis, the name
traditionally used for the most common, widespread, and frequently collected species of
Andira.
Infrageneric classification
Bentham (1837, 1839, 1860, 1862) features prominently in the taxonomic history of
Andira. After studying the specimens of Martius, Pohl, and Schott from eastern Brazil, he
produced the first major account of Andira (1837, reprinted in 1839). He placed the
species in two sections: sect. Lumbricidia, defined by a hairy ovary on a relatively short
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
stipe character (my measurements indicate that the ratio of stipe length to ovary length
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
nized 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
glabrous ovary; these subsections correspond to Bentham's sections. Section Paucifoli
olatae is defined by presence of 1-3 leaflets, and therefore contains only A. unifoliolata
and A. trifoliolata.

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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
cluding A. nitida, A. carvalhoi, and A. fraxinifolia) are small trees and shrubs abundant in
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
species (A. verm?fuga, A. carvalhoi, A. legalis, A. nitida, and-A. fraxinifolia) are able to
root-sprout (pers. obs.), often extensively. This may enable A. carvalhoi to survive on
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
though too few observations have been made to assess their taxonomic significance. Pale
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.
obs.). The orange-brown slash of A. parviflora and A. cori?cea contrasts with the more
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
solitary or clustered in radial multiples of up to four, are large (98-234 mm), widely
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 are mostly heterocelluar (A. in
ermis is homocellular). The fibers are non-septate and usually very thick-walled.

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2003 ANDIRA 5
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
is useful for species delimitation, and provides three characters for cladistic analysis
(characters 5, 8, 10).
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. 1 A; 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
from one in A. unifoliolata to up to 19 (nine pairs) in A. chigorodensis (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
lar A. 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
(Fig. 2C) of six species (such as A. legalis and A.fraxinifolia). Distinguishing these two
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

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FIG. 1. Stipules and leaves in Andira (characters 3 and 4). A. Large, persistent stipules of A. anthel
(scale bar = ca. 5 cm; photo G. P. Lewis). B. Small stipules ("s" at arrows) of A. surinamensis (ruler left-
scale = inches; right-hand scale = cm). C. Small, unifoliolate leaves of A. unifoliolata. D. Large, imparipin
leaves of A. legalis. (Based on: A, R. T. Pennington 183; B, R. T. Penningfon 463; C, W. Rodrigues 11182; D
T Penningfon 305.)

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2003 ANDIRA 7
y.y s
FIG. 2. Indumentum and venation of leaflets of Andira (characters 5 and

surface of A. inermis, showing plane secondary and tertiary veins. B. Abaxial lea
with short, straight, tightly appressed hairs, prominently raised secondary vein, a
C. Abaxial leaflet surface of A. fraxinifolia with long, flexuose, erect hairs, prom
ondary veins, and slightly raised tertiary veins. D. Pale, long, fine, tangled hairs
A. cujabensis. E. Adaxial leaflet surface of A. surinamensis with channelled prim
face of A. unifoliolata with plane primary vein. P = primary vein; s = secondary v
= 1 mm. (Based on: A, T. D. Pennington 13558; B, R. T. Penningfon 463; C, G.
Ferreira 6259; E, R. T. Penningfon 463; F, A. Ducke 35078; all from herbarium ma
plant growing at Royal Botanic Gardens, Kew.)

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to mixed eucamptodromous to entirely brochidodromous (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
uniformly curving secondary veins of A. macrothyrsa contrast with those of A. inermis,
which 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
analysis.
Inflorescence. Inflorescences are terminal and axillary panicles, which vary in size,
shape, compactness, number of branches, angle at which the branches are held, and the
density of the flowers. Intraspecific variation is often extreme (e.g., in A. fraxinifolia and
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 are small, narrow, and often early ca
ducous. There is little variation in their morphology throughout Andira. The bracteoles are
paired at the base of the calyx.
Flowers. Flower 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
sioides, 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
glabrous, and this variation shows geographical correlation, which is the basis for the de
scription of subspecific taxa in A. inermis (see discussion under the individual species).
The corolla o? Andira consists of five glabrous petals. The keel petals are not united
but are firmly attached because of 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.
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).
Androecium. In all species of Andira, the stamens are diadelphous (9 + 1); the vexil
lary stamen is free (Fig. 3). Very rarely, individuals of A. fraxinifolia and A. macrothyrsa
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

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

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B(i)
FIG. 4. Wing petal sculpturing in Andira. Species with wing petal sculpturing: A, wing petal of A. ver
m?fuga, S = lamellate sculpturing, scale bar = 1 mm; A(i) wing petal of A. fraxinifolia (16 mm long); A(ii) wing
petal of A. carvalhoi (15 mm long); A(iii) wing petal of A. nitida (11 mm long); A(iv) wing petal of A. jalis
censis (12 mm long). Species without wing petal sculpturing: B, wing petal of A. trifoliolata, scale bar = 1 mm;
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 (8 mm long). (Based on: A, J. Ratter 3595; A(i) R. T Pennington 228; A(ii) M. P. M. de
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
ganization in Andira that appears to reflect a general bee pollination mechanism.
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

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2003 ANDIRA 11
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.
Fruit. Fruit of Andira are single-seeded (occasionally 2-3-seeded) drupes with

woody endocarps, fibrously fleshy or hard mesocarps, and thin exocarps. They are diverse
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
dry, and much larger fruits up to 12 cm long and weighing to 40-300 g when dry. These
size differences reflect probable differences in dispersal biology (see "Dispersal
Syndromes").
Bentham (1860) and Lewis (1987) point to the probable utility of fruit morphology in
Andira 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 the mesocarp and ex
ocarp are useful characters. For example, the green mesocarp of A. anthelmia and A. frax
inifolia contrasts with the pale green-white mesocarp of A. ormosioides, and confirms the
distinctness of A. ormosioides. Andira anthelmia and A. ormosioides have dark brown ex
ocarps, whereas that of A. fraxinifolia is green.
Detailed field notes of exocarp and mesocarp color and structure, and preservation of
fruits in alcohol should be a priority for future collectors of Andira. Better field data might
clarify some of the outstanding problems of species delimitation in Andira (such as that
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, it was decided that mesocarp
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

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

parenchyma cells in which 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 a multistate cladistic character (Pennington & Gemein
holzer 2000; character 10). This is a refinement of the coding of Pennington (1996), where
the distinct state of the endocarp dominated by parenchyma and/or collapsed cell tissue
was not recognized.
Five species have an endocarp predominantly formed by parenchyma and/or col
lapsed cell tissue, which is most often brown due to tannin. Twelve species have an en
docarp dominated by fibers. Small parenchyma cells and sometimes squashed cell tissue
occur between the fibers. Air gaps were often observed, probably due to ripening or de
generation processes. The fibers can be arranged predominantly in layers, but in A. ver
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
docarp comprises two layers of equal thickness: an inner layer of parenchyma and col
lapsed cell tissue, and an outer layer of stone cells. This species is coded with an unique
(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
(1982) noted seed pr?dation by Cleogonus weevils in A. inermis, and I have seen fruit of
A. macrothyrsa on a herbarium specimen (A. Gentry 37168) that have been attacked by
weevils. Both of these species have weak endocarps composed of parenchymatous tissue.
Andira humilis has an endocarp of woody fibers, but this layer is thinner than in any other
species examined, perhaps making it less of a barrier against burrowing seed predators;
Handro (1969) reported seed pr?dation by Cleogonus weevils in this species. Field obser
vation and examination of preserved fruit revealed no seed pr?dation in A. 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.
These data suggest that greater seed protection may be provided by these harder, thicker
endocarps.
Seeds. Seeds of species of Andira can be considered "overgrown" (sensu Comer,

1976), because they have an undifferentiated testa and contain no endosperm. They fill the
entire cavity of the fruit, and are more or less globose to more 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
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.

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2003 ANDIRA 13
Seedlings. Germination is cryptohypogeal (de Lima 1991; pers. obs. for A. carvalhoi,
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
foliolata, pers. obs.). Leaflet nyctinasty (evening leaf movements) was not observed in
any seedling of the seven species of Andira grown during this study.
It appears possible that polyembryony?the presence of more than one embryo in an
ovule (Richards 1997)?may occur in Andira, because more than one shoot can be pro
duced from a single seed o? A. fraxinifolia (P. Griffiths, pers. comm.). This raises the pos
sibility that the complex morphological variation patterns of this species might be ex
plained in part by apomixis. Further developmental studies of this and other species are
necessary.
POLLINATION BIOLOGY
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 18 mm long. Although the larger (19-24 mm long) flowers of A.
anthelmia have superficially similar morphology and are not significantly larger in terms
of flower length, I have only seen these flowers 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
valhoi, and A. verm?fuga, where few individuals in a population flower synchronously (R.
T. Pennington, unpubl.; J. A. Ratter, pers. comm.). Two other species, A. legalis and A. 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 that they, too, have distinct pollina
tion biology, although there are no field observations 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.

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FIG. 5. A. Small, bat-dispersed fruits of A. fraxinifolia (R. T. Pennington 213). B. Large, rodent-dispersed
fruits of A. legalis (G. P. Lewis & H. C. de Lima 1196; photo G. P. Lewis.)
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
ily Phyllostomidae in Costa Rica. It seems reasonable to assume that most Andira species
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).
Eight species of Andira possess much larger, heavier fruits, up to 12 cm long and
weighing probably up to 500 g when fresh (Fig. 5B; dry fruit of A. macrocarpa weigh
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
important dispersers of large fruit in the Neotropics (Emmons 1990). Of these large
fruited species, I have only seen fresh fruit of A. carvalhoi, which are brown and odorless

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2003 ANDIRA 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
cases of rodent dispersal in the estimated 650 genera of Leguminosae are of Parkia mul
tijuga Benth. (Mimosoideae; Hopkins & Hopkins 1983) and Hymenaea courbaril L. (Cae
salpinioideae; Hallwachs 1986). Some species appear to be secondarily dispersed by
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 that mode 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
is A. inermis (see Figs. 14,15). Putatively rodent-dispersed species are often restricted en
demics, e.g., A. micrantha is confined to the vicinity of Manaus in central Amazonia, A.
grandistipula is endemic to the Pakaraima Mountains in Guyana, A. carvalhoi is endemic
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
persed by birds and primates. Unfortunately, the data in Table 1 are somewhat inconclu
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 composition of five species of Andira. All figures are % by weight of the air-dried
mesocarp; the maturity of the fruit was proven by the subsequent germination of seed from all accessions, ex
cept A. ormosioides. All vouchers are collections of R. T. Pennington and deposited at CEPEC, FHO, and K.
Fruit size and

Species presumed disperser Protein Fat Sugars Starch
A. surinamensis RTP 364 small, bat 6.6 3.4 32.6 8.9

A. fraxinifolia RTP 202 small, bat 7.3 7.2 3.3 8.6
A. fraxinifolia RTP 213 small, bat 9.5 1.6 2.2 5.6
A. ormosioides RTP 306 small, bat 5.3 11.6 2.8 13.7
A. anthelmia RTP 227 small, bat 5.3 2.0 8.8 28.7
A. carvalhoi RTP 232 large, rodent 6.8 0.6 2.3 26.2

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caution. Second, although A. carvalhoi has a particularly high starch content, which re
flects the large numbers of starch grains in the parenchyma cells of its mesocarp, the sim
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 the much stronger shoot growth of the other four
bat-dispersed species germinated.
CHROMOSOME NUMBERS
Knowledge of chromosome numbers in Andira is restricted to those reported by

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
species (not named by Goldblatt) have n = 10.
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
washed 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
drochloric acid at room temperature before staining with Feulgen (prepared after Fox,
1969) for two hours. They were then mounted in 45% acetic acid and observed under
phase contrast.
Perhaps because the root tips were not collected from entirely healthy, vigorously
growing plants, the preparations showed few cells in metaphase. Thus, the counts made
should be regarded as preliminary. Counts for A. fraxinifolia show 2n = 22, and for A. an
thelmia, 2n = 21 or 22. These results, together with those reviewed by Goldblatt (1981),
indicate that there may be two basic chromosome numbers in Andira: n = 10 and n = 11.

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2003 ANDIRA 17
INTRASPECIFTC cpDNA POLYMORPHISM AND

POTENTIAL HYBRIDIZATION IN ANDIRA
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
cific cpDNA polymorphism in A. inermis, A. humilis, A. carvalhoi, and A. verm?fuga.
Subsequently, the intraspecific cpDNA polymorphism in A. verm?fuga was dismissed, be
cause the accession R. T. Pennington 250 does not belong to this species, but is an indi
vidual of A. fraxinifolia or perhaps a new species (Pennington 1996; see notes under A.
fraxinifolia). The intraspecific variation in A. 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 in A. humilis and A. carvalhoi are phylogenetically disparate. This
type of intraspecific cpDNA polymorphism can be explained by two processes: introgres
sive hybridization and/or the maintenance of an ancestral polymorphism. Unfortunately,
the most basic information necessary to confirm introgression as a possibility in Andira?
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
species. If ancestral polymorphism were the explanation, then this polymorphism has
been successfully maintained over remarkably long periods, and perhaps we might also
expect to discover maintenance of the intermediate haplotypes.
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 in misleading 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 in more populations of A.
humilis in an attempt to discover how widespread intraspecific cpDNA polymorphism is
in this species.
Methods
All methods followed Pennington (1995). Fourteen restriction site mutations (Ap
pendix 1) diagnostic for different clades of species (Figs. 7, 9) were selected for study, a
technique recommended by Doyle et al. (1990) and Rieseberg and Brunsfeld (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.

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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
phism in Andira. These data confirm that intraspecific cpDNA polymorphism involving
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.
Andira humilis is widespread in seasonally dry cerrado vegetation in central Brazil,
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
the more mesic 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
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
sequent introgression have occurred in this area. This hypothesis is supported by the ob
servation that the individuals RTP 246 and 247 were small trees, rather than geoxylic suf
frutices, the typical growth form of A. humilis. Although A. surinamensis, a species of
Clade III, is distributed along the southern fringes of Amazonia that border the northern
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 it would be interesting to
screen populations of A. humilis from this area to test this.
The case of A. carvalhoi is more complex. Pennington (1995) screened two accessions
fully (i.e., for 37 phylogenetically informative restriction sites). One had the plastome of
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
available 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
hoi's true phylogenetic position is in Clade III and its "true" cpDNA type is that of Clade
III, it is difficult to explain the plastome type polymorphism in A. carvalhoi by recent

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

Rain Forest plastome type of Clade III
Seasonally Range of A. carvalhoi

Dry Vegetation
Atlantic Coastal A Populations of A. humilis with
Rain Forest plastome type of Plastome Group I
FIG. 6. Location of populations of Andira humilis and A. carvalhoi sampled in a study of intraspecific
cpDNA polymorphism. The plastome types present in each of the different vegetation 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. It might 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
adapted to water stress, may have been more widespread. That the cpDNA type of acces
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

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have been the source of its plastome. To unravel the reasons for the complex intraspecific
cpDNA polymorphism in A. carvalhoi clearly requires sampling of more populations and
full characterization of plastome types.
It is tempting to ascribe some of the difficulties in species delimitation in Andira, such
as the difficulties in distinguishing some specimens as A. fraxinifolia and A. ormosioides,
to the occurrence of hybrids. If morphological intermediacy may be taken as evidence for
hybridity (there are frequent exceptions; e.g., see Bennett 1994, McDade 1990), then hy
bridization may be possible, but rare. I reported having seen no morphological evidence
of hybridity amongst sympatric species of Andira in southeastern Brazilian restinga
forests (Pennington 1995). Subsequently, I have observed a single plant that appeared to
be intermediate in leaf and stipule morphology between A. anthelmia and A. fraxinifolia
in the Poco das Antas Reserve in the State of Rio de Janeiro.
CLADISTIC ANALYSES OF SPECIES RELATIONSHIPS
Introduction
Cladistic analysis of chloroplast DNA restriction site characters for species o? Andira
and two outgroup species of Hymenolobium are presented by Pennington (1995). Pen
nington (1996) argued that directly combining both cpDNA restriction site and morpho
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
analysis of morphological characters produced a highly unresolved result because of the
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
species, whilst the morphological characters provide resolution within these groups. The
same view is taken here: that a combined cladistic analysis of all available data represents
the best phylogenetic hypothesis for Andira, the hypothesis that will be used for discus
sions of infrageneric classification, character evolution, and biogeography.
Species Concepts
Species in this monograph were delimited largely using the framework of the phylo
genetic species concept (e.g., Nixon & Wheeler 1990; Luckow 1995) and die characters
and character states listed below. They are diagnosable entities, characterized by constant
or consistent differences.
Methods
Three cladistic analyses are presented that are based upon Pennington's studies (1995,
1996), with small modifications. These are: 1) morphological analysis; 2) combined mol
ecular and morphological analysis; and 3) combined molecular and morphological analy
sis, including a subset of cpDNA restriction site data for species not included by Pen
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

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 21
Table 2. Data matrix of morphological characters of Andira.
Character
Hymenolobium flavum 0
Hymenolobium nitidum 0
A. inermis subsp. inermis 0
A. inermis subsp. rooseveltii 0
A. inermis subsp. glabricalyx 0
A. jaliscensis 0
A. multistipula 0
A. cubensis 0
A. galeottiana 0
A. macrothyrsa 0
A. verm?fuga 1
A. humilis 2
A. anthelmia 0
A. carvalhoi 1
A. fraxinifolia 1
A. legalis 0
A. marauensis 0
A. /u'f?fti 1
A. ormosioides 0
A. surinamensis 0
A. cordata 0
A. cori?cea 0
A. cujabensis 0
A. grandistipula 0
A. micrantha 0
A. parviflora 0
A. praecox 0
A. trifoliolata 0
A. unifoliolata 0
A. chigorodensis 0
A. taurotesticulata 0
A. tervequinata 0
A. macrocarpa 0
here included in A. 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
among the terminal taxa, which implies that they are not environmentally related.

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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
iting states within them (Pimentel & Riggins 1987; Farris 1990; Stevens 1991; Gift &
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.
Several multistate 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
(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
parentheses) refers to its position in the data matrix of the combined analysis. 2) The
states of each character are described and assigned a code (the number in parentheses
after each character state).
Vegetative characters.
1 (39). Tree (0); able to root-sprout and form a multistemmed 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
plying Weston's (1988) modification of Nelson'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.
2A, B, C, D]?The fine, pale, tangled, erect hairs (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 to white, the standard generally with red or purple markings
(0); flowers pink to purple, the standard generally with a pale central marking, less than

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 23
18 mm 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
above. Flower length in Andira 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
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 drupe, with fibrous mesocarp con
taining air spaces (A. galeottiana only) (4).
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);
endocarp of fiber and stone cells (A. jaliscensis only) (4).?This character presents
problems, because the outgroup species of Hymenolobium lack an 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
Hymenolobium become optimized for states of an endocarp, a structure that they do not
have. For this reason the character is coded here as a single multistate character with ab
sence as one state. Moreover, coding as two characters, as recommended by Hawkins et
al. (1997), does not change the outcome of any analyses, because only the outgroup taxa
lack the structure.
Combined molecular 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.
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
agnostic mutations described above (Appendix 1) as part of the study of intraspecific
cpDNA polymorphism and potential hybridization. Where cpDNA data were not avail
able, character states were scored with question marks. The data matrix is presented in
Table 3.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
Table 3. Matrix of cpDNA restriction site and morphological data for Andira. Characters 1-38 are re
striction sites; characters 39-48 are morphological. Multiple accessions for a species included in the molecular
study are indicated by collector and number. Collectors are abbreviated: AMC = A. M. de Carvalho; CEH = C.
E. Hughes; HCL = H. Cavalcante de Lima; MC = M. Cheek; MS = M. Sugiyama; RTP = R. T. Pennington; TDP
= T. D. Pennington.
1 2
Character 12 3 4 5 6 7 90123456789012345
1 1 0 1flavum
Hymenolobium 1 1 1 1 0 0 ? ?
1 00 1 10 0 1 1 0
1 1 0 nitidum
Hymenolobium 1 1 1 1 1 001 0 0 ? 0 1 10 0 1 1 0
A. inermis 1CEH 1 0 1 11673
0 1 110 1 1 0 1 0 0 1 0 1 1 1 1
A. inermis TDP 1 1 0 1 13558
1 0 1 11? 0 ? 0 1 0 0 0 0 1 1 0 1
A. inermis MC 3579 ? 1 1 0 1 1 0 1 1 ? 1 1 0 1 0 0 0 0 1 1 0 1
A. macrothyrsa 0 0 1 0 0 1 1 1 0 1 1 1 0 1 0 1 0 1 0 0? 1 0
A. galeottiana 0 0 1 0 0 1 1 1 0 1 0 1 0 1 0 1 0 0 1 1 0
A. humilis RTP 269 0 0 1 0 0 1 1 1 0 1 0 1 0 1 0 1 0 0 1 1 0
A. fraxinifolia RTP 250 00 1 1 0 0 1 0 ? 1 0 1 0 0 1 1 0 1 0 0 1 1 0
A. verm?fuga 0 0 0 1 1 1 0 1 1 1 0 1 0 1 0 1 0 0 1 1 0
A. anthelmia RTP 227,282 0 1 0 0 1 0 0 1 0 1 0 0 1 1 0 1 0 0 1 1 0
A. carvalhoi RTP 229 0 1 0 0 1 1 1 0 1 1 1 0 0 0 1 0 1 0 0 1 1 0
A. carvalhoi RTP 233 0 1 1 0 0 1 0 0 1 0 1 0 0 1 1 0 1 0 0 1 1 0
A. fraxinifolia RTP 213 0 1 1 0 0 1 0 0 1 0 1 0 0 1 1 0 1 0 0 1 1 0
A. fraxinifolia MS 889 0 1 1 0 0 1 0 0 1 0 1 0 0 1 1 0 1 0 0 1 1 0
A. legalis RTP 307 0 1 1 0 0 1 0 0 1 0 1 0 0 1 1 0 1 0 0 1 1 ?
A. legalis HCL s.n. 0 1 1 0 0 1 0 0 1 0 1 0 0 1 1 0 1 0 0 1 1 0
A. nitida RTP 300 0 1 1 0 0 1 0 0 1 0 1 0 0 1 1 0 1 0 0 1 1 0
A. nitida RTP 292 0 1 1 0 0 1 0 0 1 0 1 0 0 1 1 0 1 ? ? 1 1 0
A. nifwfa RTP 301 0 1 1 0 0 1 0 0 1 0 1 0 0 1 1 0 1 0 0 1 1 0
A. ormosioides 0 1 1 0 0 1 0 0 1 0 1 0 0 1 ? ? ? ? 0 1 ? ?
A. surinamensis 0 1 1 0 0 1 0 0 1 0 1 0 0 1 1 0 1 0 0 1 1 0
A. cordata 0 0 0 0 1 1 1 0 0 1 0 0 0 1 1 0 1 1 0 0 1 1
A. grandistipula 0 0 0 0 1 1 1 0 1 0 0 0 0 0 0 1 1 0 0 1 1
A. parviflora 0 0 0 0 1 1 1 0 1 0 0 0 1 1 ? 1 1 0 0
A. unifoliolata 0 0 0 0 1 1 1 0 1 0 0 ? 0 1 0 1 1 0 0
A. inermis KY? 512 ? 1 0 1 1 0 1 1 0 1 ? ? ? ? 0 1 ?
A. inermis RTP 580 0 ? ? 0 ? 1 0 1 ? ? ? 0 1 ? ? ?
A. inermis RTP 589 0 1 1 1 0 1 9 9 9 0 1 ? ? 9
A. multistipula 0 1 ? 0 1 ? ? ? ? 1 ? ? ?
A. jaliscensis 0 ? ? 0 ? 9 9 9 ? ? ? 1 ? 9 9
A. taurotesticulata 0 1 0 1 1 0 1 ? 9 9? ? ? 0 1 ? ? ?
A. marauensis 1 0 1 0 1 ? 9 9 9 9 9 ? ? 0 0? 9 9
A. cujabensis 0 0 1 11 0 1 0 ? ? ? ? ? ? 1 0 ? ? ?
A. carvalhoi AMC 0 0 ? 1 ? 1 ? ? ? ? ? ? ? ? ? 0 ? ? ?
A carvalhoi RTP 217 1 ? 0 ? 1 ? 0 9 9 ? 0 ? 9 9 9 9 9 9 9
Results
Morphological analysis. Pennington (1996) showed that a cladistic analysis of mor

phological characters of a subset o? Andira species provided little phylogenetic resolution.
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
analysis resulted in 2966 equally most-parsimonious trees (the limit that the computer
could store; CI = 0.57, RI = 0.79, length = 38). The strict consensus tree is entirely unre

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 25
Table 3 continued.
2 3 4
Character_67890123456789012345678
Hymenolobiumflavum 0 1 ? 1 10?00010?0001100000
Hymenolobiumnitidum 01??????01 ?0?0001110000
A. inermis CEH 1673 010001 10010000100100011
A. inermis TDP 13558 01000100010000100100011
A. inermis MC 3579 010001 10010000100100011
A. macrothyrsa 07001 10001 1000101000011
A. galeottiana 00001 10001 1000101100142
A. ?urni/fr RTP 269 00001 10011 100210?1?0122
A. fraxinifolia RTP 250 ?00?1 10701 101 1 101100012
A. verm?fuga 00001100111001101100722
A. anthelmia RTP 227,282 00001101011010111200012
A. carvalhoi RTP 229 00001100011001100100032
A. carvalhoi RTP 233 00001101011011100100032
A.yhmm/o/??RTP213 00001101011011101100012
A.yhmm/0/uzMS889 70077777011011101100012
A. legalis RTP 307 70071107011010111200032
A. legalis UCLs.n. 00001101011010111200032
A. nitida RTP 300 00001 10101 101 1107170772
A. mttfo RTP 292 07701 10701 101 1 10?1?0??2
A. n?tida RTP 301 00001101011011107170772
A. ormosioides 7077777701 1010101200012
A. surinamensis 00001101011010101100022
A. cordata 10101100010100100011113
A. grandistipula 11101100010100111717033
A.parviflora 70177107017100101001013
A. unifoliolata 70101100010100101071013
A. inermis RTP 5\2 77777777070700100100011
A. inermis RTP 5S0 77777777070700100100011
A. intrm? RTP 589 77777777070700100100011
A. multistipula 77777777077700110100011
A. jaliscensis ????????0???00102100014
A. taurotesticulata 77777777077700101001031
A. marauensis 7777777707770010111017?
A.cujabensis 77777777070700102001013
A. carvalhoi AMC 77777777077701 100100032
A. ozraz//i0/RTP217 77777777777771100100032
solved, except for a clade comprising A. cordata, A. cori?

character of leaflet number ordered, the analysis resulte
most-parsimonious trees (2966, CI = 0.57, RI = 0.80, length
sus tree resolved another clade comprising A. unifoliolata
Combined molecular and morphological analyses. The fi

equally most-parsimonious trees of length = 76; CI = 0.72
sus is shown in Fig. 7. Of the equally most-parsimonious
cause they result from incorrect optimization of missing
for character 47. These accessions are assigned a charac
persed fruit, when they do not have this state (this speci
states of bat-dispersed fruit). Rejection of these topologi

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
-Hymenolobium flavum
-Hymenolobium n'rtidum
-A. inermis CEH 1673
-A. inermis TDP 13558 CLADE I

-A. inermis MC 3579
-A. parviflora
-A. cordata
CLADE II
-A. grandistipula
-A. unifoliolata
-A. carvalhoi RTP 229
-A. macrothyrsa
PLASTOME
-A. galeottiana GROUP I
-A. humilis RTP 269
-A. verm?fuga
-A. surinamensis
-A. ormosioides
-A. anthelmia RTP 227,282
-A. legalis RTP 307
-A. legalis HCL sn

CLADE III
-A. fraxinifolia RTP 250
-A. fraxinifolia MS 889
-A. nitida RTP 300
-A. nitida RTP 292
-A. nitida RTP 301
FIG. 7. Strict consensus cladogram of five equally most-parsimonious cladograms resulting from cladistic
analysis of combined Andira cpDNA restriction site and morphological data. Collector and number are indicated
when more than one accession for a single species was included in the molecular study.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 27
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
most-parsimonious trees reflect different arrangements of the accessions of A. fraxinifo
lia, A. nitida, and A. carvalhoi {RTP 233). The semi-strict consensus tree is more resolved,
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 a monophyletic 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
species of this clade, because it has the plesiomorphic state of restriction site characters
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 site characters.
Andira 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
increasing the numbers of equally most-parsimonious trees and decreasing the resolution
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.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
-Hymenolobium flavum
-Hymeno/obium nrtidum
-A. inermis CEH 1673
-A. inermis TDP 13558
-A. inermis MC 3579
-A. inermis RTP 512
-A. inermis RTP 580 CLADE I
-A. inermis RTP 589
-A. mu/tistipula
-A. jaliscensis
-A. taurotesticulata
-A. parvif/ora
-A. cujabensis
-A. cordata CLADE II
-A. grandistipu/a
-A. unifoliolata
-A. carvalhoi AMC
-A. macrothyrsa PLASTOME
GROUP I
-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
CLADE III
-A. n?tida RTP 300
-A. n?tida RTP 292
-A. n?tida RTP 301
FIG. 8. Strict consensus cladogram of 2996 equally most-parsimonious cladograms resulting from cladis
tic analysis of combined Andira cpDNA restriction site and morphological data, including accessions of species
not studied previously by Pennington (1995,1996). Collector and number are indicated when more than one ac
cession for a single species was included in the molecular study.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 29
CHARACTER EVOLUTION
Stipules (Fig. 9, character 41). Large persistent stipules have arisen three times inde
pendently in Andira, and are autapomorphic for A. grandistipula and A. multistipula and
synapomorphic for A. anthelmia and A. legalis. In Pisum sativum L. (pea), a single gene
mutant, stipules-reduced {st) causes a marked reduction in stipule 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,
persistent stipules that is the diagnostic character that distinguishes this species from A.
inermis. From the standpoint of the phylogenetic species concept (Nixon & Wheeler
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 the molecular 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
petals, is morphologically most similar to A. macrothyrsa and may be its sister species.
Loss of wing petal sculpturing is a synapomorphy for Clade II, and an autapomorphy for
A. taurotesticulata. It may have also been independently lost by A. chigorodensis, if this
species is most closely allied with A. macrothyrsa.
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 in Andira. 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
and endocarps of stone cells, but they share no great morphological similarity with A.
grandistipula, 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
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
creasing seed protection (Pennington & Gemeinholzer 2000; discussion above), and seed
pr?dation may have been the pressure driving this evolutionary sequence.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
r-fl-[]?Hymenolobium f/avum
?|?Hymenolobium nitidum
?A. inermis CEH 1673
1 7 13202530323642
fOttHf?
1232
10 1 110 10 0 1824 rff--A. inermis TDP 13558
CLADE I
4XH 00
0 ? ?-A. inermis MC 3579
3 1012192122283637434548
I-A. parvif/ora
4246
ttOHUOHHH
000100101013
A. cordata
16274147 CLADE M
A. grandistipu/a
0 113
0 1111 A. unifoliolata
394247
S-S? A. carvalhoi RTP 229
1?3 4348
-A. macrothyrsa
1423 0 1 PLASTOME
4647 -??A. galeottiana GROUP I
if
1 2 3439
-??A. /H//W//S /?7P 269
HHKHKtH
ft 1 1
0 0 0 10 2 ?A. verm?fuga
A. surinamensis
A. ormosioides
A. anthelmia RTP 227r282
4 6 8 3338 A. /e#3//s /?7P 307

LKK}ft
10 0 11 A. legalis HCL sn
A. fraxinifolia RTP 250

CLADE III
A. fraxinifolia RTP 213

47
?A, carvalhoi 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

cpDNA restriction site and morphological data. The bars representing character changes are labelled a
the character number and below with the character state. Solid black bars = unique changes, stippled bla
= homoplasious changes. Collector and number are indicated when more than one accession for a sing
was included in the molecular study.

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2003 ANDIRA 31
CRYPTIC CLADES
Pennington (1995) reported that none of the monophyletic groups, even the well-sup
ported clades, discovered in the molecular 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
morphological characters provide unambiguous support for some of these cryptic groups.
Clade II is supported by two unique micromorphological character states: lack of wing
petal sculpturing (character 45) and an endocarp of stone cells (character 48). All species
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
monocotyledons (Rudall 2000). In both cases, reciprocal illumination provided by new
hypotheses of grouping from molecular data is important. For example, it was 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.
INTRAGENERIC CLASSIFICATIONS IN
THE LIGHT OF THE CLADISTIC ANALYSES
The cladistic analyses show that the infrageneric classifications proposed for Andira
by Bentham (1860, 1862) and Mattos (1979) are flawed. The only section that has been
proposed that may be monophyletic is sect. Paucifoliolata Mattos, comprising only A.
unifoliolata and A. trifoliolata. This is a monophyletic group in the morphological cladis
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
(Fig. 10); thus Bentham's section (Mattos's subsection) Lumbricidia is paraphyletic and
section Euandira (Mattos's subsection Glabratae) polyphyletic.
Unfortunately, the cladistic framework presented here does not provide a clear route
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
plasious restriction site character. Moreover, both Clade III and Clade I remain cryptic,
supported by restriction site data but not by morphological characters. Whilst there is no
reason to doubt that these cryptic clades reflect true phylogeny because they lack mor
phological support (indeed, ultrastructural and micromorphological characters may well
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,
they cannot be keyed out for conventional identification. Hence, I propose that sectional
classification within Andira should be abandoned. A further factor supporting this

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
?Hymenolobium flavum = 0
-Hymeno/obium nitidum = 0
-A. taurotesticulata = 0
?A. jalisensis = 0
-A. inermis TOP 13558= 0
1-A
-A. inermis MC 3579 = 0
-A. inermis CEH 1673= 0
-A. inermisRTP 512= 0
-A. inermis RTP 580= 0
-A. inermis RTP 589 = 0
?A. multistipu/a = 0
-A. cujabensis = 0
-A. parviflora = 0
A. cordata = 1
-A. grandistipula = 0
A. unifoliolata = 0
A. carvalhoi RTP 229= 0
c A. carvalhoi AMC= 0
A. macrothyrsa= 0
A. galeottiana = 1
1
i-A. humilis RTP 269= 1
??A.. verm?fuga = 1 <?
-j?A. marauensis= 1
A. surinamensis = 0
A. ormosioides= 0
A. anthelmia RTP 227,282= 0
-r
A. legalis RTP 307 = 0
??A. legalis HCL sn = 0
A. fraxinifolia RTP 250 = 0
A. fraxinifolia RTP 213= 0
A. fraxinifolia MS 889= 0
?A. n?tida RTP 300= 0 <?
A. n?tida RTP 292= 0 <?
A. n?tida RTP 301= 0 <?
?
FIG. 10. An equally most-parsimonious cladogram resulting from cladistic analysis of combined Andira
cpDNA restriction site and morphological data showing character state changes for indumentum of the gynoe
cium. The bars represent character state changes from a hairy to a glabrous gynoecium. Terminal taxa marked
"0" have a hairy gynoecium; terminal taxa marked "1" have a glabrous gynoecium. Andira nitida and A. ver
m?fuga have both hairy and glabrous gynoecia. Collector and number are indicated when more than one acces
sion for a single species was included in the molecular study.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 33
argument is that molecular data are not available for some species whose relationships are
hard to estimate based upon morphology alone (e.g., A. cubensis, A. macrocarpa).
BIOGEOGRAPHY
Distribution and Habitats
The majority o? Andira species are endemic to South America. The exceptions are A.
cubensis (endemic to Cuba), A. jaliscensis and A. galeottiana (endemic to Mexico), and
A. inermis (widespread throughout the entire Neotropics and also present in Africa).
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
Andira 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
unresolved (Pennington et al. 2001). Furthermore, the lack of resolution in the basally di
vergent Andira clade (clade I, Fig. 8), in which all distribution areas (South America, Cen
tral America, 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
gin (cf. Lavin and Luckow 1993), but no firm conclusions can be drawn until relationships
are better resolved. The phylogenetic position of the Mexican 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
biogeographical relationships between Africa and North America in two legume groups
that diversified 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 a more 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 (Buskirk 1985; Donnelly 1988, as cited in Lavin, 1993), and therefore its

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
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
magnitude shorter than that between South America and Africa.
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 in many 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 in Mexico
(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 on M. Cheek
3579); in the Neotropics it often grows along rivers and can survive in mangrove swamps
(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 to Haiti (ca.
100 km at the closest point). Given the presence of A. inermis on the islands of the Lesser
Antilles 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
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 in man
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 species of Clade III, with the exception of A. surinamensis (an Amazonian
species), occur in the moist forests of coastal southeastern Brazil. Thus, there seems to have
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
foliolata), the Guianas (specifically Guyana; A. grandistipula), and the central Brazilian
cerrados (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
cal characters supporting this clade (lack of wing petal sculpturing, endocarp composed

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 35
H oc
H 20e
CERRADO
BIOIVJE
80? 50?
-Contact line of Andira cujabensis/cordata
-Contact line of harry & glabrous forms of

Andira humilis
.Miiimmi. in Contact line of hairy & glabrous forms of
Pterodon 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
cordata and A. cujabensis (see Fig. 39). Dashed line with dots: line of contact between hairy and glabrous for
of A. humilis. Comb: line of contact between hairy and glabrous forms of Pterodon emarginatus. Dotted lines:
lines dividing the distinct northern and southeastern cerrado sites identified by analysis of the floristic compo
tion of 98 areas of cerrado vegetation (Ratter et al. 1996)
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 th
areas, so the five species represented in the molecular and combined analyses probably ar
a representative geographic sample.
The parapatric distributions of the Clade II species A. cujabensis and A. cordata, whi
are found in the Brazilian cerrado vegetation, are especially interesting (see Fig. 11). Th

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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
vegetation 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
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 in Minas 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 the
ovary, which is approximately oval in cross section. The "upper surface" refers to the
areas on and adjacent to the adaxial suture, and the "lower surface" refers to the areas on
and adjacent to the abaxial suture; the "sides" refer to the two regions bounded by 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.
Mattos, Loefgrenia 58: 2. 1973, nom. superfl.?Type: Andira inermis (W.
Wright) DC.
Lumbricidia Vellozo, Fl. flumin. 305. 1829. Andira sect. Lumbricidia (Vellozo) Ben
tham, Comm. 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
brochidodromous or mixed eucamptodromous-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,

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 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;
mesocarp fibrously fleshy and sweet 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
tohypogeal. Chromosome number: x = 10, 11.
The order of species reflects the groups discovered by the combined analysis of
cpDNA restriction site characters and morphology. The relationships of species not
included in this analysis are estimated based upon their morphological characters. The
groups are (refer to Fig. 8): (i) "Clade I": A. inermis, A. jaliscensis, A. multistipula,
A. taurotesticulata, and A. cubensis (the last species not included in the combined analy
sis but placed in this group because of its morphological similarities to A. inermis);
(ii) "Plastome Group I": A. macrothyrsa, A. galeottiana, A. verm?fuga, A. humilis, and
A. chigorodensis (the last species not included in the combined analysis but placed
here because of its morphological similarities to A. macrothyrsa); (iii) "Clade III":
A. surinamensis, A. anthelmia, A. legalis, A. ormosioides, A. fraxinifolia, A. n?tida,
A. carvalhoi, A. marauensis, and A. macrocarpa (the last species not included in the
combined analysis but placed here because of its morphological similarities to A. suri
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
petals without sculpturing), A. parviflora, A. cordata, A. grandistipula, A. cujabensis,
and A. unifoliolata.
Keys to the Species of Andira

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

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
FIG. 13. Fruit morphology of Andira. A. ? Globose, smooth, dried fruit (ca. 3.5 cm long) of A. jaliscen
sis. B. Elongated, smooth, dried fruit (ca. 6 cm long) of A. chigorodensis. C. Elongated, wrinkled, dried fruit (ca.
4.5 cm long) of A. nitida.
measurements of petiolule length and leaflet size and shape are for the terminal pair of leaflets. Leaflet length is
from the base to the tip of the lamina and does not include the length of the petiolule. Acumen length is mea
sured from the point where the leaflet margin curves towards the apex. Wlien the acumen is broad, this point is
ill defined, so acumen lengths must be treated as approximate.
Flower length is reported for dried specimens, and is measured from the base of the calyx to the end of the
wing petals in fully open flowers. A fully open flower is one in which the standard petal is reflexed. Wing petal
sculpturing is described following the terminology of Stirton (1981).
Many species have more or less globose fruits (Fig. 13A). Other species have fruits that are referred to as
"elongated" (Fig. 13B, C). Fruit of some species dry smooth (Fig. 13 A, B) and of others distinctly wrinkled (Fig.
13C). Fruit length is measured from the tip of the stipe (where it joins the body of the fruit) to the tip of the fruit;
fruit height is the maximum measurement from the abaxial to adaxial suture; fruit width is the maximum mea
surement between the fruit walls.
I. Key emphasizing vegetative characters.
1. Leaves all uni- or trifoliolate; primary vein plane on adaxial surface.

2. Leaves trifoliolate. 28. A. irifoliolala.
2. Leaves unifoliolate. 29. A. unifoliolata.
1. Leaves all with two or more pairs of leaflets, or sometimes also some leaves
sunken on adaxial surface.
3. Geoxylic suffrutex forming mats up to 10 m in diameter (occasionally
10. A. humilis.
3. Tree or shrub.
4. Stipules generally more than 1.7 cm long and 0.5 cm wide at base, p
florescence bases and shoot apices.
5. Leaflets glabrous (very occasionally with sparse, erect hairs);
5. Leaflets abaxially hairy, sparsely hairy, or with tiny appressed in
microscope); Guianas and eastern Brazil.
6. Leaflets abaxially with tiny (<0.1 mm long) appressed straight
lens or microscope); leaflets up to 4 pairs; the Guiana
6. Leaflets abaxially with erect or ? appressed, often flexuose hai
visible to the naked eye); leaflets 3-9 pairs; eastern Brazil.
7. Stipules red-brown pubescent, glabrescent at maturity; leaf
hairs red-brown, erect; fruit 5.6-12 cm long, weighing over 10
brown when fresh and dry, odorless when fr
1. Stipules generally glabrous at maturity (red-brown appress
young); leaflets sparsely to very sparsely hairy abaxially, h
whitish, ? appressed to ? erect; fruit 3-6.2 cm long, weighing c
dark brown when fresh and dry, sweet smelling when f
4. Stipules less than 1.7 (-2.0) cm long and 0.5 cm wide at base, oft
entirely absent (rarely persistent and then paired only at leaf bases an
rescence bases or shoot apices).
8. Leaflets with indumentum abaxially (use lens or microscope).

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
ANDIRA 39
9. Leaflet undersurfaces (particularly primary and secondary veins) with erect, gener
ally flexuose hairs (>0.2-1.0 mm long); indumentum generally visible with the
naked eye and perceptible to touch.
10. Leaflets abaxially with dense indumentum of pale (occasionally red-brown), tan
gled hairs that ? obscure the epidermis; leaflets appear pale abaxially (seen with
the naked eye).
11. Leaflets coriaceous, base rounded to cordate; flowers 5.5-7 mm long;
Brazil. 21. A. cujabensis.
11. Leaflets chartaceous to subcoriaceous, base obtuse to rounded; flowers
9-1 lmm long; Mexico. 2. A. jaliscensis.
10. Leaflets abaxially with indumentum of red-brown hairs (or if pale, then indu
mentum sparse) that do not obscure the epidermis; leaflet often appearing red
brown abaxially (to the naked eye; if pale then indumentum is sparse).
12. Secondary veins plane or slightly raised (occasionally prominently raised)
abaxially, tertiary veins plane or very slightly raised abaxially; indumentum
of abaxial leaflet surfaces caducous; fruit distinctly ribbed from suture to
suture; in montane forests at 700-2000 m, occasionally in lowland forest
(Ecuador). 5. A. taurotesticulata.
12. Secondary veins prominently raised abaxially, tertiary veins prominently
raised abaxially; indumentum of abaxial leaflet surface persistent; fruit not
ribbed from 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,
sometimes retuse; fruit 2.4-6 (-10 [A. galeottiana only]) cm long,
weighing 10-20 (-125 [A. galeottiana only]) g when dry.
14. Tree; fruit 6-10 cm long, 40-125 g when dry, dried mesocarp
relatively soft with sponge-like structure; Mexico. 8. A. galeottiana
14. Shrub or tree; fruit 2.4-6 cm long, weighing 10-20 (-30) g when
dry, dried mesocarp hard (difficult to insert the point of a needle),
with hard, fibrous or granular structure; South America.
15. Secondary veins 10-15; flowers 6-7.5 mm 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
noecium glabrous or with sparse hairs along upper and
lower surfaces of ovary; dry fruits distinctly wrinkled;
Central Brazil and Bolivia (cerrado), occasionally
Amazonia. 9. A. verm?fuga.
16. Stipels 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).
17. Shrub or small tree to 12 m, with spreading crown in
open situations; flowers 13-17 mm long; fresh fruit
green with green mesocarp. 16. A. fraxinifolia.
17. Tree to 30 m, tall with small crown in open situations;
flowers 18-23 mm long; fresh fruit very dark brown
with pale greenish-white mesocarp. 15. A. ormosioides.
9. Leaflets abaxially with tiny (<0.2 mm long), tightly appressed hairs; indumentum
generally not visible to the naked eye or perceptible by touch (use lens or
microscope).
18. Tertiary veins distinctly raised on abaxial leaflet surface, secondary veins
impressed or slightly impressed on adaxial leaflet surface. 12. A. surinamensis.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
18. Tertiary veins plane or slightly raised on abaxial leaflet surface, secondary veins
plane or slightly raised on adaxial leaflet surface.
19. Shrub.
20. Leaflets 1-2 pairs; flowers 9 mm long; Venezuelan Guayana.
27. A. tervequinata.
20. Leaflets 2-4 (-5) pairs; flowers 10-13 mm long; eastern Brazil. 17. A. n?tida.
19. Tree.
21. Leaflets (1-) 2 (-3) pairs.
22. Secondary veins 6-7; leaflets widely elliptic to widely obovate, 1-2
pairs; small tree to 7 m tall; Venezuelan Guayana. 27. A. tervequinata.
22. Secondary veins 8-10; leaflets elliptic to narrowly obovate (occa
sionally widely obovate), (1-) 2 (-3) pairs; tree to 35 m tall; eastern
Brazil.
23. Stipules to 10 mm long; stipels 1-2 mm long; flowers 6.5-7 mm
long; Amap?. 26. A. praecox.
23. Stipules to 2.5 mm long; stipels absent (perhaps early
cous); flowers 9-11 mm long; Bahia. 18. A. m
21. Leaflets 3-9 (-10) pairs.
24. Leaf axis 37-55 cm long; leaflets 7-9 pairs. 7. A. ch
24. Leaf axis 6-33 (-40) cm long; leaflets 3-8 (-10) pairs.
25. Leaflets coriaceous; flowers red to purple; Cuba. 4.
25. Leaflets subcoriaceous; flowers white to yellowish, the st
marked with red or purple; Panama and South America.
26. Stipules absent (or early caducous; never seen), stipel
mm long, caducous; fruit 5-5.5 cm long, smooth, weigh
10-20 g when dry. 6. A. macroth
26. Stipules to 12 mm long, stipels 1-3 mm long, generally
sistent; fruit 5.4-7.2 cm long, distinctly ribbed from sut
to suture, weighing 100-200 g when dry. 5. A. taurot
8. Leaflets glabrous abaxially (use lens or microscope).
27. Leaflet base truncate or cordate (rarely rounded); Brazil, in cerra
27. Leaflet base obtuse to rounded (rarely ? truncate or slightly cordate); Am
Guianas, eastern Brazil, in rain forest and restinga.
28. Shrub (occasionally small tree to 10 m). 19. A
28. Tree.
29. Leaflets in 2-9 pairs (some leaves on a plant always with more than 3 pairs
of leaflets); Neotropics and Africa. 1. A. inermis.
29. Leaflets in 2-3 pairs; South America.
30. Stipules quite persistent, to 5 mm wide at base, to 15 mm long; the
Guianas. 24. A. cori?cea.
30. Stipules caducous, often entirely absent, <1 mm wide at base, to 6 mm
long; Central Amazonia, eastern Brazil.
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
dry; central Amazonia. 23. A. micrantha.
31. Stipules generally present at shoot apices (though caducous);
leaflets 2-4 (-5) pairs; petals 10-13 mm long, pinkish white to pur
ple, the standard with a central white spot; Atlantic coastal Brazil.
17. A. n?tida.
II. Key emphasizing flower and fruit characters.
1. Leaves all uni- or trifoliolate; primary vein plane on adaxial surface.

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

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 41
Leaves all with two or more pairs of leaflets, or sometimes also some leaves trifoliolate; primary vein
sunken on adaxial surface.
3. Stipules generally more than 1.7 cm long and 0.5 cm wide at the base, persistent, crowded at in
florescence base and shoot apex.
4. Flowers 19-24 mm long; eastern Brazil.
5. Fruit 5.6-12 cm long, weighing over 100 g when dry; stipules red-brown pubescent
when young, glabrescent at maturity. 14. A. legalis.
5. Fruit 3-6.2 cm long, weighing ca. 20 g when dry; stipules red-brown appressed
pubescent when young, generally glabrous at maturity. 13. A. anthelmia.
4. Flowers 11-15 mm long; Amazonia and Guianas.
6. Leaflets in (2-) 3-4 pairs, apex rounded (rarely obtuse) with an acumen to 3 mm long,
with tiny (<0.1 mm long) appressed hairs abaxially (use lens or microscope).
25. A. grandistipula.
6. Leaflets in (5-) 6-9 pairs, apex acute to obtuse with an acumen to 15 mm long, glabrous
abaxially (very occasionally with few, erect hairs). 3. A. multistipula.
3. Stipules less than 1.7 (-2) cm long and 0.5 cm wide at base, often caducous, sometimes entirely
absent (rarely persistent and then paired only at leaf bases, but not crowded at inflorescence base
or shoot apex).
7. Fruit 5-11 cm long, weighing (40-) 100-300 g when dry.
8. Leaflets glabrous abaxially.
9. Flowers 14-15 mm long; shrub or small tree to 10 m; eastern Brazil, in restinga.
19. A. carvalhoi.
9. Flowers 6-7 mm long; trees to 40 m; Amazonia and the Guianas, in rain forest.
10. Ovary glabrous; stipules quite persistent, to 15 mm long, 5 mm wide. 24. A. cori?cea.
10. Ovary sparsely pubescent; stipules early caducous (not seen). 23. A. micrantha.
8. Leaflets with indumentum abaxially.
11. Fruits ? globose; petals white, the standard marked with red; tertiary veins plane on
abaxial surface of leaflet. 5. A. taurotesticulata.
11. Fruits elongated; petals pink to violet, the standard marked with white
tiana; flowers unknown for A. macrocarpa); tertiary veins distinctly raised
ial surface of leaflet.
12. Dried mesocarp relatively soft (the point of a needle is easily inserted), f
and spongy; fruits weighing 40-125 g when dry; Mexico. 8. A.
12. Dried mesocarp hard (the point of a needle is inserted only with difficu
fibrous; fruits weighing 300 g when dry; Ecuador. H.A. m
1. Fruit 2.4-6 (-7) cm long, weighing 10-20 (-30) g when dry.
13. Petals white to yellow (occasionally pale pinkish or pale lilac), the standard
with red or purple markings.
14. Ovary glabrous. 22. A. cordata.
14. Ovary pubescent.
15. Leaflets with erect indumentum on abaxial surface.
16. Secondary veins 10-15; leaflets abaxially with red-brown indument
epidermis clearly visible; tree to 20 m, bark smooth; Brazil (Amazo
Para), in rain forest. 20. A. parviflora.
16. Secondary veins 8-10; leaflets abaxially generally with pale indumentum,
often so dense that the epidermis is hidden; tree to 12 m (occasionally a
shrub), bark thick, fissured; Brazil (Mato Grosso, Mato Grosso do Sul,
Goi?s, Para), in cerrado and gallery forest. 21. A. cujabens
15. Leaflets glabrous or with appressed indumentum on abaxial surface.
17. Leaflets 4-9 pairs; endocarp of fruit fibrous (the point of a needle is easily
inserted).
18. Flowers 9-11 mm long; leaf axis 10-33 (-40) cm long; leaflets 4-8
pairs. 6. A. macrothyrsa.
18. Flowers 5-6 (-7) mm long; leaf axis 37-55 cm long; leaflets 7-9 pairs.
7. A. chigorodensis.
17. Leaflets 1-2 (-3) pairs; endocarp of fruit very hard, amorphous (the point of
a needle is inserted only with difficulty).

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
19. Flowers 6.5-7 mm long; secondary veins 8-12 per leaflet; leaves gener
ally with 2 pairs of leaflets (occasionally with 3 pairs or trifoliolate); tree
to 30 m. 26. A. praecox.
19. Flowers 9 mm long; secondary veins 6-7 per leaflet; leaves with 1-2
pairs of leaflets; shrubs or small trees to 7 m. 27. A. tervequinata.
13. Petals red or pink to purple, the standard generally with a white or cream central mark
ing.
20. Dry fruits distinctly wrinkled (Fig. 13).
21. Leaflets with erect indumentum on abaxial surface. 9. A. verm?fuga.
21. Leaflets glabrous or with appressed indumentum on abaxial surface.
22. Geoxylic suffrutex forming mats up to 10 m in diameter (occasionally
shrublike, to 2 m tall); fresh mature fruit yellowish; central Brazil and
Paraguay, in cerrado. 10. A. humilis.
22. Shrub or tree to 40 m; fresh mature fruit green; Amazonia, the Guianas
eastern Brazil.
23. Secondary veins plane to slightly raised abaxially, tertiary veins plane;
flowers 10-13 mm long; eastern Brazil. 17. A. n?tida.
23. Secondary veins and tertiary veins prominently raised abaxially; flowers
13-18 mm long; Amazonia and the Guianas. 12. A. surinamensis.
20. Dry fruits smooth (the surface sometimes rough and warty, but not wrinkled) (Fig.
13).
24. Flowers 7-8.5 mm long; Cuba. 4. A. cubensis.
24. Flowers 9-23 mm long; Neotropics, Caribbean (excluding Cuba), and Africa.
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.
2. A. jaliscensis.
26. Flowers 13-23 mm long; upper and lower surfaces and sides of ovary
pubescent; leaflets abaxially hairy but the epidermis always visible;
Brazil.
27. Flowers 18-23 mm long; fresh fruit very dark brown with pale
greenish white mesocarp; tree to 30 m, with a small crown in open
situations. 15. A. ormosioides.
27. Flowers 13-17 mm long; fresh fruit green with green mesocarp;
shrub or small tree to 12 m, with a broad spreading crown in open
situations. 16. A. fraxinifolia.
25. Leaflets glabrous or abaxially with appressed indumentum.
28. Leaflets glabrous abaxially. 1. A. inermis.
28. Leaflets with appressed hairs abaxially.
29. Calyx with sparse appressed indumentum; leaflets 2-4 (-5) pairs;
leaflet base obtuse, or rounded to slightly cordate; flowers 10-13
mm long; ovules (1-) 2-4. 17. A. n?tida.
29. Calyx glabrous except around margins of lobes; leaflets (1-) 2 (-3)
pairs; leaflet base acute to obtuse; flowers 9-11 mm; ovules 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

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 43
brown, 5-10 mm thick; twigs sparsely hairy when young, the hairs red-brown, erect,
glabrescent. Stipules to 17 mm long, to 1 mm 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
liptic, narrowly obovate to oblanceolate, subcoriaceous to thick-chartaceous, base obtuse
(rarely acute), apex obtuse to rounded with an acumen to 15 mm long, glabrous (occa
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
nal, occasionally axillary, 10-40 cm long, hairy to sparsely hairy at branch tips, becom
ing less hairy towards the base, hairs red-brown, ? erect; bracts 2.5-3 mm long, with red
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 in A. in
ermis varies continuously from 9-19 mm, and alone forms no basis for the distinction of
intraspecific taxa; however, the Mexican 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 in Andira 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 in Mexico.
Wright (1777) first described this species as Geojfroea inermis jamaicensis. Subse
quently (Wright 1787), he reported the medicinal uses of the tree and used the binomial

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
Geoffraea inermis. Though he provided no botanical description in 1787, the name G. in

ermis is validly published, because Wright cited his previous work (1777).
Key to the Subspecies of Andira inermis

1. Calyx covered with appressed indumentum. la. Andira inermis subsp. inermis.
1. Calyx glabrous except at the base and margins of the lobes.
2. Flowers 10-11 mm long; calyx 2.5-3 mm long; Mexico. lb. Andira inermis subsp. glabricalyx.
2. Flowers 13-14 mm long, calyx 4-5 mm long; Africa. lc. Andira inermis subsp. rooseveltii.
la. Andira inermis subsp. inermis.

Andira excelsa Kunth in H. B. K., Nov. gen. sp. 6: 385.1824.?TYPE: MEXICO. Guer
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
capoua sapindoides (DC) Kuntze, Revis, gen. pi. 1: 212. 1891.?Type: Bertero
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
Polhill, Kew Bull. 23: 490. 1969.?Type: Gambia. Albreda, Perrottet 31 (holo
type: P, not located; isotype: BM!).
Glycyrhiza undulata Ruiz & Pav?n ex G. Don, Gen. syst. 2: 227. 1832.?Type:
Peru. Ruiz & Pav?n s.n. (holotype: BM!).
Andira acuminata Bentham, Comm. legum. gen. 45. 1837.?Type: Brazil. "In
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!).
Lonchocarpusl staudtii Harms in Engler, Bot. Jahrb. Syst. 26: 301. 1899.?Type:
Cameroon. Kumba [=Johann-Albrechtsh?he], Staudt 912 (holotype: B, de
stroyed; isotype: BM!).
Andira chiricana Pittier, Contr. U.S. Nati. Herb. 18(6): 235. 1917.?TYPE: PANAMA.
Chiriqu?: David, 30 Mar 1911, H. Pittier 3372 (holotype: US!; isotypes: GH!
NY!).
Flowers (9-) 10-12 (-19) mm long. Calyx covered with appressed indumentum.
Fig. 2A.
Phenology. Dry vegetation in Central America and the Caribbean coast of South
America: flowering concentrated February-May; other records scattered throughout the
year. Caribbean: flowering concentrated May-September, but scattered through the year.
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

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 45
110 105 100 95 90 85 80 75 70 65 60 55 50 45 40 35
Rico, Virgin Islands, Antigua, Guadeloupe, Dominica,

Grenada, Trinidad, Tobago); Mexico and throughout
bia, Ecuador, Peru, Bolivia, the Guianas, Paraguay, a
Africa (Gambia, Senegal, Ivory Coast, Nigeria, Camer
ferent parts of its range, A. inermis subsp. inermis
occurs in wooded savanna and rain forest. In Centra
northern South America it is common in seasonal
where in the Neotropics it grows in rain- and gallery
even mangrove.
Vernacular names. Black plum (Tobago); l'an
tinique, Guadeloupe); palo maco (Dominican Repu
(Jamaica); tololote, quiringuacua, cuautololote, c
bage bark, barley wood, cirvelillo, frijolillo
Guatemala, Honduras, Nicaragua); cabbage bark, q
dividivi, majagua gallina, palmillo, ciruelo de pl
pil?n, lumbricero, peloto, mot?n (Venezuela); ba
pisho, sacha ushuu (Peru); koraro, bat seed (Guya
(Suriname); lebi kiabici, St. Martin rouge (Frenc
(Brazil); mendubira (Argentina).

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
FIG. 15. Distribution of Andira inermis subsp. inermis and A. inermis subsp. rooseveltii in Africa.
Representative Specimens. Jamaica. Manchester: Mandeville, Donnell Smith 2389 (US). Haiti.
Nord-Ouest: Massif du Nord, Port-de-Paix, Petit Ford, Ekman 4326 (NY, US). Dominican Republic. Santo
Domingo: vie. Ciudad Trajillo, Allard 13367 (US).?SAMAN?: Finca Hacienda Nydia, 3.5 km E of village of
Las Terrenas, Zanoni & Mejia 17748 (NY).?DAJAB?N: Loma de Cabrera, banks of R?o de Masacre, Zanoni &
Mej?a 17899 (NY).?S?NCHEZ RAM?REZ: 7 km from center of Pimentel on road to Cotui, Zanoni et al. 16093
(NY). Puerto Rico. Ca. 2 km N Hwy 3, just E town of Rio Grande, Boom 6783 (GH, NY). Virgin Islands.
St. Croix: Belvedere estate, D'Arcy 4727 (GH, MO). Antigua. Wallings, Box 1385 (A, US). Guadeloupe.
Pointe Noire, Duss 3231 (NY, US). Dominica. Batalle, Ramage 221 (K). Martinique. Basse-Pointe, Sastre
7731 (GH). St. Lucia. Anse Louvet, Slane 803 (A). St. Vincent. Kingstown, Beard 1336 (F, GH, MO, NY,
US). Grenada. Balthazar, banks of Great River, Beard 157 (GH). Trinidad. Pitch Lake, vie. Brighton, Brit
ton 2906 (GH, NY, US). Tobago. Scarborough, Broadway s.n. (F, NY).
Mexico. CAMPECHE: Km 30 desviaci?n carretera Esc?rcega-Villahermosa, rumbo a Palizada, Chan & Lira
4707 (MEXU).?Chiapas: Mpio. Angel Albino Corzo, 500 m W de la salida de la colonia Quer?taro, sobre la
terraceria a Finca Prusia, Reyes Garc?a 1724 (MO).?Guerrero: Mpio. Chilpancingo de los Bravos, 4 km SE
de Tlahuizapa, camino El Ocotito-Jalenca, R?o Potrero, Mart?nez & T?llez 170 (MO).? MlCHOAC?N: El Hue
tamo, arroyo de Chapa, Soto N??ez 93 (MO).?Oaxaca: outskirts of San Pedro de Tepantepec, close to main
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
Para?so, Cowan & Maga?a 3046 (MO).
Belize. BELIZE: Belize-Sibun Road, Gentle 15 (F, NY, US).?Cayo: along Sibun River, just W of Hum
mingbird hwy, Croat 24861 (MO, US).?Stann Creek: Cockscomb Mountains, trib. of Cocoa Branch of Sit
ten River, 2 km N of Victoria Peak, Gentry 7887 (MO, US).?TOLEDO: hwy to Punta Gorda, 1 mi E of junction
with road to San Antonio, Croat 24471 (MO, US). Guatemala. Alta Verapaz: Rio Sebol, downstream from
Carrizal, Steyermark 44779 (US).?ESCUINTLA: Escuintla, Donnell Smith 2823 (GH, US).?IZABAL: Maricos,
bordering Lake Izabal, E. Contreras 7600 (MO).?JUTIARA: near Jutiapa, Standley 60583 (F).?PETEN: Laguna
Petexbat?m (Laguna M?xico) S of Sayaxch?, Steyermark 46202 (F, US).?San Marcos: Palmatto Flats, 1-2
mi N of Oc?s, Steyermark 37865 (F).?SANTA ROSA: Chiquilmulilla, Standley 79182 (F).?SUCHITEP?QUEZ:
vie. Tiquisate, Steyermark 47462 (F).?Zacapa: between R?o Honda and Santa Cruz, Standley 74130 (F).
Honduras. ATLANT?DA: vic. Tela, Standley 53570 (GH, US).?Choloteca: en la vega de Quebrada Tolobre,
matorrales y bre?ales entre Morolica y Tolobre, Standley 14219 (NY).?COL?N: Trujillo, R?o Negro, Clewell et
al. 4339 (MO).?COMAYAGUA: Pito Solo, Lake Yojoa, Edwards 429P (GH).?CORTES: Matorrales, 1 km N
Villa Nueva, Molina R. 6790 (GH, US).?ISLAS DE LA Bah?a: West End, Isla de Roat?n, Nelson & Romero 4519
(MEXU).?Olancho: Mpio. San Esteban, Santa Mar?a del Carb?n, 30 km al NE de San Esteban, Sousa et al.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 47
13369 (MEXU).?Valle: Bah?a de San Lorenzo, Molina R. 8633 (MO). El Salvador. Ahuachap?n: R?o
Paz, near Paso de Santa Cruz, Standley 20333 (US).?La Libertad: vie. La Libertad, Standley 23206 (US).?
La Uni?n: vic. La Uni?n, Standley 20861 (GH, US).?San Miguel: Laguna de Olomega, Standley 20991 (GH,
MO, NY).?San Salvador: Finca Altamira, hills S of San Salvador, Alien 7190 (US).?San Vicente: vic. San
Vicente, Standley 21277 (GH, US).?SONSONATE: vic. Sonsonate, Standley 21796 (GH, MO, US). Nicaragua.
Chinandega: vic. Chichigalpa, Standley 11189 (F).?LE?N: 3 km S El Sanee on banks of a small dry river
course, Hughes 452 (FHO, MEXU).?MANAGUA: camino a Salamina, hacienda Santa Cruz, a 6 km de la car
retera a Montelimar, Guzman & Castro 138 (MEXU).?RlVAS: R?o Lim?n, Lago da Granada, Shannon 5025
(US).?Zelaya: El Recreo, Long 229 (F). Costa Rica. ALAJUELA: Parque Alberto Manuel Brenes, San
Ram?n, Carvajal 263 (F, MO).?GUANACASTE: hwy Esparza-Ca?as at R?o Abangartos, Croat 61182 (MO).?
HEREDIA: Finca La Selva, OTS field station on Rio Puerto Viejo, Folsom 10047 (MO).?LLM?N: Rio Colorado
between Islas Buena Vista and Cerro Coronel, Davidse & Herrera 31242 (MO).?PUNTARENAS: Cant?n de Osa,
trail Palmar Norte-Ca?ablancal, R?o Terraba, Alien 5220 (F, MO, US).?San Jos?: 10 km NW Gu?piles, L. D.
G?mez 18517 (WO). Panama. CfflRlQU?: Progreso, Cooper & Slater 265 (F, GH, NY, US).?Cocl?: vic. El
Valle, Allen 1770 (GH, NY, US).?DARBEN: Rio Ucurganti, Bristan 1127 (NY, US).?HERRERA: Alto de Las
Minas, Carrasquilla 256 (F).?Panam?: R?o San Juan D?az, above Juan D?az, Alien 938 (GH).?SAN BLAS:
mainland opposite Play?n Chico, 0-3 mi from Caribbean, Gentry 6373 (F).?VERAGUAS: vic. Santa F?, Allen
4435 (GH).
Venezuela. DELTA Amacuro: Serra Imataca, Cerro La Paloma, E side R?o Cuyubini, S?eyermark 87604
(U).?DISTRITO FEDERAL: E of Caraballeda, Hacienda Juan D?az, Steyermark 62930 (F, MO).?FALC?N: Dist.
Colina, R?o Ricoa, Dos Bocas, Steyermark & A. Gonzales 113652 (MO).?Miranda: R?o Chico, Jahn 1239
(GH, NY, US).?Monagas: between La Pica and Ca?o Colorado, E Marur?n 6 km W La Ormega, Wurdack &
Monachino 39495 (NY).?T?CHIRA: Cerro Las Minas, S main road from Santa Ana, 17 km SE Santa Ana, Stey
ermark et al. (MO).?Zulla: Distrito Col?n, intersection of R?o Catatumbo and La Fria-Maracaibo road,
Davidse 18831 (MO). Guyana. West Bank Demerara River, Boom 7198 (NY, US); Pomeroon-Supernaam re
gion, Adventure, Gillespie & Persaud 1217 (MEXU, US). Suriname. Between mouths of rivers Coppename
and Coronie, Lanjouw & Lindeman 1460 (MO, U). French Guiana. Vicinity Cayenne, Broadway 611 (GH,
US). Colombia. Amazonas: P. N. Amacayac?, Cabana Pamat?, Narv?ez & Olmos 69 (COL).?Antioquia:
2 km from Barraquillita, Brand & Cogollo 95 (COL, JAUM, MO).?BOL?VAR: lands of Loba, San Mart?n de
Loba, Curran 23 (US).?Caldas: R?o Magdalena, cerca a La Dorada, confl. del R?o Purnio, Idrobo & R.
Jaramillo 2269 (COL, F, MO, NY, US).?CAQUET?: Florencia, Quebrada la Yuca, M. S?nchez & G. Mahecha
9a (UDBC).?CAUCA: Mpio. Guap?, P. N. de Isla Gorgona, camino a Pablo Sexto y Cabo de Hornos, Lozano et
al. 5245 (COL).?CHOC?: Llor?, 50 km S Quibd?, at june. R?o Atrato and R?o Andagueda, Archer 2050 (NY,
US).?C?rdoba: Planeta Rica, S. L?pez 4005 (COL).?Guajira: Sierra Nevada de Santa Marta, Sierra de San
Antonio, Bosque de La Cueva, Cuadros & Gentry 2965 (MO, NY).?MAGDALENA: valle inferior del Mag
dalena, Dugand 841 (F, US).?Meta: R?o Umea, Gavia 5122 (US).?Narl?O: Tamaco, R?o Rosario, 6 kms ar
riba de la desembocadura del Caunap?, Romero 5218 (MO, NY).?Santa Marta: near Bonda, H. H. Smith 18
(F, GH, MO, NY, US).?Santander: Puerto Araujo, Renter?a e? al. 1887 (JAUM, MO).?Valle: Isla del
Guayabal en el desembocadura del R?o Cajambre, Cua?recasas 16224 (F).?Sucre: around Verrugas, Finca La
Aguada, Bernai 158 (COL). Ecuador. Esmeraldas: San Lorenzo, 0.5 km S on walk to R?o Nadadero, Little
Jr. 6341 (F, US).??apo: La Joya de los Sachos Cant?n, Pompeya, R?o Indillana, between its mouth at the R?o
?apo and the MAXUS road, Gudi?o et al. 2125 (QCNE). Peru. AMAZONAS: Quebrada Huampami, Kayap
1061 (F, MO).?Cuzco: Pilcopta, Atalaya, Paucartambo, N??ez 6851 (MO).?Hu?NUCO: Prov. Pachitea, Dtto.
Honoria, Bosque Nacional de Iparia a lo largo de R?o Pachitea, 1 km arriba del pueblo de Tournavista, Schunke
1958 (F, GH, NY, US).?LORETO: Quebrada Cuninico, Croat 17759 (MO).?MADRE DE DIOS: Prov. Manu,
Cerro de Pantiacolla, R?o Palotoa, 10-15 km NNW Shintuya, Foster et al. 11027 (F).?PASCO: Prov. Oxapampa,
R?o Pichis, 1 hour below Puerto Bermudez, between Puerto Bermudez and Paujul, Gentry et al. 42145 (F,
MO).?SAN Mart?n: Prov. Lamas, Distr. Lamas, 1-2 km E Tabalosos, Belshaw 3370 (F, GH, MO, NY, US).?
Ucayali: Prov. Coronel Portal, Yarinacocha, Gentry & Horna 29375 (MO). Bolivia. BENI: Mamore, Tyson &
Kuns 1003 (MO).?PANDO: Prov. N Su?rez, road Cobija-Porvenir Km 2, Dom?nguez & E. Gonzales 57 (NY).?
Santa 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?: Igarap? do lago do Marac?, Mazag?o, Rabelo et al. 2238 (MO,
NY).?AMAZONAS: Mpio. Mara?, Rio Japur?, afl. do Rio Solim?es, Cid Ferreira 3380 (F, MO).?Goi?S: Serra
de Caldas Novas, Parque da Pousada do Rio Quente, margem do Rio Quente, Heringer 12228 (MO, UB).?
Maranh?o: Maracassum? River Region, Una do Trauira, Froes 1836 (GH, NY).?Mato GROSSO: Mpio.
Coxim, Rio Taquar?, Anderson 11292 (US).?Mato GROSSO DO SUL: Pantanal do Rio Negro, Fazenda Salina,
Dubs 475 (NY).?PARA: lina de Arana, Mpio. Santarem, Curral Grande, Black & Ledoux 50-10366 (GH).?

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
PARAN?: Mpio. Icaraima, Rio Paran?, Barra Rio Ivai, Hatschbach & Hass 15795 (SPF).?ROND?NIA: Forte
Principe da Beira, local Concei?ao, W. Rodrigues & Wilson 4273 (US). Paraguay. MISIONES: Ayolas, Schinini
& Vanni 25958 (GH, SPF). Argentina. Corrientes: Depto. Ituzaing?, Isla Apip? Grande, Puerto San Anto
nio, Krapovickas et al 24452 (MO).
Mali. K?m?bra, Roberty 17062 (K). Senegal. Koudougou (Saraya), Nongonierma 449 (K). Gambia.
Galam, Heudelot 380 (K). Ivory Coast. Near Kpakobo, Ak? Assi s.n. (K). Nigeria. Prov. Calabar, Dist. Itu,
Atau Eki Beach, Iyizoba s.n. (K-2 sheets). Cameroon. Mayaen, 25 km NNW Douala, Letouzey 14743 (K);
Victoria, near Victoria, Maitland 1064 (K); Marienberg, Polhill et al. 5214 (K); Douala-Edea Reserve, bank of
Sanaga River, D. W. Thomas 313 (K). Equatorial Guinea. Bioco (Fernando P?), Malabo-Rebola, Km 5, J.
Fern?ndez et al. 2823 (K).
Bentham (1860) recognized A. sapindoides based on two specimens from Antigua

and Dominica. These and other collections from the Lesser Antilles (Antigua, Dominica,
Guadeloupe, Martinique, Monserrat, and St. Lucia) have much larger flowers, up to 19
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
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
specific recognition. Subspecies inermis grows in rain forest around the Gulf of Guinea in
Cameroon and Nigeria, and has flowers of 12-13 mm long; subspecies grandiflora, from
savannas in Gambia, Senegal, Mali, Central African Republic, and Ivory Coast, has larger
flowers (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.
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 in Venezuela
and Costa Rica. The bark may be used as a vermifuge but is poisonous in high doses
(Weaver 1989). It is also taken for intermittent fevers in Mexico and Brazil (Morton 1981).
lb. Andira inermis subsp. glabricalyx R. T. Pennington, subsp. nov.?Type: Mexico.

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

floribus minoribus (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.

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2003 ANDIRA 49
Phenology. Flowering May and June, fruiting August and September, with single
fruiting records in December and March.
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
the calyx on the fruiting specimens Rose 1639,1782 are glabrous.
Additional Specimens Examined. Mexico. Nayartt: Mpio. Nayar, between La Nanchflera and Arroyo
de las Ventanas, Ben??ez-Paredes 3658 (MEXU); Arroyo Brasil, Embalse P. H. Aguamilpa, ca. 10 km E la
cortina, Calzada et al. 18688 (MEXU); between Zopilote and Santa Cruz de Guaybel, near junction for San
Miguel del Zapote, Ch?zaro et al 5474 (MEXU); Mpio. Nayar, 19 km NE San Pedro Ixcat?n, 2 km NE El
Naranjo, terracer?a San Pedro Ixcat?n-Santa Cruz Guaybal, G. Flores e? al. 1841 (MEXU); along dirt road be
tween Pochitit?n and Francisco Madero, Miller & T?llez 3181 (MEXU, MO); Mpio. Tepic, 4.3 km E junction to
la Escondida, old road Tepic-Mazatl?n, R. Ramirez & G. Flores 674 (MEXU); Mpio. Tepic, 5-7 km W.
Pochotit?n, 35 km W Tepic, T?llez 9076 (MEXU); 4 km NE Pochotit?n, road to Mojarritas, T?llez & Miller
10478 (MEXU); Mpio. Tepic, Paso de Bueyes, 40 km SE Tepic, Tenorio e? al. 16869 (MEXU).?Slnaloa: be
tween Rosario and Colomas, Rose 1639 (US); near Colomas, Rose 1782 (US).
It is tempting to speculate that morphological divergence has occurred in isolated

populations at the northern edge of the Neotropical distribution of A. inermis subsp. iner
mis, producing both A. inermis subsp. glabricalyx and A. jaliscensis. The latter occurs in
Jalisco and Michoac?n, separating the ranges of A. inermis subsp. inermis, and A. inermis
subsp. glabricalyx. More accurate information on 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 inermis subsp. rooseveltii (de Wildeman) Gillett ex Polhill, Kew Bull. 23:
490.1969. Millettia rooseveltii de Wildeman, PL Bequaert. 3: 353.1925.?Type:
Sudan. Equatoria Province: Nimule-Gondokoro, Mearns 3004 (holotype: BR;
isotype: BM!).
Ostryoderris brownii Hoyle, Kew Bull. 1932: 262. 1932.?Type: Ghana. Ashanti
Region: Jema, Feb 1931, W. T. Brown 2163 (holotype: K!).
Flowers 13-14 mm long. Calyx 4-5 mm long, glabrous except at base and margins
of lobes.
Phenology. Flowering February and March, with one record in May, fruiting
April-June.
Distribution (Fig. 15). Nigeria, Chad, Central Africa Republic, Togo, Uganda, Sudan;
in wooded savanna.
Vernacular names. Gwaska (Nigeria); Weri-dee (Central African Republic).
Additional Specimens Examined. Togo. Without locality, Kersting 554 (K). Nigeria. Gangoro Forest
Reserve, Chapman 4203 (K); northern Nigeria, Yola, Dalziel 168 (K); Bauchi, Lely P192 (K); Sokoto, Uly 832
(K); Abakaliki, Opara 609 (K); Plateau Prov., Selfwe River, Thornewill 54 (K). Chad. N of Nd?ll?, between
Golo and Mansaca, Chevalier 7773 (K). Sudan. Yirol District, around Koudogoi, F. W. Andrews 464 (K);
Imatong Mts, Kinyeti Valley, between Imeila and Hiliu, ca. 5 km S Hiliu, Friis & Vbllesen 1042 (K); without lo
cality, Schweinfurth 1875 (K). Central Africa Republic. Near Bonasse lu, 20 kms from center, Fay s.n. (K);

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2 km S Camp Koumbala along Koumbala river, Fay 4321 (K); Bouar, Mildbraed 9422 (BM, K). Uganda. W
Madi, Leya and Aiyu, river junction, Greenway & Eggeling 7252 (K).
Andira inermis subsp. rooseveltii is more 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 babies with a decoction (herb, label, Central African Republic).
2. Andira jaliscensis R. T. Pennington, sp. nov.?Type: Mexico. Jalisco: Mpio. Talpa de

Allende, camino a la mina del Cuale, 7.6 km E de la carretera Puerto Val
larta-Barra de Navidad, 30 May 1985, E. J. Lott 2544 (holotype: MEXU!; iso
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 1 mm wide, caducous, densely
hairy, hairs white, 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 {-A) cm wide, narrowly obovate or elliptic, thick-chartaceous to
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 1 mm 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 10 mm 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

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 51
when dry, green, drying brown, appearing smooth but minutely wrinkled (best seen with
lens or microscope); suture slightly raised adaxially with a narrow groove, indistinct abax
ially; stylar remnant tiny; composition of mesocarp and endocarp unknown. Chromosome
number unknown. Figs. 3E, 4(iv), 5A, 16.
Phenology. Flowering May to June with occasional records in January and March.
Distribution (Fig. 17). Mexico (Jalisco, Michoac?n); in deciduous or semi-deciduous
tropical forest, often by seasonal water courses.
Vernacular names. Manzanita (Jalisco); garrapata (Michoac?n).
Additional Specimens Examined. Mexico. Jalisco: Arroyo Chamela, Bullock 910 (MEXU); Mpio. La
Huerta, Est. de Biolog?a Chamela, Arroyo Chamela, Bullock 1313 (MEXU, MO); Estaci?n Biol?gica Chamela,
A. Delgado et al. 480 (MEXU); Mpio. La Huerta, 7 km al N del Puente Chamela, camino a Nacastillo, pasando
Arroyo Colorado, por los terrenos de la U. de G., G. Flores-Franco et al. 4795 (MEXU); Mpio. La Huerta, Ma
gallanes 421 (MEXU); Mpio. La Huerta, camino al E de Nacastillo, en el Lindero de la Universidad de Guadala
jara, Magallanes 585 (MEXU); Arroyo Chamela, Magallanes 1008 (MEXU); 3 km W de Macuautitl?n, camino
Tomatl?n-Talpa de Allende, Magallanes 1044 (NY); Mpio. La Huerta, Chamela, camino a Nacastillo, Magal
lanes 1383 (MEXU); Mpio. La Pe?a, camino Santiago Chacala-Colima, 14 km N Santiago, Magallanes 1669
(F); Mpio. Tomatl?n, Las Jarillas, carretera Puerto Vallarta-Barra de Navidad, Magallanes 1710 (MEXU);
Mpio. La Huerta, Est. de Biolog?a Chamela, Magallanes 2273 (US); Mpio. El Tuito, Las Gu?cimas, camino El
Tuito-Chacala, Magallanes 3556 (MEXU); Mpio. La Huerta, camino a Nacastillo, Magallanes 4197 (MEXU);
Mpio. La Huerta, Arroyo Chamela, Magallanes 4404 (NY); Chamela, L. A. P?rez 182 (MEXU); Chamela, L. A.
P?rez & M. Hern?ndez 848 (MEXU, NY); Mpio. El Tuito, 14 km N road El Tuito-Puerto Vallar?a in direction
of El Cuale, Solis-Magallanes 2148 (MEXU); Mpio. Cuautitl?n, 3 km N Teqesquitl?n, Solis-Magallanes 2950
(MEXU).?MICHOAC?N: Mpio. Aquila, El Tenamaste, 4 km de Aquila, Guerrero 766 (MEXU); Mpio. Aquila,
Los Tanamastes, Guerrero et al. 1262 (MEXU); Mpio. Chinicuila, 10 km NW Villa Victoria, Soto N??ez et al.
8187 (MEXU); Mpio. Tiquicheo, Paraje la Escondida el Lim?n, Verduzco s.n. (MEXU).
Andira jaliscensis is clearly distinct from A. inermis in its abaxially densely hairy
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).
All the collections of A. jaliscensis from Michoac?n come from the southwestern part of
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
patric in Michoac?n, and future fieldwork should concentrate here to see whether any
intermediate forms exist.
3. Andira multistipula Ducke, Bol. T?cn. Inst. Agron. N. 2: 30. 1944.?Type: Brazil.
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,
oxidizing darker, 4 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

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
FIG. 16. Andira jaliscensis. A. Habit B. Abaxial leaflet surface. C. Flower. D. Calyx, opened to show inner
surface. E. Standard petal, inner surface. F. Wing petal, outer surface. G. Keel petal, inner surface. H. Androe
cium. I. Gynoecium, also shown above in longitudinal section. J. Fruit. (Based on: A-I, E. J. Lott 2544; J, J. A.
S. Magallanes 4404.)

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 53
110 105 100 95 90 85 80 75 70 65 60 55 50 45 40 35
FIG. 17. Distribution of Andira jaliscensis, A. multistipul
narrowly ovate or lanceolate), chartaceous to thick-char

often slightly decurrent, apex acute to obtuse, with an
(one specimen seen with very sparse erect hairs abaxial
ially; secondary veins 10-14, plane adaxially, ? plane to
brochidodromous, tertiary veins plane adaxially and aba
illary (generally a large, terminal, ? conical inflorescen
florescences at the base, forming a single "attractiv
sparsely hairy, hairs red-brown, ? appressed, glabresce
with red-brown appressed hairs; pedicels to 0.5 mm lon
1.5-2 mm long, with red-brown appressed hairs. Flowe
yellow with red wine-colored lobes, 4-5 mm long, with
to glabrous except at the base and margins of the lobes
tuse. Petals pink to deep red-purple, the standard with
marking; standard blade 9-9.5 mm wide, 8-9 mm high,
long, 3.2-3.5 mm, claw 3 mm long, lamellate sculpturin
keel 8-8.5 mm long, 3.5-4 mm wide, claw 3.5-4 mm
ments white-pink, united for the basal 4.5-8.5 mm, fre
illary stamen 8.5 mm long. Gynoecium 11-12 mm long
sparsely hairy, the hairs red-brown, ? erect, occasionally
stipe 4-4.5 mm long, ovary reddish, 3.5-4 mm long, sty

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2.5-3.5 cm long, 2-2.5 cm high, elongated, weighing ca. 20 g or less when dry, green, dry
ing very dark brown to black, slightly wrinkled; stylar remnant barely discernible; meso
carp 1.5-2 mm thick, greyish brown, hard, granular; endocarp 1-1.5 mm thick, brown,
woody, fibrous. Chromosome number unknown.
Phenology. Flowering records throughout the year (March, April, June, October to
December).
Distribution (Fig. 17). Brazil (Amazonas, Acre), Colombia (Amazonas), Ecuador
(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
forest.
Vernacular names. Pisho, barbasco caspi, arco caspi, arco sacha (Peru).
Additional Specimens Examined. Colombia. Amazonas: Bocas del R?o Yari en el R?o Caquet?, Pab?n
& Mahecha 482 (COL). Ecuador. ?apo: La Joya de los Sachos Cant?n, Pompeya, R?o Indillama, between its
mouth at the Rio Napo and the MAXUS road, Guai?o et al. 2145 (QCNE); Parque Nacional Yasun?, MAXUS
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
Laguna Grande, Balslev e? al. 97505 (QCA, QCNE). Peru. Amazonas: Prov. Bagua, Dist. Imaza, R?o Cenepa
region, Comunidad Yamayakat, Hodges & Gorham 162 (E, NY); 1 km debajo de La Poza, bande del R?o San
tiago, Huashika? 49 (MO); R?o Santiago, 2 km atr?s de la Comunidad Caterpiza, Huashikai 925 (MO); Prov.
Bagua, Dist. Imaza, Regi?n del Mara??n, Comunidad de Yamayakat, R?o Mara??n, Jaramillo 575 (E, MO).?
HU?NUCO: Jacintillo, near Tingo Mar?a, Schunke 5725 (F, MO, US).?LORETO: Prov. Maynas, Santa Rosa, R?o
Itaya, Ayala et al. 3102 (MO); Quebrada Cuninico, Croat 17753 (MO); Quebrada Yanayac?, Croat 20320 (F,
GH, MEXU, MO, US); Prov. Coronel Portillo, Carretera Federico Basadre Km 99, Arboretum Von Humboldt,
C. D?az et al. 670 (F, MO, NY); Prov. Alto Amazonas, orillas del R?o Pastaza, entre Rimachi y R?o Witoyacu,
C. D?az 1315 (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
uitos, V?squez 1142 (F, MO).?MADRE DE Dios: Prov. Manu, Parque Nacional Manu, Zona Reservada R?o
Manu, Cocha Otoronga, Foster & Vivar 13259 (F, K).?San Mart?n: Prov. Mariscal Caceres, Dtto. Tocache '
Nuevo, camino a Shunte, Schunke 3859 (COL, GH, NY). Bolivia. PANDO: Prov. Manuripi, along R?o Madre
de Dios, 80 km NE (by air) downstream from Chiba, Nee 31530 (MO). Brazil. Acre: Mpio. Sena Madureira,
Km 1-3 on road Sena Madureira to Rio Branco, Prance e? al. 7968 (COL, GH).?AMAZONAS: Rio Javar?, Pt. 2
do RADAM, Cavalcante 3164 (RB, ?B); Mpio. Nova Japura, between Tamandar? and Manquari, Rio Japura,
Cid Ferreira & Lima 3611 (K, NY); Mpio. Limoeiro, Est. Ecol?gica do Juami-Japura, Cid Ferreira et al. 7250
(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
2731 (GH, K, NY); Alto Solim?es, Mpio. S?o Paulo de Oliven?a, S?o Luis do Pass?, H. C. de Lima et al. 2767
(GH, K, NY); Rio Juru?, Pena 608 (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 of intermediates argue against
splitting A. multistipula into two taxa.
Two specimens of the type collection (Ducke 1035) were found at RB. The larger
specimen with more stipules was chosen as the lectotype.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 55
4. Andira cubensis Bentham, J. Proc. Linn. Soc., Bot. 4, Suppl. ("A Synopsis of the Dal
bergieae"): 121. 1860. Vouacapoua cubensis (Bentham) Kuntze, Revis, gen. pl.
1: 212. 1891.?Type: Cuba. Oriente: Tinal de Nimanima, Apr 1844, J. Linden
2160 (holotype: K!; isotypes: G-2 sheets! K-2 sheets! NY-2 sheets!).
Andira microcarpa 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!).
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,
glabrescent, hairs brown, ? appressed to ? erect. Stipules to 3 mm long, to 1 mm wide, very
early caducous (only seen at shoot apices); leaf axis 6-22 (-32 cm long on sterile branches);
rhachis sparsely hairy, glabrescent, the hairs ? erect, brown, becoming or drying pale;
stipels minute, very early caducous; petiolules 1.5-5 mm long, sparsely hairy, glabrescent,
hairs ? erect, brown, becoming or drying pale; leaflets in 3-4 (-5) pairs, 5-13 cm long,
2.3-5.5 cm wide, narrowly ovate, elliptic (rarely obovate or broadly obovate), coriaceous,
base obtuse, rounded to ? cordate, apex obtuse and often slightly acuminate or rounded and
often retuse (apex of the terminal leaflet tends to be more 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
adaxially, ? plane to slightly raised abaxially, pattern brochidodromous, tertiary veins plane
adaxially and plane to slightly raised abaxially. Panicles axillary or terminal, 8-30 cm long,
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 1 mm 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
mm 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
the distal 2-3.5 mm. Gynoecium 5.5-8.5 mm long, upper and lower surfaces of the ovary
and lower half of the style hairy (ovary sides glabrous), hairs brown, ? appressed; stipe
1.5-2.5 mm long, ovary 2-3 mm long, style 1.5-3 mm long; ovules 1. Fruit 2.5^4 cm long,
2-2.8 cm high, 2.1-2.9 cm wide, 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
single, possibly immature fruit). Chromosome number: n = 11.
Phenology. Flowering April to August, concentrated in June, July.
Distribution (Fig. 17). Cuba; forest, swamp, river bank, wood pasture, savanna and
pine savanna; to ca. 1000 m.
Vernacular name. Yaba.
Additional Specimens Examined. Cuba. Habana: Isla de Pinos, Los Indios, N. L. Britton et al. 14246
(NY); near Puentes Grandes, Bro. Le?n 3347 (NY); 2 km N Nueva Gerona, Isla de Pinos, Morton 9983 (US);
Cerro Nueva Gerona, Isla de Pinos, Morton 10121,10155 (US); Isla de Pinos, Herradura Beach, Morton 10304

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(US); near Nueva Gerona, Isla de Pinos, A. H. Curtiss 525 (G, MO, NY, PR).?Las Villas: Santa Clara towards
Manicaragua, N. L. Britton & Cowell 10259 (NY); Trinidad Mountains, 14 km NW Trinidad on road to Tope de
Collantes, Harvard Course in Trop. Botany 197 (GH); Cienfuegos, between Rancho Luna and central Soledad,
Hodge s.n. (US); W of Rio San Juan, crossing of road to Trinidad, Howard et al. 379 (NY); Belmonte, Soledad,
Cienfuegos, Jack 4999 (US); Limones, Jack 5390 (GH); Belmonte, Soledad, Cienfuegos, Jack 5664 (A).?
Matanzas: hills S Matanzas, N.L. Britton & Wilson 474 (NY).?Oriente: 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
4349 (NY).?Pinar DEL R?O: San Diego de los Ba?os, Bro. L?on 4628 (GH, NY); near Vinales, Bro. Alain 4404
(GH); road to San Vicente, 21 km N Pinar del Rio, Harvard Course in Trop. Botany 10 (GH, NY).
Bentham (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, a more densely hairy gynoecium, a
single ovule, and a fruit with a more pointed apex. It also tends to have only 3^4- (-5) pairs
of leaflets compared to 2-9 in A. inermis, which have a rounded to cordate base, whereas
those in A. inermis are obtuse. Additionally, the leaflets are abaxially hairy when young.
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.
5. Andira taurotesticulata R. T. Pennington, sp. nov.?Type: Colombia. Antioquia:

Mpio. Ca?as Gordas, Boquer?n de Toyo, 1 Jul 1984, G. Lozano, G. Galeano, R.
Londo?o, R. Bernai & D. Restrepo 3963 (holotype: COL!; isotype: MO!).
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 15 mm 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
becoming brochidodromous or ? completely 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

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2003 ANDIRA 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,
weighing 100-300 g when dry, pale brown or dark brown marked with pale brown (both
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?,
Cundinamarca, 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).
Additional Specimens Examined. Panama. Dari?n: Cuasi-Cana trail between Cerro Campamento and
La Escalera, E of Tres Bocas, Kirkbride & Duke 1315 (MO, NY).? San Blas: El Llano-Carti road, Km 12, in
vicinity of Gorgas Lao Mosquito Control Project, Croat 26024 (MEXU, MO).
Colombia. Antioquia: Mpio. San Rafael, 16 km E San Rafael along Guatap?-San Rafael road, Brant &
Roldan 1533 (E, MO); Mpio. San Carlos, Correg. Alto Samana, vereda Miraflores, finca "El Despesero," Calle
jas et al. 8584 (HUA, NY); Mpio. San Luis, right bank Rio Samana, vereda el Port?n, road to the autopista
Medell?n-Bogot?, C?rdenas 2501 (COL, JAUM); Mpio. San Luis, Autopista Medell?n-Bogot?, vereda la Jose
fina, quebrada la Mariola, C?rdenas & J. G. Ram?rez 2741, 2743 (JAUM); Mpio. San Luis, Autopista Medel
l?n-Bogot?, Sector R?o Samana-R?o Claro, camino hacia la vereda Tulip?n, Cogollo & Estrada 199A (JAUM,
MO); Mpio. San Luis, Autopista Medell?n-Bogot?, Sector R?o Samana-R?o Claro, camino hacia la vereda La
Josefina, Cogollo & Estrada 270 (JAUM, MO); Mpio. San Luis, carretera hacia Aquitania, 12 km from Au
topista Medell?n-Bogot?, Cogollo et al 3762 (JAUM); Mpio. San Luis, Autopista Medell?n-Bogot?, vereda la
Josefina, quebrada la Mariola, Cogollo et al. 4306 (JAUM); Parque Nacional Natural "Las Orqu?deas" Sector
Calles, right bank of R?o Calles, Cogollo 4145 (COL, JAUM); Mpio. San Luis, Autopista Medell?n-Bogot?,
vereda la Josefina, Hoyos & J. J. Hern?ndez 286 (JAUM, MO); Mpio. San Luis, Autopista Medell?n-Bogot?,
camino de la vereda la Josefina a la vereda la Holanda, Hoyos & J. J. Hern?ndez 334 (JAUM, MO); Mpio. San
Luis, Autopista Medell?n-Bogot?, vereda la Josefina, camino al Tulip?n, ca?o la Mariola, Hoyos & J.J. Hern?n
dez 917 (JAUM, MO); Frontino-El Cerro road, 11 km above Frontino around el Aeropuerto, Luteyn & Lebr?n
Luteyn 7196 (COL, MO, NY, US); Mpio. San Luis, Quebrada "La Cristalina," J. G. Ram?rez & L?pez 950 (HUA,
JAUM, MO), 1161 (COL, HUA, JAUM, MO); Mpio. Anori, Corregimiento de Providencia, valle del R?o Anor?,
entre Dos Bocas y Anor?, Buenos Aires, 4 km from Providencia, Soejarto 3954 (GH); Mpio. Ituango, Km 9 of
road to vereda La Hundida (WSW of Ituango), Zarucchi & Betancur 6444 (HUA, MEXU, MO).?Boyac?:
near P?ez, Espinal & Montenegro 1698 (COL).?CUNDINAMARCA: Cerro Quinini, Idrobo 5337 (COL).?Meta:
Restrepo, vereda El Caney, Ca?o Agua Dulce, Forero et al. 10205 (COL).?QuiNDlO: road from Pijao to
Paramo de Chili, Gentry et al. 65327 (COL, E, MO).?SANTANDER: Mpio. Charal?, Corregimiento de Virol?n,
Vereda El Reloj, S. D?az 1279 (COL).?VALLE: Sevilla, Hacienda La Esmeralda, frente al vivero de la secretar?a
de Agricultura y Fomento, Cuadros 643 (US); Mpio. Sevilla, via a Morroazul, quebrada La Raquelita, Devia 941
(MO). Venezuela. T?CHIRA: Parque Cazadero, Quebrada Cazadero, 16 km NW San Crist?bal, Liesner &

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FIG. 18. Andira taurotesticulata. A. Habit. B. Broad leaflet. C. Narrow leaflet. D. Abaxial leaflet surface.
E. Flower. F. Calyx, opened to show inner surface. G. Standard petal, inner surface. H. Wing petal, outer sur
face. I. Keel petal, inner surface. J. Androecium. K. Gynoecium, also shown above in longitudinal section. L.
Fruit. M. Fruit in longitudinal section, showing wall structure. (Based on: A (leaf), W. Devia 941; A (inflores
cence), E-K, G. Lozano et al. 3963; B, D, S. Hoyos et al. 334; C, H. Cuadros 643; L-M, W. Devia 941.)

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2003 ANDIRA 59
A. taurotesticulata
FIG. 19. Distribution o? Andira taurotesticulata.
Guariglia 11706 (MO); Cerro Las Minas, S of main road from Santa Ana, 17 km SE Santa Ana, Steyermark et
al 119886 (MO); Cerro Las Minas, slopes leading to Cerro Azul, 18 km SE Santa Ana, Steyermark 120019
(MO); Distr. Lobatera, La Cazadora, van der Werff & Ortiz 5553f 5556, 5576 (E, MO). Ecuador. NAPO: Can
ton Orellana, Reserva El Chuncho, 1 km W Rio Payamino, NW Coca, Baker 7025 (NY); Cant?n Orellana, Via
de Zorros, Pozo de Jaguar I, Palacios 3204 (QCNE); Cant?n Orellana, Reserva El Chuncho, 1 km W R?o
Payamino, NW Coca, R. T. Pennington & M. Aulestia 524, 525 (E, K, QCA, QCNE).?PICHINCHA: Quito,
Reserva Juan Manuel Durini, Palacios 12274 (QCNE).?SUCUMBIOS: Lago Agrio, reserva Faun?stica
Cuyabeno, Laguna Grande, near Catholic University Field Station, Neill 10214 (QCNE).
Andira taurotesticulata is the only species that grows in montane forest (to 2200 m),
occurring in all three cordilleras of the Andes in Colombia, and extending along the
Cordillera Occidental to Venezuela. It is also found in the Caribbean coastal ranges in
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
morphologically similar; the only difference is an endocarp with more abundant stone
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

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
the other species are relatively smooth and lack the distinct ribs. Moreover, these other
species have restricted distributions in the Guianas (A. cori?cea) and central Brazil (A. mi
crantha), and are widely disjunct from A. taurotesticulata.
The name for this species comes from its local name in Colombia: "cojones de toro."
6. Andira macrothyrsa Ducke, Bol. T?cn. Inst. Agron. N. 2: 31. 1944.?Type: Brazil.
Amazonas: Esperan?a, 22 Oct 1942, A. Ducke 1036 (holotype: RB!; isotypes: K!
R?RB-2 sheets! US!).
Andira multistipula var. peruana N. F. Mattos, Loefgrenia 53: 2.1971.?TYPE: PERU.
R?o Huall?ga, 16 Feb 1924, J. G. Kuhlmann 1505 (holotype: RB!; isotypes: RB
2 sheets!).
Tree to 40 m tall, often a canopy emergent; buttresses slight; bark smooth and grey;
wood 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,
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
slightly raised adaxially, slightly raised abaxially, pattern eucamptodromous becoming
brochidodromous, the veins generally curving uniformly, tertiary veins plane adaxially
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 1 mm long or flowers subsessile; bracteoles 1 mm 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
acuminate. Petals white-yellow to greenish, the standard with a central red-brown to
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 long. Gynoecium 6.5-8 mm long, the ovary with dense red-brown appressed hairs,
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, Los R?os),
Peru (Loreto), Brazil (Acre, Amazonas); in terra firme and seasonally flooded forest and
also caatinga (low forest on white sand). Andira macrothyrsa is disjunct across the Andes;
all the collections from Colombia and some from Ecuador are from the Pacific coast.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 61
id - -
A. macrothyrsa
A a. chigorodensis
FIG. 20. Distribution of Andira macrothyrsa and A. chigorodensis.
Vernacular names. Uxi de morcego (Brazil, Acre); casub guesmoteig (Aw?; Ecua
Carchi).
Additional Specimens Examined. Colombia. Choc?: Parque Nacional de Utr?a, NE de la Ense?ada de

Utria, entre al Manglar y la falda de la monta?a alrededor de la quebrada la Chunga, F. Garcia C. & Agualimpia
443 (MO).?Valle: Bah?a de Malaga, near mouth of Quebrada la Sierpe, Gentry et al. 40434 (COL, JAUM,
MO). Ecuador. CARCHI: Tulcan Cant?n, Parroquia Tobar Donoso, Sector Sabalera, Reserva Ind?gena Aw?,
Tipaz et al. 1225 (E, MO, QCNE), 1345 (MEXU, QCNE).?Los R?OS: Quevedo Cant?n, Cerro Centinela, Mon
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 4818 (MO); Cant?n Loreto, Sector R?o Guataraco, ca. 1 km after R?o Guataraco
on road Holl?n-Loreto, left side road, R. T. Pennington & M. Aulestia 523 (E, K, QCA, QCNE).?SUCUMBIOS:
Reserva Faunistica Cuyabeno, 1 km N Laguna Grande, hectare plot 1, Valencia 67876 (QCA). Peru. Loreto:
Yanamono, Explorama tourist camp, halfway between Indiana and mouth of Napo, Gentry et al. 60683 (MO);
Yanamono, Explorama Tourist Camp, R?o Amazonas above mouth of Rio Napo, Gentry et al. 37168 (MEXU,
MO, NY); Iquitos, Quistacocha, T. D. Pennington & Ruiz 13550 (K); Prov. Requena, Sapuena, Bagaz?n-R?o
Ucayali, V?squez & Jaramillo 8745 (MO); Maynas Province, Iquitos, Allpahuayo, Estaci?n Experimental IIAP,
V?squez et al. 13802 (E, MO). Brazil. ACRE: Mpio. Rio Branco, estrada para Porto Acre Km 13, ramai a es
querda, 4 km da estrada Colonia Cinco Mil, Cid Ferreira & Nelson 3057 (NY); Mpio. M?ncio Lima, estrada
para o lugar Bar?o, Km 30-32, Cid Ferreira et al. 5247 (NY); Mpio. M?ncio Lima, estrada do Isac, 4 km from
city, Cid Ferreira et al. 10950 (MEXU, NY); Sub-base do Cruzeiro do Sol, Marinho 27 (RADAM), vie. Serra
da Moa, Prance et al. 12388 (US); Cruzeiro do Sol, pr?ximo ao Acampamento do Projeto RADAM, Rosa 668
(NY).?Amazonas: Alto Solim?es, Mpio. Sao Paulo de Oliven?a, estrada do Bon?im, logo ap?s a saida da
cidade de Sao Paulo de Oliven?a, H. C. de Lima et al. 2783 (K).

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
Andira macrothyrsa is morphologically most similar to A. chigorodensis (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 chigorodensis R. T Pennington, sp. no v.?Type: Colombia. Antioquia: 21

km de Apartado hacia Chigorod?, 10 Jul 1981, J. Brand & A. Cogollo 114 (holo
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,
narrowly elliptic to lanceolate, thick-chartaceous to subcoriaceous, base obtuse to
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
ally and plane to slightly raised abaxially. Panicles axillary (appearing terminal, but in the
axil of a terminal bud), 20-25 cm 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, 1 mm 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
mm, vexillary stamen 3.5 mm long. Gynoecium 4.7 mm long, ovary hairy, hairs ? ap
pressed, pale red-brown; stipe 1.5 mm 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.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA
FIG. 21. Andira chigorodensis. A. Habit. B. Abaxial leaflet surface. C. Flower. D. Calyx, opened to show
inner surface. E. Standard petal, inner surface. F. Wing petal, outer surface. G. Keel petal, inner surface.
H. Androecium. I. Gynoecium, also shown above in longitudinal section. J. Fruit. (Based on: A-I, J. Brand &
A. Cogollo 114; J, E. Renteria et al. 3579.)

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
Phenology. Flowering July (only one record); fruiting records from September, No
vember, January, February.
Distribution (Fig. 20). Colombia (Antioquia); in rain forest; to 300 m.
Additional Specimens Examined. Colombia. Antioquia: Turbo, 8 km de Carepa hacia Chigorod?,

Brand & Cogollo 215 (HUA, JAUM, MO); Mpio. Turbo, corregimiento de Lomas Aisladas, Renteria et al. 3579
(JAUM-2 sheets), MO); Mpio. Chigorod?, camino Malag?n-Chigorod?, Renteria 4634 (JAUM, MO); Mpio.
C?ceres, 13 km W of El Jardin, Finca El Amparo, Zarucchi & D. C?rdenas 4215 (COL, HUA, MO).
Andira chigorodensis most closely resembles A. macrothyrsa. It differs in its larger

leaves, 37-55 cm long, with narrowly elliptic or lanceolate leaflets, and smaller flowers,
5-6 (-7) mm long. The leaves of A. macrothyrsa are 10-35(-40) cm long and have nar
rowly ovate, elliptic, or narrowly obovate leaflets, and the flowers are 9-11 mm long.
Andira chigorodensis is endemic to the Chigorod? region of Antioquia, Colombia.
8. Andira 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 1 mm 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
dard blade 11-14 mm high, 9-12 mm wide, claw 4 mm; wing 8-10 mm long, 4^.5 mm
wide, claw 5 mm long, lamellate sculpturing present; keel 8-10.5 mm long, 4^.5 mm
wide, claw 5.5-6 mm 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
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.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 65
110 105 100 95 90 85 80 75 70 65 60 55 50 45 40 35
FIG. 22. Distribution o? Andira galeottiana and A.
Phenology. Flowering April to July.

Distribution (Fig. 22). Mexico (Veracruz, Oaxaca
banks, seasonally flooded zones, also in non-inundat
Vernacular names. Macayo/a (Veracruz, Oaxaca
Representative SPECIMENS EXAMINED. Mexico. Chiapas: T

s.n. (MEXU); Palenque, Fuentes s.n. (MEXU); 1 km SW juncti
c?rcega, Grether & Quero 1579 (MEXU); La Arena, Palenque
Campo Petrolero Cactus, Tenorio 19498 (MEXU).?OAXACA: L
chit?n, 6.6 km W Mago?? por camino a Guichieovi, H. M. Hern
Cnimalapa, Santa Maria, H. M. Hern?ndez 418 (MO); Mpio. Tuxt
MEXU, MO); Mpio. Tuxtepec, Ejido Benito Ju?rez, T. D. Pennin
Figueroa, Salazars.n. (MEXU, US); Dist. Choapain, San Juan La
tepec, Las Pinas, Sousa 3667 (MEXU, US); Ubero, L. Williams
F, U).?TABASCO: R?o Rosario, Huimanguillo Savanna A
Zapata-Tenonique, Ch?velas & P?rez J. ES892 (NY); Km 35 de
Rueda, C. Cowan et al 2294 (MO, NY); Balanc?n, Matuda 312
Vieja, cerca de Luis Gil P?rez, S Villahermosa, Novelo & Ram
La Venta, Laguna Yucateco, edge of lake, near Rio Chicozapo
junction to Tenosique on road to Balanc?n, T?llez & E. Mart?ne
Triunfo, on road to Balanc?n, T?llez & E. Mart?nez 948 (MEX
ago Jalahui, carretera a San Juan del R?o, Calzada 16846

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
de Ocampo, Cedillo & Calzada 17 (F); San Lorenzo Tenochtitl?n, Ch?velas et al. ES-2396 (MEXU); Mpio.
Playa Vicente, near Huaxpala, Ch?velas et al ES-4301 (MEXU); Mpio. Catemaco, Agaltepec Island, Lake
Catemaco, Faden et al. 76/125 (F, K); along E edge Lago Catemaco, 1.5 km N Tabanca, 10 km E Catemaco,
Hanson & Nee 7637 (F, MO); Catemaco, Laguna de Sontecomap?n (R?o Coscoapan), Men?ndez 121 (MO);
Barra de Sontecomap?n, Nevling & G?mez-Pompa 171 (GH, MEXU-2 sheets); 2 km Adelante Panga, R?o
Teschaacu, Nevling & G?mez-Pompa 840 (GH, MEXU, US); P. Est. Chocam?n, Chocam?n, ca. de C?rdoba,
Salazar s.n. (MEXU); Region de los Tuxtlas, R?o Tecesapa, al E de Seteapan, Sousa 3267 (MEXU); Mpio.
Catemaco, Lago Catemaco, a 2 km de Coyame sobre camino a Tebanca, orilla de Lago Catemaco, Sousa et al.
13407 (MEXU); Fortuno, Coatzacoalcos river, Williams 8483 (F, US); village Tenochtitl?n on path to orange
grove, Wing 25 (GH).
Despite their wide geographical separation, A. galeottiana and A. verm?fuga are

closely related; their plastomes differ by only one restriction site. Vegetatively and flo
rally, the two species are remarkably similar, but the large fruit of A. galeottiana, with its
soft and somewhat spongy mesocarp, is unique in the genus. These fruits, especially when
the mesocarp 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
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 and Martius, Reise Bras. 2: 788. 1828. Vouacapoua ver
m?fuga (Martius) Kuntze, Revis, gen. pi. 1: 212. 1891.?Type: Brazil. Minas
Gerais: "Inter fr?tices ad Salgado, et in campis editis Taboleiro," Martius s.n.
(holotype: M!).
Geoffroea 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
prov. Minarum campis editis siccis ad Contendas," Martius s.n. (holotype: M!).
Andira paniculata Bentham, Comm. legum. gen. 45. 1837. Vouacapoua paniculata
(Bentham) Kuntze, Revis, gen. pl. 1: 212. 1891.?Type: Brazil. "Barbacena et
Ponte d'Erva," Pohl s.n. (holotype: K!; isotypes: K! M, PR!, photos of M iso
type: G! GH!).
Andira kuhlmannii N. F. Mattos, Loefgrenia 40: 2. 1970.?TYPE: Brazil. S?o Paulo:
Moji-Mirim, margem da rodovia para Campinas, pr?ximo ? entrada da Fazenda
Holambra, M. Kuhlmann 3945 (holotype: HB; isotypes: K! SP).
Tree to 12 (-25) m tall with spreading crown, occasionally a shrub, able to root
sprout; buttresses very slight or absent; bark thick, rough with deep vertical fissures;
slash pale brown to red-brown to orange-brown, generally with red ex?date; wood
buff/cream; twigs often swollen, densely to sparsely hairy, glabrescent, the hairs ?

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 67
appressed, red-brown; bark cracking on older twigs; lenticels absent. Stipules to 5 mm

long, to 1 mm 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
biculate to narrowly obovate), subcoriaceous to coriaceous, dark green, shiny adaxially,
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
adaxially; secondary veins 5-11, slightly sunken adaxially, raised abaxially, pattern eu
camptodromous becoming brochidodromous (occasionally completely brochidodro
mous), tertiary veins plane adaxially and raised abaxially. Panicles terminal (more
rarely axillary), densely covered red-brown erect hairs at branch tips, glabrescent to
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, 1 mm 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
wide, 9-12 mm high, claw 3.5-6 mm long; wing 8-10 mm long, 3.5-6 mm wide, claw
4-6.5 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
red-brown bases; stipe 4-6.5 mm long, ovary 3-5.5 mm long, style 2.5-4.5 mm long;
ovules (2-) 4-6. Fruits 2.4-4 cm long, 1.2-2.5 cm high, 1.7-2.5 cm wide, elongated,
weighing ca. 20 g or less when dry, green, strongly scented, drying brown (more rarely
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.
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 Gerais,
Piau?, R?ndonia, Sao Paulo); in cerrado and gallery forest.
Vernacular names. Angelim preto, mata barata, angelim branco (Brazil).
Representative Specimens Examined. Peru. Pasco: Cerro de Pasco, central selva, Palcazu Valley,
Hartshorn et al. 2918 (MO). Bolivia. PANDO: Manuripi, 35 km N Puerto America, A. Jardim 1050 (K,
MO).?Santa Cruz: Prov. Chiquitos, Serran?a de Santiago, near Santiago de Chiquitos, 5 km ENE Santiago,
Daly et al. 6272 (NY); Prov. Chiquitos, Serran?a de Santiago, Santiago de Chiquitos, camino a la cueva, Ma
mani & A. Jardim 1291 (MEXU). Brazil. Acre: near mouth of Rio Macauhan (tributary of Rio Yaco),
Krukoff 5483 (A, F, U, US).?AMAZONAS: Mpio. Humait?, Janssen 139 (K); Transamaz?nica, 150 km SE Hu
mait?, 22 km E Bodoc?, Sanaiotti 287 (E, INPA).?Bah?a: 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);
Mpio. Barreiras, ca. 2 km along road to airport, R. T. Pennington & Brito 265 (CEPEC, FHO, K); Mpio. tt>oti
rama, rd. Ibotirama-Seabra (BR-242) Km 8, R. T. Pennington & Brito 270, 271, 272, 273 (CEPEC, FHO, K);
Mpio. Correntina, Fazenda Jatob?, Rezenda et al. 20 (?B).?Cear?: Serra de Ibiapaba, Campo Grande,
Dahlgren 880 (F).?DISTRITO FEDERAL: Bacia do Rio S?o Bartolomeu, Heringer et al. 5450 (US); Chapada
da Contagem, Irwin et al. 7954 (MO, US); Fazenda Agua Limpa, near Vargem Bonita, 18 km SSW Brasilia

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
TV tower, Ratter et al 3595 (E, F, K, UB).?Goi?s: Mpio. Vian?polis, Alvarenga et al 194 (UB); Chapada
dos Veadeiros, 11 km E Cavalcante, W. R. Anderson 7294 (NY); Chapada dos Veadeiros, Fazenda Breij?o,
Bridgewater et al. 199 (E, UB); Mpio. Goi?s, S?o Joaquim, Gibbs et al. 2713 (K, NY); Mpio. Goi?s, Serra
Dourada, Gibbs 2782 (K, NY, UB); near C?rrego Estrema, ca. 38 km NE Formosa, Irwin et al. 15191 (F,
MEXU, NY); Mpio. Presidente Kennedy, rd from BR-153-Itapor?, 12 km W of village of Presidente
Kennedy, Fazenda Primavera along Riber?o Fe?nho, Plowman e? al. 8298 (GH, NY, US).?MARANH?O:
arredores de Bar?o do Graja?, Riacho Grande, L. Duar?e & Castellanos 602 (NY); Mpio. Lor?to, between
R?os Balsas and Parna?ba, 35 km S Lor?to, Eiten & Eiten 4513 (NY); Mpio. Carolina, Transamazonian High
way, BR-230 and BR-010, Pedra Caida, 35 km N Carolina, Serra da Baleia, E.L. Taylor et al. E1248 (K, NY,
US).?Mato Grosso: Mpio. Santa Terezinha, Serra do Vi?va, 12 km da Cidade, Cid Ferreira et al. 6416
(NY, US); Brotas, 30 mi NE Cuiab?, Collenette 155 (K); Mpio. Barra do Gar?as, 152 km NNE Xavantina, 4.5
km S C?rrego Tangur?, Eiten & Ei?en 9849 (NY, US); watershed between Amazonas and Rio Paraguaya, 1-10
km W Alto Araguaia, Brasilia-Acre highway, Maguire 56241 (NY, UB); 10 km W S?o Felix, Richards 6482
(K, NY, UB); road MT-170, track west, 3 km N of junction with BR-364, Fazenda Itamarati, transect 2,
Sanaiotti 449, 460, 461 (E, INPA).?Minas G?rais: Porto Buriti, margem esquerda do Rio Paracatu, de F.
Almeida 140 (K); Patos de Minas, A. P. Duarte 3282 (NY); Felixlandia, Heringer 7120 (NY, UB); Cristalina,
Paracatu, barranco do Rodo via, Heringer 17740 (US); Mpio. Unai, margem da rod. BR-251, perto da entrada
da Fazenda Paulista, B. A. S. Pereira 904 (SP).?PlAU?: Alencar s.n. (RB).?ROND?NIA: Rd. Vilhena-Col
orado, 20 km from Vilhena, Zarucchi et al 2810 (GH, NY, US).?S?O PAULO: Araraquara, Usina Tamoio,
Fazenda Santa Joana, Felippe 94 (US); Mpio. Botucatu, 18 km N Botucatu, 14 km E de S?o Manuel, along
road S?o Manuel to Piracicaba, near Est. "13 Maio" of the old railroad, /. S. Gottsberger & C. J. Campos 12
221174 (UB); Mpio. Brotas, Ja?, Leit?o Filho et al. 12927 (US).
Andira verm?fuga is most closely related to A. galeottiana (no. 8; see notes under that
species for differential characters).
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.
These two forms overlap somewhat geographically, though the majority of the thin
twigged collections are from the northeast of the range of A. verm?fuga (Bahia and
Maranh?o). No differences can be found to separate these groups in leaf, flower, or fruit
characters, and there are some collections in which the twigs appear intermediate (e.g.,
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
slender and leptocaul, suggesting that the underlying basis for the differences may be
environmentally related, e.g., repeated burning may induce formation of thick twigs.
This is corroborated by personal observations of a population of A. verm?fuga (at
Fazenda Agua Limpa, Distrito Federal, Brazil), which showed a 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
sociation. Prance et al. 8719 (Rond?nia, Brazil), Krukoff 5843 (Acre, Brazil) and
Hartshorn et al. 2918 (Pasco, Peru) all represent large trees to 25 m tall, but with flow
ers 12.5-14 mm long, which is somewhat small for A. verm?fuga. They were collected
in seasonal "transition forest" (sensu Ratter, 1992) on the edge of Amazonia, extending
the range of A. verm?fuga into this ecosystem. Prado and Gibbs (1993) and Pennington
et al. (2000) give several examples of this type of distribution of a woody species, which

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 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
been collected in the isolated Amazonian savanna of Humait? (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 in the synonymy of A.
verm?fuga, which therefore takes priority.
10. Andira humilis Martius ex Bentham, Comm. legum. gen. 45. 1837. Vouacapoua hu
milis (Bentham) Kuntze, Revis, gen. pi. 1: 212. 1891.?Type: Brazil. Minas
Gerais: "ad Chapado do Par?nan," Martius s.n. (holotype: M!).
Andira laurifolia Bentham, Comm. legum. gen. 45. 1837. Vouacapoua laurifolia
(Bentham) Kuntze, Revis, gen. pl. 1: 212. 1891.?Type: Brazil. Minas Gerais:
Fa?ado, Pohl s.n. (holotype: K!; isotype: W, photo: G!).
Andira pauciflora Bentham, Comm. legum. gen. 45. 1837.?Type: Brazil. Minas
Gerais: "Ad Barbacena et Ponte d'Erva," Pohl s.n. (lectotype, here designated:
K!).
Andira humilis var. cordata N. F. Mattos, Loefgrenia 40: 3. 1970.?TYPE: Brazil.
S?o Paulo: Itahy, 10 Dec 1929, /. /. de Lima s.n. (holotype: RB!).
Geoxylic suffrutex forming mats up to 10 m in diameter, aerial shoots to 50 cm tall

(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
vate, oblanceolate, ovate to lanceolate), subcoriaceous to coriaceous, shiny, dark green
adaxially, paler and matt abaxially, the venation paler, base obtuse, rounded (occasionally
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,
2-2.8 cm high, 2-2.8 cm wide, elongated, weighing ca. 20 g or less when dry, green

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FIG. 23. Distribution of Andira humilis.
turning yellowish when ripe, sweet smelling, drying wrinkled, dark brown; stipe 5-10 mm
long, but many fruits falling off without a stipe; suture raised adaxially with a central
groove, less obvious below; stylar scar tiny; mesocarp 0.5-2 mm thick, pale yellow and
soft when ripe, fibrous with oily granules; endocarp 0.5-1 mm thick, brown, woody, fi
brous. Chromosome number unknown.
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 (Minas Gerais); angelim rasteiro (Maran
h?o); manga do campo (S?o Paulo).
Representative Specimens. Brazil. Bah?a: Espig?o Mestre, ca. 100 km WSW Barreiras, W. R. An
derson et al. 36865 (NY); Ibiquara, ca. 25 km N Barra da Estiva, Harley 26967 (K); Mpio. Morro do Chap?u,
Serra do Trombador, 6 km S Morro do Chap?u, Irwin et al. 32461 (MO, NY, US); Mpio. Feira de Santana,
Campus da UEFS, R. T. Pennington & Brito 239, 240, 241, 242 (CEPEC, FHO, K); Mpio. Angical, road Bar
reiras-Ibotirama (BR-242) Km 35, R. T. Pennington & Brito 266, 267, 268, 269 (CEPEC, FHO, K).?

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2003 ANDIRA 71
Distrito Federal: Fazenda Agua Limpa, Heringer 16202 (SP); 30 km S Brasilia on road to Belo Horizonte,
Irwin & Soderstrom 5662 (NY).?Goi?S: Chapada dos Veadeiros, ca. 20 km by road N Alto Para?so, W. R.
Anderson 6231 (NY); Serra Geral da Paran?, 1 km E S?o Jo?o da Alian?a, W. R. Anderson 7895 (NY); Serra
do Caiap?, ca. 60 km S Caiaponia on road to Jatai, Irwin & Soderstrom 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
Goi?s on road to Teresina de Goi?s, R. T. Pennington et al 499, 500, 501, 502, 503; Chapada dos Veadeiros,
5 km from Alto Para?so de Goi?s on road to the National Park, R. T. Pennington et al. 505, 506, 507, 508,
509.?Maranh?O: Mpio. Lor?to, 45-50 km S Lor?to on road between Santa Estev?o and Fazenda Santa Es
tev?o, G. Eiten & L. T. Eiten 5457 (NY); Grajah?, Lisboa s.n. (RB).?MATO GROSSO: 15-120 km beyond
Alto Araguaia, road to Cuiab?, Maguire et al. 56291 (NY, UB, US); Santa Anna das Chapadas, Malme s.n.
(F).?Mato Grosso do Sul: road between Aquiduana and Miranda, Bridgewater e? al 354 (E, UB); on road
to Porto Murtinho from Bonito, Bridgewaler e? al 363, 364 (E, UB).?Minas Gerais: Mpio. Santa Luzia,
Lagoa Santa, B?rrelo 5491 (F); Joaquim Felicio, Serra do Cabrai, Cavalcan?i e? al s.n. (SPF); Mpio. Caet?,
Fazenda Geriza, Felippe 36 (US); Mpio. Lavras, Heringer e? al. 15968 (US); 27 km SW Diamantina on road
to Goveia, Irwin e? al 22146a (F, MO, NY, RB, UB, US); Morro das Pedras, ca. 40 km NE Patrocino, Irwin
e? al. 25699 (MO, NY, US); between Sete Lagoas and C?rvelo, R. S. Sanios & Casiellanos 24071 (K).?Para:
Serra do Cachimbo, road Cuiab?-Santar?m, Km 856, Prance e? al 25053 (US).?Paran?: Jaguariahyva,
Dombrowski 9799 (US); Rio das Cinzas, Barra do Perdizes, Haischbach 7200 (US); Mpio. Campo Mour?o,
Campo Mour?o, Haischbach 15930 (F, US); Mpio. Arapoti, Haischbach 19955 (F, NY).?PERNAMBUCO:
Sitio de Lima, /. L. S. de Lima 01 (RB).?Rio GRANDE DO NORTE: road Cear? Mirim-Touros, Km 25, Bamps
5078 (NY); Natal, Moacyr Alvarenga 8 (MO).?ROND?NIA: 4 km from Vilhena, Vieira e? al 622 (NY).?
S?O PAULO: Morro Pellado, Edwall 5650 (SP); Mpio. Moji-Gua?u, Campos das Sete Lagoas, Faz. Campin
inha, just N Rio Moji-Gua?u, G. Eiten & L.T. Eiten 2363 (F, US); Mpio. Pirassununga, estrada Pirassu
nunga-Emas, Forero 8278 (SP); Mpio. S?o Carlos, N of city of S?o Carlos, de Freitas Campos 31 (UB, US);
Mpio. Botacatu, 18 km N Botacatu, 14 km E S?o Manuel, along S?o Manuel-Piracicaba road near old rail
road station, Gottsburger 2131 (UB); Mpio. Botacatu, ? margem do rodovia municipal, 5 km de Vitoriana,
Hernandes Bicuda et al 1538 (SP); 37 km de Avare, rodovia Avar?-Sao Manoel, /. Mattos 14530a (SP);
Sarapui, Yano 05 (SP). Paraguay. Amambay: Parque Nacional de Cerro Cora, Fern?ndez Casas & Molero
4079 (NY).?CAAGUAZ?: between Caaguaz? and Yh?, Fern?ndez Casas & Molero 6362 (MO); entre Yh? y
San Blas, Fern?ndez Casas & Molero 3851 (NY); 3 km S Arroyo Taruma, Zardini & Vel?squez 25331 (K); 2
km N Arroyo Guaranungua, Zardini & Vel?zquez 25676 (MO).?SAN PEDRO: 4 km S de R?o Verde, P?rez et
al. 1497 (MO); Dist. Lima, Estancia Carumbe, Pedersen 8510 (A, K); Alto Paraguay, Primavera, Woolston
734 (K, U).
Andira humilis is easy to recognize by its geoxylic suffrutex growth form, which is
unique in the genus and was described in detail by Warming (1892). It appears that occa
sionally A. humilis (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
Paulo, Goi?s, Paran?, and Distrito Federal (Fig. 11), whereas specimens with sparsely
hairy calyces are found to the northeast of these areas. I have not recognized these spec
imens as subspecif?c taxa, because there are intermediates (particularly in Paraguay and
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

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the synonymy of A. humilis, but maintained A. laurifolia. Handro (1969) recognized only
one species, A. humilis, and therefore this name takes priority.
11. Andira macrocarpa R. T. Pennington, sp. nov.?Type: Ecuador. Napo: Aguarico,

Reserva ?tnica Huaorani, Maxus road and pipeline construction project, Km 98,
23 Jul 1994, N. Pitman, A. Dik & C. Aulestia 631 (holotype: QCNE!; isotypes:
E! MO!).
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
cipue mesocarpiis duris, solidis (in A. galeottiana mesocarpiis mollibus, spongiosis)
recedit.
Emergent tree more than 30 m tall, 1 m 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
nently raised abaxially, pattern eucamptodromous becoming brochidodromous, tertiary
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).
Andira macrocarpa is known only from the type collection gathered in rain forest in
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
carp morphology that is soft and somewhat spongy with numerous air gaps when dry. The
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
apex, and the venation on the undersurfaces of the leaflets of A. macrocarpa is more
prominently raised.

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2003 ANDIRA 73
FIG. 24. Andira macrocarpa. A. Habit. B. Adaxial leaflet surface. C. Abaxial leaflet surface. D. Fruit. E.
Fruit in section, showing wall structure. (Based on Pitman et al. 631.)
I attempted to re-collect this species in December, 1994. The tree from which the type
collection was made grew alongside a road and had been cut down because of concerns
about the damage that falling fruit (which must weigh over 500 g when fresh) might cause
to passing vehicles.
12. Andira surinamensis (Bondt) Splitgerber ex Amshoff, Meded. Bot. Mus. Herb. Rijks
Univ. Utrecht 52: 60. 1939. Geoffroea surinamensis Bondt, Dissertatio medica
inauguralis de cortice Geoffraeae surinamensis, 8, figs 1-8. 1788. Geoffrea ob
tusifolia Stokes, Bot. Mat. Med. 4: 46. 1812, nom superfl. Andira retusa ? suri
namensis (Bondt) DC, Prodr. 2: 476. 1825. Vouacapoua surinamensis (Bondt)

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Kuntze, Revis, gen. pi. 1: 212. 1891?TYPE: SURINAME. Bondt s.n. (holotype:
L!).
Geoffroea 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
type, here designated: P!, isolectoype: P!).
Andira retusa var. oblonga Bentham, J. Proc. Linn. Soc, Bot. 4, Suppl. ("A Synop
sis of the Dalbergieae"): 121.1860.?Type: Brazil. Para: Santarem, Spruce 914
(lectotype, here designated: K! isolectotypes: K! RB!).
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
short, golden-brown to red-brown, appressed; stipels 1-3 mm long; petiolules 3-5 mm
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
men to 7 mm long (to 20 mm long on sapling leaflets), glabrous adaxially, sparsely hairy
abaxially, hairs short (<0.2 mm long), appressed, golden-brown to red-brown; primary
vein channelled adaxially; secondary veins 7-13, sunken or slightly sunken adaxially,
raised abaxially, pattern eucamptodromous becoming brochidodromous, tertiary veins
plane to slightly impressed adaxially and raised abaxially. Panicles terminal or occasion
ally axillary, 11-28 cm long, with brown to pale brown hairs at the branch tips, glabres
cent towards the base, hairs short, appressed; bracts 2 mm long, hairy to sparsely hairy,
hairs golden-brown to red-brown; pedicels 2-4 mm long; bracteoles 1 mm long, indu
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 12 mm 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, fibrous, brown, drying paler. Chro
mosome number unknown. Figs. IB, 2B, E.
Phenology. Flowering year-round.
Distribution (Fig. 25). Trinidad; Colombia (Amazonas, Meta, Vichada), Venezuela

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2003 ANDIRA 75
FIG. 25. Distribution 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,
Para, Roraima, Maranh?o, Cear?, Mato Grosso); in forest, especially along rivers and
gallery forest in savannas, also as an isolated tree in savanna (pers. obs. in Guyana).
Vernacular names. Yatuinay (Guahibo; Colombia); arisoru (Warao), canelito negro,
pilon, sob? (Yekuana), winka, wonka (Panare), palo blanco (Venezuela); koraro, bat seed,
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 Specimens. Trinidad. Toco Road, Valencia, N. L. Britton et al. 1776 (NY); Arena Re
serve, Gill 12621 (NY).
Colombia. Amazonas: bocas del Rio Yari en el R?o Caquet?, Pab?n & Mahecha 478 (COL, COAH).?
META: Villa Vecencio, cerca al terminal de transportes, Qui?ones 2764 (COL).?Vichada: Los Llanos, R?o
Orinoco, Puerto Carre?o, Cuatrecasas 4079 (US). Venezuela. AMAZONAS: Depto. Atabapo, Ca?o Negro, r?o
arriba desde la confluencia con el R?o Cuncunuma, Steyermark et al. 126212 (NY).?ANZOATEGUI: R?o Mapire,
afl. norte del R?o Orinoco medio, estaci?n Musinacio, Rosales & Valles 79 (NY).?Apure: Distritos Pedro
Camejo y Achaguas, galer?as del Cinaruco entrando por la Laguna de Calceta, R. G?mez et al. 588 (NY).?

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BOL?VAR: Dist. Cede?o, vie. Panare, village of Corozal, 6 km from Maniapure toward Caicura, Boom & Grillo
6460 (COL, NY). Guyana. NW slopes of Kanuku Mts. in drainage of Moku-moku Creek, A. C. Smith 3447
(A, MO, U); District 8, Armai, R. T. Pennington 433,462,463. Suriname. Schulz 7321 (COL); Sectie O, Sta
hel 57 (MEXU). French Guiana. Crique Jacques, Wachenheim s.n. (US); Cayenne, Martin s.n. (US).
Ecuador. Napo: Reserva Faunistica Cuyabeno, Laguna Cocha, Palacios et al. 7747 (E, MO, QCNE). Peru.
Loreto: Prov. Maynas, Nina Rimi, a orillas del R?o Nanay, 25 km SW Iquitos, Torres 160 (MO, NY).?Madre
DE Dios: Tambopata Reserve, junction of R?os La Torre and Tambopata, Gentry & Jaramillo 58003 (MO).?
UCAYALl: Bosque Nacional von Humboldt, road Pucallpa-Tingo Mar?a Km 88, Gentry et al. 36390 (MO, NY).
Bolivia. Pando: Cobija, camino a Puerto Rico, San Francisco, Meneces & Hartshorn 2093 (NY).?SANTA
Cruz: Velasco, Reserva Ecol?gica El Refugio, Killeen & P?rez 6682 (K, MO). Brazil. Amap?: Rio Araguar?,
cachoeira do Pared?o, Froes & Black 27684 (INPA).?AMAZONAS: Rio Negro, Santa Isabel, Ducke 145 (A)?
Cear?: Fortaleza, estrada do aeroporto, Ducke 2432 (US).?Maranh?o: Island of S?o Luis, Froes 11805 (A,
GH, K, NY, U, US).?MATO GROSSO: Aripua?a, Km 245 da BR-174, M. G. Silva & A. Pinheiro 4327 (F).?
Para: Santarem, Km 35 da estrada do Palh?o, acampamento do Igarap? Curupira, M. Silva & R. Sousa 2284
(COL, US).?Rond?NIA: Mpio. Ji-Paran?, Rio Machado, F. E. Miranda et al. 296 (INPA).?RORAMA: estrada
Manaus-Caracari, Km 515, bank Igarap? Dias, Steward et al. 135 (GH).
Andira surinamensis might be confused with A. verm?fuga, A. humilis, A. galeottiana,

and A. macrocarpa. Andira macrocarpa and A. galeottiana differ in their much larger
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
namensis, but it is a geoxylic suffrutex rather than a tree.
Most previous workers (e.g., Mattos 1979) have cited this species as A. surinamensis
(Bondt) Splitgerber ex Pulle; however, J. C. Lindeman (pers. comm.) pointed out that
Pulle (1906) made this combination in synonymy, which is inadmissible under terms of
the ICBN (Art 34.1; Greuter et al. 2000). The correct attribution is A. surinamensis
(Bondt) Splitgerber ex Amshoff.
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
raima, Brazil, a decoction of the bark of this tree is taken as a vermifuge, as reported by
Bondt (1788) in the first description of the species, and also as a malaria remedy.
13. Andira 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
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!).
Andira stipulacea var. bahiensis Bentham, J. Proc. Linn. Soc, Bot. 4, Suppl. ("A
Synopsis of the Dalbergieae"): 119. 1860. Andira legalis var. bahiensis (Ben
tham) N. F. Mattos, Loefgrenia 40: 3. 1970.?Type: Brazil. Bahia: Blanchet
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
sent; bark grey-brown to brown, Assuring vertically and scaling on larger trees; slash pale

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 77
salmon pinkish, oxidizing red-brown, with clear ex?date and small amount of red ex?date;
wood 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
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
long; petiolules 1.5^4.5 mm long, indumentum like that of rhachis; leaflets in (3-) 4-6
(-7) pairs, (2.2-) 4.1-12.5 (-20.5) cm long, (1-) 2.2-6.5 (-7.5) cm wide, elliptic, nar
rowly obovate (rarely narrowly ovate, narrowly elliptic, broadly obovate to oblanceolate),
thick-chartaceous (rarely chartaceous or subcoriaceous), shiny dark green adaxially, matt
abaxially, base obtuse or rounded, often slightly decurrent, apex obtuse or rounded (rarely
acute), often slightly retuse or with a short acumen to 10 mm long, glabrous adaxially ex
cept the groove of the primary vein very sparsely hairy, glabrescent, the hairs erect,
sparsely to very sparsely hairy abaxially, hairs ? appressed to ? erect, red-brown to
whitish, >0.2-1.0 mm long; primary vein channelled adaxially; secondary veins 8-13, ?
plane to slightly sunken 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.
Gynoecium 15.5-21 mm long, ovary hairy, stipe and style sparsely hairy to upper and
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.
Fruits 3-6.2 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 12 mm long, but fruit usually breaking off without a stipe; suture raised
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
restinga forest and rain forest. I have also seen this species growing along river banks in
Bahia, and in a seasonally inundated area in Rio de Janeiro; it can clearly tolerate water
logging, and it seems likely that its fruits may be secondarily dispersed by water.
Vernacular names. Angelim coco, angelim preto (Bahia); angelim coco (Rio de
Janeiro).

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FIG. 26. Andira anthelmia and A. legalis. A. A. anthelmia, habit. B. A. anthelmia, abaxial leaflet surface.
C. A. legalis, 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,
also shown above in longitudinal section. M. A. anthelmia, gynoecium, also shown above in longitudinal sec
tion. N. A. anthelmia, fruit. O. A. legalis, fruit, partially sectioned to show wall structure. (Based on: A-B, R. T.
Pennington 281; C-F, R. T. Pennington 307; F-L, A. Lima 51-944; M, E. Pereira et al 4156; N, R. T. Pen
nington, 183; O, G. P Lewis & H. C. de Lima 1196.)

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 79
55 50 45 40 35
FIG. 27. Distribution of Andira anthelmia.
Additional Specimens Examined. Brazil. Alagoas: with

Estrada Mara?-Ubaitaba, Bel?m 1879 (NY, UB); Coaraci, cocoa pla
Mpio. Ilh?us, Fazenda Serra Grande, W of CEPEC cacao plantatio
h?us, estrada Ilh?us-Itabuna, pr?x. a CEPLAC, H. C. de Lima et
h?us road, 2 km from CEPLAC, R. T. Pennington et al. 183 (CE
Antonio de Baixo, Km 16 on road from BR-101 to Palmira, then 1 km
286 (CEPEC, FHO, K); Mpio. Marau, road Mara?-Ubaitaba Km 4,
K); Mpio. Ubaitaba, banks of Rio de Contas, by road Ubaitaba-Mar
FHO, K); Mpio. Ilh?us, road Ilh?us-Itabuna, 5 kms before CEPL
(CEPEC, FHO, K); Mpio. Jussari, Junction of BR-101 and road to
valho 294 (CEPEC, FHO, K); Mpio. Porto Seguro, BR-101, bank of
de Carvalho 297 (CEPEC, FHO, K); Mpio. Prado, 4 km along track o
R. T. Pennington ? F. A. de Carvalho 312 (CEPEC, FHO, K); Mpio
Alian?a, R. S. Pinheiro 250 (US); estrada Santa Ines-Rio Bahia, K
Santo: estrada do Pan?as, Collatina, /. G. Kuhlmann 360 (RB).?M
Gomes 2864 (RB, UB).?R?o DE JANEIRO: Praia de Grumari, D.
Cim, D. Ara?jo & Maciel 4044 (GUA); Silva Jardim, Poco d'Anta
deirantes, Morro do Calemb?, Lanna Sobrinho 649 (US); Resting
Restinga de Jacarepagu?, Pereira et al. 4156 (NY); Goitacazes, Lago
Segadas-Vianna et al 291 (US).

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
Andira anthelmia is most closely related to A. legalis (Fig. 9), which has similar
large, persistent stipules. The 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
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
frondosa (Martius ex Bentham) Kuntze, Revis, gen. pi. 1: 212. 1891.?TYPE:
Brazil. Rio de Janeiro: "In arenosis sylvarum ad Capo Frio et alibi circa oram
Brasiliae 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
hairs, glabrescent; 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
rhachis; leaflets in 4-7 (-9) pairs, 5.5-15 (-30) cm long, 2.3-7 (-11) cm wide, elliptic,
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
ondary veins 8-13, slightly sunken adaxially, raised abaxially, pattern eucamptodromous
becoming brochidodromous, 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-) 15 mm 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.
Gynoecium (14.5-) 17.2-18 mm long, ovary sparsely hairy, stipe and style very sparsely
hairy to upper (basal half only) and lower surface of ovary hairy with a few hairs extend
ing down the stipe, hairs red-brown, ? appressed; stipe 5-6 mm long, ovary (5.5-) 7 mm
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

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2003 ANDIRA 81
55 50 45 40 35
FIG. 28. Distribution of Andira legalis.
dark brown (both fresh and dry); stipe to 5 mm lo

ally, slightly raised abaxially; stylar remnant not o
brown, hard, finely granular; endocarp 3-18 mm th
some number unknown. Figs. ID, 26.
Phenology. Flowering August to November.
Distribution (Fig. 28). Brazil (Bahia, Espirito San
de Janeiro); in restinga and rain forest.
Vernacular names. Angelim preto, angelim ro
Janeiro).
Additional Specimens Examined. Brazil. Bah?a: Mara?, Bel?m 1830 (CEPEC, MEXU, NY, UB, US);
Mpio. Uru?uca, Dist. Serra Grande, 7.3 km na estrada Serra Grande-Itacar?, Fazenda Lagoa do conjunto
Fazenda Santa Cruz, A. M. de Carvalho et al 3508, 3509 (CEPEC, NY); Mpio. Porto Seguro, junction on right
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
Santa Cruz, 1 km up this road, R. T. Pennington & Lewis 191 (CEPEC, FHO, K); Mpio. Mara?, road
Mara?-Ubaitaba Km 6, R. T. Pennington et al 214, 215 (CEPEC, FHO, K); Mpio. Alcoba?a, 1 km N do centro

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
da cidade, R. T. Pennington & F. A. de Carvalho 305,307,308,310 (CEPEC, FHO, K); Teixeira de Freitas, Vale
do Rio Alcoba?a, dos Santos 2108 (US); Mpio. Uru?uca, 7.3 km N Serra Grande on road to Itacar?, Fazenda
Lagoa do Conjunto e Fazenda Santa Cruz, W. W. Thomas et al 8533 (CEPEC, NY).?ESPIRITO SANTO: Reserva
Florestal da CVRD, Linhares, DA.F. 128 (RB); Vitoria, Fazenda Maruhype, J. G. Kuhlmann 478 (NY).?Per
NAMBUCO: Mata de Dois limaos, margem estrada para o acuda do Prata, Lima 51-944 (K).?R?o DE JANEIRO:
Guanabara, Marambaia, Almeida de Jesus 2044 (NY); Mpio. Maca?, near Brejo de Bezerra, D. Araujo 4612
(GUA); Mpio. Saguarema, near Sambaqui da Beirada, D. Araujo 8079 (GUA); Mpio. Cabo Fri?, dune system
at Dama Branca, D. Araujo et al. 8326 (GUA); Mpio. Saguarema, loteamento da Praia de Itauna, D. Araujo et
al. 8597 (GUA); Mpio. Saguarema, Comoros da Lagoa Vermelha, D. Araujo & Mauro 8614 (GUA); Mpio. Ar
rail do Cabo, restinga de Massambaba, Zacara, D. Araujo & Maciel 8700 (GUA); Recreio dos Bandeirantes, W.
Hoehne 5718 (SP), 5895 (COL, SP); Goitacazes, Rio Doce, J. G. Kuhlmann 06529 (RB); Mpio. Marica, Barra
de Marica, pr?x. a Lagoa de Marica, Lewis & H. C de Lima 1196 (K); Mpio. Cabo Fri?, Cabo Fr?o, Arraial do
Cabo, L. B. Smith 6554 (US).
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).
15. Andira ormosioides Bentham, Comm. legum. gen. 44. 1837. Andira anthelmia var.
ormosioides (Bentham) Bentham, J. Proc. Linn. Soc, Bot. 4, Suppl. ("A Synop
sis of the Dalbergieae"): 120. I860.?Type: Brazil. Tingua, Schott s.n. (holo
type: K!; isotype: F!).
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; leaf axis 9-30 (-35) cm
long; rhachis with red-brown erect hairs, glabrescent; stipels 2-3 (-6) mm long; petiolules
2-5 mm long, indumentum like that of rhachis; leaflets in (3-) 4-5 pairs, 4.7-15.5 (-19
on sterile shoots) cm long, 1.8-7 cm wide, elliptic to narrowly 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
mentum most dense on veins, hairs red-brown, erect, >0.2-1.0 mm long; primary vein
channelled adaxially; secondary veins 8-11, slightly sunken adaxially, raised abaxially,
pattern eucamptodromous becoming brochidodromous, tertiary veins plane to slightly im
pressed adaxially and raised abaxially. Panicles 10-30 cm long, terminal (more rarely ax
illary), with red-brown, erect hairs; bracts 3-7 mm long, with red-brown, ? appressed
hairs; pedicels 3-7 mm long; bracteoles 1-2.5 mm long, indumentum like that of bracts.
Flowers 18-23 mm long. Calyx 9-10 mm long, with red-brown, ? appressed to ? erect
hairs; lobes 0.8-2 mm long, obtuse to acute. Petals pink to purple; standard blade 17-20
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,
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,

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 83
55 50 45 40 35
FIG. 29. Distribution of Andira ormosioides. The collec

mosioides (see discussion); if so, the locality is a considera
smooth, very dark brown with green beneath

pearing smooth but minutely tuberculate (best
mm long; suture slightly raised adaxially, flan
raised abaxially; stylar remnant raised at ap
greenish white, drying pale brown, granular; en
fibrous. Chromosome number unknown. Fig. 3
Phenology. Flowering June to December.
Distribution (Fig. 29). Brazil (Bahia, Espirit
S?o Paulo); in restinga, rain forest, and semi-d
Vernacular names. Angelim pedra (Espirito S
gelim amargoso (Rio de Janeiro); Jacaranda r
Additional Specimens Examined. Brazil. Bahia: Mpio.

& F. A. de Carvalho 306, 309 (CEPEC, FHO, K).?Espir
164/79 (RB).?Minas Gerais: Patos de Minas, A. P. Duar

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
de Fora, Heringer 2700 (RB); Carangola, Rio Carangola, Leoni & Leoni 856 (K); 400 km E Belo Horizonte,
vicinity of Rio Manhuac?, Valley of Rio Doce, Estac?o Biol?gica Caralinga, Fazenda Montes Claros, Lopes &
Andrade 850 (K, RB).?Rio DE Janeiro: Serra da Estrela, de Almeida s.n. (RB); Barreira de Soberbo, Dias s.n.
(R); Rio de Janeiro, Horto Florestal s.n. (RB 54881) (RB); Rio de Janeiro, Horto Florestal s.n. (RB 81810) (RB);
Avellar, /. G. Kuhlmann 29 (RB); Guanabara, /. G. Kuhlmann 528 (F, NY, RB, UB); Mpio. Mage, Cachoeiras
de Macacu, Para?so, near CPRJ, banks of Rio Para?so, H. C. de Lima et al. 4253 (RB); estrada do Recreo dos
Bandeirantes, Lutz 1493 (R); Restinga da Trjuca, Machado s.n. (RB 76106), Machado s.n. (RB 75296) (RB);
Mpio. Mag? (IIIo), Dist. Para?so, Centro de Primatologia do RJ, Martinelli & Pessoa 10503 (RB); Mpio. Mag?,
Para?so, Sampaio 6 (RB); Gavea, E. Ule LVII (R).?SAO PAULO: Aeto da Serra, de Andrade s.n. (R); Ubatuba,
pr?ximo a Base Norte do Instituto Ocean?grafico, da Cruz 13 (NY, SP).
Field observation in Bahia of growth form and fruit indicates that A. ormosioides is
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
carp of A. fraxinifolia and A. anthelmia). Andira ormosioides has a long bole and small
crown, even in open situations, whereas A. fraxinifolia and A. anthelmia have a short bole
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
sioides occurs in the semi-deciduous forests of the Rio Doce valley of eastern Minas
Gerais, which are transitional to the coastal rain forests, whereas A. legalis grows strictly
in rain forests and restingas. The Duarte specimen from the dry area of Patos de Minas is
more likely to be A. ormosioides and is therefore included here.
16. Andira fraxinifolia Bentham, Comm. legum. gen. 44. 1837. Vouacapoua fraxinifolia
(Bentham) Kuntze, Revis, gen. pi. 1: 212. 1891.?Type: Brazil. Minas Gerais:
prope Itamb?, Pohl s.n. (lectotype, here designated: K!; isolectotype: K!).
Andira parvifolia Martius ex Bentham, Comm. legum. gen. 44. 1837. Vouacapoua
parvifolia (Martius ex Bentham) Kuntze, Revis, gen. pi. 1: 212. 1891.?TYPE:
Brazil. Minas Gerais: "in campis altis Serro Frio," Martius s.n. (holotype: M!).
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 ad Mucuri," 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!).
Andira micans Taubert ex Glaziou, Bull. Soc. Bot. France 52, M?moire 3: 151.
1906.?Type: Brazil. Rio de Janeiro: Nova Friburgo, Glaziou 20274 (holotype:
P!; isotypes: BR, K!; photo of B isotype: G!).

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 85
Andira handroana N. F. Mattos, Loefgrenia 40: 1. 1970.?Type: BRAZIL. Bahia: be

tween Len?ois and Itaberaba, 15 Sep 1956, E. Pereira 2064 (holotype: R!; iso
type: RB!).
Andira fraxinifolia var. latifoliolata N. F. Mattos, Loefgrenia 40: 2. 1970.?TYPE:
Brazil. Rio de Janeiro: Restinga da Gavea, 13 Aug 1949, O. Machado s.n.
(holotype: RB!).
Andira anthelmia var. gracilis N. F. Mattos, Loefgrenia 40: 3. 1970.?Type: Brazil.
S?o Paulo: Limeira, Navarro de Andrade 167 (holotype: R; isotype: RB!).
Andira bahiensis N. F. Mattos, Loefgrenia 45: 2. 1970.?Type: Brazil. Bahia: Km
80 between Betanha and Canavieiras, 13 Jul 1964, N. T. Silva 58414 (holotype:
UB; isotype: CEPEC!).
Andira pernambucensis N. F. Mattos, Loefgrenia 53: 1. 1970.?TYPE: BRAZIL. Per
nambuco: Rio Formoso, Engenho S?o Manoel, 3 Sep 1954, J. Falc?o, Engler &
E. Pereira 944 (holotype: RB!; isotypes: RB-2 sheets!).
Andira fraxinifolia var. lanceata N. F. Mattos, Loefgrenia 58: 3. 1973. Brazil. S?o
Paulo: 2 km N of Atibaia, 24 Aug 1963, J. Mattos & N. F. Mattos 8383 (holo
type: SP!).
Andira pisonis var. emarginata N. F. Mattos, Loefgrenia 58: 2. 1973.?Type:
Brazil. Bahia: Igua?u, 30 Dec 1922, P. Campos Porto s.n. (lectotype, here des
ignated: RB!).
Andira pisonis war. puberula N. F. Mattos, Loefgrenia 58: 2. 1973.?Type: Brazil.
Espirito Santo: Linhares, Lagoa do Dur?o, Rio Doce, Apr 1934, /. G. Kuhlmann
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
cream/buff; twigs brown to dark brown, older twigs paler, often buff/whitish, hairy to
sparsely hairy, glabrescent, hairs erect. Stipules 2-9 mm long, to 1 mm 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,
oblanceolate (more rarely lanceolate to broadly obovate), thick-chartaceous to subcoria
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
veins, hairs pale to red-brown, erect, >0.2-l .0 mm long; primary vein channelled adaxially;
secondary veins 6-10, ? plane to slightly 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.
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

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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
mm, vexillary stamen 9-12 mm long. Gynoecium 13.5-18 mm long, ovary hairy, stipe and
style sparsely hairy to upper and lower surfaces of ovary sparsely hairy with scattered hairs
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,
sweet-smelling, 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.
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).
Representative Specimens. Brazil. Alagoas: Fleixeiras, Fazenda S?o Louren?o, Andrade-Lima e? al

7 (F).?Bah?a: Mpio. Sa?de, Cachoeira do Paulista, 6 km along road starting at Km 10 of road Sa?de-Jacobina,
Amorim et al 1055 (CEPEC, E); Serra Jacobina, Blanchet 2723 (E, BM, F, K, NY); Mpio. Jacobina, Serra do
Tombador, ca. 8 km SW da sede do municipio, na estrada para Morro do Chap?u, A. M. de Carvalho et al 6141
(E, NY); Mpio. Saude, Cachoeira do Paulista, 6 km along road starting at Km 10 of road Sa?de-Jacobina, /. G.
Jardim 71 (CEPEC, FHO); Mpio. Ilh?us, Oliven?a, 100 m up hill from Club Tomoromba, R. T. Pennington 194
(CEPEC, FHO, K); Mpio. Ilh?us, ca. 3 km N Oliven?a, road to Pontal, junction on left, about 300 m up this
track, R. T. Pennington 200 (CEPEC, FHO, K); Mpio. Ilh?us, Agua da Oliven?a, R. T. Pennington 201, 202
(CEPEC, FHO, K); Mpio. Jussari, Fazenda Teimoso, R. T. Pennington & A. M. de Carvalho 204, 228 (CEPEC,
FHO, K); Mpio. Jussari, road from BR-101 to Palmiras, ca. Km 15, R. T. Pennington & A. M. de Carvalho 207
(CEPEC, FHO, K); Mpio. Mara?, road Mara?-Ubaitaba Km 5, R. T. Pennington et al. 211, 213 (CEPEC, FHO,
K); Mpio. Una, road to Independencia Km 4 (off road to Pedras from main road Oliven?a-Una), R. T. Pen
nington e? al 234, 236 (CEPEC, FHO, K); Mpio. Jacobina, Km 20 along road Jacobina to Morro do Chap?u
(BR-324), R. T. Penningion & Brito 248,249,250; Mpio. Morro do Chap?u, 18 km E Morro do Chap?u on BR
052, R. T. Penningion & Brito 251 (CEPEC, FHO, K); Mpio. Len?ois, 5 km from Len?ois on road to BR-242,
R. T. Pennington & Brito 274, 275, 276, 278, 279; Mpio. Alcoba?a, ca. 2 km N Alcoba?a, R. T. Pennington &
F. A. de Carvalho 302 (CEPEC, FHO, K); Mpio. Itua?u, arredores do Morro da Mangabeiras, Queiroz et al.
1610 (K, NY).?Cear?: Santana, Poranga, S. J. Filho 63 (RADAM).?Distrito Federal: Campus of Uni
versity of Brasilia, Heringer 10064 (UB); Fazenda Agua Limpa, near Vargem Bonita, ca. 18 km SSW Brasilia
TV Tower, Ratter et al 3791 (UB).?ESPIRITO SANTO: Lagoa do Dur?o, Linha Res., Rio Doce, J. G. Kuhlmann
163 (NY); Mpio. Linhares, Reserva Florestal da CVRD, H. C de Lima 1711 (RB).?GOI?S: Mpio. Santo An
tonio Descoberto, Corrego da Fazenda Cabteiro do Para?so, Elias de Paula 3147 (UB).?MINAS GERAIS: San
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,
US); Concei?ao de Mata Verde, estrada para Lamban, E. Pereira 5788 (RB); Rio Vermelho, Pedra Menina,
Fazenda Vargem da Ang?lica, Morro da Virada do Mato Virgem, Pirani et al s.n. (SP).?Paran?: Guaratuba,
Morro da entrada da Baia, Braga 2364 (RB); Itacar?, opp. Marungaia, Dus?n 16511 (A); Mpio. Trjucas do Sol,
Matul?o, Haischbach 15117 (NY, US); Mpio. Antonica, Rib. Lagoa Vermelha Haischbach 18102 (NY, US);
Mpio. Morretes, Col. Florestal, Hatschbach 20206 (RB); Mpio. Senges, Rio do Funil, Hatschbach 27144
(NY).?Pernambuco: Recife, Mata de Dois Irm?os, Lima 48-126 (RB).?Rio de Janeiro: Cabo Fri?, Arraial
do Cabo, Graziela 7 (NY); Guanabara, Reserva Biol?gica de Jacarepagu?, Lanna 124 (RB); Mpio. Maca?,

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 87
FIG. 30. Andira fraxinifolia. A. Habit. B. Node of rhachis with stipels. C. Abaxial leaflet surface. D.
Flower. E. Calyx, opened to show inner surface. F. Standard petal, inner surface. G. Outer surface of wing petal
with lamellate sculpturing. H. Detail of lamellate sculpturing. I. Keel petal, inner surface. J. Androecium. K. Gy
noecium, also shown above in longitudinal section. L. Fruit. M. Fruit in section, showing wall structure. O. Seed.
(Based on: A-K, R. T. Pennington 228; L-M, R. T. Pennington 202)

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
75 70 65 60 55 50 45 40 35
FIG. 31. Distribution 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
1173 (RB); Mpio. Parati, Parati-Mirim, H. C. de Lima 3673 (RB).?Santa Catarina: N Joinvile, J. Mattos
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
Cardoso, Morro do Pereirinha, de Barros 2238 (SP); S?o Vicente, W. Hoehne s.n. (SPF); near Moji-Mirim, M.
Kuhlmann 1477 (NY); Juqui?, Capueira, na margem do Rio Juqui?, M. Kuhlmann 4694 (SP); Ubatuba, Praia de
Perequer?-Ass?, J. Mattos 13802 (SP); Mpio. Iguape, esta?ao ecol?gica de Jureia, Pirani 818 (SPF); Mpio.
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,
whereas those o? A. fraxinifolia are 13-17 mm long. Andira 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
only, whereas in A. fraxinifolia it is uniformly hairy. The fruits of A. verm?fuga dry with
a wrinkled surface, whereas in A. 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.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 89
Vegetative characters, particularly leaf and leaflet size and the amount and color of
the indumentum, are variable in A. 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.
17. Andira n?tida Martius ex Bentham, Comm. legum. gen. 45. 1837.?Type: Brazil.
Espirito Santo: Rio Doce, 1827, Wied-Neuwied s.n. (holotype: BR!; isotypes:
BM! K! M, NY!; photo of M isotype: F!).
Multi-stemmed shrub to tree 25 m tall; buttresses absent or very small; bark brown,
Assuring vertically and 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
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 10 mm long, to 1 mm 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;
leaflets in 2-4 (-5) pairs, 2.4-11.5 (-14) cm long, 1.7-4.5 (-6) cm wide, elliptic, broadly
elliptic, narrowly ovate, ovate (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, or ? completely brochidodromous, tertiary
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

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
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 thick when 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 (to May in Espirito Santo).
Distribution (Fig. 33). Brazil (Bahia, Espirito Santo, Pernambuco, Rio de Janeiro); in
open restinga scrub on white sand, closed restinga forest, and rain forest.
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,
US); Sa?da de Itaju do Colonia para Itap?, Km 20, Almeida 356 (US); Mpio. Uru?uca, Dist. Serra Grande,
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. Ilh?us, ca. 7 km along road Oliven?a-Vila Brasil, A. M. de Carvalho et al.
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
(RB); Mpio. Salvador, pr?ximo ao aeroporto, M. L. Guedes 972 (RADAM); 16 km S Cumuruxatiba on road to
Prado, Harley et al. 18086 (K); Mpio. Ilh?us, grounds of CEPLAC, J. G. Jardim 756 (CEPEC, E, NY); Mpio.
Prado, 12 km S Prado, estrada para Alcoba?a, G. P. Lewis & A. M. de Carvalho 791 (K); Mpio. Nova Vi?osa,
Copuva, G. Hatschbach & J. M. Silva 48748 (K); Mpio. Mara?, 1 km S Porto de Campinhos, estrada para
Ubaitaba, G. P. Lewis & A. M. de Carvalho 1029 (K, MEXU); Mpio. Valen?a, Km 13 da estrada Valen?a para
Guaibim, G. P. Lewis & A. M. de Carvalho 1050 (K); Mpio. Belmonte, estrada Belmonte-Itapebi, Km 26, Mat
tos Silva & Hage 585 (CEPEC, K, UB); Mpio. Ilh?us, just S of Oliven?a, R. T. Pennington 186 (CEPEC, FHO,
K); Mpio. Santa Cruz da Cabr?lia, 10 km N Porto Seguro on road to Santa Cruz Cabr?lia, R. T. Pennington &
G. P. Lewis 188 (CEPEC, FHO, K); Mpio. Ilh?us, ca. 3 km W Oliven?a, R. T. Pennington 192,198 (CEPEC,
FHO, K); Mpio. Buerarema, ca. 15 km S Itabuna on road BR-101, R. T. Pennington et al. 209 (CEPEC, FHO,
K); Mpio. Marau, road Porto de Campinhos-Ubaitaba, Km 9, R. T. Pennington et al. 221 (CEPEC, FHO, K);
Mpio. Una, near Pedras, Km 5 on road to Independencia, R. T. Pennington et al. 235 (CEPEC, FHO, K); Mpio.
Salvador, dunas de Itapua, R. T. Pennington et al. 288, 290, 291, 292 (CEPEC, FHO, K); Mpio. Alcoba?a, Km
4 on road Alcoba?a-Teixeira de Freitas, R. T. Pennington & F. A. de Carvalho 300 (CEPEC, FHO, K); Mpio.
Alcoba?a, ca. 2 km N Alcoba?a, 1 km from the sea, R. T. Pennington & F. A. de Carvalho 301 (CEPEC, FHO,
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.
Itacar?, "Campo Cheiroso," 14 km N Serra Grande, off road to Itacar?, W. W. Thomas et al 9469 (CEPEC);
Mpio. Belmonte, Barrol?ndia, Est. Experimental "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
& Medeiros-Costa 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
ney 2930 (RB); Mpio. Santo Amaro de Brotas, Sap?-Fejeunde Arauari, near Torre de Embratel, Farney &
Gomes 2994 (RB); Mpio. Estancia, Rod. Abais-BRIOl, 1 km W Abais, Mattos Silva et al. 3027 (CEPEC).

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 91
FIG. 32. Andira nitida. A. Habit. B. Abaxial leaflet surface. C. Flower. D. Calyx, opened to show inner
surface. E. Standard petal, inner surface. F. Outer surface of wing petal with lamellate sculpturing and detail of
sculpturing. G. Keel petal, inner surface. H. Androecium. I. Gynoecium, also shown below in longitudinal sec
tion. J. Fresh fruit. K. Wrinkled dried fruit. L. Fruit in section, showing wall structure. M. Seed. (Based on: A,
B, R. T. Pennington 288 (leaves), H C. de Lima 1668 (inflorescence); C-I, A. Ribeiro 62; J, L, R. T. Pennington
292; K, R. T. Pennington 291.)

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
55 50 45 40 35
FIG. 33. Distribution o? Andira nitida.
Andira nitida is most likely to be confused with

valhoi. Andira humilis is distinguished by its uniqu
this is not noted on herbarium specimens, the two
tida's smaller flowers (10-13 mm as compared to
more, A. humilis is a cerrado species that is not sym
to the rain forests and restingas of coastal Brazil.
valhoi in Bahian restingas, but the latter is clearly
cm long, 4.8-8 cm high, 4.5-9 cm wide), brown f
are provided by the larger flowers (14-15 mm) an
Andira nitida is also sympatric with A. marauensi
ferences between these two species are outlined u
Field observations in Bahia revealed two sympa
(also growing in damp areas in restinga) with gene
and a smaller, smooth fruit, which dries wrinkled.
can tolerate dry restinga habitats; it has coriaceou
larger fruit, which is ridged from the lower to th

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 93
extensive collections (mostly sterile) of both growth forms in Bahia and found them
distinct morphologically. I initially considered them as possibly distinct species (A. n?tida
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
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 marauensis N. F. Mattos, Loefgrenia 45: 1. 1970.?Type: Brazil. Bahia:

Mara?, 12 Jan 1967, R. P. Belem & R. S. Pinheiro 3089 (holotype: UB!; 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 1 mm 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
long, 1.1-3.7 cm wide, elliptic, narrowly obovate (rarely broadly obovate), terminal
leaflets often the most distinctly obovate, subcoriaceous, base acute to obtuse, slightly de
current, apex obtuse, rounded (rarely slightly retuse or with an acumen to 2 mm long),
glabrous adaxially, very sparsely hairy abaxially, hairs red-brown, short (<0.2 mm long),
appressed; primary vein channelled adaxially; secondary veins 8-10, ? plane adaxially,
slightly raised abaxially, pattern brochidodromous, tertiary veins plane adaxially and
plane or slightly raised abaxially. Panicles 5-11 cm long, terminal and axillary, with
sparse, short, appressed red-brown hairs at branch tips, glabrescent towards the base;
bracts narrow, caducous, ca. 2 mm long, sparsely hairy, hairs red-brown, ? appressed;
pedicels 1-2 mm long; bracteoles caducous, not seen. Flowers 9-11 mm long. Calyx
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
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 names. Angelim.
Additional Specimens Examined. Brazil. Bah?a: Mpio. Una, Reserva Biolog?a do Mico Le?o, estrada
no Km 46 da Rod. BA-001 (Ilh?us-Una), Amorim et al. 1411,1238,1908 (CEPEC, NY); Mpio. Mara?, Mara?,
R. P. Bel?m & Pinheiro 3091 (NY, UB), 3142 (MEXU, UB, US); Mpio. Uru?uca, Dist. de Serra Grande, 7.3 km
na estrada Serra Grande-Itacor?, A. M. de Carvalho et al. 3510 (CEPEC, NY); Mpio. Una, Reserva Biolog?a do
Mico Le?o, estrada no Km 46 da Rod. BA001 (Ilh?us-Una), /. G. Jardim et al. 90 (CEPEC, FHO); Mpio. Una,
Maruim, border of Fazendas Maruim and Dois de Julho, 33 km SW Oliven?a on road Oliven?a-Buerarema,

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
55 50 45 40 35
FIG. 34. Distribution of Andira marauensis.
Mori et al 13800 (NY); Mpio. Ilh?us, Fazenda Barra do Manguinho, Km

102 (CEPEC, K-2 sheets).
Andira marauensis is morphologically similar to rain for

differs in its smaller leaves, its obovate, blunt-ended leaflet
smaller flowers, and fewer ovules. It must be considered e
of suitable rain forest habitat remaining in southern Bahia.
19. Andira carvalhoi R. T. Pennington & H. C. Lima, Ke

Brazil. Bahia: Mpio. Ilh?us, estrada Oliven?a-Vila Br
13 Jan 1981, A. M. de Carvalho et al. 491 (holotype
Shrub, often multi-stemmed with creeping rhizomes, o

10 m tall with spreading branches; buttresses absent; bark g
vertically and flaking slightly; slash 2-4 mm thick, pale bro

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
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 1 mm 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 the margins 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
plane adaxially and abaxially. Panicles 5-30 cm 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
hairs; pedicels 1.5-3 mm long; bracteoles 1.5-2 mm long, narrow, moderately persistent,
indumentum like that of bracts. Flowers 14-15 mm 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 11 mm
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: Mpio. Mara?, rod. BR-030, Porto de
Campinhos-Mara?, Km II, A. M. de Carvalho et al. 178 (CEPEC, K); Mpio. Ilh?us, estrada Ilh?us-Canavieiras,
Km 33, A. M. de Carvalho 622 (CEPEC); Mpio. Ilh?us, estrada Oliven?a-Maruim, Km 9, A. M. de Carvalho &
Faria 2547 (CEPEC, UEFS); Mpio. Dh?us, 7 km on road Oliven?a-Vila Brasil, A. M. de Carvalho et al. 3308
(CEPEC); Mpio. Ilh?us, estrada Pontal-Oliven?a, Km 10, Gentry & Zardini 50011 (CEPEC); Mpio. Ilh?us,
estrada Oliven?a-Maruim, Km 5-8, M. P. M. de Lima et al. 20 (CEPEC, RB-2 sheets); Mpio. Ilh?us, estrada
Oliven?a-Maruim, Km 7-10, Martinelli et al. 11102 (CEPEC, RB); Mpio. Ilh?us, estrada Oliven?a-Una, Km 5,
Fazenda Jairi, Mattos Silva et al. 1199 (CEPEC, K); Mpio. Ilh?us, Fazenda Barra do Manguinho, ramai com en
trada no Km 10 da estrada Pontal-Oliven?a, Km 10, Mattos Silva et al. 1393 (CEPEC, K); Mpio. Ilh?us,
Fazenda Guanabara (junto ? Fazenda Barra do Manguinho), ramai com entrada no Km 10 da estrada
Ilh?us-Oliven?a, Mattos Silva et al 1877 (CEPEC, K); Mpio. Ilh?us, dirt road Oliven?a-Maruim, Km 6, R. T.
Pennington et al 181 (CEPEC, FHO, K); Mpio. Ilh?us, just S Oliven?a, R. T. Pennington 184, 185, 229
(CEPEC, FHO, K); Mpio. Ilh?us, ca. 3 km N Oliven?a, R. T. Pennington 197 (CEPEC, FHO, K); Mpio. Mara?,

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
FIG. 35. Andira carvalhoi. A. Habit. B. Abaxial leaflet surface. C. Inflorescence. D. Flower. E. Calyx,
opened to show inner surface. F. Standard petal, inner surface. G. Outer surface of wing petal with lamellate
sculpturing and detail of sculpturing. H. Keel petal, inner surface. I. Androecium. J. Gynoecium, also shown
below in longitudinal section. K. Fruit. L. Fruit in section, showing wall structure. M. Seed. (Based on: A, B, R.
T Pennington 233; C, A. M. de Carvalho 491; D-J, M. P. M. de Lima et al. 20; K-L, R. T. Pennington 197.)

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 97
55 50 45 40 35
FIG. 36. Distribution o?Andira carvalhoi.
road Mara?-Porto de Campinhos Km 20, R. T. Pennington et

Oliven?a-Una, Km 42, ca. 0.5 km down road to Pedras, R. T
Mpio. Ilh?us, Road Oliven?a-Maruim, 8.9 km from Oliven?a,
Ilh?us, 10 km S Hh?us airport on road to Oliven?a, then 3 km W
Ilh?us, 8.9 km SW Oliven?a on road to Maruim, W. W. Thom
Andira carvalhoi is only likely to be confused wit

its much larger, brown fruits. It also has larger flow
of A. n?tida are 10-13 mm long and the leaf axis r
Although A. carvalhoi is abundant in some sites
considered endangered because of its restricted rang
is under great pressure for development and is hig
Moreover, the fruits of A. carvalhoi are dispersed b
1995) and may not be distributed adequately in
been demonstrated for Hymenaea courbaril (Leg
al. 1999). It seems likely that many areas of restin

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
support healthy populations of large rodents, given the high levels of disturbance of the
vegetation and proximity to centers of population.
20. Andira parviflora Ducke in Arq. Inst. Biol. Veg. 2(1): 47. 1935.?Type: Brazil.
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!).
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
ally, pattern eucamptodromous becoming brochidodromous, the brochidodromous veins
anastamosing very close to the margin, 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 1.5 mm 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
brown, granular, hard; endocarp 1.5-4 mm thick (thickened along upper side), cream to
pale brown, hard, non-fibrous. Chromosome number unknown. Fig. 37.
Phenology. Flowering February to July.
Distribution (Fig. 38). Brazil (Amazonas, Para); in terra firme forest on sandy and
clay soil, and also in low forest on white sand.
Vernacular names. Sucupira vermelha, acupu rana, sucupira chorona (Amazonas);
andira uchi (Para).
Additional Specimens Examined. Brazil. Amazonas: Manaus, Distrito Agropecuario da Suframa,

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

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 99
V?rSeUAAM yC?ltf_s
FIG. 37. Andira parviflora. A, B. Habit. C. Abaxial leaflet surface. D. Adaxial leaflet surface. E. Flower.
F. Calyx, opened to show inner surface. G. Standard petal, outer surface. H. Wing petal, outer surface. I. Keel
petal, inner surface. J. Androecium. K. Gynoecium, also shown below in longitudinal section. L. Entire fruit
(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.)

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
FIG. 38. Distribution of Andira parviflora, A. cujabensis, 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
do Taruma, Ducke RB 23866 (RB); Mpio. Manaus, 90 km NNE Manaus, Distrito Agropecuario da S?frama, re
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
(COL, F, GH, K); estrada Manaus-Caracara?, Km 60, Prance & Ramos 23563 (US); estrada Manaus-Itacoat
iara, Km 130, ?rvore XIV-104 do inventario florestal, W. Rodrigues 7990 (INPA); estrada Manaus-Itacoatiara,
Km 85, W. Rodrigues 8484 (INPA); Manaus, WWF/INPA DBFF reserve Km 51, W. Rodrigues et al 11179
(FHO, INPA, K); estrada Manaus-Caracara?, Km 30, M. F. Silva et al. 91 (F); Rio Uaup?s, island above rapids
at Ipanor?, D. W. Stevenson et al 958 (K).?Para: C.PT ?as proximidades do aeroporto, L. S. Co?lho et al. 270
(INPA); Porto Trombetas, estrada da barragem do Caran?, pr?ximo ao igarap?, E. Soares 104 (INPA).
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!).

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 101
Andira 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!).
Tree (occasionally a shrub) to 12 m; buttresses absent; bark thick, fissured; slash
honey-brown; wood 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
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
rowly obovate), coriaceous, base rounded to cordate, apex obtuse (rarely acute or
rounded), often with a short blunt acumen to 5 mm long or slightly retuse, glabrous adax
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
adaxially, raised abaxially, pattern eucamptodromous becoming brochidodromous or ?
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, 1 mm wide, caducous, with long, pale red-brown
hairs; bracteoles 1 mm 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 acuminate. Petals dirty white, the standard with mauve-purple
markings; standard blade 5-7 mm wide, 4-5 mm high, claw 2 mm long; wing 3.5-4.5 mm
long, 1.5-2.2 mm wide, sculpturing absent; keel 2.5-4 mm long, 1-2 mm wide, claw
2.5-3 mm long. Stamens 4.5-5.5 mm long, filaments united for the basal (1.8-) 2.8-3.5
mm, free for the distal 1.5-2 mm; vexillary stamen 4 mm long. Gynoecium 5-5.75 mm
long, ovary hairy on its lower surface or on both the upper and lower surfaces, or the en
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
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 to March (occasional records as early as November or
as late as May).
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, sucupira (Mato
Grosso); angelim amargoso (Goi?s); andira (Para).

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
Additional Specimens Examined. Brazil. Goi?s: 8 km N Terezina by road, W. R. Anderson 7150 (NY);
11 km E of Cavalcante by road, W. R. Anderson 7293 (NY, QCA); Serra Dourada, ca. 15 km S of Goi?s Velho,
W. R. Anderson 10051 (NY); Urua?u, Elias de Paula 3277 (UB); arredores de Guarai, Hatschbach & Kummrow
38339 (K); Serra do Caiap?, 50 km S Caiap?nia, road to Jata?, Irwin et al 17962 (INPA); Chapada dos Vead
eiros, ca. 8 km S Cavalcante, Irwin et al. 23984 (F, G, UB); 8 km S Niquel?ndia, Irwin et al. 34872 (MO, NY,
UB, US); Mpio. Piren?polis, Mac?do 4352 (US); margem direita do Rio Tocantins, canteiro de obras da Usina
Hidrel?trica da Serra da Mesa, 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 9691 (NY); Mpio. Coxim, Rio Taquar?, W. R. Anderson
11293 (SPF); Mpio. Chapada dos Guimar?es, at western edge of Chapada dos Guimar?es, W. R. Anderson 11361
(NY); Corrego de Maribondo, 22 km S of Royal Geographical Society Base Camp, de Castro R2102 (E, K, U,
UB); Mpio. Sinop, estrada para Porto dos Gauchos a 25 km da BR-163, Fazenda Mission?ria, Rio Teles Pires,
Cachoeira Cachoeir?o, Cid Ferreira et al. 6259 (F); Mpio. Cuiab?, Rio Caxipozinho, pr?ximo a Cachoeira V?u
de Noiva, Cid Ferreira et al. 6541 (INPA); Mpio. Nova Andradina, Casa Branca, Hatschbach 31876 (NY);
Mpio. Cuiab?, Parque Aguas Quentes, Hatschbach 34085 (MEXU, US); 1 km NE Garap?, Irwin & Soder str?m
6551 (F, US); Serra do Roncador, Rio Turvo, 210 km N Xavantina, Irwin et al 16179 (F, G); 15 km S Xavan
tina, Irwin et al. 16873 (F, GH); Km 197 on road from Caceres-S of Serra do Agaupei, J. H. Kirkbride & Lieras
3024 (F, US); 8 km S Xavantina, M. C. G. Kirkbride 1597 (UB); Livramento, Fazenda Rozalina, near old road,
M. Mac?do & Assump??o 2265 (UB); 100 km N Cuiab? on road to Diamantino, Maguire et al. 56394 (FHO,
NY, UB, US); Luciara, Santa Terezinha, Fazenda Santa Terezinha, perto do Lago Portugu?s, /. Mattos 15539
(SP); vicinity of Veu de Noiva, Chapada dos Guimar?es, Prance et al 18965 (K, U, US); Fazenda Santa Filom
ena, Prance & Schaller 26266 (GH, NY); 270 km N Xavantina, Royal Geographical Society Base Camp, Ramos
& 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 (K).?Mato Grosso do Sul: Mpio. Nova Andradina, MS-134 a 30 km de Nova Andradina, Leite & Klein
51 (RADAM, RB); Mpio. Nova Andradina, MS-134 a 22 km de Nova Andradina, Pastore & Klein 77
(RADAM).?Para: margems do Rio Araguaia, pr?ximo ? Santana do Araguaia, Lemes & Mileski 158
(RADAM, RB); near Reden??o, Fazenda Prof. Getulino, Ratter et al. 6869V (E); Rio Araguaia, Fazenda do Rio
Inaj?, N.T. Silva 4808 (NY, US).
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 cordata Arroyo ex R. T. Pennington & H. C. Lima, Kew Bull. 50: 562.
1995.?Type: Brazil. Bahia: 30 km W Barreiras, 12 Jan 1977, G. Hatschbach
39477 (holotype: MBM!; isotypes: G! K! MEXU!).
Tree 6 (-15) m tall, occasionally a shrub; buttresses absent; bark thick (to 1 cm),
brown 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
(or sparsely hairy); leaflets in 2-4 pairs, 2-7 cm long, 1.4^- cm wide (the lower leaflets

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 103
to 5 cm wide), broadly elliptic to ovate, (rarely suborbiculate, elliptic, narrowly ovate to

broadly ovate), 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
ally, slightly raised abaxially, pattern eucamptodromous becoming brochidodromous, ter
tiary veins plane adaxially and plane to slightly raised abaxially. Panicles 5-25 cm long,
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 the margins 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
distal 1.5-2.5 mm; vexillary stamen 3.5-4 mm long. Gynoecium 5.5-6.5 mm long,
glabrous, stipe 1.75-2.2 mm long, ovary 2-2.5 mm long, style 1.75-2 mm long; ovules 2.
Fruit 3-3.6 cm long, 2.5-2.9 cm high, 2.5-2.9 cm wide, ? globose, weighing ca. 20 g or
less when dry, green with pale yellowish specks when young, drying dark brown, almost
black, appearing smooth but minutely tuberculate (best seen with lens or microscope);
stipe 3.5 mm long; sutures and stylar remnant not apparent; mesocarp 1-2 mm thick, gran
ular, hard; endocarp 2^- mm thick. Chromosome number unknown. Figs. 3A, 4B(i), 39.
Phenology. Flowering January to April.
Distribution (Fig. 38). Brazil (Maranh?o, Bahia, Goi?s, Tocantins); in cerrado.
Vernacular name. Gr?o de galo (Bahia).
Additional Specimens Examined. Brazil. Bahia: Espig?o Mestre, ca. 100 km WSW Barreiras, W. R.
Anderson et al 36607 (NY), 36797 (NY, UB); Mpio. Correntina, Faz. Jatob?, margem da estrada que da acesso
a guarita, Aparecida da Silva 1352 (UB); Mpio. Barreiras, Elias de Paula 3122 (UB); Mpio. Correntina, Fazenda
Formoso do Guara, Rio Formoso, M. M. Fern?ndez & Collares 17 (RADAM, RB); drainage of Rio Corrente,
Rio Piau, ca. 150 km WSW Barreiras, Irwin et al 14855 (F, G); Mpio. Barreiras, 5 km NW Barreiras, Irwin et
al. 31521 (UB); Mpio. Barreiras, ca. 2 km up road to airport, R. T. Pennington & Brito 261, 262, 263, 264
(CEPEC, FHO, K); Mpio. Cocos, Lagoa do Pratud?o, S. B. da Silva & M. G. de Lima 372 (NY); Mpio. Formoso
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,
RB); Mpio. Codo, Fazenda Canto do Ro?a, /. Go?tsberger & G. Gottsberger 15-9482 (NY); Mpio. S?o
Raimundo das Mangabeiras, em dire?ao ? Balsas, C. A. Miranda & J. A. Ferreira 355 (RADAM, RB); duas
leguas de Carolina, J. M. Pires &G.A. Black 1630a (NY); near Carolina on road N to Estreito, transect area 2,
Ratter & V. P. de Lima 6716 (E); Mpio. Carolina, Transamazonian hwy, BR-230 and BR-010, 35 km N of Car
olina, Ponto Tur?stico W towards Serra de Baleia, E. L. Taylor et al E1259 (NY).?TOCANTTNS: Mpio.
Taguatinga, estrada Taguatinga-Mimoso do Oeste, Km 14, B. A. S. Pereira 1604 (NY).
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,

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
FIG. 39. Andira cordata. A. Bark of trunk. B. Portion of twig with lenticels twig. C. Habit. D. Node of
rhachis with stipels. E. Flower. F Calyx, opened to show inner surface. G. Standard petal, inner surface. H. Wing
petal, outer surface. I. Keel petal, inner surface. J. Androecium. K. Gynoecium, also shown at right in longitu
dinal section. L. Fruit. (Based on: A, R. T. Pennington 264; B-D, R. T. Pennington 262, inflorescence of C and
E^K, G. Hatschbach 39477; L, J. M. Pires 1630a.)

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 105
p. 275) with it. The combined phylogenetic analysis indicates that these taxa are closely
related (Fig. 8). Andira cordata is distinguished from A. cujabensis by its lack of indu
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.
23. Andira micrantha Ducke, Arq. Inst. Biol. Veg. 2(1): 48.1935.?Type: Brazil. Ama
zonas: Manaus, estrada do Aleixo, 10 May 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
brown with clear 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
flowers dissected 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
long, 6.5 cm high, 6-6.3 cm wide, elongated, weighing 100-300 g when dry, green ripen
ing grey to grey-brown, with mesocarp purple (at least on outer edge), drying very dark
brown, almost black, shallowly wrinkled and minutely tuberculate, with pale spots
(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
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.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
70 65 60 55 50 45 40 35
FIG. 40. Distribution of Andira micrantha, A. cori?cea, and A. grandistipula.
Additional Specimens Examined. Brazil. Amazonas: Manaus, Reserva Florestal

(INPA); Manaus, Ducke RB 21365 (F); Manaus, estrada do Taruma, Ducke RB 23401 (G,
estrada do Aleixo, Ducke RB 35077 (G, INPA); Manaus, terreno da SIDERAMA, Loureir
Mpio. Manaus, 90 km NNE Manaus, Distrito Agropecuario, reserve 1501 ("Km 41") of
Oliveira et al. 420 (K); Manaus, Reserva Florestal Ducke, quadra 23, O. Pires 29 (INPA); M
ter Egler, road Manaus-Itacoatiara, Km 64, Prance ei al 9049 (U); Manaus, estrada Mau?,
(MEXU, NY, U); Basin Rio Negro, Rio Cuieras just below mouth Rio Brancinho, Pra
Igarap? da Cachoeira Alta, W. Rodrigues & Chagas 1534 (US); Manaus, Reserva Florest
?rvore no. 1578, W. Rodrigues & Osmarino 7929 (COL, INPA); estrada Manaus-Itacoatiara
V-13 do inventario florestal, W. Rodrigues 7992 (INPA); Manaus, Cachoeira Baixa do Taru
L. Co?lho 8557 (INPA).
Andira micrantha is a rain forest species and is most likely to be confu

ri?cea and A. taurotesticulata, which also grow in this habitat and posses
These two species are not, however, sympatric with A. micrantha, which g
Amazonia in the vicinity of Manaus. Andira micrantha may be distinguis
ri?cea by the persistent stipules of the latter. These reach 15 mm long a
whereas stipules of A. micrantha are caducous (not present on any specim
and presumably tiny; no stipule scars have been observed. Andira tau
fruit that are ribbed from suture to suture and leaflets with sparse hairs
face, whereas the fruit of A. micrantha are smooth and the leaflets glabr
24. Andira cori?cea Pulle, Rec. Trav. Bot. N?erl. 6: 267. 1909.?Ty
Boschreserve Sectie O, tree number 61, 15 May 1907, Boschbeh
(lectotype, here designated: U!).

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 107
Andira wachenheimii R. Benoist, Bull. Mus. Hist. Nat. (Paris) 25(4): 296. 1919.?
Type: French Guiana. Maroni, Wachenheim 79 (lectotype, here designated: P!;
isolectotype: P!).
Tree 25 (^10) m tall, with small buttresses; bark grey-brown, Assuring vertically,
flaking; slash orange-brown; twigs glabrous, very dark brown, almost black, with nu
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 becoming brochidodromous; tertiary veins
plane adaxially and abaxially. Panicles 10-22 cm long, axillary and terminal, branch
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 at margins. 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. Flowering May to October (May, Suriname; August-October, French
Guiana).
Distribution (Fig. 40). French Guiana and Suriname; in rain forest.
Vernacular names. Kobon, roode kabbes, reddie kabbes, koeraroe (Arawak), akoeli
tjerere (Suriname); St. Martin rouge, lebi kiabici/kiabici lebi (Par?maca; French Guiana).
Additional Specimens Examined. Suriname. Sectie O, B. W. 1909 (COL, U, US); Sectie O, tree 61,
B. W. 3933 (U); Forest Reserve Zanderij 1, tree 5, B. W. 4092 (U); Zanderij I, B.W. 5450 (U); Brownsberg,
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 & Jonker 868 (U); Sabanpassie in bosje op te savanna ten O. van de Brinck
heuvel, Heyligeri 158 (U); via secta ab Moengo 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);
Sabanpasi-savannecomplex, Naturreservaat Brinkheuvel, Teunissen & Wildschut 11830 (U); Dist. Broko
pondo, 8 km ESE of Brownsberg, van Donselaar 1743 (U). French Guiana. St. Laurent, BAFOG 5IN (U);
Cayenne, BAFOG 1201 (U); Cayenne, BAFOG 1264 (U); route de l'Est, near the bridge on the Comte River,
ca. 52 km S Cayenne, Champagne 56 (NY); piste de St. Elie-interfluve Sinnamary/Counamama Km 15.7,
Orstom camp, Pr?vost 3046 (B, CAY, K); route de Saint Laurent ? Cayenne, Km 12, 50M (U); route de
Charvein ? 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 et en bordure de la route, 7699 (U); route de l'Acarouany, Km 4.300, c?t? gauche et en
bordure de la route, 7733 (U).

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
Andira cori?cea is most likely to be confused with A. taurotesticulata (no. 5) and A.

micrantha (no. 23), the other rain forest species with large fruit; see those species for a
discussion of their differences.
CTFT (1989) reported that the timber of A. cori?cea is used in the construction of
wooden houses and has commercial potential for the manufacture of items such as
snooker cues and umbrella handles.
Pulle (1909) cited two collections with his description of A. cori?cea. The flowering
specimen, collected on 15 May 1906, is selected here as the lectotype.
25. Andira grandistipula Amshoff, Bull. Torrey Bot. Club 75(4): 394. 1948.?Type:
Guyana. Kaieteur Plateau, Kaieteur Savannas, 6 May 1944, B. Maguire & D. B.
Fanshawe 23273 (holotype: NY!; isotypes: A! F! K! U!).
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
black, 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
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
mous, tertiary veins plane adaxially and plane to slightly raised abaxially. Panicles
17-32 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.
15 mm 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. 10 mm 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
ripe, drying pale brown and dark brown, some areas somewhat yellow, appearing
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);
mesocarp 5-12 mm thick, greyish brown, hard, finely granular, often splitting along
upper suture when dry; stylar remnant minute; endocarp 3-13 mm thick, cream-buff,
non-fibrous, very hard. Chromosome number unknown.
Phenology. Flowering in May (single record); two fruiting records in June and
November.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 109
Distribution (Fig. 40). Guyana, restricted to the Pakaraima Mountains; in scrub in

areas of savanna.
Additional Specimens Examined. Guyana. Mazaruni-Potaro Region: Pakaraima Mts, Imbaimadai,

Hoffman 1992 (COL, K, US); Pakaraima Mts, Imbaimadai Creek W of Imbaimadai, Pipoly 7972 (U, US);
Pakaraima Mts, Mt. Membaru, Maas & Westra 4299 (K, U); Upper Mazaruni River Basin, Partang Rapids, near
mouth Partang River, Maguire et al 43878 (NY-2 sheets).
Andira grandistipula is distinguished by its large, persistent stipules from the other
eight species of clade II of Andira (defined by an endocarp of stone cells). None of the
collections examined had fully mature flowers, and the petal color is unknown.
26. Andira praecox Arroyo ex R. T. Pennington, sp. nov.?Type: Brazil. 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!).
Ab Andira tervequinata floribus majoribus et statura altiore recedit (arbor grandis

usque ad 30 m alta, non frutex vel arbuscula usque ad 7 m alta).
Tree to 30 m 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 1 mm wide, caducous, with appressed, red-brown hairs;
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,
thick-chartaceous to subcoriaceous, base obtuse (rarely acute or rounded), slightly decur
rent, rounded (rarely obtuse), apex slightly 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,
plane adaxially, raised abaxially, pattern eucamptodromous becoming brochidodromous,
or ? completely brochidodromous, 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. 3 mm 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
black, slightly wrinkled (with brown pock-marks where the surface is broken) and
minutely tuberculate with a slight waxy bloom (best seen with lens or microscope); stipe
to 9 mm 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
fibrous, very hard. Chromosome number unknown. Fig. 41.

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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.)

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 111
FIG. 42. Distribution of Andira praecox, A. tervequinata, A. trifoliolaia, and A. unifoliolata.
Phenology. Flowering in September (single record); two fruiting records in Decem

ber and January.
Distribution (Fig. 42). Brazil (Amap?); in terra firme forest.
Vernacular names. Andiroba jaruba, sucupira vermelha.
Additional Specimens Examined. Brazil. Amap?: Mpio. Mazag?o, BR-156, incomplete road toward
Monte Dourado, 75-80 km WSW Macap?, 5-10 km SW Rio Preto, Daly & Cardoso 3923 (K, NY); Mpio.
Macap?, road Cupixi-Rio Vila Nova, 18 km SSW Cupixi, Rabelo et al 3199 (ULM, US); Serra do Navio, Ro
drigues 2917 (TNPA).
Andira praecox is most similar to A. 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.
praecox, leaflets are narrowly obovate (rarely elliptic), but in A. tervequinata they are
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

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black, whereas the fruit of A. tervequinata is larger (5.5 cm long) and generally more red
brown with pale, raised specks.
27. Andira tervequinata R. T. Pennington, G. Aymard & N. Cuello, Novon 7: 72.

1997.?Type: Venezuela. Bol?var: Distrito Heres, west bank of R?o Trueno
Alto, 35 km W of Chiguao, high plateau, ecotone between gallery forest and sa
vanna, 23 Mar 1985, Huber 10345 (holotype: NY!; isotypes AAU, E! HBG,
INPA, K, MO, MY, MYF! P, PORT, RB, US, VEN).
Small tree or shrub to 7 m tall; presence of buttresses unknown; bark and slash un
known; twigs very dark brown with raised elongated lenticels, outer bark very dark brown
(almost black), this layer often flaking to reveal red-brown beneath, with sparse, red
brown, ? appressed hairs, glabrescent. Stipules to 7 mm long, to 1 mm 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 1 mm 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
high, ? globose, weighing ca. 20 g or less when dry, green or grey-green, drying dark
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 in March and with young and mature fruits in May.
Distribution (Fig. 42). Venezuela (Bol?var: Distrito Heres and Distrito Piar); in
gallery forest, in shrub and tree islands in savanna dominated by Trachypogon plumosus
(G. Aymard, pers. comm; this vegetation was on conglomerated 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 Tepui, Aymard 6147 (MO, NY, PORT); Distrito Piar, R?o Aparam?n at Kambay
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).

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

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Andira tervequinata might be confused with A. trifoliolata (no. 28) and A. praecox
(no. 26); see notes under the latter. Differences from A. trifoliolata were discussed in de
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
short, appressed hairs on the undersurface, a channelled primary vein on the upper sur
face, and a retuse or emarginate apex.
28. Andira trifoliolata Ducke, Arq. Inst. Biol. Veg. 4(1): 22. 1938.?Type: Brazil.
Amazonas: Rio Curicuriare, 20 Feb 1936, A. Ducke RB 35079 (lectotype, here
designated: RB!; isolectotypes: G! K! U!).
Tree to 20 (-30) m tall; presence of buttresses unknown; bark dark brown, slash
yellow; wood hard, yellow, fibrous; twigs dark brown (or almost black), sparsely hairy,
glabrescent; occasional pale, elongated lenticels. Stipules to 9 mm long, to 1 mm wide,
brown, caducous, with appressed, red-brown hairs at tips, glabrous near base; leaves
pinnately trifoliolate, leaf axis 2.5-9 (-12) cm long. Rhachis dark brown, this layer
peeling to reveal pale brown beneath, glabrous; stipels 1 mm 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
brochidodromous or ? eucamptodromous or mixed, tertiary veins plane adaxially and
abaxially. Panicles axillary or terminal, 6.5-17.5 cm 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,
vexillary stamen 4.5-5 mm long. Gynoecium 7.2-8.5 mm long, the ovary sparsely hairy
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, ? globose, weighing ca. 20 g or less when dry, green, drying dark
brown or almost black, appearing smooth but minutely wrinkled (best seen with lens or
microscope), with pale pock-marks where surface is broken; stipe 15-25 mm long; su
ture a thin raised line adaxially, not visible abaxially; stylar remnant tiny; mesocarp
1.5-2 mm thick, pale to dark brown (or greenish), hard, granular; endocarp 2-3 mm
thick, pale buff, very hard, non-fibrous. Chromosome number unknown. Fig. 4B.
Phenology. Flowers recorded in February, June, July, and October.
Distribution (Fig. 42). Colombia (Guainia); Venezuela (Amazonas); Brazil (Ama
zonas); along rivers, but also in terra firme and secondary forest.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 115
Additional Specimens Examined. Colombia. Guain?a: R?o Atabapo, Region Cacaual, R. Rodrigues &
Acero 221 (COL, INPA, TJDBC-2 sheets). Venezuela. Amazonas: San Carlos de R?o Negro, forest near mu
nicipal playing field, Christenson et al 1432 (US); Depto. Casiquiare, R?o Casiquiare, Laguna del Pucibe and
area, Colella et al 2020 (FHO, NY); Depto. Atabapo, La Esmeralda, 5 km N village, near savanna "morichal,"
Coomes 81 (K); Depto. Rio Negro, Mucuriapi, small laja along lower part of R?o Baria, Davidse & Miller 26752
(NY); Depto. Atabapo, L. Delgado 551 (FHO); road from San Fernando de Atabapo to Santa Barbara, 12^0
km from San Fernando, Gentry & Tillett 10889 (MO); Depto. Atabapo, R?o Cunucunuma, entre las Comu
nidades de Culebra y Huachamacari, Steyermark et al. 125856 (NY); Rio Casiquiare, near Solano, Wurdack &
Adderley 43368 (F, NY, U, US). Brazil. AMAZONAS: Barcelos, Ducke RB 17293 (RB, K); Rio Negro, Rio
Curicuari, O.C. Nascimento 764 (NY); estrada Manaus-Porto Velho, trecho entre os Rios Castanho e Tupana,
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).
29. Andira unifoliolata Ducke, Arq. Inst. Biol. Veg. 4(1): 22. 1938.?Type: Brazil.
Amazonas: Manaus, estrada do Aleixo, 3 Mar 1937, A. Ducke RB 35078 (holo
type: RB!; isotypes: G! K! INPA! RB-5 sheets! U!).
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
beneath; stipels to 2 mm long (seen only on seedlings); petiolule 3.5-6 mm long,
glabrous; unifoliolate (1-2 pairs leaflets in seedlings), leaflet 4.7-13 cm long, 1.3-5 cm
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,
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,

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
i\??M?^ L?\r\^
FIG. 44. Andira unifoliolata. A. Habit. B. Abaxial leaflet surface. C. Flower. D. Calyx, opened to show
inner surface. E. Standard, inner surface. F. Wing petal, outer surface. G. Keel petal, inner surface. H. Androe
cium. I. Gynoecium, also shown below in longitudinal section. J. Fruit. K. Endocarp. (Based on: A-I, A. Ducke
61 A; J. A. Ducke 30578.)

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2003 andira 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,
turning yellow, sweet-smelling, very dark brown, almost black, with a slight waxy bloom,
very slightly wrinkled; stipe 3-4 mm long; suture a thin line adaxially, not detectable
abaxially; stylar remnant tiny; mesocarp pale reddish brown, hard, granular; endocarp un
known. Chromosome number unknown. Figs. IC, 2F, 44.
Phenology. Flowering February to March.
Distribution (Fig. 42). Brazil (Amazonas, Para, Roraima); in terra firme rain forest on
sandy and clay soils.
Vernacular names. Sucupira amarela, sucupira chorona, sucupira vermelha (Ama
zonas); andira-uchi (Para).
Additional Specimens Examined. Brazil. Amazonas: Manaus, Reserva Florestal Ducke, Adair s.n.
(INPA); Manaus, Reserva Florestal Ducke, ?rvore do fenel?gico 20, Alu?sio s.n. (INPA); Mpio. Mau?s, Rio
Apoquitaua ?cima do lugar S?o Sebasti?o, Cid Ferreira 4268 (INPA); Manaus, estrada do Aleixo, Ducke 674
(F, GH, K, NY, US); Manaus, Reserva Florestal Ducke, Lourival s.n. (INPA), Mac?do s.n. (INPA), F. C. Mello
et al. s.n. (INPA); Manaus, Reserva Florestal Ducke; estrada Manaus-Itacoatiara, Km 133, Monteiro & F.
Mello s.n. (INPA); Manaus, Reserva Florestal Ducke, W. Rodrigues & Loureiro 5953 (INPA); Manaus,
Reserva Florestal Ducke, ?rvore do fenel?gico 13, W. Rodrigues & D. Co?lho 7554 (INPA); estrada Man
aus-Itacoatiara, 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);
estrada Manaus-Porto Velho, Rio Castanho, margem direita, estrada para o Careiro, M. F. Silva e? al. 373
(INPA); Manaus, Reserva Florestal Ducke, estudo de associa?ao, J. A. C. Souza s.n. (INPA).?Para: Rio
Trombetas, entre as cachoeiras de Tremalhetinha e Jandi?, L. S. Co?lho 103 (INPA); Porto Trombetas, estrada
do Caran?, entrada para a Barragem Velha, E. Soares 5 (INPA).?Roraima: UHE de Samuel, are? do futuro
reservatorio, Maciel 104 (INPA, RB, UB).
Unifoliolate leaves are unique to A. unifoliolata, and this species cannot be confused
with any other of Andira. Morphologically it is most similar to A. trifoliolata. The two
species are easily distinguished vegetatively: A. trifoliolata is uniformly trifoliolate,
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
timber.
Doubtful and Excluded Names
Andira sect. Aristobulia Bentham, Comm. legum. gen. 43. 1837.?LECTOTYPE, here des
ignated: Andira amazonum Martius ex Bentham. = Watairea guianensis Aublet, Hist,
pi. Guiane 2: 755, t. 302. 1775.
Andira amazonum Martius ex Bentham, Comm. legum. gen. 43. 1837.?TYPE: BRAZIL.
Rio Negro, Martius s.n. (holotype: BR!; isotype: M). = Vatairea guianensis Aublet,
Hist. pi. Guiane 2: 755, t. 302. 1775; see also de Lima (1982).
Andira 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. =

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
Vataireopsis araroba (Aguair) Ducke, Ann. Acad. Brasil. Ciencias 8: 26. 1936; see
also de Lima (1980).
Andira aubletii Bentham, Comm. legum. gen. 44. 1837, nom superfl. = Vouacapoua
americana Aublet, Hist. pi. Guiane, suppl. 9. 1775.
Andira bracteosa Bentham, Comm. legum. gen. 43. 1837.?Type: Brazil. 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 32 (holotype: P!).

= Aganope gabonica (B?illon) Polhill, Kew Bull. 25: 269. 1971.
Andira horsfieldii Leschenault, Ann. Mus. Nati. Hist. (Paris) 16: 481, t.12. 1810.?TYPE:
Leschenault, Ann. Mus. Nati. Hist. (Paris) 16: 481, t.12.1810. = Euchresta horsfieldii
(Leschenault) Bennett, PL jav. rar. 148, pi. 31. 1838.
Andira racemosa Lamarck ex J. St. Hilaire, Diet. Sei. Nat. 2: 137. 1804, nom. superfl. =
Vouacapoua americana Aublet, Hist. pi. Guiane, suppl. 9. 1775.
Andira villosa Kleinhoonte, Rec. Trav. Bot. Ne?rl. 22: 404.1925.?Type: Suriname. Am
Coppenameflusse bei dem Raleighstrom, Aug 1920, Pulle 319 (holotype: U!).?The
type does not belong Andira, and the floral morphology reveals that it does not clearly
fit any genus of Papilionoideae. It matches a specimen from Peru (Loreto: Iquitos,
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 zehntneri Harms, Repert. Spec. Nov. Regni Veg. 17: 443. 1921.?Type: Brazil.
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 supported initially by the Science and Engineering Research Council (SERC)
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
Caddick. Fieldwork in Guyana was supported by The Commonwealth and Government of Guyana Iwokrama
Rain Forest Programme. I am grateful to my Ph.D. advisers Roger Polhill, Robert Scotland, Chris Leaver, and
Mike Bennett, who supervised the start of this research. I particularly acknowledge the guidance of Brian Styles,
who very sadly died before my thesis work was complete. I also thank: Maureen Warwick (artwork), Robert Mill

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 119
(Latin diagnoses), Mark Chase (advice on molecular work and cladistic analysis), Tony Cox (molecular work),
Peter Gasson (fruit anatomy), Bente Klitgaard (pollen data), Iain Prance (biogeography), David Harris (numer
ous discussions and reading an entire draft), David Mabberley, Richard Brummitt, Mark Watson, and Barbara
MacKinder (nomenclatural issues), Denis Filer (databasing), Peter Hollingsworth (constructive comments on in
traspecific cpDNA polymorphism), Colin Hughes (whose monograph of Leucaena was an inspiration for this
work, and who collected Andira inermis in Mexico), Serena Marner, Anne Sing, Alison Strugnell (help with ad
ministrative problems), Gwilym Lewis (advice on all topics and particularly for invaluable help on my arrival
in Brazil), Lourdes Rico (who collected leaves and fruit of Mexican species), Clive Foster, Phil Griffiths (who
expertly grew species of Andira), Ary Oliveira-Filho (discussion of Brazilian species), Eimear Nie Lughadha,
Charlie Stirton (arrangements for fieldwork in Guyana), Mario Sousa (hospitality in Mexico, discussion of Mex
ican species), Martin Cheek (who collected Andira inermis in Africa), Jeff Doyle (molecular systematics), Larry
Kelly, Anne Bruneau (cladistics and critically reading portions of this manuscript), Jane Doyle, Paul Manos,
Beth Dickson, Karen Hanson, Robert Soreng, Stephan Maumont (molecular systematics), Kevin Nixon (cladis
tic advice and copies of Clados and Dada). For my fieldwork, I particularly thank Andr? Carvalho for help and
enormous hospitality, and also Haroldo Cavalcante de Lima, Alvaro Cogollo, Jim Ratter, Sam Bridgewater, and
Felipe Ribeiro. For help with fieldwork I also thank Fernando ("Guga") Carvalho, Herminio Brito, Taiman dos
Santos, Lenise Guedes, Isolde Ferraz, W. Wayt Thomas, Bruce Nelson, William Rodrigues, Niro Higuchi, Jo?o
Alu?sio, Alex Monro, Vanessa Plana, Fernando Montenegro, Milton Aulestia, Lino Veloz, Terry Pennington,
David Neill, Tamara N??ez, Efrain Frere, Mar?a Margarida Fiuza de Mel?, M. Sugiyama, Nelson Zamora, Ma
noel Claudio da Silva, Jes?s Idrobo, Jos? Luis Fern?ndez, Bruce Hoffman (especially for collecting Andira gran
disiipula), and Isaac Johnson. All botanical drawings were provided by Rosemary Wise. The following herbaria
provided material and/or facilities for this study: A, BM, BR, CEPEC, COL, CR, E, F, FHO, G, GH, HUEFS,
INBIO, INPA, JAUM, K, M, MEXU, MO, NY, OXF, P, PR, PRC, QCA, QCNE, R, RB, RADAM, SP, U, UB,
US, USP.
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Wojciechowski, M. F, M. J. Sanderson, B. G. Baldwin, and M. J. Donoghue. 1993. Monophyly of aneuploid As
tragalus (Fabaceae): evidence from nuclear ribosomal DNA internal transcribed spacer sequences. Amer.
J. Bot. 80:711-722.
Wright, W. 1777. Description and use of the cabbage-bark tree of Jamaica. Philos. Trans. 67: 507-512.
-1787. An account of the medicinal plants growing in Jamaica. Lond. Med. J. 8: 217-295.

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
APPENDIX 1
Restriction Sites
Restriction site characters used to assess phylogenetic position of species of Andira not included in the
study by Pennington (1995).
Character2 Enzyme Probe regionb Mutation0 Site Diagnostic For
3 Asel MB8+9 3.6=1.8+1.7 Clade H (site absence)

4 Asel MB8+9 1.8 = 1.05+ n.d. Clade III (site presence)
5 Asel MB11 2.0=1.7 + n.d. Clade I (site presence)
6 Asel MB11 3.7 = 2.0+1.7 Clade m (site absence)
7 Asel MB 13 1.0 = 0.6 + 0.4 Clade I (site absence)
8 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
19 HindHI MB2 5.2 = 4.0+1.2 Clade H (site presence)
20 Hindm MB8+9 3.6 = 3.2 + n.d. Clade I (site presence)
34 XmnI MB3 1.8 = 1.2 + 0.6 A. verm?fuga; A. humilis
RTP 269 (site presence)
35 XmnI MB11 0.96 = 0.6 + n.d. Clade I and Clade H
(homoplasy; site absence)
37 XmnI MB13 3.5 = 3.4 + n.d. Clade m (site presence)
Character numbers of those of Pennington (1995, Table 2).

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

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 125
APPENDIX 2
Inferred Plastome Types of Andira Accessions
The species are listed alphabetically and the accessions of each species ordered numerically; collectors are
abbreviated: AMC = A. M. de Carvalho; CEH = C. E. Hughes; HCL = Haroldo Cavalcante de Lima; JR = J. A.
Ratter; LR = L. Rico; MC = M. Cheek; MCh = Mark Chase; MdS = Man l Claudio da Silva; MS = M.
Sugiyama; RTP = R. T. Pennington; TDP = T. D. Pennington. Accessions marked with an asterisk are those
screened in this study; the remainder are those screened previously by Pennington (1995). Notations in the col
umn marked "Clade": I = restriction site mutations characteristic of clade I; II = restriction site mutations char
acteristic of clade II; m = restriction site mutations characteristic of clade III; PI = restriction site mutations char
acteristic of plastome group I; PITH restriction site mutations characteristic of the monophyletic group of
plastome group I and clade II [these accessions were screened previously by Pennington (1995), but no muta
tions diagnostic for clade II were screened].
Species Voucher Herbarium Population Clade Locality

A. anthelmia RTP 227 CEPEC, FHO, K ffl Brazil: Bahia;
14?19'S, 39?21'W
A. anthelmia RTP 281 CEPEC, FHO, K m Brazil: Bahia;
14?51'S, 39?04'W
14?51'S, 39?04'W
15?06'S, 39?33'W
15?05'S, 39?33'W
A. anthelmia RTP 294 CEPEC, FHO, K in Brazil: Bahia;
15?06'S, 39?24'W
A. carvalhoi RTP 217 CEPEC, FHO, K pi/m Brazil: Bahia;
13?58'S, 39?01,W
A. carvalhoi RTP 229 CEPEC, FHO, K pi Brazil: Bahia;
14?57'S, 39?01'W
A. carvalhoi RTP 233 CEPEC, FHO, K m Brazil: Bahia;
15?10'S, 39?06'W
A. carvalhoi AMC* CEPEC pi Brazil: Bahia;
14?57'S, 39?01'W
A. cordata RTP 262 CEPEC, FHO, K pi/n Brazil: Bahia;
12?07'S, 45?06'W
A. cordata RTP 263 CEPEC, FHO, K ii Brazil: Bahia;
12?07'S, 45?06'W
A. cordata RTP 264 CEPEC, FHO, K n Brazil: Bahia;
12?0rS, 45?06'W
A. cujabensis RTP 467* E,UB n Brazil:Goi?s;
16?44'S, 52^7^
A. cujabensis RTP 474* E,UB ii Brazil: Goi?s;
16?52'S, 52^0^
A. cujabensis RTP 478* E,UB n Brazil: Goi?s;
16?30'S, 52^3^
A. cujabensis RTP 485* E,UB ii Brazil: Goi?s;
16?44'S, 52?37'W

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A. cujabensis RTP 486* E,UB 1 n Brazil: Goi?s;

16?44,S, 52?37'W
16?44'S, 52?37'W
13?55'S, 47*23^
A. cujabensis RTP 498* E,UB 2 ii Brazil: Goi?s;
13?55'S, 47?23'W
A. cujabensis RTP 503* E,UB 2 ii Brazil: Goi?s;
13?55'S, 47?23'W
A. fraxinifolia RTP 213 CEPEC, FHO, K 1 in Brazil: Bahia;
14?12'S, 39?03'W
15?10'S, 39?06'W
A. fraxinifolia RTP 274 CEPEC, FHO, K 3 m Brazil: Bahia;
12?35'S, 41?16'W
12?32'S, 41?14'W
14?56'S, 39?01'W
A. fraxinifolia RTP 318 CEPEC, FHO, K 4 ni Brazil: Bahia;
14?55'S, 39?01'W
15?05'S, 39?33'W
15?05'S, 39?33,W
17?13'S, 39?14'W
17?30'S, 39?!^
17?30'S, 39?!!^
A. fraxinifolia RTP 249* CEPEC, FHO, K 7 m Brazl: Bahia;
llo02'S,40?43'W
11?02'S,40?43,W
A. fraxinifolia MS 889 K,SP 8 ni Brazil: S?o Paulo;
23?S, 38?W
A. galeottiana LR s.n. K 1 pi Mexico: Oaxaca;
17?N, 95?W
A. galeottiana MCh s.n.* E 2 pi Mexico: Chiapas;
17?N, 91?W
A. humilis RTP 239 CEPEC, FHO, K 1 in Brazil: Bahia;
12?15'S, 38^7^
A. humilis RTP 246 CEPEC, FHO, K 1 m Brazil: Bahia;
11?09'S,40?32,W
A. humilis RTP 247 CEPEC, FHO, K 1 m Brazil: Bahia;
llo09,S,40?32/W

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2003 ANDIRA 127

Ahumilis RTP 267* CEPEC, FHO, K 2 PI Brazil: Bahia;
12?10'S, 44^3^
Ahumilis RTP 268* CEPEC, FHO, K 2 PMI Brazil: Bahia;
12?10'S, 44?43'W
Ahumilis RTP 269 CEPEC, FHO, K 2 PI Brazil: Bahia;
12?10'S, 44?43'W
Ahumilis RTP 499* E, UB 3 PI Brazil: Goi?s;
13?55'S, 47?23/W
A. humilis RTP 500* E, UB 3 PI Brazil: Goi?s;
13?55'S, 47^3^
13?55'S, 47?23'W
13?55'S, 47?23'W
14?07,S,47?31/W
14?07,S,47?31'W
14?07'S, 47?31'W
14?07'S, 47?31'W
A. humilis MdS 1* E 5 PI Brazil: D. F.;
15?55'S, 47?50'W
Ahumilis MdS 2* E 5 PI Brazil: D. F;
15?55'S, 47?5(TW
Ahumilis MdS 3* E 5 PI Brazil: D. F. ;
15?55'S, 47?5(TW
Ahumilis MdS 4* E 5 PI Brazil: D. F.;
15?55'S, 47?5(W
Ahumilis MdS 5* E 5 PI Brazil: D. F.;
15?55'S, 47?50/W
A. inermis TDP13558 K 1 I Costa Rica: Puntarenas;
09?07'N, 83?19/W
A. m^w CEH1673 FHO, K 2 I Mexico: Oaxaca;
16?22/N,94?11'W
A. miroi? MC 3579 K 3 I Cameroon, SW Province;
03?58'N, 09?16'E
A. inermis RTP 512* E, QCNE 4 I Ecuador: Napo;
00?33'S, 77?50^
A. merm?5 RTP 580* E, INBIO 4 I Costa Rica: Puntarenas;
08?42,N, 83^0^
A. ?n*/row RTP 589* E, INBIO 5 I Costa Rica: Puntarenas;
08?42'N, 83?3(TW
A. te^a/w RTP 305 CEPEC, FHO, K 1 HI Brazil: Bahia;
17?30'S, 39?!^
Alegalis RTP 307 CEPEC, FHO, K 1 ffl Brazil: Bahia;
17?30'S, 39?!!^

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A. legalis RTP 308 CEPEC, FHO, K ffl Brazil: Bahia;
17?30,S,39?11'W
A. legalis HCL s.n. E m Brazil: Rio de Janeiro;
22?55'S, 43?10'W
A. macrothyrsa TDP 13550 K PI Peru: Loreto;
00?S, 73?W
A. macroihyrsa RTP 523* E,QCNE PI Ecuador: Napo;
00?40'S, 77?10'W
A. marauensis AMC s.n. CEPEC ffl Brazil: Bahia;
15?09'S, 39?05'W
A. nilida RTP 300 CEPEC, FHO, K ffl Brazil: Bahia;
17?30'S, 39?13'W
A. niiida RTP 304 CEPEC, FHO, K ffl Brazil: Bahia;
17?30'S, 39?!^
A. niiida RTP 311 CEPEC, FHO, K ffl Brazil: Bahia;
17?30'S, 39?!^
A. niiida RTP 313 CEPEC, FHO, K III Brazil: Bahia;
17?13'S, 39?14'W
A. nitida RTP 288 CEPEC, FHO, K ffl Brazil: Bahia;
13?02'S, 38^3^
13?02'S, 38?23'W
13?02'S, 38?23"W
17?30'S, 39?13/W
15?06'S, 30?\1^
15?10'S, 39?06'W
A. nitida AMC 3309 CEPEC ffl Brazil: Bahia;
15?18'S, 39?04'W
A. surinamensis RTP 433 FHO, K, U, US ffl Guyana:
0A?\3^, 58^8^
04?13'N, 58*58^
0A?\3f"H, 58^8^
04?09,N, 59^2^
A. verm?fuga RTP 256* CEPEC, FHO, K PI Brazil: Bahia;
11?15/S,42?52,W
11?15,S,42?52,W
ll015'S, 42^2^
A. verm?fuga RTP 259 CEPEC, FHO, K PI Brazil: Bahia;
11?15'S, 42^2^

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2003 ANDIRA 129
A. verm?fuga RTP 265 CEPEC, FHO, K 2 PI Brazil: Bahia;

12?07'S, 45?06'W
A. verm?fuga RTP 270 CEPEC, FHO, K 3 PI/H Brazil: Bahia;
12?07'S, 43?15'W
A verm?fuga RTP 271 CEPEC, FHO, K 3 PI Brazil: Bahia;
12?07'S, 43?15'W
A. verm?fuga RTP 272 CEPEC, FHO, K 3 PI/II Brazil: Bahia;
12?07'S, 43?15'W
A. verm?fuga RTP 273 CEPEC, FHO, K 3 PITH Brazil: Bahia;
12?07'S, 43?15'W
A. verm?fuga RTP 465* E,UB 4 PI Brazil: Goi?s;
16?52'S, 52?20'W
16?52'S, 52^0^
16?52'S, 52^0^
A verm?fuga RTP 471* E,UB 4 PI Brazil: Goi?s;
16?52'S, 52?20'W
14?07'S, 4713^
A. verm?fuga JR 7232V* E 5 PI Brazil: Goi?s;
14?07'S, 47?13'W
NUMERICAL LIST OF SPECIES
1. A. inermis (subspecies not determined) 14. A. legalis

la. A. inermis subsp. inermis 15. A. ormosioides
lb. A. inermis subsp. glabricalyx 16. A. fraxinifolia
lc. A. inermis subsp. rooseveltii 17. A. nitida
2. A. jaliscensis 18. A. marauensis
3. A. multistipula 19. A. carvalhoi
4. A. cubensis 20. A. parviflora
5. A. taurotesticulata 21. A. cujabensis
6. A. macrothyrsa 22. A. cordata
7. A. chigorodensis 23. A. micrantha
8. A. galeottiana 24. A. cori?cea
9. A. verm?fuga 25. A. grandistipula
10. A. humilis 26. A. praecox
11. A. macrocarpa 27. A. tervequinata
12. A. surinamensis 28. A. trifoliolata
13. A. anthelmia 29. A. unifoliolata

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INDEX TO NUMBERED COLLECTIONS EXAMINED
The numbers in parentheses refer to the corresponding species in the text and in the Numerical List of
Species presented above.
Acevedo, P., & A. Sinca 3775 (la). Aubreville, A., & W. Rodrigues 681 (12).
Acevedo, R., & J. Sosa 1070 (lb). Ayala, F., et al. 3102(3).
Acevedo-Rodr?guez, P., & J. A. Cede?o 7430 (3). Aymard, G. 6147 (27); 7991 (12); 9263 (28).
Acevedo-Rodr?guez, P., et al. 8127 (28); 8357 (28). Aymard, G., ?fe L. Delgado 8233 (12).
Ackerly, D., et al. INPA/WWFl302.4500.2 (20). Azuara, B. 133 (la).
Adams, C. D. 7843 (la). Azurdia, C, et al. 700 (la).
Aguilar, R. 228 (la). BAFOG 51N (24); 1183 (12); 1201 (24); 1264 (24).
Alain 4404 (4). Bailey, L. H., <fc E. Z. Bailey 52 (la); 149 (la); 321
Albert de Escobar, L., & J. I. Santa 3527 (la). (la); 1352 (12); 1657 (12).
Albert de Escobar, L., et al. 1848 (5). Baitello, J. B., & J. R. Guilamon 7751 (16).
Alencar (RB 54889) (12); (RB 61016) (9). Baker, M. A. 7025 (5).
Allard, H. A. 13367 (la); 13885 (la). Balslev, H. et al., 84821 (la); 97505 (3).
Allen, P. H. 873 (la); 938 (la); 1770 (la); 2955 (la); Bamps, P. 5078 (10).
4435 (la); 4470 (la); 4482 (la); 5220 (la); 5458 Bangham, W. N. 393 (la); 432 (la).
(la); 5609 (la); 7190 (la). Barbosa, C. 1090 (la).
Almeida, J. 12 (13); 356 (17); 2044 (14). Barbosa, C, et al. 1856 (la).
Almeida, J., & T. S. dos Santos 103 (17); 130 (17). Barkley, F. A., & M. Hern?ndez 40602 (la).
Alu?sio, J. 115 (20); 224 (29); 226 (29). Barlow, F. D., 30/186D (8).
Alvarenga, D., et al. 194 (9). Barreto, M. 937 (16); 1515 (16); 1782 (16); 5485 (10);
Amaral, D. L. 145 (21). 5491 (10); 5590 (16); 5592 (16); 5608 (16);
Amorim, A. M. A., et al. 347 (17); 348 (17); 1055 5764 (16); 10096 (16).
(16); 1238 (18); 1411 (18); 1896 (16); 1908 (18). Bartlett, H. H., ?fe T. Lasser 16319 (la).
Anderson, C. W. 332 (12). Bautista, H. P. 533 (17); 1145 (17); 1368 (17).
Anderson, W. R. 6231 (10); 6456 (10); 6771 (21); Beaman, J. H. 6332 (la).
7150 (21); 7293 (21); 7294 (9); 7895 (10); 9560 Beard, J. S. 157 (la).
(21); 9691 (21); 10051 (21); 11292 (la); 11293 Beard, P. 1336 (la).
(21); 11361 (21); 36797 (22). Beck, S. G. 18686 (la).
Anderson, W. R., et al. 7159 (10); 36024 (16); 36607 Beck, S. G., et al. 19556 (3).
(22); 36865 (10). Becker, R. M. 42 (21).
Andrade-Lima & Medeiros-Costa 105 (17); 154 (17). Belem, R. P. 1830 (14); 1879 (13).
Andrade-Lima et al. 7 (16). Belem, R. P., ?fe R. S. Pinheiro 2445 (16); 2550 (16);
Andrews, F. W. 464 (le). 2591 (16); 2807 (17); 2949 (13); 2988 (17);
Andrews, L. M. 3-7 (la). 3089 (18); 3091 (18); 3142 (18).
Antonio, N., & M. Heinrich GUI224 (8). Belshaw, C. M. 3370 (la).
Antonio, T. 3616 (la). Bena,P. 1167(12).
Aparecida da Silva, M., et al. 1352 (22); 2738 (9); Benitez-Paredes, A. 3658 (lb); 3796 (lb).
3441 (9). Bernai, H. Y. 158 (la).
Arana, A. 27 (la). Black, G. A. 54-17372 (la).
Araujo, D. 1843 (13); 3553 (16); 4612 (14); 5289 Black, G. A., & Klein 54-17197 (la); 54-17372 (la).
(16); 6176 (16); 6598 (16); 7390 (16); 8079 Black, G. A., & P. Ledoux 50-10366 (la); 50-10386
(14); 8295 (16); 9687 (16); 9763 (16). (12).
Araujo, D., & R. Henriques 4874 (16). Black, G.A., et al. 54-16454 (12)
Araujo, D., & N. C. Maciel 3683 (16); 4044 (13); Blanchet, J. S. 607 (13); 2723 (16); 3089 (16); 3137
4259 (16); 4465 (16); 7514 (16); 8700 (14). (10); 3672 (16).
Araujo, D., & A. Magnanini 6031 (16). Boege, W. 1244 (8).
Araujo, D,. & J. Mauro 8614 (14). Bohrer, C. 47 (16).
Araujo, D., & A. L. Peixoto 324 (17). Bonder, G. 2465 (16).
Araujo, D., et al. 3461 (16); 8326 (14); 8597 (14). Bonpland 1599 (la); 3901 (la).
Archer, W. A. 2050 (la); 2563 (la). Boom, B. M. 6738 (la); 6791 (la); 7198 (la).
Aristeguieta, L. 2805 (la). Boom, B. M., ?fe A. L. Weitzman 5233 (la).
Atchinson, E. 276 (4). Boom, B. M., <& M. Grillo 6460 (12).

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 131
Boschbeheer, (B.W.) 45 (12); 57 (12); 61 (24); 1909 Casta?eda, R. R. 103 (la); 584 (la); 5218 (la); 6136
(24); 1944 (12); 2216 (12); 2458 (12); 2579 (12); (la); 6430 (la).
3547 (12); 3933 (24); 4092 (24); 4210 (24); Castellanos, A. 25336 (9).
5245 (24); 5252 (24); 5450 (24); 6870 (24). Castillo, A. 2353 (12).
Bowie, ?fe Cunningham N0.3 28 (16). Castillo, G. 1115 (la).
Box, H. E. 1385 (la). Castillo, J. 830 (la).
Braga, P. I. S. 2364 (16). Cavalcante, P. 238 (la); 3164 (3).
Brand, J., ?fe A. Cogollo 114 (7); 178 (la); 215 (7). Cavalcanti, T. B., et al., 1738 (9).
Brand, J., ?fe M. Gonz?lez 988 (6). Cedillo, T., & J. I. Calzada 17 (8).
Brant, A. E., <& J. Roldan 1533 (5). Champagne, H. 56 (24).
Bravo, H. 8 (8); 142 (8). Chan, F, & R. Lira 4707 (la).
Braz?o, J. E. M. 257 (9). Chapman, J. D. 4203 (le).
Breedlove, D.E. 37711 (la). Chavarria, U. 581 (la).
Brenes, A. M. 12619 (la); 15595 (la). Ch?velas, J., & L. A. P?rez 80 (8); ES-892 (8).
Bridgewater, S., ?fe A. Ib??ez-Garc?a S435 (16); S447 Ch?velas, J., & C. Zamora ES-4742 (la); ES4890 (8).
(16). Ch?velas, J., et al. ES-2396 (8); ES-4301 (8).
Bridgewater, S., ?fe E. Cardoso S243 (9). Ch?vez, D. 412 (la).
Bridgewater, S., ?fe J. Fonseca Filho S354 (10); S363
Ch?zaro, M., et al. 5474 (lb).
(10); S364 (10). Chevalier, A. 7773 (le); 24917 (1).
Bridgewater, S., et al. S241 (21); S246 (21). Chiang, F., et al. 765 (la).
Bristan,N. 1127(3) (la). Choussy, F. 1583 (la).
Christenson, G. M., et al. 1432 (28).
Britton, E. G., & D. W. Marble 2328 (la).
Cid Ferreira, C. A. 3380 (la); 3382 (la); 4268 (29);
Britton, N. L. 2906 (la); 4848 (4).
5247 (6).
Britton, N. L., ?fe J. F. Cowell 1018 (la); 10259 (4).
Cid Ferreira, C. A., & J. Lima 3611 (3).
Britton, N. L., ?fe P. Wilson 474 (4).
Cid Ferreira, C. A., & B. W. Nelson 3057 (6).
Britton, N. L., et al. 1776 (12); 14246 (4).
Cid Ferreira, C. A., et al. 6259 (21); 6270 (la); 6416
Broadway, W. E. 611 (la); 3170 (la); 6667 (la); 6944
(9); 6541 (21); 7250 (3); 9948 (la); 10824 (3);
(la).
10896 (la); 10950 (6).
Brooke, W. M. A. 5825 (la).
Claussen, P. 12 (10); 62 (10); 236 (10).
Brooks, R. J. 12630 (la).
Clewell, A. J., et al. 4339 (la).
Brunt, M. 2243 (la).
Cobra, L. A., & J. Oliveira 69 (9).
Bullock, S. H. 910 (2); 1313 (2).
Co?lho, D., & C. Mota 621 (3).
Bunting, G. S. 7707 (la); 7795 (la); 7976 (la).
Co?lho,L. 115 (20); 394 (23).
Bunting, G. S., et al. 4125 (12).
Co?lho, L., et al. 394 (3).
Burchell, W. J. 8491 (21); 8544-2 (21).
Co?lho, L. S., et al. 103 (29); 270 (20).
Burger, W., ?fe G. Matta 4767 (la).
Cogollo, A., & C. C. Estrada 270 (5); 199A (5).
C. A. C. 2728 (la).
Cogollo, A., et al. 6036 (5); 6346 (5); 6353 (5); 6390
Cabrera, E., ?fe H. de Cabrera 7028 (8); 7378 (la).
(5); 6400 (5); 6486 (5); 6521 (5); 6583 (5); 6907
Cabrera Salge, N. 13 (la). (5); 6958 (5); 7281 (5).
Calder?n, S. 317 (la). Colella, M., et al. 2020 (28).
Callejas, R., et al. 8584 (5). Collares, J. E. R. 91 (17).
Calzada, J. I. 19 (8); 988 (8); 14929 (8); 16846 (8); Collenette, C. L. 155 (9).
16880 (8); 19293 (la). Colonnello, G. 953 (la).
Calzada, J. L, et al. 11500 (la); 18688 (lb). Constantino, A. 133 (16).
Campos, A. 1352 (la); 1381 (la). Contreras, E. 7083 (la); 7600 (la); 10139 (la).
Campos, J. M. 9 (16). Coomes, D. 81 (28).
Campos Porto, P. 2650 (16). Cooper, G. P., & G. M. Slater 9 (la); 147 (la); 215
Candida, M., et al. 168 (16). (la); 265 (la).
Capucho 481 (la). Coradin, L., et al. 6510 (16).
Carauta, J. P. P., et al. 2593 (14). Cordeiro, L, et al. 825 (16).
C?rdenas, D. 1300 (la); 1485 (la); 2501 (5). Cordovil, S., et al. 119(9).
C?rdenas, D., ?fe J. G. Ram?rez 2741 (5); 2743 (5). Correa, J. A. 22 (16).
C?rdenas, M. 1346 (la). Cowan, C, & M. A. Magana 3046 (la).
Carlson, M. C. 511 (la); 1136 (la); 1300 (la). Cowan, C, et al. 2294 (8).
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Navarro de Andrade 167 (16). (13); 300 (17); 301 (17); 302 (16); 303 (16); 304
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Nieustedt, E. F. 125 (9). 224 (17); 227 (13); 231 (16); 232 (19); 233 (19);
Noblick, L. R. 2921 (10); 3415 (10); 3534 (10). 234 (16); 235 (17); 236 (16); 237 (17); 288 (17);
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179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
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Roberty, G. 17062 (la).
Pr?vost, M. F. 648 (la); 1192 (la); 3046 (24).
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Rabelo, B. V, et al. 2238 (la); 2789 (la); 3199 (26);
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179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
2003 ANDIRA 139
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Schwacke, C. A. W. 4239 (13); 4527 (10); 5033 (16); Sousa, M., ?fe A. S. Magallanes 7382 (la).
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Simpson, D. R. 675 (12). Stevenson, D. W. 958 (20).
Sintenis, P. 3356 (la). Stevenson, J. A. 2194 (la).
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179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00
Sucre, D., et al. 6583 (16); 10228 (16). van der Werff, H., & R. Ortiz 5553 (5); 5556 (5); 5576
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V?squez, R., et al. 5919 (la); 12814 (la); 13802 (6).
T?llez, O. 4733 (8); 5477 (la); 5521 (la); 9076 (lb).
Velazco, J. 1285 (12).
T?llez, O., & E. Mart?nez 905 (8); 948 (8); 8079 (la).
Veloso, H. P. 98 (16).
T?llez, O., & J. Miller 10478 (lb).
Ventura, E., ?fe E. L?pez 1140 (la); 3365 (la).
Tenorio, P. 19498 (8).
Ventura, F. 12803 (8).
Tenorio, P., & R. Torres 212 (la).
Viana-Freise, C. 287 (16).
Tenorio, P., et al. 395 (la); 3108 (la); 16869 (lb);
Vieira, M. G., et al. 622 (10).
19415 (la).
Tessmann, G. 3437 (la); 3836 (3). Voeks, R. 102 (18)
Teunissen, R A., & J. T. Wildschut 11830 (24). Wagner, R. J. 280 (la); 1644 (la); 1804 (la).
Thomas, D. W. 313 (la). Walter, B. M. T., et al. 228 (22); 1481 (21); 2733 (21).
Thomas, J., et al. 1546 (la). Wanderley, M. G. L., et al. 105 (16).
Thomas, W. W, et al. 3962 (21); 4312 (21); 8533 (14); Wasshausen, D. C, ?fe F. Encarnaci?n 1057 (la).
8969 (19); 9469 (17); 9726 (19); 9881 (17); Webster, G. L., ?fe K. I. Miller 8718 (la).
10055 (16); 10950 (19); 11196 (13). Wedel, H. von 1905 (la); 2017 (la).
Thomsen, K. 594 (la); 607 (la). Weddell, M. H. 2898 (la); 3420 (9).
Thornewill, A. S. 54 (le). Wendt, T., ?fe A. Villalobos 4463 (la).
Tillet, S. S. 756-393 (la). Wendt, T., et al. 3416 (la).
Tipaz, G., et al. 1225 (6); 1345 (6); 2215 (la). West, E., & Arnold L. 877 (la).
Toepffer, A. 124 (la). White, S., ?fe W. S. Alverson 459 (la); 470 (la).
Tonduz, A. 6511 (la); 6874 (la); 9771 (la). Wilhams, L. O. 2421 (la); 4727 (la); 6457 (la); 8483
Torres, J. 160 (12). (8); 9451 (8); 9484 (8); 9612 (8).
Torres, R., & R. Cedillo 2783 (la). Williams, L. O., <& V. Assis 7430 (10): 7453 (10).
Torres, R., & C. Mart?nez 6005 (8). Williams, L. O., ?fe A. Molina 14628 (la).
Torres, R., & M. A. Mart?nez 6606 (la). Williams, L. O., ?fe T. P. Williams 24614 (la).
Torres, R., & P. Tenorio 210 (la); 220 (la). Williams, R. S. 419 (la); 420 (la).
Torres, R., et al. 1279 (la); 2618 (la); 5361 (8); 5362 Wing, E. S. 25 (8).
(la); 9729 (la).
Woodworm, R. H. 189 (la).
Treacy, J., & J.B. Alcorn 477 (la).
Woolston, A. L. 734 (10).
Triana,J. 1131 (la).
Woytkowski, F. 7269 (la); 35016 (la).
Trigos, R. C. 556 (la); 737 (la); 1070 (la); 1495 (la).
Wright, C. 1160 (4); 1187 (4); 1188 (4); 2356 (4);
Trigos, R. C, & R. Torres 1483 (la).
2358 (4).
Tuerckheim, H. von 7834 (la).
Wurdack, J. J., ?fe L. S. Adderley 43368 (28).
Tuez, & Cochran 29269 (16).
Tupinamba 22 (16). Wurdack, J. J., ?fe J. V. Monachino 39345 (la).
Turner, L. 108 (1). Yano, T. 05 (10).
Tyson, E., & M. Kuns 1003 (la). Yuncker, T. G. 18268 (la).
Tyson, E., et al. 4849 (la). Zabelo, H. 50 (12).
Ule, E. 7627 (12); LV11(15). Zamora, N., ?fe M. Castrillo 2031 (la).
Underwood, L. M., & R. F. Griggs 711 (la). Zanoni, T., ?fe R. Garcia 26927 (la).
Urban, I. 6178 (la). Zanoni, T., ?fe M. Mejia 17748 (la); 17899 (la).
Uribe, L. 6650 (la). Zanoni, T., et al. 16093 (la); 21028 (la); 28788 (la).
Usteri, P. A. 104 (16). Zardini, E., ?fe U. Velazquez 25531 (10); 25676 (10).
Valcarcel, P. F. J. 125-5/B (la). Zarucchi, J. L. 2810 (9).
Valencia, R., et al. 67876 (6). Zarucchi, J. L., & J. Betancur 6444 (5).
Valerio, M. 522 (la). Zarucchi, J. L., ?fe D. C?rdenas 4215 (7).
Valeur, E. J. 665 (la). Zentner 376 (9).

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2003 ANDIRA 141
INDEX TO SCIENTIFIC NAMES
Accepted names are in roman type; the main entry for each is in boldface. Synonyms are in italics.
Aglaia 15 var. lanceata N. F. Mattos 85

Aganope var. latifoliolata N. F. Mattos 85
gabonica (Baill.) Polhill 118 var. rosea (Mart, ex Benth.) Benth. 84
Agouti frondosa Mart, ex Benth. 80
paca 14 var. longifoliolata N. F. Mattos 76
Amerimnum gabonica Baill. 118
affine Spreng. 44 galeottiana Standl. 12, 15, 21, 23, 24, 25, 26, 28,
Andira Juss. 3 30, 32, 33, 34, 37, 39, 41, 64-66, 68, 72, 76,
Andira Lam. 36-37 126, 129
sect. Aristobulia Benth. 117 grandiflora Guillem., Perr. ?fe A. Rich. 44
sect. Euandira 3, 31, 36 grandistipula Amshoff 5, 15, 21, 24, 25, 26, 28,
sect. Lumbricidia (Veil.) Benth. 3, 22, 31, 36 29, 30, 32, 34, 37, 38, 41,106,108-109,129
sect. Paucifoliolatae N. F. Mattos 3, 22, 31, 36 handroana N. F. Mattos 85
subsect. Lumbricidia (Veil.) N. F. Mattos 3, 31, 36 horsfieldii Lesch. 118
subsect. Glabratae N. F. Mattos 3, 31, 36 humilis Mart, ex Benth. 4, 5, 8, 12, 14, 16, 17, 18,
acuminata Benth. 44 19, 21, 23, 24, 25, 26, 28, 30, 32, 33, 35, 37,
amazonum Mart, ex Benth. 117 38, 42, 43, 69-72,76, 92, 103,126, 127, 129
anthelmia (Veil.) Benth. 4, 5, 6, 11, 12, 13, 15, 16, var. cordata N. F. Mattos 69
18, 20, 21, 22, 24, 25, 26, 27, 28, 29, 30, 32, inermis (W. Wright) DC. 1, 2, 3, 4, 5, 7, 8,10, 12,
33,34,37,38,41,76-30,82,84,88,125,129 13, 14, 15, 16, 17, 24, 25, 26, 27, 28, 30, 32,
var. acuminata Benth. 76 33, 34, 36, 37, 40, 42-50, 51, 53, 56, 62,
var. gracilis N. F. Mattos 85 124, 127, 129
var. ormosioides (Benth.) Benth. 82 subsp. inermis 4, 21, 29, 33, 43, 44-48, 49, 50,
anthelmia (Veil.) J. F. Macbr. 76 62,129
anthelmia (Veil.) Toledo 76 subsp. glabricalyx R. T. Penn. 21, 33,43,44,45,
anthelmintica Benth. 76 4*49,129
araroba Aguiar 117 subsp. grandiflora (Guillem., Perr. ?fe A. Rich.)
aubletii Benth. 118 Gillett ex Polhill 44,48
bahiensis N. F. Mattos 85 subsp. rooseveltii (de Wild.) Polhill 4, 21, 33,
bracteosa Benth. 118 43, 44,45, 49-50,129
carvalhoi R. T. Perm. & H. C. Lima 4, 9, 10, 12, var. riedelii Benth. 44
13, 14, 15, 16, 17, 18, 19, 20, 21, 24, 25, 26, jaliscensis R. T. Penn. 5, 9, 10, 12, 21, 22, 23, 24,
27, 28, 30, 32, 33, 34, 37, 40, 41, 92, 94-98, 25, 27, 28, 32, 33, 37, 38, 39, 42, 49, 50-52,
125, 129 53, 129
chigorodensis R. T. Penn. 5, 8, 9, 10, 21, 23, 29, jamaicensis Urban 42
37, 38, 40, 41, 61, 62-64,129 kuhlmannii N. F. Mattos 66, 68
chiricana Pittier 44, 48 lanei N. F. Mattos 101
cinerascens Mart. 118 laurifolia Benth. 69, 71, 72
cordata Arroyo ex R. T. Penn. & H. C. Lima 4, 5, legalis (Veil.) Toledo 4, 5, 6, 12,13, 14, 18, 21,
9, 10, 18, 20, 21, 24, 25, 26, 28, 30, 32, 33, 24, 25, 26, 27, 28, 29, 30, 32, 33, 36, 37, 38,
34, 35, 37, 40, 41, 100,102-105,125, 129 41, 78, 80-82, 84, 88, 127, 128, 129
cori?cea Pulle 4, 5, 13, 16, 21, 27, 29, 35, 37, 40, var. bahiensis (Benth.) N. F. Mattos 76
41, 56, 59, 60,10?-108,129 macrocarpa R. T. Penn. 14, 21, 29, 33, 37, 39, 41,
cubensis Benth. 8, 16, 21, 23, 33, 34, 37, 40, 42, 72-73, 75, 76, 129
53, 55-56, 129 macrothyrsa Ducke 4, 5, 8, 10, 12, 13, 21, 24, 25,
cujabensis Benth. 4, 5, 7, 12, 13, 17, 18, 21, 22, 26, 28, 29, 30, 32, 37, 40, 41, 56, 59, 60-62,
23, 24, 25, 27, 28, 32, 33, 34, 35, 37, 39, 41, 64, 128, 129
100-102,103,105,125,126,129 marauensis N. F Mattos 5, 21, 23, 24, 25, 27, 28,
excelsa Kunth 44 32, 37, 40, 42, 92, 93-94,128, 129
fraxinifolia Benth. 2, 4, 5, 7, 8, 9, 10, 11, 12, 13, micans Taub, ex Glaz. 64
14, 15, 16, 17, 18, 20, 21, 24, 25, 26, 27, 28, micrantha Ducke 5,15, 16, 21, 25, 27, 29, 35, 37,
30, 32, 33, 37, 39, 42, 84-S9,126, 129 40, 41, 56, 59, 60,105-106, 108, 129

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microcarpa Griseb. 55 Dalbergieae 1, 2, 33

multistipula Ducke 5, 8, 21, 23, 24, 25, 27, 28, 29, Dasyprocta 14
32, 37, 38, 41, 43, 51,5^-54,129 Diptychandra
var. peruana N. F. Mattos 60 aurantica Tul. 36
nitida Mart, ex Benth. 4, 5, 9,10, 11, 12,13, 18, subsp. aurantica 36
21, 24, 25, 26, 27, 28, 30, 32, 33, 37, 38, 40, subsp. epunctata (Tul.) Lima, Carvalho & Costa
42, 89-93, 94, 97, 128, 129 36
ormosioides Benth. 4, 5, 8, 9,11, 12, 13, 15, 20, Dussia Krug & Urb. ex Taub. 4
21, 24, 25, 26, 27, 28, 30, 32, 33, 37, 39, 42, Euchresta
80, 82-S4, 88, 129 horsfieldii (Lesen.) Benn. 118
paniculata Benth. 66 Geoffroea Jacq. 2, 4
parviflora Ducke 4, 5, 21, 24, 25, 26, 28, 30, 32, acutifolia Stokes 42
34, 37, 39, 41, 98-100,129 inermis W. Wright 42, 44
parvifolia Mart, ex Benth. 84 obtusifolia Stokes 72
pauciflora Benth. 69, 71 pubescens Rich. 74
pernambucensis N. F. Mattos 85 retusa Poir. 74
pisonis Mart, ex Benth. 84 spinulosa Mart. 66, 69
var. emarginata N. F. Mattos 85 surinamensis Bondt 73
var. puberula N. F. Mattos 85 verm?fuga Mart. 66, 69
praecox Arroyo ex R. T. Penn. 5, 21, 27, 29, 35,
Glycyrrhiza
37, 40, 42,109-112, 114, 115, 129 undulata Ruiz & Pav. ex G. Don 44
racemosa Lam. ex J. St. Hil. 2, 3, 118 Guttiferae 71
retusa (Poir.) DC. 74
Hymenaea
var. oblonga Benth. 74
courbaril L. 15, 97
var. surinamensis (Bondt) DC. 73
Hymenolobium Benth. 1, 2, 20, 23, 33, 118
riparia Kunth 44
flavum 21, 24, 25, 26, 28, 30, 32
rosea Mart, ex Benth. 84
nitidum 21, 24, 25, 26, 28, 30, 32
sapindoides (DC.) Benth. 44, 48 velutinum Ducke 118
spinulosa (Mart.) Benth. 66
Inocarpus J. R. Forst. & G. Forst. 1
stipulacea Benth. 76
var. bahiensis Benth. 76 Kielmeyera
rubriflora71
surinamensis (Bondt) Splitg. ex Amshoff 2, 4, 5,
6, 7, 13, 15, 16, 18, 21, 24, 25, 26, 28, 30,
Leguminosae 15, 36, 49, 97
Lennea
32, 34, 37, 39, 42, 72, 73-76, 128, 129
viridiflora Seem. 49
var. ovatifoliolata N. F. Mattos 74
taurotesticulata R. T. Penn. 9, 10, 16, 21, 23, 24, var. novogaliciensis Lavin & Sousa 49
var. viridiflora 49
25, 27, 28, 29, 32, 37, 39, 40, 41, 56-60, 72,
106, 108, 129 Lonchocarpus
staudtii Harms 44
tervequinata R. T. Penn., Aymard ?fe N. Cuello 5,
Lumbricidia Veil. 36
21, 27, 29, 37, 40, 42, 109, 111,112-114,
129 anthelmia Veil. 76
trifoliolata Ducke 3, 5, 10, 21, 22, 25, 27, 29, 31, legalis Veil. 36, 80
35, 37, 38, 40, 111, 114-115, 117, 129
Meliaceae 15
unifoliolata Ducke 3, 5, 6, 7, 13, 21, 22, 24, 25,
Millettia
26, 28, 30, 31, 32, 34, 36, 37, 38, 40, 111, rooseveltii de Wild. 49
115-117,129 Mimosoideae 15
verm?fuga (Mart.) Benth. 4, 5, 10, 12, 13, 17, 18, Myoprocta 14
19, 21, 24, 25, 26, 28, 30, 32, 33, 37, 39, 42, Ostryoderris
65, 66-69,72, 76, 88, 102, 103, 128, 129 brownii Hoyle 49
villosa Kleinhoonte 118 Papilioniodeae 1, 36, 49, 118
wachenheimii Benoist 107 Parkia
zehntneri Harms 118 biglobosa 50
Caesalpinoideae 15, 97 multijuga Benth. 15
Cleogonus 12 Pisum
Coursetia sativum L. 29
caribaea (Jacq.) Lavin Poeppigia
var. caribaea 34 procera C. Presl 69

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2003 ANDIRA 143
Pterocarpus Jacq. 4 anthelmia (Veil.) Kuntze 76

sapindoides DC. 44 cubensis (Benth.) Kuntze 55
Pterodon cujabensis (Benth.) Kuntze 100
emarginatus Vogel 35, 36 fraxinifolia (Benth.) Kuntze 84
Robineae 49 frondosa (Mart, ex Benth.) Kuntze 80
Skolemora humilis (Benth.) Kuntze 69
pernambucensis Arruda 118 inermis (W. Wright) Lyons 42
Sophoreae 33 laurifolia (Benth.) Kuntze 69
Trachypogon legalis (Veil.) Kuntze 80
plumosus 112 paniculata (Benth.) Kuntze 66
Vatairea pisonis (Mart, ex Benth.) Kuntze 84
guianensis Aubl. 117, 118 retusa (Poir.) Kuntze 74
Vataireopsis sapindoides (DC.) Kuntze 44
araroba (Aguair) Ducke 118 spinulosa (Mart.) Lyons 66
Vigna 124 surinamensis (Bondt) Kuntze 73
Vouacapoua Aubl. 2 verm?fuga (Mart.) Kuntze 66
americana Aubl. 2

179.236.175.49 on Mon, 20 Nov 2023 12:55:07 +00:00

Pennington MonographAndiraLeguminosaePapilionoideae 2003

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Monograph of Andira (Leguminosae-Papilionoideae)

Author(s): R. Toby Pennington

Stable URL: https://www.jstor.org/stable/25027903

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Andira is a genus of 29 woody species (including three subspecies of A. inermis) dis

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

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FIG. 2. Indumentum and venation of leaflets of Andira (characters 5 and

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to mixed eucamptodromous to entirely brochidodromous (terminology of Hickey, 1979).

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Fruit. Fruit of Andira are single-seeded (occasionally 2-3-seeded) drupes with

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

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

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These observations of pollination biology have influenced the coding of character 7,

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Fruit size and

A. surinamensis RTP 364 small, bat 6.6 3.4 32.6 8.9

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Knowledge of chromosome numbers in Andira is restricted to those reported by

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INTRASPECIFTC cpDNA POLYMORPHISM AND

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Amazonian Populations of A. humi/is with

Seasonally Range of A. carvalhoi

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CLADISTIC ANALYSES OF SPECIES RELATIONSHIPS

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Table 2. Data matrix of morphological characters of Andira.

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Morphological analysis. Pennington (1996) showed that a cladistic analysis of mor

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solved, except for a clade comprising A. cordata, A. cori?

Combined molecular and morphological analyses. The fi

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-A. inermis CEH 1673

-A. inermis TDP 13558 CLADE I

-A. carvalhoi RTP 229

-A. anthelmia RTP 227,282

-A. legalis RTP 307

-A. legalis HCL sn

-A. carvalhoi RTP 233

-A. fraxinifolia RTP 213

-A. fraxinifolia MS 889

-A. nitida RTP 300

-A. nitida RTP 292

-A. nitida RTP 301

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