Monograph of Andira

Monograph of Andira (Leguminosae-Papilionoideae) Author(s): R. Toby Pennington Source: Systematic Botany Monographs, Vol. 64, Monograph of Andira (LeguminosaePapilionoideae) (Mar. 31, 2003), pp. 1-143 Published by: American Society of Plant Taxonomists Stable URL: http://www.jstor.org/stable/25027903 Accessed: 02/12/2009 12:12 Your use of the JSTOR archive indicates your acceptance of JSTOR's Terms and Conditions of Use, available at http://www.jstor.org/page/info/about/policies/terms.jsp. JSTOR's Terms and Conditions of Use provides, in part, that unless you have obtained prior permission, you may not download an entire issue of a journal or multiple copies of articles, and you may use content in the JSTOR archive only for your personal, non-commercial use. Please contact the publisher regarding any further use of this work. Publisher contact information may be obtained at http://www.jstor.org/action/showPublisher?publisherCode=aspt. Each copy of any part of a JSTOR transmission must contain the same copyright notice that appears on the screen or printed page of such transmission. JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected].

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

species,

cessions

A.

inermis,

of A. humilis with plastomes characters provides

12 morphological of cpDNA

restriction

A cladistic analysis of from probable introgressive hybridization. little phylogenetic resolution in comparison with a simultaneous analysis some of the clades discovered site data and morphology. Although by this analysis are derived

characters and have no morphological the "cryptic," i.e., they are supported only by molecular synapomorphies, results provide a reasonable estimate of the phylogeny of Andira and are used as the basis of discussions of char acter evolution, and biogeography. Previous of Andira classification, infrageneric infrageneric classifications were based on the nature of the indumentum of the gynoecium, but glabrous ovaries have evolved in parallel at least three times. Detailed

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

into dispersal

ation of species of Andira are illustrated. Five new macrocarpa,

A. praecox,

are fully described and their ranges are mapped, and 15 are proposed: A. chigorodensis, A. jaliscensis, A. subspecies and A. inermis subsp. glabricalyx.

in eastern Brazil. All

species

and one new

species A. taurotesticulata,

INTRODUCTION is a genus of 29 woody species (including three subspecies of A. inermis) dis tributed throughout tropical America; A. inermis also occurs in Africa. Andira is most Benth. (Bentham 1860; Polhill 1981), a genus of 10-12 closely related toHymenolobium species confined to South America. They are similar florally and vegetatively, and they Andira

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

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systematic studies (Doyle et al. 1997; Lavin et al. 2001) indicate that Dal This confirms earlier suggestions, based upon consideration bergieae is not monophyletic. that Dalbergieae are not a natural group (de Lima of fruit, seed, and seedling morphology,

molecular

1991). This taxonomic revision

is based upon examination of ca. 3,000 herbarium specimens in five Neotropical countries from 1991 to 1997. Species de inAndira has been regarded as problematic, and the only recent revision (Mat

and field studies conducted

limitation tos 1979), covering the Brazilian species, is inadequate for species identification, particu coastal rain forest and coastal restinga forest of larly for taxa from the Atlantic southeastern Brazil (Lewis 1987), where Andira is both diverse (eight species) and abun dant. Three new species have been described recently from eastern Brazil (Pennington & de Lima 1995) and the Venezuelan Guayana (Pennington et al. 1997). Four of the five new species included in this monograph area of diversification for Andira.

grow around the fringes of Amazonia,

which

is an

frameworks for Andira based upon both chloroplast DNA (cpDNA) re Phylogenetic striction site data and morphology have been presented by Pennington (1995, 1996). The are drawn from these studies, but in this monograph phylogenetic analyses presented in coding of some characters are proposed based upon new observations of modifications pollination syndromes and fruit anatomy. Furthermore, new cpDNA restriction site data have provided phylogenetic placements for five more species and helped resolve some of the problems of intraspecific cpDNA polymorphism discussed by Pennington (1995, are used to re-evaluate infra The refined here 1996). phylogenetic hypotheses presented in Andira, to present the first biogeographic generic classification the evolution and diversification of the genus, and to investigate phological features, such as stipules, flowers, and fruits.

hypotheses to explain the evolution of mor

TAXONOMICHISTORY to the attention of European science through the use of extracts from the bark and seeds as anthelminthic drugs by indigenous peoples in the Neotropics. as an anthelminthic in Per Piso (1648) described the use of the seeds of A. fraxinifolia Andira

first came

nambuco, Brazil, and named the species "Andira Ibaiariba." The first species descriptions were included in accounts of the medicinal use of the bark and seed of A. inermis in Ja maica (Wright 1777) and A. surinamensis in Suriname (Bondt 1788). Both authors placed their species in Geoffroea Jacq., which shares Andira's unusual drupaceous fruit, but is distinct inmany floral and vegetative characters. the first valid publication of the name Andira has been problematic. It Determining had generally been assumed to be by Lamarck (1783), and as such was proposed (Harms 1904) and accepted (Briquet 1906) for conservation against Vouacapoua Aubl. Although these two genera are now considered taxonomically quite distinct, Lamarck had included Vouacapoua in his original concept of Andira, making the latter name illegitimate. Yet, as part of the Rickett and Stafleu revision of the lists of Nomina Conservanda, Cowan (1959) noted that Lamarck's

(1783) description of Andira racemosa provided no generic diagno sis, and since Lamarck cited Vouacapoua americana Aubl. (1775) in synonymy his genus and species could not be treated as new. Cowan decided that the long-standing provisions of the International Code of Botanical Nomenclature (ICBN) permitting a "descriptio to validate simultaneously the generic and specific names could not generico-specifica"

3

ANDIRA

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apply. He therefore concluded that neither Andira nor the species name, A. racemosa, were validly published by Lamarck, and proposed that the conservation of Andira be from its next usage, that by A. L. de Jussieu (1789), with A. racemosa Lam. ex J. St.-Hil., ap parently the first validation of that name, as type. R. K. Bmmmitt (pers. com.) noted that Lamarck did indeed validly publish Andira but as an illegitimate replacement (a nomen novum) of Vouacapoua Aubl., because he treated

Vouacapoua

as a synonym

of his

generic

name

Andira.

His

species

name

A.

race

is also illegitimate, as a nomen novum for V. americana. Under Art. 7.5 of the cur rent ICBN (Greuter et al. 2000), the type of Andira Lam. is that of Vouacapoua (i.e., the type of V. americana Aubl.), but because Vouacapoua is now recognized as a distinct cae salpiniaceous genus, this typification would be extremely disruptive.

mosa

The current conservation is of "Andira Juss.," not "Andira Lam." Jussieu (1789) cited no species in his account, and Cowan (1959) considered Andira Juss. to be typified by A. racemosa Lam. ex J. St. Hil.; however, Jaume St. Hilaire (1804) was merely using Lamarck's A. racemosa, so there is no case for treating his name as a later homonym under Art. 48.1 of the ICBN. If "Andira Juss." were to be maintained as the conserved name, the type requires correction to A. racemosa Lam., which is automatically typified by V. americana. Therefore, themaintenance of Andira Juss. would defeat the purpose of the conservation of the name Andira. For these reasons, a formal proposal to change the authorship of Andira to Andira Lam. nom. cons., and to provide it with a conserved type, A. inermis, was submitted and accepted (Pennington 2002). The name Andira inermis (Wright) DC. has been widely ap plied to the species that includes A. racemosa Lam., the type of the conserved Andira (i.e., excluding the Vouacapoua element). Thus, itwas appropriate to propose, under Art. 14.9 of the ICBN, that the type of A. inermis be the conserved type of Andira Lam. If the name A. racemosa Lam. were conserved with a new type, itwould displace A. inermis, the name traditionally used for the most

common, widespread,

and frequently

collected

species of

Andira.

Infrageneric Bentham

classification

1839, 1860, 1862) features prominently in the taxonomic history of studying the specimens of Martius, Pohl, and Schott from eastern Brazil, he the first major account of Andira (1837, reprinted in 1839). He placed the produced two in sections: sect. Lumbricidia, defined by a hairy ovary on a relatively short species (1837,

Andira. After

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

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

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

The fibers are non-septate

and usually very thick-walled.

ANDIRA

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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 for species delimitation, (characters 5, 8, 10). is useful

and provides

three characters

for cladistic

analysis

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

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

analysis). Venation

pattern varies continuously

from entirely eucamptodromous

6

SYSTEMATICBOTANYMONOGRAPHS

FIG. 1. Stipules and leaves = ca. 5 cm; (scale bar photo G. P. scale = inches; right-hand scale = leaves of A. legalis. (Based on: A,

in Andira (characters 3 and 4). A. Large, persistent stipules of A. anthelmia Lewis). B. Small stipules ("s" at arrows) of A. surinamensis (ruler left-hand leaves of A. unifoliolata. D. Large, imparipinnate cm). C. Small, unifoliolate R. T. Pennington 183; B, R. T. Penningfon 463; C, W. Rodrigues 11182; D, R.

T Penningfon

305.)

VOLUME 64

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y.y

7

s

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

at Royal

Botanic

Gardens,

Kew.)

SYSTEMATICBOTANYMONOGRAPHS

8

VOLUME 64

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

which

analysis.

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

and

bracteoles.

Bracts

and

bracteoles

ducous. There is little variation in theirmorphology paired at the base of the calyx.

are

small,

narrow,

and

often

early

throughout Andira. The bracteoles

ca

are

length varies continuously from 5 to 23 mm (Fig. 3).Within species, the range of variation is relatively narrow, approximately 2 mm for small-flowered species and 6 mm for large-flowered species. Flower length is a useful character for separating similar pairs of species, such as A. inermis and A. cubensis, A. fraxinifolia and A. ormo Flowers.

sioides,

Flower

and A. macrothyrsa

and A.

chigorodensis.

The angle and depth of the calyx lobes varies widely in several species and is thus of limited utility in species delimitation; however, the calyx indumentum (character 8 in cladistic analysis) provides more useful characters. For example, the densely hairy calyx of A. cubensis distinguishes this species from A. inermis. In A. inermis, A. multistipula, and A. humilis, the calyx is either sparsely hairy (the hairs short and appressed) or and this variation shows of scription subspecific taxa inA. The corolla o? Andira consists but are firmly attached because of

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

Lamellate

wing petal sculpturing (Fig. 4A; character 9 in cladistic analysis) is present in all species with flowers greater than 9 mm long, and also in A. cubensis where the flow ers are 7 to 8.5 mm long. Lamellate wing petal sculpturing is absent from the other small flowered species (Fig. 4B).

In all species of Andira, the stamens are diadelphous (9 + 1); the vexil Androecium. and A. macrothyrsa lary stamen is free (Fig. 3). Very rarely, individuals of A. fraxinifolia have one side of the vexillary filament united to the other nine. The filaments are unequal in length; those closest to the standard are shorter and united for one half to two-thirds of their length. The ratio of the length of the united and the length of the free filaments varies

ANDIRA

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FIG.

9

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

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SYSTEMATICBOTANYMONOGRAPHS

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B(i)

in Andira. Species with wing petal sculpturing: A, wing petal of A. ver FIG. 4. Wing petal sculpturing = lamellate S scale bar = 1mm; A(i) wing petal of A. fraxinifolia (16 mm long); A(ii) wing m?fuga, sculpturing, (15 mm long); A(iii) wing petal of A. nitida (11 mm long); A(iv) wing petal of A. jalis petal of A. carvalhoi censis (12 mm long). Species without wing petal sculpturing: B, wing petal of A. trifoliolata, scale bar = 1mm; B(i) wing petal of A. cordata (5.5 mm long); B(ii) wing petal of A. chigorodensis (5 mm long); B(iii) wing petal of A. taurotesticulata 228; A(ii) M. P. M. de (8 mm long). (Based on: A, J. Ratter 3595; A(i) R. T Pennington Lima et al. 20; A(iii) A. J. Ribeiro 62; A(iv) E. J. Lott 2544; B, J. J. Wurdack & L. S. Adderley 43368; B(i) G. . Hatschbach 39477; B(ii) /. Brand & A. Cogollo 114; B(iii) G. Lozano & G. Galeano 3963.)

little across all Andira species, which is further proof of the highly conservative floral or in Andira that appears to reflect a general bee pollination mechanism.

ganization

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

11

ANDIRA

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will provide taxonomic characters within Andira. The pollen grains of the two species ex amined have an even, minutely reticulate tectum, are 3-colporate, with the colpi extend ing only partly to the poles, and have granular colpal membranes. Gynoecium. The ovary of Andira is borne on a stipe and contains 1-8 ovules (Fig. 3). The fruit of Andira is usually single-seeded (see "Fruit" below), the result of ovule abor tion. The style is curved and is included within the keel petals. The stigma is ciliate (sensu Lavin and Delgado, 1990). The proportions of the stipe, style, and ovary are more or less invariant. The extent of the indumentum of the gynoecium is, however, very variable (Fig. 3; character 10 in cladistic analysis) and is a critical character for species delimitation. are single-seeded 2-3-seeded) (occasionally drupes with or and hard thin exocarps. They are diverse mesocarps, endocarps, fibrously fleshy

Fruit. woody

Fruit of Andira

in the color

of

exocarp

and mesocarp,

scent,

mesocarp

structure,

and

endocarp

structure.

There are two general size classes of fruit: 6 cm long or less, weighing ca. 10-20 g when to 40-300 g when dry. These dry, and much larger fruits up to 12 cm long and weighing in dispersal biology size differences reflect probable differences (see "Dispersal Syndromes"). Bentham

in (1860) and Lewis (1987) point to the probable utility of fruit morphology species delimitation and the lack of adequate fruit collection. Field observations and the inferences made from alcohol preserved fruit confirm this. I have made field ob servations of fruit of seven species, which suggest that coloration of themesocarp and ex ocarp are useful characters. For example, the green mesocarp of A. anthelmia and A. frax

Andira

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

A. ormosioides, and confirms the ormosioides have dark brown ex

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

species discovered by cpDNA data (Pennington 1995). Pennington and Gemeinholzer (2000) examined the pericarp anatomy of 25 species in search of further taxonomic characters, and a summary of this work is provided here. The mesocarp of all Andira species examined comprises parenchyma cells in which stone cells are embedded, either individually or in patches. The mesocarp of seven species is homogeneous, whereas in 20 species it is heterogeneous, having either patches of stone cells close to the exocarp, fibers near the endocarp, and/or ergastic materials distributed unevenly throughout the parenchyma. Despite this variation, itwas decided thatmesocarp structures could not be used cladistically, because intraspecific variation was too great. Starch grains are abundant, occurring either scattered throughout the tissue, as greyish cell content or as "tyloses-like" features (probably tightly packed starch grains in highly packed parenchyma cells with squashed cell walls). At maturity, the endocarp is divided into a thicker hard layer and a thin papery layer within it, which is the remnant of the "seed cushion." This latter structure, described by

12

SYSTEMATICBOTANYMONOGRAPHS

VOLUME 64

(1914), is characteristic of legume fruits and consists of thin-walled, juicy parenchyma cells inwhich the developing seeds are embedded. At fruit maturity it shrinks and dries out, forming flake-like shreds. The hard, thick part of the endocarp is variable and provides phylogenetically useful information. Three main structures are present: (i) dominated by parenchyma and/or col lapsed cell tissue; (ii) dominated by fibers; (iii) dominated by stone cells. This variation can be coded as amultistate cladistic character (Pennington & Gemein holzer 2000; character 10). This is a refinement of the coding of Pennington (1996), where

Fucsk?

the distinct was

not

state of the endocarp dominated

by parenchyma

and/or collapsed

cell tissue

recognized.

formed by parenchyma and/or col species have an endocarp predominantly most to cell often tannin. Twelve species have an en which is brown due tissue, lapsed dominated fibers. Small cells and sometimes squashed cell tissue by parenchyma docarp occur between the fibers. Air gaps were often observed, probably due to ripening or de in layers, but in A. ver generation processes. The fibers can be arranged predominantly Five

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

of parenchymatous tissue. is thinner than in any other burrowing seed predators;

in this species. Field obser of preserved fruit revealed no seed pr?dation inA. fraxinifolia, A. anthelmia, A. ormosioides, A. nitida, and A. carvalhoi, which have thicker, tougher, woody fibrous endocarps and in A. cujabensis, which has an endocarp of stone cells. vation and examination

These data suggest

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

endocarps.

Seeds of species of Andira can be considered "overgrown" (sensu Comer, because 1976), they have an undifferentiated testa and contain no endosperm. They fill the entire cavity of the fruit, and are more or less globose tomore elongated, 2-8 cm long and 1.5-6 cm wide. The testa is thickly chartaceous, generally dark reddish brown, and often remains attached to the walls of the seed cavity (overlying the remains of the seed cush Seeds.

ion) when the seed is removed. The cotyledons are virtually united, and a thin line can be seen in transverse section where they meet. The hypocotyl-radicle axis is small and ap parent as a fold at the apical end of the seed.

2003

ANDIRA

13

(de Lima 1991; pers. obs. for A. carvalhoi, Seedlings. Germination is cryptohypogeal A. cujabensis, A. fraxinifolia, A. nitida, A. surinamensis, A. cori?cea, A. anthelmia). Nu merous spirally arranged scale-like prophylls are produced before the first true leaves, which are trifoliolate or with two or more pairs of leaflets (even in the unifoliolate A. uni was not observed in pers. obs.). Leaflet nyctinasty (evening leaf movements) foliolata, seven of the of Andira this any seedling grown during species study. It appears possible that polyembryony?the presence of more than one embryo in an occur in Andira, because more than one shoot can be pro (Richards 1997)?may duced from a single seed o? A. fraxinifolia (P.Griffiths, pers. comm.). This raises the pos that the variation patterns of this species might be ex sibility complex morphological ovule

plained

in part by apomixis.

studies of this and other species are

Further developmental

necessary.

POLLINATIONBIOLOGY Frankie et al. (1976) recorded numerous species of bees visiting the flowers of A. in ermis in Costa Rica in large numbers. Other observations of floral visitors are anecdotal. I have made similar observations of a variety of bee species visiting A. fraxinifolia and A. carvalhoi (R. T. Pennington, unpubl.) in Bahia, Brazil. These are two species with pink purple flowers, less than 18mm long. Although the larger (19-24 mm long) flowers of A. and are not significantly larger in terms visited by large xylocopid bees, which suggests a different pollination biology. This also appears to be reflected in the phenology of A. anthelmia, where a high percentage of individuals were observed to be synchro nously flowering in one area (vicinity of Ilh?us, Bahia, Brazil, October 1994; R. T. Pen nington, unpubl.) in contrast to smaller-flowered species, such as A. fraxinifolia, A. car anthelmia of flower

have superficially similar morphology length, I have only seen these flowers

valhoi, and A. verm?fuga, where T. Pennington,

unpubl.;

J. A.

few individuals

Ratter,

pers.

comm.).

in a population Two

other

flower synchronously

species,

A.

legalis

and A.

(R. or

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

that they, too, have distinct pollina to confirm this. James Ratter (herb,

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

SYSTEMATICBOTANYMONOGRAPHS

14

FIG. 5. A. Small, fruits of A.

legalis

fruits of A. fraxinifolia (R. T. Pennington bat-dispersed (G. P. Lewis & H. C. de Lima 1196; photo G. P. Lewis.)

VOLUME 64

213). B. Large,

rodent-dispersed

These observations of pollination biology have influenced the coding of character 7, which now has three states, rather than the two described by Pennington (1996).

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

ANDIRA

2003

15

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

Table 1.Mesocarp of five species of Andira. All figures are % by weight of the air-dried composition the maturity of the fruit was proven by the subsequent germination ex of seed from all accessions, mesocarp; All vouchers are collections of R. T. Pennington and deposited at CEPEC, FHO, and K. cept A. ormosioides. Fruit Speciespresumed A. surinamensis A. fraxinifolia A. fraxinifolia

RTP

364

RTP 202 RTP

213

size and disperser

Starch Sugars

6.6

3.4 32.6

7.3

7.23.3

9.5 5.3

11.6 2.8

5.3

2.0

6.8

A. anthelmia

RTP 227

A. carvalhoi

RTP 232

large, rodent

RTP 306

Fat

small, bat small, bat small, bat small, bat small, bat

A. ormosioides

Protein

8.9

8.6

1.62.2 5.6

2.3 26.2

0.6

13.7 28.7 8.8

SYSTEMATICBOTANYMONOGRAPHS

16

VOLUME 64

caution. Second, although A. carvalhoi has a particularly high starch content, which re the sim flects the large numbers of starch grains in the parenchyma cells of itsmesocarp, ilar value for A. anthelmia indicates that high starch content is not a unique feature of large fruit. Overall, there is little difference in composition between the two fruit types, and the inclusion of more rodent-dispersed species and more accessions of individual species are necessary to further investigate the small differences found. A notable aspect of these data is the high sugar content of the mesocarp of A. suri namensis. This sample was analyzed four weeks after collection in Guyana, whereas the other accessions were a year old; however, the sugars should remain more or less stable unless dried fruit suffer microbial contamination, of which there was no evidence. It ap pears that these fruit are much sweeter than the others. It is possible that these sweet fruits are attractive to different bat species from those that disperse other species of Andira. The fruit of A. humilis has a completely different scent from the fruit of the five other bat-dis persed species that I have collected, which may also be attractive to alternative bats. Andira fruit generally contain a single seed, so the large, putatively rodent-dispersed fruit contain a correspondingly large seed. It is possible that large seeds allow seedlings to tolerate shade for long periods in the rain forest understory (Leishman & Westoby 1994), which may explain why several rain forest species o? Andira (e.g., A. micrantha, A. cori?cea, A. taurotesticulata) have large fruit. In restinga vegetation, the large seed of A. carvalhoi may permit the immediate development of an extensive root system, which helps the seedlings to survive in the easily eroded, rapidly drained, white sandy soils. Seed of this species, grown at the Royal Botanic Gardens, Kew, produced extensive roots, and little above-ground growth, in contrast to themuch stronger shoot growth of the other four bat-dispersed

species germinated.

CHROMOSOMENUMBERS of chromosome numbers in Andira is restricted to those reported by Knowledge Goldblatt (1981). Both n = 10 and n = 11 have been reported in A. inermis. One of these counts (Fritsch 1970; n = 11) is from a tree from Cuba, where A. inermis is absent, and is therefore more likely to be of A. cubensis, the only Andira species in Cuba. Two other = 10. species (not named by Goldblatt) have n Further chromosome counts were attempted for A. anthelmia {R. T. Pennington 282) and A. fraxinifolia {R. T. Pennington 236), two species for which living plants were avail able at the Royal Botanic Gardens, Kew and Edinburgh. Root tips were collected and in distilled water before pre-treatment in a saturated solution of paradichloroben zene for four hours to overnight at room temperature. They were then fixed in ethanol:acetic acid (3:1 vol:vol). The root tips were hydrolyzed for 40 minutes in 5N hy

washed

acid at room temperature before staining with Feulgen (prepared after Fox, two for hours. They were then mounted in 45% acetic acid and observed under 1969)

drochloric

phase

contrast.

Perhaps because the root tips were not collected from entirely healthy, vigorously growing plants, the preparations showed few cells inmetaphase. Thus, the counts made should be regarded as preliminary. Counts for A. fraxinifolia show 2n = 22, and for A. an = 21 or 22. These results, together with those reviewed by Goldblatt (1981), thelmia, 2n indicate that there may be two basic chromosome numbers in Andira: n = 10 and n = 11.

ANDIRA

2003

17

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

likely explanation than the maintenance of ancestral polymorphism be cause of the large number of restriction sites (7-10) that differentiate the haplotypes in A. carvalhoi and A. humilis, and the lack of intermediate haplotypes in these or other extant then this polymorphism has species. If ancestral polymorphism were the explanation, been successfully maintained over remarkably long periods, and perhaps we might of the intermediate haplotypes. expect to discover maintenance

also

Whatever the explanation for this infraspecific cpDNA polymorphism involving phy logenetically disparate haplotypes, it is vital to assess whether it is widespread, because processes of hybridization with subsequent introgression and lineage sorting (where an ancestral species is polymorphic for cpDNA, and these cpDNA types sort in subsequent species) can result inmisleading interpretations of phylogenetic relationships based upon cpDNA data (Doyle 1992). Fieldwork in Central Brazil, Costa Rica, and Ecuador pro vided a valuable opportunity to extend this survey to other species from different geo graphical areas. In particular, cpDNA variation was examined inmore populations of A. humilis in an attempt to discover how widespread is intraspecific cpDNA polymorphism in this species. Methods All methods

followed

(1995). Fourteen restriction site mutations Pennington (Ap for different clades of species (Figs. 7, 9) were selected for study, a pendix 1) diagnostic and Brunsfeld technique recommended by Doyle et al. (1990) and Rieseberg (1992). DNAs of accessions included in this previous study (A. humilis RTP 239, 269; A. ver m?fuga RTP 265; A. fraxinifolia RTP 213; A. inermis C. E. Hughes 1673) were included to facilitate identification of banding patterns. Forty-one accessions of eight species were screened (Appendix 2). These included 14 accessions from three populations of A. hu milis, 11 accessions from three populations of A. verm?fuga, and eight accessions from two populations of A. cujabensis.

18

SYSTEMATICBOTANYMONOGRAPHS

VOLUME 64

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

folia, A. nitida, A. anthelmia, A. legalis), which are common elements of vegetation in coastal southeastern Brazil. It is thus tempting to hypothesize that hybridization and sub area. in This have occurred this sequent introgression hypothesis is supported by the ob servation that the individuals RTP 246 and 247 were

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

hoi's true phylogenetic position is in Clade III and its "true" cpDNA type is that of Clade in A. carvalhoi by recent III, it is difficult to explain the plastome type polymorphism

2003

andira

19

PLASTOMES: CLADEI CLADE II CLADE III(one species) PLASTOME GROUP I (one species)

-10?

PLASTOMESr CLADE II PLASTOME GROUP I

H 20? PLASTOME: CLADE IIIONLY

50?

Amazonian Rain Forest

Populations of A. humi/is with plastome type of Clade III

Seasonally Dry Vegetation

Range

FIG.

6. Location

polymorphism.

of populations The plastome

of Andira

humilis

types present

carvalhoi

A Populations of A. humilis with plastome type of Plastome Group

Atlantic Coastal Rain Forest

cpDNA

of A.

and A. carvalhoi

in each of the different

sampled vegetation

I

in a study of intraspecific types are indicated.

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

VOLUME 64

SYSTEMATICBOTANYMONOGRAPHS

20

have been the source of its plastome. To unravel the reasons for the complex intraspecific inA. carvalhoi clearly requires sampling of more populations and cpDNA polymorphism full characterization of plastome types. It is tempting to ascribe some of the difficulties in species delimitation inAndira, such some specimens as A. fraxinifolia as the difficulties in distinguishing and A. ormosioides, of hybrids. Ifmorphological intermediacy may be taken as evidence for see are Bennett 1994, McDade 1990), then hy frequent exceptions; e.g., hybridity (there evidence bridization may be possible, but rare. I reported having seen no morphological of hybridity amongst sympatric species of Andira in southeastern Brazilian restinga forests (Pennington 1995). Subsequently, I have observed a single plant that appeared to to the occurrence

between A. anthelmia be intermediate in leaf and stipule morphology in the Poco das Antas Reserve in the State of Rio de Janeiro.

and A. fraxinifolia

CLADISTICANALYSES OF SPECIESRELATIONSHIPS Introduction Cladistic analysis of chloroplast DNA restriction site characters for species o? Andira are presented by Pennington and two outgroup species of Hymenolobium (1995). Pen site and that both restriction directly combining cpDNA morpho nington (1996) argued logical characters in a single cladistic matrix and analyzing both simultaneously provided the best phylogenetic hypothesis for Andira. Cladistic analysis of cpDNA restriction site characters alone failed to resolve relationships between closely related species. Cladistic characters produced a highly unresolved result because of the analysis of morphological lack of characters with discrete states that are suitable for cladistic analysis in Andira. In contrast, combining these data sets directly produced good phylogenetic resolution, be cause the different character sets appear to be providing resolution at different hierarchi cal levels: the cpDNA characters appear to be evolving slowly and delimit groups of characters provide resolution within these groups. The species, whilst the morphological same view is taken here: that a combined cladistic analysis of all available data represents the best phylogenetic sions of infrageneric

hypothesis for Andira, the hypothesis that will be used for discus classification, character evolution, and biogeography. Species

Concepts

were delimited largely using the framework of the phylo 1990; Luckow 1995) and die characters genetic species concept (e.g., Nixon & Wheeler and character states listed below. They are diagnosable entities, characterized by constant or consistent differences. Species

in this monograph

Methods Three cladistic analyses are presented that are based upon Pennington's studies (1995, These are: 1)morphological 1996), with small modifications. analysis; 2) combined mol ecular and morphological and combined molecular and morphological 3) analysis; analy a of subset restriction site data for species not included by Pen sis, including cpDNA nington (1995, 1996). It should be noted that "Andira sp. nov. 1" and "Andira sp. nov. 2" (Pennington 1995, 1996) have subsequently been named A. carvalhoi and A. cordata, re spectively (Pennington & de Lima 1995). "Andira sp. nov. 3" (Pennington 1995, 1996) is

2003

21

ANDIRA

Table

2. Data matrix

of morphological

characters

of Andira.

Character flavum nitidum

Hymenolobium

Hymenolobium A. inermis subsp.

inermis

0 0 0

subsp. rooseveltii

0

subsp. glabricalyx 0 A. jaliscensis A. multistipula 0

0

A.

inermis

A.

inermis

0

A. cubensis A. galeottiana A. macrothyrsa

0 0

A. verm?fuga 1 A. humilis 2 A. anthelmia

0

A. carvalhoi

1

A. fraxinifolia A. legalis 0

1

A. marauensis

0

A.

/u'f?fti 1

A. ormosioides

0

A. surinamensis A. cordata

0

0

A. cori?cea

0

0 A. cujabensis A. grandistipula A. micrantha

0

0

A. parviflora 0 A. praecox 0 A. trifoliolata A. unifoliolata

0 0

A. chigorodensis A. taurotesticulata A. tervequinata A. macrocarpa

0 0 0 0

here included inA. nitida, which is now polymorphic for several character states (see dis cussion under that species), and "Andira sp. nov. 4" (Pennington 1996) is included in A. fraxinifolia (see notes under that species). Data matrices were handled using Dada (Nixon 1993). Parsimony analyses were car ried out using Hennig86 (Farris 1988), using the mh*bb* command, and PAUP* 4.0, beta 2 (Swofford 1999). Character optimization and tree printing was achieved using Clados 1.2 (Nixon

1992).

Morphological analysis. Characters and character states. The data matrix is shown in Table 2. The criteria for selecting morphological characters used in the cladistic analysis follow those of Lavin (1993), and include the assumption that they are functionally and developmentally independent of one another, and that they are intrinsic attributes of the taxa. The characters should also show uniform and consistent occurrence or absence related. among the terminal taxa, which implies that they are not environmentally

22

SYSTEMATICBOTANYMONOGRAPHS

VOLUME 64

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

treated as unordered, thus min imizing the assumptions built into the analysis. In the list of characters and character states, notes have only been added where characters were not presented or where their coding differs from that described by Pen nington (1996). These differences are: (i) anatomical studies of fruit have refined the coding of the endocarp character; (ii) observation of the pollination of A. anthelmia has suggested a re-coding of the petal color character based upon presumed pollinator; and Several multistate

(iii) including all species (rather than just those for which cpDNA data are available) al lows a character state to be added to the leaflet indumentum character (5) and two new characters to be included: leaflet number (4) and nature of the primary vein (6). The fol lowing conventions are adopted: 1) The first number of each character refers to its po sition in the data matrix for the morphological cladistic analysis; the second number (in to refers its in the data matrix of the combined analysis. 2) The parentheses) position states of each character are described and assigned a code (the number in parentheses after

each

Vegetative

character

state).

characters.

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

characters.

7 (43). Flowers yellow flowers (0); pink to purple,

to white,

the standard generally with red or purple markings the standard generally with a pale central marking, less than

ANDIRA

2003

23

18mm

long, pollinated by numerous species of small bees (1); flowers pink to purple, the standard with a pale central marking, 18-23 mm long, pollinated by large xylocopid bees (2).?The coding of this character reflects inferences of pollination biology described length inAndira varies continuously (Fig. 3), and alone it does not provide a good basis for the delimitation of character states. 8 (44). Calyx with indumentum (0); calyx glabrous (1). 9 (45). Wing petal sculpturing present (0); wing petal sculpturing absent (1). 10 (46). Gynoecium with indumentum (0); gynoecium totally glabrous or the ovary

above. Flower

with very few (1-3)

scattered hairs (1).

Fruit characters 11 (47). Fruit a samara (0); fruit a small, bat-dispersed drupe that dries smooth (1); fruit a small, bat-dispersed drupe that dries wrinkled (2); fruit a large, putatively rodent dispersed drupe, with non-fibrous mesocarp (large drupes with mesocarp drying finely granular) (3); fruit a large, putatively rodent-dispersed taining air spaces (A. galeottiana only) (4).

con

drupe, with fibrous mesocarp

12 (48). Endocarp absent (0); endocarp dominated by parenchyma and/or collapsed cell tissue (1); endocarp dominated by fibers (2); endocarp dominated by stone cells (3); character presents endocarp of fiber and stone cells (A. jaliscensis only) (4).?This an of because the lack outgroup species Hymenolobium problems, endocarp. Hawkins et al. (1997) argued that the most theoretically consistent way of coding such characters is by re-conceptualizing them as two characters (the first coding the presence and ab sence of the structure, the second coding the different types of structure, with the ter minal taxa that lack the structure scored as missing data). This is not a perfect solution, because it introduces problems of optimization?for the second character, species of for states of an endocarp, a structure that they do not is coded here as a single multistate character with ab coding as two characters, as recommended by Hawkins et

become optimized Hymenolobium have. For this reason the character sence as one state. Moreover, al. (1997), does not change lack the structure.

the outcome

of any analyses,

because

only the outgroup

taxa

and morphological analyses. Species for which cpDNA data are lacking (e.g., A. cubensis, A. chigorodensis) were not included in these analyses. The first analysis is identical to that presented by Pennington (1996) but includes the re-cod ing of the fruit characters and flower color character described above. The accession A. Combined

molecular

humilis RTP 239 has been identified as possessing a "foreign" plastome, probably derived from introgressive hybridization (see above), and is thus excluded from this and the fol lowing cladistic analysis. The data matrix is presented in Table 3. The second analysis includes a subset of cpDNA restriction site data for species not included by Pennington (1995, 1996). These are accessions o? A. jaliscensis, A. multistip ula,

A.

taurotesticulata,

A.

cujabensis,

and A. marauensis

that were

screened

for

the di

(Appendix 1) as part of the study of intraspecific and cpDNA polymorphism potential hybridization. Where cpDNA data were not avail were states character scored with question marks. The data matrix is presented in able, Table 3. agnostic mutations

described

above

SYSTEMATICBOTANYMONOGRAPHS

24

VOLUME 64

data for Andira. Characters 1-38 are re 3. Matrix of cpDNA restriction site and morphological a for accessions included in the molecular sites; characters 39-48 are morphological. species Multiple are abbreviated: AMC = A. M. de Carvalho; CEH = C. study are indicated by collector and number. Collectors de Lima; MC = M. Cheek; MS = M. Sugiyama; RTP = R. T. Pennington; TDP E. Hughes; HCL = H. Cavalcante = T. D. Pennington. Table

striction

Character

Hymenolobium

flavum nitidum Hymenolobium A. inermis CEH 1673 A.

inermis TDP

A.

inermis MC

A. macrothyrsa A. galeottiana

13558 3579 0 0

A. humilisRTP 269 A. fraxinifolia RTP 250 A. verm?fuga 0 A. anthelmia RTP 227,282 A. carvalhoi

RTP 229

A. carvalhoi

RTP 233

A. fraxinifolia A. fraxinifolia A. A.

RTP 213 MS

889

legalis RTP 307 legalis HCL s.n.

A. nitida RTP

300

A. nitida RTP 292 A. nifwfa RTP

301

A. ormosioides A. surinamensis A. cordata A. grandistipula A. parviflora A. unifoliolata inermis KY?

A.

512

A.

inermis RTP 580

A.

inermis RTP 589

A. multistipula A. jaliscensis A. taurotesticulata A. marauensis A. cujabensis A. carvalhoi AMC A

carvalhoi

RTP 217

12

3 4 5 6 7

1 1 1 1 1 1 ? 1 0 0 0 0 0 0

1 1 1 1 1 1 1 1 1

0

0

0

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ?

1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 1

0 0 0 0 0 0 0 0 1 0 1 0 1 1 1 1 1 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 1

1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 ? 1 1 ? 1 0 0 0 ?

1 1 1 1 1 1 1 1 0 1 0 1 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 ? 1

1 1 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0

0 1 1 1 ? 1 0 ?

1 90123456789012345 1 1 1 1 1 1 1 1 0 1 0 1 0 0 0 0 0 0 0 0 0 0 1 1 1 1 1 ?

1 0 1 ? 1

1 1 1 1 1 1

0 0 0 ? ? 1

1 1 1 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 0 0 0 1 1 1 1 ?

0 0 1 ? 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ?

0 0 0 0 0 1 1 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ?

? ? 1 1 1 0 0 0 1 0 1 0 1 1 1 1 1 1 1 1 1 1 1 0 1 0

0 0 0 ? 1 0 0 1 1 0 1 0 0 1 1 0 1 0 0 1 0 0 1 0 0 1 0 0 1 0 0 1 0 0 1 0 0 1 0 0 1 0 0 1 0 0 0 1 0 1 0 1 0 1 ? ? 1 0 1 0 ? ? 9 9 1 0 ? 0 ? ? 9 9 ? 0 1 0 1 ? 9 9? 1 ? 9 9 9 9 0 1 0 ? ? ? 1 ? ? ? ? ? ? 0 9 9 ? 0

2

? 0 0 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 ? 1 1 0 1 1

1 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 ? 0

1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 ? 1 1 1 1 1

? ? ? ? ?

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 ? 0 1 1 1 1 0 0 0 ? ? 0 0 1 ?

0 1 0 1 1 1 1 1 1 1 0? 0 1 0 1 0 1 0 1 0 1 0 1 0 1 0 1 0 1 0 1 0 1 0 1 ? 1 0 1 0 1 0 1 0 0 0 0 0 0 0 0 1 ? 1 ? 1 ? 1 ? 1 ? 1 ? 0? 0 ? 0 ?

9

9

9

?

? 9

? 9

? ? 9 ? ? ? ? ? ? 9

9

1 1 1 0 0 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 ? 1 1 1

0 0 1 1 1 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 ? 0 1 1

? ? ?

9

? ? 9

9

? ? 9

9

? ? ? ? 9

9

Results Morphological analysis. Pennington (1996) showed that a cladistic analysis of mor characters of a subset o? Andira species provided little phylogenetic resolution. phological A cladistic analysis of all species, using the twelve morphological characters listed above, also produced little phylogenetic resolution. This lack of resolution reflects the low num ber of characters and the high levels of homoplasy. With all characters run unordered, the trees (the limit that the computer analysis resulted in 2966 equally most-parsimonious could store; CI = 0.57, RI = 0.79, length = 38). The strict consensus tree is entirely unre

ANDIRA

2003

Table

25

3 continued.

2

4

3

Character_67890123456789012345678

Hymenolobiumflavum Hymenolobiumnitidum A. inermisCEH 1673 A. inermisTDP 13558 A. inermisMC 3579 A. macrothyrsa A. galeottiana A. ?urni/frRTP 269 A. fraxinifolia RTP 250 A. verm?fuga A. anthelmia A. carvalhoi

RTP 227,282 RTP 229

A. carvalhoiRTP 233 A.yhmm/o/??RTP213 A.yhmm/0/uzMS889 A. legalisRTP 307 A.

legalis UCLs.n.

A. nitidaRTP 300 A. mttfo RTP 292 A. n?tidaRTP 301

0 1 ? 1 10?00010?0001100000 ?0?0001110000 01??????01 10010000100100011 010001 01000100010000100100011 10010000100100011 010001 1000101000011 10001 07001 1000101100142 10001 00001 10011 100210?1?0122 00001 1 101100012 ?00?1 10701 101 00001100111001101100722 00001101011010111200012 00001100011001100100032

00001101011011100100032 00001101011011101100012 70077777011011101100012 70071107011010111200032 00001101011010111200032

00001 10101 07701 10701 00001101011011107170772

101 101

A. ormosioides

7077777701

1010101200012

A. surinamensis

00001101011010101100022

A. cordata

10101100010100100011113

A. grandistipula

11101100010100111717033

A.parviflora

70177107017100101001013

A. unifoliolata

70101100010100101071013

A. inermisRTP 5\2 A. inermisRTP 5S0 A. intrm?RTP 589

77777777070700100100011 77777777070700100100011 77777777070700100100011

A. multistipula

77777777077700110100011

A. jaliscensis

????????0???00102100014

A.

77777777077700101001031

taurotesticulata

A. marauensis

1107170772 1 10?1?0??2

7777777707770010111017?

A.cujabensis

77777777070700102001013

A. carvalhoiAMC

77777777077701

A. ozraz//i0/RTP217

77777777777771100100032

100100032

solved, except for a clade comprising A. cordata, A. cori?cea, and A. micrantha. With the character of leaflet number ordered, the analysis resulted in the same number of equally trees (2966, CI = 0.57, RI = 0.80, length = 38), but the strict consen most-parsimonious sus tree resolved another clade comprising A. unifoliolata and A. trifoliolata. Combined molecular and morphological analyses. The first analysis resulted in five trees of length = 76; CI = 0.72; RI = 0.88. The strict consen equally most-parsimonious sus is shown in Fig. 7. Of the equally most-parsimonious trees, two can be rejected be cause they result from incorrect optimization of missing values in accessions of A. nitida for character 47. These accessions are assigned a character state for large, rodent-dis for the two persed fruit, when they do not have this state (this species is polymorphic states of bat-dispersed fruit). Rejection of these topologies does not change the topology

26

VOLUME 64

SYSTEMATICBOTANYMONOGRAPHS

-Hymenolobium

flavum

-Hymenolobium

n'rtidum

-A.

inermis

CEH

1673

-A.

inermis

TDP

13558

-A.

inermis MC

CLADE I

3579

-A. parviflora -A.

cordata

-A.

grandistipula

-A.

unifoliolata

-A.

carvalhoi

CLADE II

RTP 229

-A. macrothyrsa

PLASTOME GROUP I

-A.

galeottiana

-A.

humilis

-A.

verm?fuga

-A.

surinamensis

-A.

ormosioides

-A.

anthelmia

-A.

legalis

-A.

legalis HCL sn

-A.

fraxinifolia

-A.

carvalhoi

-A.

fraxinifolia

-A.

fraxinifolia

-A.

nitida

RTP 300

-A.

nitida

RTP 292

RTP 269

RTP 227,282

RTP 307

CLADE III RTP 250 RTP 233 RTP 213 MS

889

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

27

ANDIRA

2003

of the strict consensus tree. When the same analysis was run on PAUP 4.0b2, where the fruit character (47) can be coded as polymorphic for A. nitida, 26 equally most-parsimo nious trees resulted, with an identical strict consensus. The greater numbers of equally trees reflect different arrangements of the accessions of A. fraxinifo most-parsimonious A. A. and carvalhoi {RTP 233). The semi-strict consensus tree ismore resolved, lia, nitida, with the accessions of A. nitida grouping with A. carvalhoi RTP 233. The only difference between this analysis and that presented by Pennington (1996) lies in the extra resolution in the strict consensus tree in Clade III (Fig. 7). Andira ormo sioides, A. anthelmia, and A. legalis form amonophyletic group, supported by the synapo morphy of larger flowers pollinated by xylocopid bees. Andira fraxinifolia, A. nitida, and A. carvalhoi are a monophyletic group diagnosed by the ability to root-sprout and form a multistemmed shrub. The combined analysis that included a subset of cpDNA restriction site data for species not included by Pennington (1995, 1996) resulted in ca. 2700 trees (the limit that the computer could store; length = 87; CI = 0.65; RI = 0.87). The strict consensus is shown in Fig. 8 and demonstrates that A. taurotesticulata, A. multistipula, and A. jaliscensis are members of Clade I, which previously only comprised accessions of A. inermis (Pen nington 1995, 1996). The affinities of A. multistipula and A. jaliscensis with A. inermis were suggested by their close overall similarity in morphology. Andira taurotesticulata, with its white-yellow flowers and large ridged fruit, is more divergent morphologically. Yet, it does have in common with A. inermis and A. multistipula an endocarp of parenchy matous

tissue, shown in the equally-most parsimonious tree (Fig. 9, character 48, state 1) to be the plesiomorphic state for Andira. Similarly, Pennington (1994) and Pennington and Gemeinholzer (2000) predicted that all species with an endocarp of stone cells would be members of Clade II. This is corroborated by the placement of A. cujabensis in this clade, and it is likely that A. cori?cea, A. trifoliolata, A. tervequinata, A. micrantha, and A. prae cox also belong here. Andira marauensis is a member of Clade III, placed as sister species to all other of this because it has the plesiomorphic state of restriction site characters clade, species 33 and 38. Clade III remains "cryptic" in the sense of Wojciechowski et al. (1993), be cause it is not marked by any morphological characters but well supported by cpDNA re striction Andira

site

characters. jaliscensis,

A.

multistipula,

A.

taurotesticulata,

A.

cujabensis,

and

A.

ma

rauensis have incomplete restriction site data, and therefore many missing values. Termi nal taxa with missing values are liable to occupy different topological positions in indi vidual equally most-parsimonious trees (Nixon & Davis 1991; Platnick et al. 1991), thus trees and decreasing the resolution increasing the numbers of equally most-parsimonious of consensus trees. Therefore, in an attempt to gain a more accurate idea of the relation ships of each of these species, separate analyses were run where all except one was ex cluded. In only one case, that of A. taurotesticulata, did this result in a different placement for the species in comparison with Fig. 8. In the analysis where A. taurotesticulata was in cluded alone, it is resolved as a sister species to the different accessions of A. inermis in Clade I in a strict consensus cladogram, because it possesses the plesiomorphic state for character 42 (leaflet undersurface indumentum). The results of this combined analysis that includes five extra species is used as the basis for the following discussion of character evolution and biogeography, despite the problem of the large number of equally-most parsimonious trees.

28

SYSTEMATICBOTANYMONOGRAPHS

VOLUME 64

-Hymenolobium

flavum

-Hymeno/obium

nrtidum

-A.

inermis

CEH

1673

-A.

inermis

TDP

13558 3579

-A.

inermis MC

-A.

inermis

RTP 512

-A.

inermis

RTP 580

-A.

inermis

RTP 589

CLADE I

-A. mu/tistipula -A.

jaliscensis

-A.

taurotesticulata

-A.

parvif/ora

-A.

cujabensis

-A.

cordata

-A.

grandistipu/a

-A.

unifoliolata

-A.

carvalhoi

RTP 229

-A.

carvalhoi

AMC

CLADE II

PLASTOME GROUP I

-A. macrothyrsa -A.

galeottiana

-A.

humilis

-A.

verm?fuga

-A. marauensis -A.

surinamensis

-A.

ormosioides

-A.

anthelmia

RTP 227,282

-A.

legalis

RTP 307

-A.

legalis

HCL sn

-A.

fraxinifolia

RTP 250

-A.

fraxinifolia

RTP 213

-A.

fraxinifolia

MS

CLADE III

889

-A.

n?tida RTP 300

-A.

n?tida RTP 292

-A.

n?tida RTP 301

-A.

carvalhoi

RTP 233

-A.

carvalhoi

RTP 217

FIG. 8. Strict consensus cladogram of 2996 equally most-parsimonious cladograms resulting from cladis tic analysis of combined Andira cpDNA restriction site and morphological of species data, including accessions not studied previously by Pennington Collector and number are indicated when more than one ac (1995,1996). cession for a single species was included in the molecular study.

ANDIRA

2003

29

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

arisen three times inde and A. multistipula and L. (pea), a single gene size from the large, fo

liaceous wild-type stipules (Marx 1987). It is tempting to ascribe the apparent lability of this character in Andira to a similarly simple genetic switch. Indeed, Andira could be a good model system to investigate the evolutionary significance of such a developmental mutation, especially in the case of A. multistipula, where it is the presence of these large, this species from A. persistent stipules that is the diagnostic character that distinguishes inermis. From the standpoint of the phylogenetic (Nixon & Wheeler concept species 1990), it was the fixation of the trait of large stipules in a population that represented the speciation

event.

Flowers (Fig. 9, character 43). White petals are shown to have arisen at least three times. They are a synapomorphy for Clade II, and an autapomorphy diagnosing both A. macrothyrsa and A. taurotesticulata. Other species not sampled in themolecular study (A. cori?cea, A. micrantha, A. praecox, A. tervequinata, A. trifoliolata) are known to have an endocarp composed of stone cells and to lack wing petal sculpturing, both synapomor phies for Clade II. Hence, all these species probably belong in this clade, which would then be defined by white flowers. Andira chigorodensis, the remaining species with white most to is A. similar and may be its sister species. petals, morphologically macrothyrsa Loss of wing petal sculpturing is a synapomorphy for Clade II, and an autapomorphy for A. taurotesticulata. Itmay have also been independently most is species closely allied with A. macrothyrsa.

lost by A. chigorodensis,

if this

Fruit type and dispersal (Fig. 9, character 47). The new restriction site data for A. tau rotesticulata demonstrate that it is a member of Clade I, and that there have been at least five independent origins of rodent dispersal from bat dispersal inAndira. This is one more independent origin than reported by Pennington (1996), confirming the prediction that four origins was an underestimate. Of the remaining species with large fruit, A. cori?cea and A. micrantha are likely to be members of Clade II based upon their floral characters of stone cells, but they share no great morphological similarity with A. the other species of that clade with large fruit. Andira macrocarpa has a woody endocarp, and its affinities are hard to guess, because it has never been collected in flower. Again, I suggest that further independent origins of rodent dispersal may have and endocarps

grandistipula,

occurred during the diversification

o? Andira.

Endocarp (Fig. 9, character 48). Examining the sequence of character state change on the cladogram (Fig. 9) shows a transition from weak endocarps of parenchyma to woody fibrous endocarps to endocarps composed of stone cells. This represents a sequence of in 2000; discussion above), and seed creasing seed protection (Pennington & Gemeinholzer have the been this pr?dation may pressure driving evolutionary sequence.

SYSTEMATICBOTANYMONOGRAPHS

30

r-fl-[]?Hymenolobium ?|?Hymenolobium

VOLUME 64

f/avum nitidum

1 7 13202530323642

fOttHf? 10 1 110

10 0

?A.

inermis

1673

CEH

1232 1824 rff--A.

4XH 0 ?

inermis

00

?-A.

CLADE I

13558

TDP 3579

inermis MC

parvif/ora 4246

I-A.

3 1012192122283637434548

ttOHUOHHH 000100101013

A.

cordata

CLADE M

16274147 0 113

A. grandistipu/a

A. unifoliolata

0 1111 394247 S-S? A.

RTP 229

carvalhoi

1? 3

4348

1423

0 1 4647 if 1 2

-A. macrothyrsa -??A. 3439

/H//W//S /?7P 269

-??A.

ft 1 1

HHKHKtH 0 0 0 10 2

PLASTOME GROUP I

galeottiana

?A.

verm?fuga

A. surinamensis A.

ormosioides A.

anthelmia

RTP 227r282 /?7P 307

4 6 8 3338

A.

/e#3//s

LKK}ft 10 0 11

A.

legalis HCL sn

A.

fraxinifolia

RTP 250

A.

fraxinifolia

RTP 213

-A. fraxinifolia 47 ?A,

MS

carvalhoi

CLADE III

889 RTP 233

A. n?tida RTP 300 A. n?tida RTP 292 A. n?tida RTP 301 FIG. 9. An equally most-parsimonious cladogram resulting from cladistic analysis of combined Andira data. The bars representing character changes are labelled above with cpDNA restriction site and morphological the character number and below with the character state. Solid black bars = unique changes, stippled black bars = and number are indicated when more than one accession for a single species changes. Collector homoplasious was included in the molecular study.

2003

ANDIRA

31

CRYPTICCLADES Pennington (1995) reported that none of themonophyletic groups, even the well-sup ported clades, discovered in themolecular analysis o? Andira had been recognized by pre vious workers. They lack major morphological innovations and hence are "cryptic" clades sensu Wojciechowski et al. (1993). The combined analysis demonstrates that new micro characters provide unambiguous support for some of these cryptic groups. morphological Clade II is supported by two unique micromorphological character states: lack of wing an and of stone cells petal sculpturing (character 45) (character 48). All species endocarp of Clade I have an endocarp of parenchyma, but this maps as a plesiomorphy for Andira rather than a synapomorphy. At a higher taxonomic level, micromorphological characters have also proved congruent with other clades that are well supported by molecular data in (Rudall 2000). In both cases, reciprocal illumination provided by new monocotyledons hypotheses of grouping from molecular data is important. For example, itwas only when the cpDNA restriction site data suggested the Clade II grouping of Andira species that I noticed

that endocarps of these species appeared somewhat paler and harder than in other species, which prompted the anatomical study reported above. Other clades, such as clade III, remain cryptic, supported by restriction site data but not by morphological characters. This raises the question of whether they should be rec ognized in formal classifications (Wojciechowski et al. 1993; Pennington & Gemeinholzer 2000), such as an infrageneric classification o? Andira.

INTRAGENERICCLASSIFICATIONSIN THE LIGHTOF THE CLADISTICANALYSES The cladistic analyses show that the infrageneric classifications proposed for Andira Bentham (1860, 1862) and Mattos (1979) are flawed. The only section that has been by that is sect. Paucifoliolata Mattos, comprising only A. may be monophyletic proposed A. amonophyletic and This is cladis unifoliolata trifoliolata. group in themorphological tic analysis, if the character of leaflet number is treated as ordered. The gynoecium indu mentum character used by Bentham to define his sections Lumbricidia and Euandira, and by Mattos to define corresponding subsections in her section Lumbricidia is homoplasious is paraphyletic and (Fig. 10); thus Bentham's section (Mattos's subsection) Lumbricidia section Euandira (Mattos's subsection Glabratae) polyphyletic. the cladistic framework presented here does not provide a clear route Unfortunately, to a new infrageneric classification o? Andira based upon monophyly. There are four well supported clades (Fig. 9): Clade I, Clade II, Clade III, and the group of Clade II, Clade III and Plastome Group I; however, giving these clades formal taxonomic recognition leaves the weakly supported "Plastome Group I," supported as monophyletic by a single, homo restriction site character. both Clade III and I remain cryptic, Clade Moreover, plasious restriction site but not data characters. Whilst there is no supported by by morphological reason to doubt that these cryptic clades reflect true phylogeny because they lack mor ultrastructural and characters may well phological support (indeed, micromorphological be found to support them, as was the case for Clade II), I question the practical utility of giving them formal taxonomic recognition. Because of the lack morphological characters, identification. Hence, I propose that sectional they cannot be keyed out for conventional classification within Andira should be abandoned. A further factor supporting this

SYSTEMATICBOTANYMONOGRAPHS

32

?Hymenolobium -Hymeno/obium

flavum nitidum

VOLUME 64

= 0 = 0 -A.

= 0

taurotesticulata ?A.

= 0

jalisensis

-A. 1-A -A.

inermis

TOP

-A.

inermis

-A.

inermisRTP

CEH

1673=

inermis

RTP 580=

-A.

inermis

RTP 589

?A.

= 0

0

512=

-A.

0

13558=

3579

inermis MC

0 0 = 0

= 0

multistipu/a -A.

= 0

cujabensis

= 0

-A. parviflora

cordata

=

-A.

grandistipula

A.

A.

c

A.

carvalhoi

RTP

A.

carvalhoi

AMC=

-j?A.

0

i-A.

=

humilis .

RTP =

verm?fuga

ormosioides= A.

A.

RTP

legalis

RTP 307

legalis

HCL

fraxinifolia

RTP

250

fraxinifolia

RTP

213=

fraxinifolia

MS

A.

n?tida RTP

A.

carvalhoi A.

0 0

0


0
A. n?tida RTP 301=

?

sn = 0

889=

292=

= 0

= 0

A.

n?tida RTP 300=

0

227,282=

A. A.

1

0

anthelmia

-r ??A.

?A.

269= 1
= 0

surinamensis A.

1

1

marauensis= A.

= 0

0

A. galeottiana

??A.

unifoliolata

= 0

0

229=

A. macrothyrsa=

1

1

0


RTP

carvalhoi

233=

RTP

217=

0 0

FIG. 10. An equally most-parsimonious cladogram resulting from cladistic analysis of combined Andira data showing character state changes for indumentum of the gynoe cpDNA restriction site and morphological taxa marked Terminal cium. The bars represent character state changes from a hairy to a glabrous gynoecium. "1" have a glabrous gynoecium. Andira nitida and A. ver terminal taxa marked "0" have a hairy gynoecium; and number are indicated when more than one acces m?fuga have both hairy and glabrous gynoecia. Collector sion for a single

species was

included

in the molecular

study.

ANDIRA

2003

33

argument is thatmolecular data are not available for some species whose relationships hard to estimate based upon morphology alone (e.g., A. cubensis, A. macrocarpa).

are

BIOGEOGRAPHY Distribution

and Habitats

The majority o? Andira species are endemic to South America. The exceptions are A. and A. galeottiana and (endemic to Cuba), A. jaliscensis (endemic toMexico), A. inermis (widespread throughout the entire Neotropics and also present in Africa). cubensis

Most species grow in rain forests. Four species, A. cordata, A. cujabensis, A. humilis, and A. verm?fuga grow in the savanna woodlands ("cerrados") of central Brazil. Andira inermis subsp. rooseveltii occurs in a similar wooded savanna habitat in Africa. Andira carvalhoi is entirely restricted to the sandy coastal restinga scrub and forests of eastern Brazil. Andira fraxinifolia, A. legalis, A. anthelmia, A. ormosioides, and A. nitida grow both in restinga and adjacent rain forests in eastern Brazil. Andira jaliscensis is a species of seasonally dry tropical forests, as is A. inermis subsp. glabricalyx. Andira inermis subsp. inermis also occurs in seasonally dry tropical forest in Central America and on the Caribbean coasts of Venezuela and Colombia, but elsewhere in the Neotropics, and in Africa, it grows in rain forests. Origin

and Transatlantic

Distribution

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

(Pennington et al. 2001). Furthermore, the lack of resolution in the basally di Andira clade (clade I, Fig. 8), inwhich all distribution areas (South America, Cen vergent tralAmerica, the Caribbean, and Africa) are represented, means that little can be inferred about the early diversification o? Andira. The placement of A. inermis and A. jaliscensis in this basal lineage is intriguing and not inconsistent with a northern Boreotropical ori

unresolved

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

SYSTEMATICBOTANYMONOGRAPHS

34

VOLUME 64

current biota must have the ability to disperse over water. The distance of Jamaica from the nearest island where A. inermis is present (Haiti; ca. 200 km) is, of course, an order of shorter than that between South America and Africa. magnitude Nothing is known of the potential of Andira fruits for long-distance dispersal by float ing in sea water. Four observations are pertinent: (i) the distributions of A. surinamensis and A. anthelmia often appear to follow river courses, a pattern seen inmany taxa known to be dispersed by water (data from herbarium labels; pers. obs.; H. C. Lima, pers. comm.); (ii) the fruits of A. galeotiiana are often found washed up on beaches inMexico (Gunn & Dennis 1976), and this species is characteristic of river and lake banks, and flooded areas (Pennington & Sarukh?n 1968); (iii) Andira seeds, protected by their hard endocarps, have remarkable viability; seeds of A. carvalhoi that were not excised from their endocarps germinated at Kew Gardens 14 months after collection; (iv) Andira iner mis grows on the strand line in Cameroon (M. Cheek, pers. comm.; see notes onM. Cheek it often grows along rivers and can survive inmangrove swamps 3579); in the Neotropics (in Colombia; notes on H. Murphy & G. Parra 723). The low frequency of long distance dispersal events may be indicated by the absence of A. inermis from Cuba (where only A. cubensis occurs) despite the proximity toHaiti (ca. 100 km at the closest point). Given the presence of A. inermis on the islands of the Lesser and all the other major islands of the Greater Antilles, this absence from Cuba is puzzling, although a similar pattern occurs in Coursetia caribaea (Jacq.) Lavin van carib aea (Lavin 1988). No other species of Andira have ranges as wide as that of A. inermis. For example, in the Caribbean, A. surinamensis (widespread in South America) is present only on Trinidad and Tobago, which are extremely close to the South American mainland. It is

Antilles

enigmatic that long-distance dispersal of these species is less effective, because there are no apparent differences in their dispersal biology in comparison with that of A. inermis. The explanation may lie in the ecology of A. inermis, such as its ability to survive inman grove swamps and on beach strand-lines, which appears unique in the genus. Recent

Patterns

in South America

The lack of cpDNA divergence within the species of clades I, II, and III (Fig. 9) is con sistent with the origin of most South American species in recent times, possibly as late as the Pleistocene. Morphological change has occurred, but the slowly evolving cpDNA has accumulated few, or in the case of Clade III, no differences among species. Such patterns are characteristic of recent radiation on oceanic islands (reviewed by Bateman, 1999). All

(an Amazonian species of Clade III, with the exception of A. surinamensis occur seems to have in moist the forests of coastal southeastern Brazil. there Thus, species), been a recent burst of speciation in this area. Andira surinamensis and the other species of Clade III are separated by the wide "dry diagonal" of cerrado and caatinga vegetation that runs across central and northeastern Brazil. The close relationship of A. surinamensis and the species of Clade III is suggestive of previous links between the mesic vegetation of southeastern Brazil and Amazonia, which are more often evidenced by disjunctions of the same species between these two areas (Mori et al. 1981; Prance 1985, 1987). The species of Clade II are from the central Amazonian region (A. parviflora, A. uni the Guianas (specifically Guyana; A. grandistipula), and the central Brazilian (A. cordata, A. cujabensis). The lack of divergence between the plastomes of these species also suggests a relatively rapid radiation of these species. The morphologi

foliolata), cerrados

cal characters

supporting

this clade (lack of wing

petal sculpturing,

endocarp composed

2003

ANDIRA

35

H oc

H 20e CERRADO BIOIVJE

50?

80? -Contact

line of Andira

cujabensis/cordata

-Contact line of harry & glabrous Andira humilis .Miiimmi.

inContact

line of

Pterodon

.

hairy

& glabrous

forms of forms

of

emarginatus

Lines dividing

distinct

northern and

south-eastern

cerrado

sites

FIG.

11. Cerrado biogeography. Dashed line: line of contact between the parapatric distributions of Andira and A. cujabensis line with dots: line of contact between hairy and glabrous forms (see Fig. 39). Dashed of A. humilis. Comb: line of contact between hairy and glabrous forms of Pterodon lines: emarginatus. Dotted cordata

lines dividing the distinct northern and southeastern cerrado tion of 98 areas of cerrado vegetation (Ratter et al. 1996)

sites identified

by analysis

of the floristic

composi

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

SYSTEMATICBOTANYMONOGRAPHS

36

VOLUME 64

line of contact of these species' distributions through the states of Goi?s and Tocantins closely matches the lines dividing the distinct northern and southeastern sites in the cerrado discovered by TWINSPAN (two-way indicator species analysis) of the floristic composition of 98 areas of cerrado and Amazonian savanna vegetation (Fig. 11 ;Ratter et al. 1996). This is also the line of contact for the glabrous and hairy forms of A. humilis (Fig. 11; see account under that species), and similar distributions are shown by hairy and glabrous forms of Pterodon emarginatus Vogel (Leguminosae, Papilionoideae; Fig. 11; R. T. Pen nington, unpubl.). The subspecies of Diptychandra aurantica Tul. also have similar distrib vegetation

utions, with

subsp. aurantica occurring in the southeast of the cerrado biome, and subsp. epunctata (Tul.) Lima, Carvalho & Costa occurring in the north, though in this case, the ranges only abut at their southern ends inMinas Gerais (de Lima et al., 1990). These con gruent patterns are suggestive of a common underlying historical factor. A possibility is re

cent separation in the late Quaternary of the now continuous cerrado vegetation by more mesic vegetation, such as that suggested by the palaeopalynological data of Ledru (1993).

TAXONOMY Note: The following terms are used to describe the placement of indumentum on ovary, which is approximately oval in cross section. The "upper surface" refers to areas on and adjacent to the adaxial suture, and the "lower surface" refers to the areas and adjacent to the abaxial suture; the "sides" refer to the two regions bounded by

the the on the

adaxial and abaxial sutures. Andira

Lamarck, Encycl. 1: 171. 1783, nom. conserv. Andira sect. Euandira Bentham, Comm. legum. gen. 45. 1837. Andira sect. Lumbricidia subsect. Glabratae N. F. 58: 2. 1973, nom. superfl.?Type: Andira inermis (W. Mattos, Loefgrenia

Wright) Lumbricidia

DC.

Vellozo, Fl. flumin. 305. 1829. Andira sect. Lumbricidia (Vellozo) Ben Comm. tham, legum. gen. 43. 1837. Andira sect. Lumbricidia subsect. Lumbri cidia (Vellozo) N. F.Mattos, Loefgrenia 58: 2.1973.?Type: Lumbricidia legalis Vellozo [=Andira legalis (Vellozo) Toledo]. Andira sect. Paucifoliolatae N. F. Mattos, Loefgrenia 58: 3. 1973.?TYPE: Andira unifoliolata

Ducke.

Trees, shrubs, or rarely geoxylic suffrutices. Bark often producing small amounts of red ex?date when cut. Indumentum of simple, red-brown or occasionally pale whitish hairs, or absent. Stipules large and persistent or small, narrow and early caducous. Leaves spirally arranged, imparipinnate, with up to 9 pairs of leaflets, occasionally 3-foliolate or 1-foliolate; rhachis canaliculate; leaflets subtended by stipels (occasionally absent); peti olules stout, swollen, canaliculate; leaflets glabrous adaxially, glabrous or hairy abaxially, primary vein channelled or plane adaxially, prominently raised abaxially, secondary veins or mixed eucamptodromous-brochidodromous. brochidodromous Inflorescences axillary and terminal, paniculate, with pale brown, brown, or red-brown, simple indumentum, the branches subtended by caducous bracts. Flowers pedicellate or occasionally sessile, each subtended by a caducous bract; paired caducous bracteoles at base of calyx. Calyx shal lowly to deeply and subequally 5-lobed, the upper lobes broader than the lower lobes, all lobes with simple, red-brown indumentum, or glabrous. Petals 5, free, clawed, purple,

ANDIRA

2003

37

pink, or white; wing petals with or without lamellate sculpturing; keel petals overlapping, firmly attached, but not united. Stamens 10, the filaments united for at least half of their length, the vexillary stamen free. Ovary distinctly stipitate, with simple hairs or glabrous; ovules 1-8. Fruit a 1 (-2-3)-seeded drupe, ? globose to elongated; sutures (along abaxial and adaxial surfaces of the fruit) raised or sunken or indistinct; stylar remnant present or indistinct; exocarp glabrous, green, brown, or occasionally yellow when ripe and fresh; sweet and mesocarp fibrously fleshy smelling (drying hard, fibrous and granular) or hard, non-fibrous, and odorless (drying hard and finely granular); endocarp very hard, woody, or woody and fibrous. Seeds white, filling the entire cavity, the hypocotyl-radicle axis a small fold at the apex; testa dark reddish brown, adhering to the endocarp. Seedlings cryp number: x = 10, 11. tohypogeal. Chromosome The order of species reflects the groups discovered by the combined analysis of The relationships of species not cpDNA restriction site characters and morphology. characters. The included in this analysis are estimated based upon their morphological A. multistipula, groups are (refer to Fig. 8): (i) "Clade I": A. inermis, A. jaliscensis, A. taurotesticulata, and A. cubensis (the last species not included in the combined analy to A. inermis); similarities sis but placed in this group because of its morphological A. A. "Plastome I": A. A. humilis, and (ii) macrothyrsa, galeottiana, verm?fuga, Group not included in A. chigorodensis last the combined (the analysis but placed species to A. macrothyrsa); similarities here because of its morphological (iii) "Clade III": A.

A.

surinamensis,

anthelmia,

A.

legalis,

A.

A.

ormosioides,

fraxinifolia,

A.

n?tida,

A. carvalhoi, A. marauensis, and A. macrocarpa (the last species not included in the of combined but here because its similarities to A. suri analysis morphological placed namensis);

(iv)

"Clade

II": A. praecox,

A. micrantha,

A.

cori?cea,

A.

trifoliolata,

A.

ter

vequinata (these species not included in the combined analysis but included here be cause they share two features that define this clade: an endocarp of stone cells and wing A. cujabensis, petals without sculpturing), A. parviflora, A. cordata, A. grandistipula, and A.

unifoliolata.

Keys

to the Species

of Andira

of Andira are easy to identify if complete material of leaves, flowers, and fruit is available, Specimens one emphasizing to distinguish when sterile. Two keys are presented: but they are often very difficult vege use of the keys, especially of the first key, re tative characters and the other flowers and fruits. Successful the three different among quires distinguishing leaflets (refer to Fig. 2 and Fig. 12). Understanding ical (refer to Fig. 13).

used to describe leaf morphology terminology is measured from the joint of the stem and petiole

The length

surface of the found on the abaxial types of indumentum of the terminology used to describe fruit shape is also crit

FIG.

long), dense,

fine, pale,

axis All

types of the abaxial leaflet surface o? Andira. A. Short (ca. 0.1 mm long), appressed erect hairs of A. fraxinifolia. B. Long (ca. 0.5 mm long), flexuose, C. Long (ca. 1mm (See also Fig. 2B, C, D.) tangled hairs of A. jaliscensis.

12. Indumentum

hairs of A. surinamensis.

follows the guidelines of Hickey (1979). Leaf to the joint of the rhachis and terminal petiolule.

SYSTEMATICBOTANYMONOGRAPHS

38

VOLUME 64

of Andira. A. ? Globose, FIG. 13. Fruit morphology smooth, dried fruit (ca. 3.5 cm long) of A. jaliscen C. Elongated, wrinkled, sis. B. Elongated, dried fruit (ca. smooth, dried fruit (ca. 6 cm long) of A. chigorodensis. 4.5 cm long) of A. nitida.

of petiolule length is length and leaflet size and shape are for the terminal pair of leaflets. Leaflet to the tip of the lamina and does not include the length of the petiolule. Acumen length is mea the acumen is broad, this point is sured from the point where the leaflet margin curves towards the apex. Wlien

measurements

from the base ill defined, Flower wing

petals

sculpturing Many "elongated" 13C). Fruit

lengths must be treated as approximate. and ismeasured from the base of the calyx to the end of the length is reported for dried specimens, in fully open flowers. A fully open flower is one in which the standard petal is reflexed. Wing petal is described following the terminology of Stirton (1981).

so acumen

have more or less globose fruits (Fig. 13A). Other (Fig. 13B, C). Fruit of some species dry smooth (Fig. 13A, from the tip of the stipe (where it joins length ismeasured from the abaxial to adaxial is the maximum measurement species

fruit height surement between

the fruit walls.

I. Key emphasizing 1. Leaves

vegetative

all uni- or trifoliolate;

2.

Leaves

trifoliolate.

2.

Leaves

unifoliolate.

1. Leaves 3.

Geoxylic

3.

Tree or shrub.

5.

vein plane on adaxial

primary

surface. 28.

A. unifoliolata. pairs of leaflets, or sometimes

A.

irifoliolala.

also some

leaves

trifoliolate;

primary

shrub-like,

to 2 m

suffrutex

forming mats

more

up to 10 m

in diameter

(occasionally

than 1.7 cm long and 0.5 cm wide

at base, persistent,

crowded

glabrous

Leaflets 6.

7.

1.

at in

3. A. multistipula. (use lens or

tiny (<0.1 mm long) appressed straight hairs beneath (use 25. A. grandistipula. leaflets up to 4 pairs; the Guianas. with erect or ? appressed, often flexuose hairs (0.2-1.0 mm long; to the naked eye); leaflets 3-9 pairs; eastern Brazil.

Leaflets abaxially lens or microscope); 6. Leaflets abaxially visible

tall). 10. A. humilis.

and shoot apices.

with sparse, erect hairs); Amazonia. (very occasionally indumentum abaxially hairy, sparsely hairy, or with tiny appressed Guianas and eastern Brazil. microscope); Leaflets

vein

surface.

Stipules generally florescence bases 5.

characters.

29.

two or more

all with

sunken on adaxial

4.

species have fruits that are referred to as B) and of others distinctly wrinkled (Fig. the body of the fruit) to the tip of the fruit; suture; fruit width is the maximum mea

with

at maturity; leaflets hairy abaxially, Stipules red-brown pubescent, glabrescent over 100 g when dry, pale hairs red-brown, erect; fruit 5.6-12 cm long, weighing 14. A. brown when fresh and dry, odorless when fresh. legalis. when (red-brown appressed-pubescent Stipules generally glabrous at maturity to leaflets sparsely to very sparsely hairy abaxially, hairs red-brown young); ca. 20 g when dry, whitish, ? appressed to ? erect; fruit 3-6.2 cm long, weighing 13. A. anthelmia. dark brown when fresh and dry, sweet smelling when fresh.

4.

sometimes less than 1.7 (-2.0) cm long and 0.5 cm wide at base, often caducous, entirely absent (rarely persistent and then paired only at leaf bases and not crowded at inflo rescence bases or shoot apices).

Stipules

8.

Leaflets

with

indumentum

abaxially

(use lens or microscope).

ANDIRA

9.

39

undersurfaces (particularly primary and secondary veins) with erect, gener visible with the hairs (>0.2-1.0 mm generally ally flexuose long); indumentum to touch. naked eye and perceptible

Leaflet

tan red-brown), abaxially with dense indumentum of pale (occasionally leaflets appear pale abaxially (seen with gled hairs that ? obscure the epidermis; the naked eye). to cordate; flowers 11. Leaflets base rounded 5.5-7 mm coriaceous, long; 21. A. cujabensis. Brazil.

10. Leaflets

to subcoriaceous, to rounded; flowers base obtuse Mexico. 2. A. jaliscensis. long; abaxially with indumentum of red-brown hairs (or if pale, then indu leaflet often appearing red sparse) that do not obscure the epidermis;

11. Leaflets

chartaceous

9-1 lmm 10. Leaflets mentum brown

is sparse). abaxially (to the naked eye; if pale then indumentum 12. Secondary veins plane or slightly raised (occasionally raised) prominently indumentum abaxially, tertiary veins plane or very slightly raised abaxially; of abaxial suture;

leaflet

surfaces

in montane

forests

caducous; fruit distinctly ribbed from suture to at 700-2000 in lowland forest m, occasionally (Ecuador).

12. Secondary veins raised abaxially; ribbed from

prominently indumentum

5. A.

taurotesticulata.

raised

abaxially, tertiary veins prominently of abaxial leaflet surface persistent; fruit not

suture to suture, or with

broad,

indistinct ribs;

lowland

forests

and savanna. 13. Leaflets

2-3 pairs, the apex rounded, often slightly retuse; fruit 10-11 cm long, weighing 300 g when dry. H.A. macrocarpa. 13. Leaflets (2-) 3-7 pairs, the apex obtuse to rounded, often acuminate, cm long, sometimes retuse; fruit 2.4-6 (-10 [A. galeottiana only]) 10-20

(-125 [A. galeottiana only]) g when dry. cm long, 40-125 6-10 g when dry, dried mesocarp 8. A. galeottiana. structure; Mexico. relatively soft with sponge-like 14. Shrub or tree; fruit 2.4-6 cm long, weighing 10-20 (-30) g when

weighing 14. Tree;

fruit

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

Brazil

and

Bolivia

(cerrado), occasionally Amazonia. 9. A. verm?fuga.

1-9 mm long; stipules to 16 mm long, caducous or moderately persistent; upper and lower surfaces and walls of ovary hairy; dry fruits smooth; eastern Brazil (restinga, rain forest, campo rupestre, and secondary vegetation).

16. Stipels

17. Shrub or small

tree to 12 m, with flowers

spreading

crown

in

13-17 mm

long; fresh fruit A. fraxinifolia. 16. green with green mesocarp. 17. Tree to 30 m, tall with small crown in open situations; open

situations;

18-23 mm long; fresh fruit very dark brown 15. A. ormosioides. pale greenish-white mesocarp. tiny (<0.2 mm long), tightly appressed hairs; indumentum to the naked eye or perceptible by touch (use lens or flowers

with

9.

Leaflets generally

abaxially with not visible

microscope). 18. Tertiary impressed

raised on abaxial leaflet surface, distinctly or slightly impressed on adaxial leaflet surface.

veins

veins secondary 12. A. surinamensis.

SYSTEMATICBOTANYMONOGRAPHS

40

VOLUME 64

18. Tertiary veins plane or slightly raised on abaxial leaflet surface, plane or slightly raised on adaxial leaflet surface. 19. Shrub. 20. Leaflets

1-2 pairs; flowers

20. Leaflets

2-4

9 mm

long; Venezuelan

secondary

Guayana. 27. A.

10-13 mm

(-5) pairs; flowers

veins

long; eastern Brazil.

tervequinata. 17. A. n?tida.

19. Tree. 21. Leaflets

(1-) 2 (-3) pairs. 22. Secondary veins 6-7; leaflets widely elliptic to widely pairs; small tree to 7 m tall; Venezuelan Guayana. 22. Secondary

veins

to 10 mm

23. Stipules

long; stipels

1-2 mm

long; flowers 26.

long; Amap?. to 2.5 mm

23. Stipules

long; stipels absent 9-11 mm long; Bahia.

1-2

27. A. tervequinata. obovate (occa

elliptic to narrowly (1-) 2 (-3) pairs; tree to 35 m

leaflets

8-10; obovate),

sionally widely Brazil.

obovate,

(perhaps

tall; eastern 6.5-7 mm A. praecox. early cadu 18. A. marauensis.

cous); flowers 3-9 (-10) pairs. 24. Leaf axis 37-55 cm long; leaflets 7-9 pairs. 7. A. chigorodensis. 24. Leaf axis 6-33 (-40) cm long; leaflets 3-8 (-10) pairs. 25. Leaflets coriaceous; flowers red to purple; Cuba. 4. A. cubensis.

21. Leaflets

25. Leaflets marked

flowers white to yellowish, the standard subcoriaceous; red or purple; Panama and South America.

with

26. Stipules absent (or early caducous; never seen), stipels to 2 mm long, caducous; fruit 5-5.5 cm long, smooth, weighing 10-20 g when dry. 6. A. macrothyrsa. to 12 mm long, stipels 1-3 mm long, generally per cm long, distinctly ribbed from suture sistent; fruit 5.4-7.2 to suture, weighing 100-200 g when dry. 5. A. taurotesticulata. (use lens or microscope).

26. Stipules

8.

Leaflets

glabrous abaxially 27. Leaflet base truncate or cordate 27. Leaflet

base obtuse

eastern Brazil,

Guianas, 28. Shrub

22. A. cordata. (rarely rounded); Brazil, in cerrado. ? or truncate the Amazonia, cordate); (rarely slightly in rain forest and restinga.

to rounded

(occasionally

small tree to 10 m).

19.

A.

carvalhoi.

28. Tree. in 2-9 pairs (some leaves on a plant always with more of leaflets); Neotropics and Africa. 1. inermis. A. 29. Leaflets in 2-3 pairs; South America. 29. Leaflets

30. Stipules

to 5 mm

quite persistent,

wide

at base,

than 3 pairs

to 15 mm Guianas.

30. Stipules caducous, often long; Central Amazonia,

entirely absent, <1 mm wide eastern Brazil.

at base,

long; the 24. A. cori?cea. to 6 mm

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

II. Key emphasizing 1. Leaves

flower and fruit characters.

all uni- or trifoliolate; primary vein plane on adaxial trifoliolate. 28. A. trifoliolata.

2.

Leaves

2.

Leaves

unifoliolate.

29. A. unifoliolata.

surface.

2003 Leaves

all with

two or more

sunken on adaxial 3.

41

ANDIRA

pairs of leaflets, or sometimes

also some leaves

Stipules generally more than 1.7 cm long and 0.5 cm wide florescence base and shoot apex. 4.

19-24 mm

Flowers 5.

trifoliolate;

primary

vein

surface.

cm

at the base, persistent,

crowded

long; eastern Brazil. over long, weighing

100 g when dry; stipules red-brown at maturity. 14. A. legalis. young, glabrescent cm long, weighing ca. 20 g when Fruit 3-6.2 dry; stipules red-brown 13. pubescent when young, generally glabrous at maturity. Fruit

5.6-12

at in

pubescent

when 5.

4.

Flowers 6.

11-15 mm

and Guianas. long; Amazonia in (2-) 3-4 pairs, apex rounded (rarely obtuse) with an acumen to 3 mm long, tiny (<0.1 mm long) appressed hairs abaxially (use lens or microscope). 25. A. grandistipula.

Leaflets with

pairs, apex acute to obtuse with an acumen to 15 mm long, glabrous A. with few, erect hairs). 3. multistipula. (very occasionally abaxially sometimes entirely Stipules less than 1.7 (-2) cm long and 0.5 cm wide at base, often caducous, base absent (rarely persistent and then paired only at leaf bases, but not crowded at inflorescence 6.

3.

appressed A. anthelmia.

in (5-) 6-9

Leaflets

or shoot apex). 7. Fruit 5-11 cm long, weighing (40-) 8. Leaflets glabrous abaxially. 9.

Flowers

9.

Flowers

14-15 mm

g when

dry.

long; shrub or small tree to 10 m; eastern Brazil,

in restinga. 19. A. carvalhoi.

6-7 mm

10. Ovary 10. Ovary 8.

100-300

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

Leaflets

with indumentum abaxially. 11. Fruits ? globose; petals white, the standard marked with red; tertiary veins plane on A. abaxial surface of leaflet. 5. taurotesticulata.

11. Fruits

elongated; petals tiana; flowers unknown

pink to violet, the standard marked with white (A. galeot for A. macrocarpa); tertiary veins distinctly raised on abax

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

300 g when dry; Ecuador. fibrous; fruits weighing 10-20 (-30) g when dry. (-7) cm long, weighing 13. Petals white to yellow (occasionally pale pinkish or pale

H.A.

macrocarpa.

Fruit 2.4-6

lilac),

the standard generally

red or purple markings. 14. Ovary glabrous. 22. A. cordata.

with

14. Ovary pubescent. 15. Leaflets with

on abaxial

erect indumentum

16. Secondary

veins

epidermis clearly Para), in rain

surface.

leaflets abaxially tree to 20 m, visible; forest.

10-15;

with

red-brown

the indumentum, smooth; Brazil (Amazonas, A. 20. parviflora. with generally pale indumentum,

bark

veins 8-10; leaflets abaxially 16. Secondary often so dense that the epidermis a is hidden; tree to 12 m (occasionally do Sul, shrub), bark thick, fissured; Brazil (Mato Grosso, Mato Grosso 21. A. cujabensis. Goi?s, Para), in cerrado and gallery forest. glabrous or with appressed indumentum 17. Leaflets 4-9 pairs; endocarp of fruit fibrous

15. Leaflets

inserted). 18. Flowers

9-11 mm

18. Flowers

5-6

17. Leaflets a needle

long;

(-7) mm

leaf axis

on abaxial

surface.

(the point of a needle

10-33

long; leaf axis 37-55

is easily

cm

long; leaflets 4-8 6. A. macrothyrsa. pairs. cm long; leaflets 7-9 pairs.

(-40)

1-2 (-3) pairs; endocarp of fruit very hard, amorphous is inserted only with difficulty).

7. A. chigorodensis. (the point of

SYSTEMATICBOTANYMONOGRAPHS

42

19. Flowers

VOLUME 64

6.5-7 mm

ally with

long; secondary veins 8-12 per leaflet; leaves gener tree 2 pairs of leaflets (occasionally with 3 pairs or trifoliolate); to 30 m. 26. A. praecox.

9 mm long; secondary veins 6-7 per leaflet; leaves with 1-2 27. A. tervequinata. pairs of leaflets; shrubs or small trees to 7 m. red or pink to purple, the standard generally with a white or cream central mark 19. Flowers

13. Petals

ing. 20. Dry fruits

distinctly wrinkled (Fig. with erect indumentum

13). on abaxial

surface. 9. A. verm?fuga. glabrous or with appressed indumentum on abaxial surface. 22. Geoxylic suffrutex forming mats up to 10 m in diameter (occasionally to 2 m tall); fresh mature fruit yellowish; central Brazil and shrublike,

21. Leaflets 21. Leaflets

10. A. humilis. Paraguay, in cerrado. 22. Shrub or tree to 40 m; fresh mature fruit green; Amazonia, eastern Brazil.

the Guianas,

23. Secondary veins plane to slightly raised abaxially, tertiary veins 17. A. flowers 10-13 mm long; eastern Brazil.

and

plane; n?tida.

raised abaxially; flowers 23. Secondary veins and tertiary veins prominently 12. A. surinamensis. 13-18 mm long; Amazonia and the Guianas. not and but 20. Dry fruits smooth (the surface sometimes wrinkled) warty, rough (Fig. 13). 24. Flowers

7-8.5 mm

24. Flowers

9-23 mm

A. 4. cubensis. long; Cuba. Caribbean (excluding Cuba), and Africa. long; Neotropics, 25. Leaflets with erect, spreading indumentum on abaxial surface. 26. Flowers 9-11 mm long; only the upper surface of ovary hairy; leaflets abaxially densely hairy and the epidermis often not visible; Mexico. 26. Flowers

13-23 mm

pubescent; Brazil. 27. Flowers greenish 27. Flowers

leaflets

2. A. jaliscensis. long; upper and lower surfaces and sides of ovary abaxially hairy but the epidermis always visible;

18-23 mm long; fresh fruit very dark brown with pale tree to 30 m, with a small crown in open white mesocarp; situations. 15. A. ormosioides. 13-17 mm

shrub or small

long; fresh fruit green with green mesocarp; tree to 12 m, with a broad spreading crown in open situations.

16. A. fraxinifolia.

indumentum. glabrous or abaxially with appressed 1. A. inermis. 28. Leaflets glabrous abaxially. 28. Leaflets with appressed hairs abaxially.

25. Leaflets

29. Calyx with leaflet base

leaflets 2-4 (-5) indumentum; sparse appressed obtuse, or rounded to slightly cordate; flowers mm long; ovules (1-) 2-4. 17. A. 29. Calyx glabrous except around margins of lobes; leaflets (1-) pairs; leaflet base acute to obtuse; flowers 9-11 mm; ovules

pairs; 10-13 n?tida. 2 (-3) 1-2.

18. A. marauensis.

1. Andira

inermis (W.Wright) DC, Prodr. 2: 475. 1825. Geoffraea inermis W. Wright, Lond. Med. J. 8: 256. 1787. Geoffrea acutifolia Stokes, Bot. mat. med. 4: 46. 1812, nom superfl. Vouacapoua inermis (W.Wright) Lyons, PL nam. 396. 1900. Andira jamaicensis Urban, Symb. antill. 4(2): 298. 1905, nom. superfl.?TYPE: Jamaica. W. Wright s.n. (holotype: BM!).

Tree to 35 m tall with broad spreading crown (in open situations in the Neotropics; trees in the African savannas may have ascending branches and a pyramidal crown; see Polhill, 1969); buttresses slight; bark rough, scaling, grey to dark brown; slash very pale

2003

ANDIRA

43

5-10 mm thick; twigs sparsely hairy when young, the hairs red-brown, erect, glabrescent. Stipules to 17 mm long, to 1mm wide, occasionally persistent; leaf axis 6-34 cm long; rhachis sparsely to very sparsely hairy, hairs red-brown, erect; stipels 1-9 mm long, often caducous; petiolules 2-7 mm long, sparsely to very sparsely hairy, hairs red brown, erect; leaflets 2-9 pairs, 3.5-13.5 cm long, 1.4-6 cm wide, narrowly elliptic, el subcoriaceous to thick-chartaceous, base obtuse liptic, narrowly obovate to oblanceolate, an acumen to 15 mm long, glabrous (occa to rounded with obtuse acute), apex (rarely brown,

sionally very sparsely hairy abaxially, the hairs erect) except the primary vein adaxially and abaxially very sparsely hairy, the hairs red-brown, erect, >0.2-1.0 mm long; primary vein channelled adaxially, raised abaxially, secondary veins 10-14 (-16), plane adaxially and abaxially or slightly raised abaxially, pattern brochidodromous, often with the basal few veins eucamptodromous, tertiary veins plane adaxially and abaxially. Panicles termi cm long, hairy to sparsely hairy at branch tips, becom 10-40 nal, occasionally axillary, the less hairs towards red-brown, ? erect; bracts 2.5-3 mm long, with red base, ing hairy brown appressed hairs; pedicels to 4 mm long or flowers subsessile; bracteoles 1.5 mm long, indumentum like that of bracts. Flowers (9-) 10-12 (-19) mm long. Calyx 4-7 mm long, with appressed, red-brown indumentum or glabrous, except at the base and margins of lobes; lobes 0.3-1 mm long. Petals pale pink to purple; standard blade 7-14 mm high, 9-11 mm wide, claw 1-3 mm long; wing 6-13 mm long, 3^.5 mm wide, claw 2.5-5.5 mm

long, lamellate sculpturing present; keel 6-13 mm long, 3^.5 mm wide, claw 2.5-6 mm long. Stamens 10-18 mm long, filaments united for the basal 5-11 mm, free for the distal 3-7 mm, vexillary stamen 7-13 mm long. Gynoecium 9-20 mm long, usually with sparse, red-brown, ? appressed hairs only on the upper surface of the ovary, but more oc casionally on both the upper and lower surfaces of the ovary, base of the style, and upper surface of the top of the stipe; stipe 3-10 mm long, ovary 3-6 mm long; style 3-5 mm long; ovules 2-4. Fruit 3-6 cm long, 2.5-^.3 cm high, 2-4.3 cm wide, ? globose to ? elon gated, weighing ca. 20 g or less when dry, green, drying dark brown to black, smooth or somewhat rough and warty, occasionally slightly wrinkled; suture slightly raised adaxially and below; stylar remnant insignificant or absent; mesocarp 1-3 mm thick, hard, granu lar; endocarp 2-5 mm thick, brown, woody, fibrous. Chromosome number: n = 10. Andira inermis, occurring in both the Neotropics and in Africa, is a widespread and variable species.The key characters that differentiate the subspecies are the presence or absence of indumentum on the calyx and the size of the flowers. The most widespread subspecies, subsp. inermis, has a calyx with indumentum covering its entire outer surface, whereas subspecies glabricalyx and rooseveltii have calices that are glabrous, except at the base and on the margins of the lobes. Similar intraspecific variation in calyx indu mentum is found in A. humilis and A. multistipula, but in these species the extremes are linked by intermediates, and therefore subspecies are not recognized. Flower size inA. in ermis varies continuously from 9-19 mm, and alone forms no basis for the distinction of intraspecific taxa; however, theMexican specimens with a glabrous calyx (subsp. glabri calyx) have flowers from 10-11 mm long, whereas the African specimens with a glabrous calyx (subsp. rooseveltii) have larger flowers (13-14 mm long). Flower length therefore provides the diagnostic character to separate these two subspecies. There is clear geographic partitioning of variation inAndira inermis, with subsp. roo seveltii restricted to wooded savanna in Africa, and subsp. glabricalyx found in Nayarit and Sinaloa at the edge of the neotropical distribution of A. inermis inMexico. inermis jamaicensis. Subse Wright (1777) first described this species as Geojfroea quently

(Wright 1787), he reported the medicinal

uses of the tree and used the binomial

SYSTEMATICBOTANYMONOGRAPHS

44

VOLUME 64

inermis. Though he provided no botanical description in 1787, the name G. in Geoffraea ermis is validly published, because Wright cited his previous work (1777). Key to the 1. Calyx

inermis

Andira

with

glabrous except 2. Flowers 10-11 mm

2. Flowers

inermis

of Andira

la. indumentum. appressed at the base and margins of the lobes. long; calyx 2.5-3 mm long; Mexico. 13-14 mm long, calyx 4-5 mm long; Africa.

covered

1. Calyx

Subspecies

lb. Andira lc. Andira

inermis inermis

subsp.

inermis.

subsp. glabricalyx. subsp. rooseveltii.

la. Andira

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

Andira riparia Kunth in H. B. K., Nov. gen. sp. 6: 386. 1824.?Type: COLOMBIA. Santander: Prope La Boca del R?o Opon, in ripa fluminis Magdalenae, Bonpland 1599 (holotype: P!). Pterocarpus sapindoides DC, Prodr. 2: 419. 1825. Amerimnum affine Sprengel, Syst. veg. 3: 192. 1826, nom. superfl. Andira sapindoides (DC) Bentham, J. Proc. Linn. Soc, Bot. 4, Suppl. ("A Synopsis of the Dalbergieae"): 123. 1860. Voua Bertero capoua sapindoides (DC) Kuntze, Revis, gen. pi. 1: 212. 1891.?Type: s.n. (holotype: G-DC!; isotype: TO). Andira grandiflora Guillemin, Perrottet & A. Richard, FI. Seneg. tent. 254. 1832. Andira inermis subsp. grandiflora (Guillemin, Perrottet & A. Richard) Gillett ex Gambia. Albreda, Perrottet 31 (holo Polhill, Kew Bull. 23: 490. 1969.?Type: not located; isotype: BM!). type: P, Glycyrhiza undulata Ruiz & Pav?n ex G. Don, Gen. syst. 2: 227. 1832.?Type: Peru. Ruiz & Pav?n s.n. (holotype: BM!). acuminata Bentham, Comm. legum. gen. 45. 1837.?Type: "In Brazil. sylvis ad flumen Solimaen provinciae Rio Negro," Martius 1157 (holotype: K!; isotype: M). Andira inermis var. riedelii Bentham, J. Proc. Linn. Soc. Bot. 4, Suppl. ("A Synopsis of the Dalbergieae"): 123. I860.?Type: Brazil. Mato Grosso: Rio Coxim, Riedel 402 (lectotype, here designated: K!). Andira

Lonchocarpusl Cameroon.

staudtii Harms Kumba

in Engler, Bot. Jahrb. Syst. 26: 301. 1899.?Type: Staudt 912 (holotype: B, de [=Johann-Albrechtsh?he],

stroyed; isotype: BM!). Andira chiricana Pittier, Contr. U.S. Nati. Herb. Chiriqu?: David, NY!). Flowers Fig. 2A.

(9-)

10-12

30 Mar

(-19) mm

1911, H. Pittier

long. Calyx

18(6): 235. 1917.?TYPE: PANAMA. 3372 (holotype: US!; isotypes: GH!

covered with

appressed

indumentum.

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

ANDIRA

2003

110

105

100

95

90

85

80

75

70

45

65

60

55

50

45

40

35

Rico, Virgin Islands, Antigua, Guadeloupe, Dominica, Martinique, St. Vincent, St. Lucia, Grenada, Trinidad, Tobago); Mexico and throughout Central America; Venezuela, Colom bia, Ecuador, Peru, Bolivia, the Guianas, Paraguay, and northern Argentina (Corrientes); Africa (Gambia, Senegal, Ivory Coast, Nigeria, Cameroon, Equatorial Guinea). In the dif ferent parts of its range, A. inermis subsp. inermis grows in various habitats. In Africa, it occurs in wooded savanna and rain forest. In Central America and the Caribbean coast of it is common in seasonally dry tropical forests, whereas else northern South America it grows in rain- and gallery forest, including inundated areas and where in the Neotropics even

mangrove.

Black plum (Tobago); l'angelin, angelim, bois palmis (Mar palo maco (Dominican Republic); cabbage bark, bastard cabbage tinique, Guadeloupe); cuilimbuca, maquilla cuautololote, (Mexico); cab (Jamaica); tololote, quiringuacua, almendro cirvelillo, (El Salvador, (Belize); frijolillo bage bark, barley wood, Guatemala, Honduras, Nicaragua); cabbage bark, quira, harino, cocu (Panama); congo, Vernacular

names.

(Colombia); majagua gallina, palmillo, ciruelo de playa, bolombolo, mot?n (Venezuela); barbasco caspi, montera caspi, numi, pil?n, lumbricero, peloto, mot?n pisho, sacha ushuu (Peru); koraro, bat seed (Guyana); reddie kabbes, vremoessoe noto rouge (French Guiana); sapupira (Suriname); lebi kiabici, St. Martin lombrigueira, (Brazil); mendubira (Argentina). dividivi,

46

SYSTEMATICBOTANYMONOGRAPHS

FIG.

15. Distribution

Representative Massif

Nord-Ouest:

of Andira

Specimens.

inermis

Jamaica.

subsp.

inermis and A.

Manchester:

du Nord,

Petit Ford, Ekman Port-de-Paix, Domingo: vie. Ciudad Trajillo, Allard 13367 (US).?SAMAN?: Las Terrenas, Zanoni & Mejia Loma 17748 (NY).?DAJAB?N:

inermis

VOLUME 64

subsp. rooseveltii

in Africa.

Donnell Smith 2389 (US). Haiti. Mandeville, 4326 (NY, US). Santo Dominican Republic. Finca Hacienda Nydia, 3.5 km E of village of

de Cabrera, banks of R?o de Masacre, Zanoni & on road to Cotui, Zanoni et al. 16093 RAM?REZ: 7 km from center of Pimentel (NY).?S?NCHEZ Puerto Rico. Ca. 2 km N Hwy 3, just E town of Rio Grande, Boom 6783 (GH, NY). Islands. (NY). Virgin St. Croix: Belvedere Box 1385 (A, US). estate, D'Arcy 4727 (GH, MO). Antigua. Wallings, Guadeloupe. Pointe Noire, Duss 3231 (NY, US). Dominica. Sastre Batalle, Ramage 221 (K). Martinique. Basse-Pointe, 17899

Mej?a

7731 (GH). St. Lucia. Anse Louvet, Slane 803 (A). St. Vincent. Kingstown, Grenada. Trinidad. Balthazar, banks of Great River, Beard 157 (GH). US). s.n. (F, NY). ton 2906 (GH, NY, US). Tobago. Scarborough, Broadway carretera Esc?rcega-Villahermosa, Mexico. CAMPECHE: Km 30 desviaci?n

Beard

1336

Pitch Lake,

(F, GH, MO, NY, vie. Brighton, Brit

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

4707

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

BELIZE: Belize-Sibun

hwy, Croat 24861 mingbird ten River, 2 km N of Victoria with

Road,

Gentle

(MO, US).?Stann

15 (F, NY, US).?Cayo: along Creek: Cockscomb Mountains,

Sibun River, just W of Hum trib. of Cocoa Branch of Sit

Peak, Gentry 7887 (MO, US).?TOLEDO: hwy to Punta Gorda, 1mi E of junction Croat 24471 (MO, US). Alta Verapaz: Rio Sebol, downstream Guatemala. from 44779 Smith 2823 (GH, US).?IZABAL: Escuintla, Donnell Maricos, (US).?ESCUINTLA:

road to San Antonio,

Carrizal,

Steyermark near Jutiapa, Standley 60583 (F).?PETEN: Laguna 7600 (MO).?JUTIARA: bordering Lake Izabal, E. Contreras S of Sayaxch?, Steyermark 46202 Petexbat?m Marcos: Palmatto Flats, 1-2 (F, US).?San (Laguna M?xico) mi N of Oc?s, Steyermark 37865 (F).?SANTA ROSA: Chiquilmulilla, Standley 79182 (F).?SUCHITEP?QUEZ: vie. Tiquisate, and Santa Cruz, Standley 47462 between R?o Honda 74130 (F).?Zacapa: (F). Steyermark en la vega de Quebrada Tolobre, ATLANT?DA: vic. Tela, Standley 53570 (GH, US).?Choloteca: entre Morolica et y Tolobre, Standley 14219 (NY).?COL?N: Trujillo, R?o Negro, Clewell al. 4339 (MO).?COMAYAGUA: Pito Solo, Lake Yojoa, Edwards 1 km N 429P (GH).?CORTES: Matorrales, DE LA Bah?a: West End, Isla de Roat?n, Nelson & Romero 4519 Villa Nueva, Molina R. 6790 (GH, US).?ISLAS

Honduras.

matorrales

y bre?ales

(MEXU).?Olancho:

Mpio.

San Esteban,

Santa Mar?a

del Carb?n,

30 km al NE

de San Esteban,

Sousa

et al.

ANDIRA

2003

47

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

Standley 21277 (GH, US).?SONSONATE: 11189 Chinandega: vic. Chichigalpa, Standley 452 (FHO, MEXU).?MANAGUA: course, Hughes

Vicente,

retera a Montelimar,

& Castro

Guzman

138

San Brenes, (F). Costa Rica. ALAJUELA: Parque Alberto Manuel at R?o Abangartos, Croat 61182 (MO).? Carvajal 263 (F,MO).?GUANACASTE: hwy Esparza-Ca?as Rio Colorado 10047 (MO).?LLM?N: HEREDIA: Finca La Selva, OTS field station on Rio Puerto Viejo, Folsom Cant?n de Osa, between Islas Buena Vista and Cerro Coronel, Davidse & Herrera 31242 (MO).?PUNTARENAS: El Recreo,

(US).?Zelaya:

229

Long

Ram?n,

R?o Terraba, Alien 5220 (F,MO, US).?San Jos?: 10 km NW Gu?piles, L. D. vic. El Panama. CfflRlQU?: Progreso, Cooper & Slater 265 (F, GH, NY, US).?Cocl?: (WO). Alto de Las Rio Ucurganti, Bristan 1127 (NY, US).?HERRERA: Valle, Allen 1770 (GH, NY, US).?DARBEN: 256 (F).?Panam?: R?o San Juan D?az, above Juan D?az, Alien 938 (GH).?SAN BLAS: Minas, Carrasquilla trail Palmar Norte-Ca?ablancal, 18517

G?mez

mainland

opposite

Play?n

Chico,

0-3 mi

from Caribbean,

Gentry

6373

(F).?VERAGUAS:

vic. Santa F?, Allen

4435 (GH). DELTA Amacuro: Serra Imataca, Cerro La Paloma, E side R?o Cuyubini, Venezuela. S?eyermark 87604 FEDERAL: E of Caraballeda, Hacienda Juan D?az, Steyermark 62930 (F,MO).?FALC?N: Dist. (U).?DISTRITO 113652 (MO).?Miranda: R?o Chico, Jahn 1239 Colina, R?o Ricoa, Dos Bocas, Steyermark & A. Gonzales between La Pica and Ca?o Colorado, E Marur?n 6 km W La Ormega, Wurdack & (GH, NY, US).?Monagas: Monachino

Davidse

Cerro Las Minas, S main road from Santa Ana, 17 km SE Santa Ana, Stey (NY).?T?CHIRA: of R?o Catatumbo and La Fria-Maracaibo Distrito Col?n, intersection road, re (MO). Guyana. West Bank Demerara River, Boom 7198 (NY, US); Pomeroon-Supernaam

39495 et al.

ermark

(MO).?Zulla:

18831

Between mouths & Persaud 1217 (MEXU, US). Suriname. Gillespie gion, Adventure, and Coronie, Lanjouw & Lindeman 1460 (MO, U). French Guiana. Vicinity Cayenne, Amazonas: P. N. Amacayac?, Colombia. Cabana Pamat?, Narv?ez & Olmos 69 US). 2 km from Barraquillita, lands of 95 (COL, JAUM, MO).?BOL?VAR: Brand & Cogollo Curran

Loba,

23

(US).?Caldas:

R?o Magdalena,

cerca

a La Dorada,

of rivers Coppename 611 (GH, Broadway (COL).?Antioquia: Loba, San Mart?n

de

del R?o Purnio, Idrobo & R. la Yuca, M. S?nchez & G. Mahecha

confl.

2269 (COL, F, MO, NY, US).?CAQUET?: Florencia, Quebrada 9a (UDBC).?CAUCA: camino a Pablo Sexto y Cabo de Hornos, Lozano et Mpio. Guap?, P. N. de Isla Gorgona, al. 5245 (COL).?CHOC?: Archer 2050 (NY, Llor?, 50 km S Quibd?, at june. R?o Atrato and R?o Andagueda, Planeta Rica, S. L?pez 4005 (COL).?Guajira: Sierra Nevada de Santa Marta, Sierra de San US).?C?rdoba: Jaramillo

de La Cueva, Cuadros & Gentry 2965 (MO, NY).?MAGDALENA: valle inferior del Mag Bosque 841 (F, US).?Meta: R?o Umea, Gavia 5122 (US).?Narl?O: dalena, Dugand Tamaco, R?o Rosario, 6 kms ar near Bonda, H. H. Smith 18 del Caunap?, Romero 5218 (MO, NY).?Santa riba de la desembocadura Marta: Puerto Araujo, Renter?a e? al. 1887 (JAUM, MO).?Valle: Isla del (F, GH, MO, NY, US).?Santander:

Antonio,

en el desembocadura del R?o Cajambre, Cua?recasas 16224 (F).?Sucre: around Verrugas, Finca La Guayabal Esmeraldas: Ecuador. San Lorenzo, 0.5 km S on walk to R?o Nadadero, Little Aguada, Bernai 158 (COL). Jr. 6341 (F, US).??apo: La Joya de los Sachos Cant?n, Pompeya, R?o Indillana, between its mouth at the R?o et al. 2125 (QCNE). and the MAXUS Peru. AMAZONAS: Quebrada Huampami, road, Gudi?o ?apo Kayap 1061

(F,MO).?Cuzco: Pilcopta, Atalaya, Paucartambo, N??ez de Iparia a lo largo de R?o Pachitea, Bosque Nacional Croat GH, NY, US).?LORETO: (F, Quebrada Cuninico,

Honoria, 1958

6851 (MO).?Hu?NUCO: Prov. Pachitea, Dtto. 1 km arriba del pueblo de Tournavista, Schunke DE DIOS: Prov. Manu, 17759 (MO).?MADRE

10-15 km NNW Shintuya, Foster et al. 11027 (F).?PASCO: Prov. Oxapampa, between Puerto Bermudez and Paujul, Gentry et al. 42145 Bermudez, (F, Mart?n: 1-2 km E Tabalosos, Belshaw 3370 (F, GH, MO, NY, US).? Prov. Lamas, Distr. Lamas, MO).?SAN Prov. Coronel Portal, Yarinacocha, Ucayali: Bolivia. BENI: Mamore, Gentry & Horna 29375 (MO). Tyson & Cerro de Pantiacolla, R?o Palotoa, 1 hour below Puerto R?o Pichis,

Kuns

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

Santa

de Caldas

da Pousada

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

Maranh?o:

Novas,

Parque Maracassum?

River

SYSTEMATICBOTANYMONOGRAPHS

48

VOLUME 64

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

Victoria,

(1860) recognized A. sapindoides based on two specimens from Antigua These and other collections from the Lesser Antilles (Antigua, Dominica, and St. Lucia) have much larger flowers, up to 19 Guadeloupe, Martinique, Monserrat, mm long, than specimens from other parts of the range. The existence of specimens with flowers of intermediate size, such as H. E. Box 1385 (Antigua), appears to confirm Ben Bentham

and Dominica.

tham's (1860) doubts that A. sapindoides may only represent a well-marked variant of A. inermis, and the name is therefore placed in synonymy. The type of A. chiricana {Pittier 3372; Panama: Chiriqui) has hairy ovaries and leaflets that are sparsely hairy abaxially, but it otherwise matches A. inermis subsp. iner mis. Ovary and leaflet indumentum appear sporadically in populations throughout the range of subsp. inermis. Polhill (1969) provided an excellent summary of the variation displayed by A. iner mis in Africa. He recognized two subspecies, which are geographically separate and have a calyx bearing indumentum: subsp. inermis and subsp. grandiflora. If the African mate rial is considered in isolation, then these morphological differences appear to merit sub inermis in rain forest around the Gulf of Guinea in grows specific recognition. Subspecies Cameroon and Nigeria, and has flowers of 12-13 mm long; subspecies grandiflora, from savannas inGambia, Senegal, Mali, Central African Republic, and Ivory Coast, has larger (to 15 mm long). These two variants agree in all other respects with Neotropical populations of subsp. inermis. Because the range of flower size in the Neotropics encom passes the range of variation in Africa, subsp. grandiflora cannot be considered distinct, and the name is therefore placed in synonymy. flowers

Andira inermis subsp. inermis has hard, heavy timber, which is used in the Neotrop ics for furniture and cabinetwork, and also for construction, railway sleepers, fence posts, etc. Because the wood has virtually no resonance, it is particularly suitable for radio and television cabinets (Weaver 1989). It is also used as an ornamental street tree inVenezuela and Costa Rica. The bark may be used as a vermifuge but is poisonous in high doses It is also taken for intermittent in fevers Mexico and Brazil (Morton 1981). (Weaver 1989). lb. Andira

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

Ab A.

inermi subsp. inermi calycibus glabris differt. Ab A. inermi subsp. rooseveltii (10-11 mm longis, non 13-14 mm) differt. Flowers 10-11 mm long. Calyx 2.5-3 mm long, glabrous except at base and margins of lobes.

floribus minoribus

2003

ANDIRA

49

Phenology. Flowering May and June, fruiting August fruiting records in December and March.

and September,

with

single

Distribution (Fig. 14). Mexico (Nayarit and Sinaloa); deciduous and semi-deciduous tropical forest. No flowering specimens have been collected in Sinaloa, but the remains of are glabrous. the calyx on the fruiting specimens Rose 1639,1782 Additional de

Specimens

las Ventanas,

Examined.

Ben??ez-Paredes

Mexico.

3658

Nayartt:

(MEXU); Arroyo between Zopilote

La Nanchflera and Arroyo ca. 10 km E la P. H. Aguamilpa, near junction for San and Santa Cruz de Guaybel, NE km San Pedro 19 Ixcat?n, 2 km NE El Nayar, Mpio.

Brasil,

Nayar, Embalse

between

et al. 18688 (MEXU); et al 5474 (MEXU); Mpio. del Miguel Zapote, Ch?zaro terracer?a San Pedro Ixcat?n-Santa Cruz Guaybal, G. Flores e? al. 1841 (MEXU); along dirt road be Naranjo, tween Pochitit?n and Francisco Madero, Miller & T?llez 3181 (MEXU, MO); Mpio. Tepic, 4.3 km E junction to la Escondida, R. Ramirez & G. Flores 674 (MEXU); Mpio. old road Tepic-Mazatl?n, Tepic, 5-7 km W. 35 km W Tepic, T?llez 9076 (MEXU); 4 km NE Pochotit?n, road to Mojarritas, Pochotit?n, T?llez & Miller cortina, Calzada

10478

(MEXU); Mpio. Tepic, Paso de Bueyes, 40 km SE Tepic, Tenorio e? al. 16869 tween Rosario and Colomas, Rose 1639 (US); near Colomas, Rose 1782 (US).

It is tempting to speculate that morphological populations at the northern edge of the Neotropical mis, producing both A. inermis subsp. glabricalyx Jalisco and Michoac?n, separating the ranges of A. accurate information on More subsp. glabricalyx.

(MEXU).?Slnaloa:

be

divergence has occurred in isolated distribution of A. inermis subsp. iner and A. jaliscensis. The latter occurs in inermis subsp. inermis, and A. inermis the phylogenetic relationships of these

taxa is necessary to corroborate this hypothesis, especially for A. jaliscensis, whose affini ties are unresolved (Fig. 8). A similar geographic pattern is seen in Lennea viridiflora Seem. (Leguminosae, Papilionoideae, tribe Robinieae), where L. viridiflora var. novo galiciensis Lavin & M. Sousa is restricted to Colima, Jalisco, and adjacent Michoac?n, whereas L. viridiflora var. viridiflora is widespread throughout Central America (Lavin & Sousa 1995). lc. Andira

(de Wildeman) Gillett ex Polhill, Kew Bull. 23: subsp. rooseveltii Millettia rooseveltii de Wildeman, PL Bequaert. 3: 353.1925.?Type: Mearns 3004 (holotype: BR; Equatoria Province: Nimule-Gondokoro,

inermis

490.1969. Sudan.

isotype: BM!). brownii Hoyle, Kew Bull. 1932: 262. 1932.?Type: Region: Jema, Feb 1931, W. T. Brown 2163 (holotype: K!).

Ostryoderris

13-14 mm

Flowers

long. Calyx 4-5 mm

long, glabrous

Ghana.

Ashanti

except at base and margins

of lobes. Phenology.

Flowering

April-June. Distribution in wooded

February

(Fig. 15). Nigeria,

and March,

with

one

record

Chad, Central Africa Republic,

in May,

Togo, Uganda,

fruiting Sudan;

savanna.

names. Gwaska

Vernacular Additional

(Nigeria); Weri-dee

(Central African Republic).

Specimens

Examined. Togo. Without locality, Kersting 554 (K). Nigeria. Gangoro Forest 168 (K); Bauchi, Lely P192 (K); Sokoto, Uly 832 (K); northern Nigeria, Yola, Dalziel 54 (K). Chad. N of Nd?ll?, between (K); Abakaliki, Opara 609 (K); Plateau Prov., Selfwe River, Thornewill Golo and Mansaca, Chevalier 7773 (K). Sudan. Yirol District, around Koudogoi, F. W. Andrews 464 (K); Reserve,

Chapman

Imatong Mts, Kinyeti cality, Schweinfurth

4203

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

VOLUME 64

SYSTEMATICBOTANYMONOGRAPHS

50

2 km S Camp Koumbala along Koumbala river, Fay 4321 (K); Bouar, Mildbraed 7252 (K). Leya and Aiyu, river junction, Greenway & Eggeling

9422

(BM, K).

Uganda.

W

Madi,

Andira inermis subsp. rooseveltii ismore or less parapatric with the African savanna form of A. inermis subsp. inermis (Fig. 15) in the wide belt of wooded savanna running across central Africa from Gambia to Sudan and Uganda. Their ranges abut between Ivory Coast (the most eastern record of the savanna form of A. inermis subsp. inermis) and Ghana (the most western record of A. inermis subsp. roosveltii). Hopkins (1983) demon strated that variation of leaf characters in Parkia biglobosa is clinal in these woodlands, and thus did not provide a basis for recognition of intraspecific taxa. From the paucity of herbarium specimens A. inermis appears a rare species throughout these African savan nas, and Boulvert (1977) reported it as very rare in the Central African Republic. It is pos sible that further collections from the savanna zone between the ranges of subsp. inermis and subsp. rooseveltii (in southern Mali and Burkina Faso) might reveal intermediates, and thus there may be a clinal situation similar to that found in Parkia biglobosa. Andira inermis subsp. rooseveltii is reported to be used for children's gera, or sever ance illness, by washing 2. Andira

babies with a decoction

(herb, label, Central African Republic).

Jalisco: Mpio. Talpa de jaliscensis R. T. Pennington, sp. nov.?Type: Mexico. camino a la mina del Cuale, 7.6 km E de la carretera Puerto Val Allende, iso larta-Barra de Navidad, 30 May 1985, E. J. Lott 2544 (holotype: MEXU!; type: MO!).

A.

inermi primo aspectu maxime similis sed foliolis subter dense pubescentibus. Tree to 25 m tall; presence of buttresses unknown; bark and slash unkown; twigs brown to dark brown with numerous pale elongated lenticels, sparsely hairy, the hairs pale, erect, glabrescent. Stipules to 10 (-20) mm long, to 1mm wide, caducous, densely erect; leaf axis 13-25 cm long; rhachis densely hairy, hairs white, erect, tangled, glabrescent, especially towards the base; stipels minute (0.5 mm long), caducous; petiolules 2-4 mm long, indumentum like that of rhachis; leaflets in 3-5 (-6) pairs, 3-10 cm long, (1.5-) 2.2-3.5 to {-A) cm wide, narrowly obovate or elliptic, thick-chartaceous hairy, hairs white,

subcoriaceous, base obtuse to rounded, apex obtuse to rounded, often with a short acumen to 4 mm long, glabrous adaxially, densely hairy abaxially and the epidermis often not vis ible, hairs >0.2-1.0 mm long, erect, tangled, pale (the leaflets appear white abaxially); primary vein channelled adaxially, raised abaxially; secondary veins 10-12, ? plane adax ially, slightly raised abaxially, pattern brochidodromous with the first 1-2 eucamptodro mous, tertiary veins plane adaxially and abaxially. Panicles terminal, 20-35 cm long, densely hairy at branch tips, becoming less hairy towards the base, the hairs red-brown becoming white with age (or drying?), erect, tangled; bracts 2 mm long with pale brown caducous hairs; bracteoles 1mm long with pale brown caducous hairs. Flowers 9-11 mm long, subsessile. Calyx 5 mm long, hairy, the hairs tangled, pale brown; lobes 0.5-0.75 mm

long, acute to obtuse. Petals pink to purple; standard blade 10mm wide, 7 mm high, claw 2.5 mm long; wing 7.5-8 mm long, 3.5 mm wide, claw 3 mm long, lamellate sculp turing present but poorly developed; keel 6 mm long, 4 mm wide, claw 4 mm long. Sta mens 10 mm long, the filaments united for the basal 4.5-5 mm, free for the distal 3.5-5 mm. Gynoecium 10-10.5 mm long, the upper surface of the ovary hairy, hairs red-brown, ? erect; stipe 3.5-4 mm long, ovary 3 mm long, style 3.5 mm long; ovules 2-3. Fruit 3.2-3.5 cm long, 2.8-2.9 cm high, 2.7-3 cm wide, ? globose, weighing ca. 20 g or less

2003

ANDIRA

51

when dry, green, drying brown, appearing smooth but minutely wrinkled (best seen with suture slightly raised adaxially with a narrow groove, indistinct abax lens or microscope); remnant tiny; composition of mesocarp and endocarp unknown. Chromosome ially; stylar number unknown. Figs. 3E, 4(iv), 5A, 16. Phenology. Flowering May to June with occasional records in January and March. in deciduous or semi-deciduous Distribution (Jalisco, Michoac?n); (Fig. 17).Mexico tropical forest, often by seasonal water courses. Vernacular names. Manzanita (Jalisco); garrapata (Michoac?n). Examined. Mexico. Jalisco: Additional Specimens Arroyo Chamela, Bullock 910 (MEXU); Mpio. La Chamela, Huerta, Est. de Biolog?a Chamela, Arroyo Chamela, Bullock 1313 (MEXU, MO); Estaci?n Biol?gica A. Delgado et al. 480 (MEXU); Mpio. La Huerta, 7 km al N del Puente Chamela, camino a Nacastillo, pasando et al. 4795 (MEXU); Mpio. La Huerta, Ma Arroyo Colorado, por los terrenos de la U. de G., G. Flores-Franco en el Lindero de la Universidad de Guadala (MEXU); Mpio. La Huerta, camino al E de Nacastillo, W 1008 km de camino 585 3 Chamela, Macuautitl?n, (MEXU); (MEXU); Magallanes jara, Magallanes Arroyo 1044 (NY); Mpio. La Huerta, Chamela, camino a Nacastillo, Magal de Allende, Magallanes Tomatl?n-Talpa 14 km N Santiago, Magallanes 1669 lanes 1383 (MEXU); Mpio. La Pe?a, camino Santiago Chacala-Colima, gallanes

421

de Navidad, Magallanes 1710 (MEXU); Tomatl?n, Las Jarillas, carretera Puerto Vallarta-Barra (F); Mpio. La Huerta, Est. de Biolog?a Chamela, Magallanes 2273 (US); Mpio. El Tuito, Las Gu?cimas, camino El a La 4197 3556 camino Tuito-Chacala, Huerta, Nacastillo, (MEXU); (MEXU); Magallanes Magallanes Mpio. 4404 (NY); Chamela, L. A. P?rez 182 (MEXU); Chamela, L. A. Mpio. La Huerta, Arroyo Chamela, Magallanes

Mpio.

P?rez & M. Hern?ndez

Vallar?a in direction (MEXU, NY); Mpio. El Tuito, 14 km N road El Tuito-Puerto 2148 (MEXU); Mpio. Cuautitl?n, 3 km N Teqesquitl?n, 2950 Solis-Magallanes 4 km de Aquila, Guerrero 766 (MEXU); Mpio. Aquila, (MEXU).?MICHOAC?N: Mpio. Aquila, El Tenamaste, Los Tanamastes, Guerrero et al. 1262 (MEXU); Mpio. Chinicuila, 10 km NW Villa Victoria, Soto N??ez et al. of El Cuale,

8187

848

Solis-Magallanes

(MEXU); Mpio.

Tiquicheo,

Paraje

la Escondida

el Lim?n,

Verduzco

s.n. (MEXU).

is clearly distinct from A. inermis in its abaxially densely hairy Andira jaliscensis leaflets. In specimens from Michoac?n this indumentum is less dense than the material from Jalisco, but they are separated from specimens of A. inermis from this area, which have entirely glabrous leaflets, even as very young seedlings (e.g., Soto N??ez ?tal. 8016). come from the southwestern part of All the collections of A. jaliscensis from Michoac?n this state, near Jalisco, except one, A. A. Verduzco s.n., which was collected near the bor der with Guerrero and M?xico. This raises the possibility that A. jaliscensis may be dis tributed across Michoac?n. Andira inermis and A. jaliscensis certainly appear to be sym and future fieldwork should concentrate here to see whether any patric in Michoac?n, intermediate 3. Andira

forms exist.

Brazil. Ducke, Bol. T?cn. Inst. Agron. N. 2: 30. 1944.?Type: multistipula Amazonas: S?o Paulo de Oliven?a, 2 Nov 1942, A. Ducke 1035 (lectotype, here designated: RB!; isotypes: K! R! RB! US!).

Tree to 20 m

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

52

FIG.

16. Andira

surface. E. Standard cium. I. Gynoecium, S. Magallanes 4404.)

SYSTEMATICBOTANYMONOGRAPHS

VOLUME 64

A. Habit B. Abaxial leaflet surface. C. Flower. D. Calyx, opened to show inner jaliscensis. petal, inner surface. F. Wing petal, outer surface. G. Keel petal, inner surface. H. Androe also shown above in longitudinal section. J. Fruit. (Based on: A-I, E. J. Lott 2544; J, J. A.

ANDIRA

2003

110

100

105

FIG.

narrowly

ovate

or

95

90

85

17. Distribution

lanceolate),

80

of Andira

chartaceous

75

70

jaliscensis,

53

65

60

A. multistipula,

to thick-chartaceous,

55

50

andA.

base

45

40

35

cubensis.

obtuse,

rarely

acute,

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

SYSTEMATICBOTANYMONOGRAPHS

54

VOLUME 64

cm long, 2-2.5 cm high, elongated, weighing ca. 20 g or less when dry, green, dry dark brown to black, slightly wrinkled; stylar remnant barely discernible; meso very ing mm thick, greyish brown, hard, granular; endocarp 1-1.5 mm thick, brown, 1.5-2 carp

2.5-3.5

Chromosome

fibrous.

woody,

number

unknown.

the year (March, April, June, October to Phenology. Flowering December). Distribution (Amazonas), Ecuador (Fig. 17). Brazil (Amazonas, Acre), Colombia (Napo, Sucumbios), Peru (Amazonas, Loreto, San Mart?n, Hu?nuco), Bolivia (Pando); in seasonally flooded forest and on river banks, with occasional records from secondary records throughout

forest.

Vernacular

names. Pisho, barbasco

caspi, arco caspi, arco sacha (Peru).

Amazonas: Bocas del R?o Yari en el R?o Caquet?, Pab?n La de los Sachos Cant?n, Pompeya, R?o Indillama, between its (COL). Joya et al. 2145 (QCNE); Parque Nacional Yasun?, MAXUS mouth at the Rio Napo and the MAXUS road, Guai?o road Km 24, by bridge over R?o Payamino, R. T. Pennington & M. Aulestia 537 (E, K, QCA, QCNE); Parque Nacional Yasun?, A?angu, SEF 8533 (QCA).?SUCUMBIOS: Reserva Faunistica Cuyabeno, R?o Cuyabeno above Additional

& Mahecha

Specimens

482

Examined.

Ecuador.

Colombia.

?apo:

Peru. Amazonas: Prov. Bagua, Dist. Imaza, R?o Cenepa Laguna Grande, Balslev e? al. 97505 (QCA, QCNE). & Gorham 1 km debajo de La Poza, bande del R?o San 162 (E, NY); region, Comunidad Yamayakat, Hodges 49 (MO); R?o Santiago, 2 km atr?s de la Comunidad 925 (MO); Prov. tiago, Huashika? Caterpiza, Huashikai de Yamayakat, R?o Mara??n, Comunidad Jaramillo 575 (E, MO).? Bagua, Dist. Imaza, Regi?n del Mara??n, Prov. Maynas, Jacintillo, near Tingo Mar?a, Schunke 5725 (F,MO, US).?LORETO: et al. 3102 (MO); Quebrada Cuninico, Croat 17753 (MO); Quebrada Yanayac?, GH, MEXU, MO, US); Prov. Coronel Portillo, Carretera Federico Basadre Km 99, Arboretum HU?NUCO:

Itaya, Ayala

C. D?az

et al. 670

C. D?az

1315

Santa Rosa, R?o Croat

20320

(F,

Von Humboldt,

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

a Shunte,

Schunke 3859 (COL, GH, NY). PANDO: Prov. Manuripi, Bolivia. along from Chiba, Nee 31530 (MO). Brazil. Acre: Mpio. Sena (by air) downstream Km 1-3 on road Sena Madureira to Rio Branco, Prance e? al. 7968 (COL, GH).?AMAZONAS: Rio do RADAM, Cavalcante 3164 (RB, ?B); Mpio. Nova Japura, between Tamandar? and Manquari,

Nuevo,

de Dios,

80 km NE

R?o Madre Madureira, Javar?, Pt. 2 Rio

Japura, do Juami-Japura, Cid Ferreira et al. 7250 (K, NY); Mpio. Limoeiro, Est. Ecol?gica (F, K, NY, US); Alto Solim?es, Rio Bora, afluente do Rio Jutai, L. Co?lho et al 394 (INPA); Rio Javar?, Esper an?a, Ducke 1822 (F, GH, NY, US); ?cima do lago Jacapar?, left bank Rio Solim?es, H. C. de Lima & J. Guedes

Cid Ferreira

2731

& Lima 3611

(GH, K, NY); Alto

(GH, K, NY);

Rio

Solim?es,

Juru?, Pena

608

Mpio.

S?o Paulo

de Oliven?a,

S?o Luis do Pass?, H. C. de Lima

et al. 2767

(NY).

The flowers and fruit of A. multistipula are very similar to those of A. inermis; how ever, it is clearly distinct because of its large, persistent stipules. Ducke (1944) claimed that these stipules distinguish A. multistipula from any other species of the entire Ama zonian

flora.

All

the specimens from Peru have a calyx with appressed indumentum, in contrast to the majority of the Brazilian material in which the calyx is glabrous, except at the base and on the margins of the lobes. The calyx is sparsely hairy in two specimens from Brazil (Prance et al. 7968, Acre; Cid Ferreira et al. 7250, Amazonas). The lack of other char acters differentiating these two groups and the presence into two taxa. splitting A. multistipula Two

specimens specimen with more

of intermediates

of the type collection (Ducke 1035) were stipules was chosen as the lectotype.

argue against

found at RB. The

larger

'

55

ANDIRA

2003

4. Andira

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

Grisebach, PL wright. 179. 1860.?TYPE: Cuba. Oriente: "In praeruptis prope Monte Verde," Jan-Jul 1865, C. Wright 1187 (lectotype, here designated: GH!; isolectotypes: G-3 sheets! GH! K! MO! NY-3 sheets! US!).

Andira microcarpa

Tree to 20 m tall, crown often described as spreading; presence of buttresses unknown; bark grey; slash unknown; twigs dark brown with pale raised lenticels, sparsely hairy, mm long, to 1mm wide, very glabrescent, hairs brown, ? appressed to ? erect. Stipules to 3 seen at shoot apices); leaf axis 6-22 (-32 cm long on sterile branches); early caducous (only rhachis sparsely hairy, glabrescent, the hairs ? erect, brown, becoming or drying pale; stipels minute, very early caducous; petiolules 1.5-5 mm hairs ? erect, brown, becoming or drying pale; leaflets 2.3-5.5 cm wide, narrowly ovate, elliptic (rarely obovate base obtuse, rounded to ? cordate, apex obtuse and often often retuse (apex of the terminal leaflet tends to be more

long, sparsely hairy, glabrescent, in 3-4 (-5) pairs, 5-13 cm long, or broadly obovate), coriaceous, slightly acuminate or rounded and rounded), occasionally acute and

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

mm

the distal 2-3.5 mm. Gynoecium and lower half of the style hairy 1.5-2.5 mm long, ovary 2-3 mm 2-2.8 cm high, 2.1-2.9 cm wide,

5.5-8.5 mm

long, upper and lower surfaces of the ovary sides (ovary glabrous), hairs brown, ? appressed; stipe long, style 1.5-3 mm long; ovules 1. Fruit 2.5^4 cm long, elongated, the apex somewhat pointed, weighing ca. 20 g

or less when dry, drying dark brown spotted with pale brown (probably green when fresh though reported on herbarium labels as red and as brown), smooth; stylar remnant insignif icant; mesocarp 2-4 mm thick, pale brown, quite soft, granular; endocarp 1-2 mm thick, brown, woody, hard, not very fibrous (note: these notes of fruit structure are based upon a = 11. single, possibly immature fruit). Chromosome number: n Phenology. Flowering April to August, concentrated in June, July. Distribution (Fig. 17). Cuba; forest, swamp, river bank, wood pasture, ca. to 1000 m. pine savanna; Vernacular

name.

savanna and

Yaba.

Examined. Additional Specimens Cuba. Habana: Isla de Pinos, Los Indios, N. L. Britton et al. 14246 Isla de Pinos, Morton 9983 (US); (NY); near Puentes Grandes, Bro. Le?n 3347 (NY); 2 km N Nueva Gerona, Cerro Nueva Gerona, Isla de Pinos, Morton 10304 10121,10155 (US); Isla de Pinos, Herradura Beach, Morton

SYSTEMATICBOTANYMONOGRAPHS

56

VOLUME 64

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

Cienfuegos, Matanzas:

hills

(GH); road to San Vicente,

21 km N Pinar del Rio, Harvard

Course

in Trop. Botany

10 (GH, NY).

(1860) considered A. cubensis to be a doubtful species, but, upon close in spection, many characters separate it from the similar A. inermis, which appears to be ab sent from Cuba. Andira cubensis has smaller flowers, amore densely hairy gynoecium, a single ovule, and a fruit with amore pointed apex. It also tends to have only 3^4- (-5) pairs of leaflets compared to 2-9 inA. inermis, which have a rounded to cordate base, whereas Bentham

the leaflets are abaxially hairy when young. those in A. inermis are obtuse. Additionally, This Cuban endemic could not be included in the molecular phylogenetic analysis, because no recently collected leaf material was available. It seems most likely that its closest relative is A. inermis, because of their morphological similarities, and therefore A. cubensis is placed with the species of the basally divergent clade I (Fig. 8). It is intrigu ing that A. inermis, which is present throughout the Caribbean islands, appears to be ab sent from Cuba. R. T. Pennington, Colombia. sp. nov.?Type: Antioquia: 1 G. Ca?as de Jul G. Galeano, R. Gordas, Lozano, 1984, Mpio. Boquer?n Toyo, Londo?o, R. Bernai & D. Restrepo 3963 (holotype: COL!; isotype: MO!).

5. Andira

taurotesticulata

Species bene distincta, quae dum florens Andira macrothyrsa et in fructu vel A. cori ?cea vel A. micrantha fortasse confusa possit. Ab A. macrothyrsa differt ovario in paginis superioribus inferioribusque tantum (haud uniformiter ubique) dense pubescenti. Ab A. cori?cea et A. micrantha fructibus non laevibus sed de sutura ad suturam costatis, foliolis inferne sparse pubescentibus, non glabris bene insignis. Tree to 25 m tall; buttresses absent; bark smooth; slash pale red brown, oxidizing darker, 7 mm thick; twigs brown to grey-brown, bark cracking on older twigs, red-brown, with ? appressed hairs, glabrescent. Stipules to 12 mm long, to 1 mm wide, with red brown appressed hairs; leaf axis 9-30 cm long; rhachis sparsely hairy, glabrescent, hairs red-brown, ? erect; stipels 1-3 mm long, persistent; petiolules 3-8 mm long, indumentum like that of rhachis; leaflets in 3-5 (-10) pairs, 4.5-15 cm long, 2-7.3 cm wide, subcoria ceous, elliptic, narrowly elliptic, or lanceolate to narrowly obovate, base obtuse to rounded, apex obtuse to rounded (rarely acute), generally with an acumen to 15mm long, mucronate becoming slightly retuse, glabrous adaxially, sparsely hairy abaxially, glabres cent, hairs pale to red-brown, ? erect to ? appressed, to 1.0 mm long; primary vein chan nelled adaxially; secondary veins 8-11 (-14), ? plane to slightly sunken adaxially, ? plane to slightly raised abaxially (occasionally prominently raised), pattern eucamptodromous or ? completely brochidodromous, becoming brochidodromous tertiary veins plane adax ially and abaxially. Panicles axillary and terminal, 6-25 cm long, with red-brown ? ap pressed hairs at branch tips, glabrescent towards the base; bracts 1.5-2 mm long, with red brown ? appressed hairs; pedicels 0.5 mm long or flowers sessile; bracteoles 1-1.2 mm long, indumentum

like that of bracts. Flowers

7-10 mm

long. Calyx

3-4 mm

long, hairy

ANDIRA

2003

57

to very sparsely hairy, hairs red-brown, ? appressed; lobes 0.2-0.5 mm long, obtuse, shal low. Petals white, the standard marked with red; standard blade 7-10 mm wide, 5.5-7.5 mm high, claw 1.5-3 mm long; wing 4-6 mm long, 2-3.5 mm wide, claw 3-4 mm long, sculpturing absent; keel 4-5 mm long, 2.5-3 mm wide, claw 4 mm long. Stamens 8-8.5 mm

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

(Fig. 19). Panama (Dari?n, San Bias), Colombia (Antioquia, Boyac?, Meta, Quindio, Santander, Valle), Venezuela (T?chira), Ecuador (Pichin cha, Sucumbios, Napo); in montane forest and its transition to rain forest (700-2000 m) in Colombia and Venezuela, and in lowland rain forest in Ecuador. Vernacular name. Cojones de toro (Antioquia).

Cundinamarca,

Additional La Escalera,

Specimens

Examined.

E of Tres Bocas, Kirkbride

Dari?n:

Panama. & Duke

1315

Cuasi-Cana

trail between

San Blas: (MO, NY).? Croat 26024 (MEXU, MO).

Cerro Campamento and road, Km 12, in

El Llano-Carti

Control Project, vicinity of Gorgas Lao Mosquito Colombia. 16 km E San Rafael along Guatap?-San Rafael road, Brant & Antioquia: Mpio. San Rafael, Roldan 1533 (E, MO); Mpio. San Carlos, Correg. Alto Samana, vereda Miraflores, finca "El Despesero," Calle San Luis, right bank Rio Samana, vereda el Port?n, road to the autopista jas et al. 8584 (HUA, NY); Mpio. C?rdenas 2501 (COL, JAUM); Mpio. San Luis, Autopista Medell?n-Bogot?, vereda la Jose Medell?n-Bogot?, C?rdenas fina, quebrada laMariola, Sector R?o Samana-R?o

l?n-Bogot?,

MO); Mpio. Josefina,

& J. G. Ram?rez Claro,

San Luis, Autopista Medell?n-Bogot?, & Estrada 270 (JAUM, MO);

Cogollo

2743 (JAUM); Mpio. San Luis, Autopista Medel la vereda Tulip?n, Cogollo & Estrada 199A (JAUM, Sector R?o Samana-R?o Claro, camino hacia la vereda La 12 km from Au San Luis, carretera hacia Aquitania, Mpio.

camino

2741,

hacia

et al 3762 (JAUM); Mpio. San Luis, Autopista Medell?n-Bogot?, vereda la topista Medell?n-Bogot?, Cogollo et al. 4306 (JAUM); Parque Nacional Natural "Las Orqu?deas" Sector quebrada laMariola, Cogollo 4145 (COL, JAUM); Mpio. San Luis, Autopista Medell?n-Bogot?, Calles, right bank of R?o Calles, Cogollo Josefina,

vereda

la Josefina, Hoyos & J. J. Hern?ndez 286 (JAUM, MO); Mpio. San Luis, Autopista Medell?n-Bogot?, de la vereda la Josefina a la vereda la Holanda, Hoyos & J. J. Hern?ndez 334 (JAUM, MO); Mpio. San vereda la Josefina, camino al Tulip?n, ca?o laMariola, Hoyos & J.J. Hern?n Luis, Autopista Medell?n-Bogot?, camino

dez 917

Cerro road, 11 km above Frontino around el Aeropuerto, (JAUM, MO); Frontino-El Luteyn & Lebr?n 7196 (COL, MO, NY, US); Mpio. San Luis, Quebrada "La Cristalina," J. G. Ram?rez & L?pez 950 (HUA, de Providencia, valle del R?o Anor?, JAUM, MO), 1161 (COL, HUA, JAUM, MO); Mpio. Anori, Corregimiento entre Dos Bocas y Anor?, Buenos Aires, 4 km from Providencia, Soejarto 3954 (GH); Mpio. Ituango, Km 9 of

Luteyn

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

SYSTEMATICBOTANYMONOGRAPHS

58

18. Andira

FIG. face.

Fruit. M. cence),

taurotesticulata.

A. Habit.

B. Broad

leaflet. C. Narrow

F. Calyx, opened to show inner surface. G. Standard K. Gynoecium, I. Keel petal, inner surface. J. Androecium.

E. Flower.

leaflet. D. Abaxial

leaflet surface.

inner surface. H. Wing petal, outer sur section. L. shown above in longitudinal

in longitudinal section, showing wall structure. (Based on: A et al. 334; C, H. Cuadros G. Lozano et al. 3963; B, D, S. Hoyos

Fruit

E-K,

petal, also

VOLUME 64

(leaf), W. Devia 941; A (inflores 643; L-M, W. Devia 941.)

ANDIRA

2003

59

A. taurotesticulata FIG.

19. Distribution

o? Andira

taurotesticulata.

et 11706 (MO); Cerro Las Minas, S of main road from Santa Ana, 17 km SE Santa Ana, Steyermark Guariglia 120019 al 119886 18 km SE Santa Ana, Steyermark (MO); Cerro Las Minas, slopes leading to Cerro Azul, van der Werff & Ortiz 5553f 5556, 5576 (E, MO). Ecuador. NAPO: Can (MO); Distr. Lobatera, La Cazadora, ton Orellana, Reserva El Chuncho, 1 km W Rio Payamino, NW Coca, Baker 7025 (NY); Cant?n Orellana, Via 1 km W R?o de Zorros, Pozo de Jaguar I, Palacios 3204 (QCNE); Cant?n Orellana, Reserva El Chuncho, & M. Aulestia 524, 525 (E, K, QCA, QCNE).?PICHINCHA: Quito, Payamino, NW Coca, R. T. Pennington reserva Faun?stica Reserva Juan Manuel Palacios 12274 Durini, (QCNE).?SUCUMBIOS: Lago Agrio, Field Station, Neill 10214 (QCNE). Laguna Grande, near Catholic University Cuyabeno,

is the only species that grows inmontane forest (to 2200 m), Andira taurotesticulata and extending along the in Colombia, occurring in all three cordilleras of the Andes to Venezuela. It is also found in the Caribbean coastal ranges in Cordillera Occidental Panama (in Dari?n and San Bias), but in Ecuador it is found at lower altitudes (to 200 m). It is clearly distinct by virtue of its small flowers, large fruit, and leaflets that are sparsely hairy abaxially. There is wide variation in leaflet size, shape, and indumentum, in flower size, and the extent of the gynoecial indumentum, which reflect this species' wide eco logical and geographical range. The variation appears continuous, and the fruit are uni form. The Ecuadorian specimens are somewhat disjunct from the rest of the range, but are similar; the only difference is an endocarp with more abundant stone morphologically cells (Gemeinholzer

1995). Flowering specimens of A. taurotesticulata might be confused with A. macrothyrsa. The wing petals of A. taurotesticulata lack sculpturing, and the ovary is hairy on its upper and lower surfaces with the sides of the ovary sparsely hairy, or only the upper and lower surfaces of the ovary are hairy. In contrast, A. macrothyrsa has wing petal sculpturing, and its ovary is uniformly densely hairy. In fruit, A. taurotesticulata might be confused with A. cori?cea and A. micrantha, the other rain forest species o? Andira with large fruit. The fruits of A. taurotesticulata are distinctly ribbed from suture to suture, whereas the fruit of

SYSTEMATICBOTANYMONOGRAPHS

60

VOLUME 64

the other species are relatively smooth and lack the distinct ribs. Moreover, these other in Guianas and central Brazil have restricted distributions the (A. cori?cea) (A. mi species are A. and from taurotesticulata. crantha), widely disjunct The name for this species comes from its local name in Colombia:

"cojones de toro."

Brazil. Ducke, Bol. T?cn. Inst. Agron. N. 2: 31. 1944.?Type: macrothyrsa Amazonas: Esperan?a, 22 Oct 1942, A. Ducke 1036 (holotype: RB!; isotypes: K! R?RB-2 sheets! US!). PERU. Andira multistipula var. peruana N. F. Mattos, Loefgrenia 53: 2.1971.?TYPE: 16 Feb 1924, J. G. Kuhlmann 1505 (holotype: RB!; isotypes: RB R?o Huall?ga,

6. Andira

2 sheets!). Tree to 40 m tall, often a canopy emergent; buttresses slight; bark smooth and grey; dark brown; slash pale red-brown, slowly oxidizing red-brown; twigs dark brown, often with the outer bark splitting to reveal paler bark beneath, densely hairy, glabrescent,

wood

hairs brown, short, appressed; older twigs with pale, raised, elongated lenticels. Stipules early caducous (not seen); leaf axis 10-33 (-40) cm long; rhachis with appressed brown hairs, glabrescent, hairs short; stipels to 2 mm long, caducous; petiolules 3-9 mm long, indumentum like that of rhachis; leaflets in 4-8 pairs, 4.5-11.5 cm long, 1.8?4 cm wide, narrowly ovate, elliptic, or narrowly obovate (rarely narrowly elliptic), subcoriaceous, base obtuse (rarely acute); apex obtuse, often retuse or with an acumen to 10 mm long, glabrous adaxially, hairy to sparsely hairy abaxially, hairs short, appressed, red-brown, <0.2 mm

long; primary vein channelled adaxially; secondary veins 8-12, plane to very becoming slightly raised adaxially, slightly raised abaxially, pattern eucamptodromous the veins generally curving uniformly, tertiary veins plane adaxially brochidodromous, and plane to slightly raised abaxially. Panicles terminal or axillary (axillary inflorescences

are smaller), 8-30 cm long, with dense red-brown appressed hairs at branch tips, glabres cent towards the base; bracts 2 mm long, early caducous, with dense appressed red-brown hairs; pedicels to 1mm long or flowers subsessile; bracteoles 1mm long, early caducous, indumentum as bracts. Flowers 9-11 mm long, unpleasantly scented. Calyx 4-5 mm long, with dense, red-brown, appressed hairs; lobes 0.5-1.2 mm long, acute or obtuse, the apex to greenish, the standard with a central red-brown to acuminate. Petals white-yellow carmine marking; standard blade 5.5-8 mm wide, 4.5-5 mm high, claw 2-2.5 mm long; wing 4-5 mm long, 1.5-2 mm wide, claw 3-3.5 mm long, lamellate sculpturing present; keel 2.5^ mm long, 1-1.5 mm wide, claw 3.2^1 mm long. Stamens 6-7 mm long, fila ments united for the basal 2-4.5 mm, free for the distal 2-3 mm, the vexillary stamen 4.5 mm

6.5-8 mm long, the ovary with dense red-brown appressed hairs, long. Gynoecium the indumentum extending to the top of the stipe and base of the style; stipe 2.5-3.5 mm long, ovary 2.5-3 mm long, style 1.5 mm long; ovules 1-2. Fruit 5-5.5 cm long, 3^1 cm high and wide, elongated, weighing 20-30 g when dry, green and strongly scented, dry ing smooth, brown to red-brown; mesocarp 2-2.5 mm thick, reddish brown, granular; en docarp 1.5 mm thick, woody with coarse, pale fibers. Chromosome number unknown.

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

Los R?os), forest and the Andes; coast.

2003

id

ANDIRA 61

- -

A. macrothyrsa A a. chigorodensis FIG. 20. Distribution

Vernacular

names. Uxi de morcego

of Andira macrothyrsa

and A. chigorodensis.

(Brazil, Acre);

casub guesmoteig

(Aw?; Ecuador,

Carchi). Additional Specimens Examined. Colombia. Choc?: de Utr?a, NE de Parque Nacional alrededor de la quebrada la Chunga, F. Garcia C. Utria, entre alManglar y la falda de lamonta?a near mouth of Quebrada 443 (MO).?Valle: Bah?a de Malaga, la Sierpe, Gentry et al. 40434 Ecuador. CARCHI: Tulcan Cant?n, Parroquia Tobar Donoso, Sector Sabalera, Reserva MO).

la Ense?ada

de

& Agualimpia (COL, JAUM,

Ind?gena Aw?, Cerro Centinela, Mon

1345 (MEXU, QCNE).?Los R?OS: Quevedo Cant?n, Tipaz et al. 1225 (E,MO, QCNE), ta?as de Da, 10 km E Patricia Pilar, Palacios & Freir? 7444 (QCNE).??apo: Carretera Holl?n-Loreto Km 75, cerca de R?o Guantaraco, Neill et al. 8056 (K,MO, QCA, QCNE); Cant?n Orellana, Est. Exp. INIAP-Payamino, 5 km NW de Coca, Palacios ca. 1 km after R?o Guataraco 4818 (MO); Cant?n Loreto, Sector R?o Guataraco, on road Holl?n-Loreto, left side road, R. T. Pennington & M. Aulestia 523 (E, K, QCA, QCNE).?SUCUMBIOS: Reserva

1 km N Laguna Grande, hectare plot 1, Valencia 67876 (QCA). Peru. Loreto: Cuyabeno, tourist camp, halfway between Indiana and mouth of Napo, Gentry et al. 60683 (MO); above mouth of Rio Napo, Gentry et al. 37168 Explorama Tourist Camp, R?o Amazonas (MEXU,

Faunistica

Yanamono, Yanamono,

Explorama

T. D. Pennington & Ruiz 13550 (K); Prov. Requena, Iquitos, Quistacocha, Sapuena, Bagaz?n-R?o Estaci?n Experimental V?squez & Jaramillo 8745 (MO); Maynas Province, IIAP, Iquitos, Allpahuayo, Brazil. ACRE: Mpio. Rio Branco, estrada para Porto Acre Km 13, ramai a es V?squez et al. 13802 (E, MO). 3057 (NY); Mpio. M?ncio querda, 4 km da estrada Colonia Cinco Mil, Cid Ferreira & Nelson Lima, estrada et al. 5247 (NY); Mpio. M?ncio Lima, estrada do Isac, 4 km from para o lugar Bar?o, Km 30-32, Cid Ferreira et al. 10950 (MEXU, NY); Sub-base do Cruzeiro do Sol, Marinho 27 (RADAM), vie. Serra city, Cid Ferreira

MO,

NY);

Ucayali,

Prance et al. 12388 (US); Cruzeiro do Sol, pr?ximo ao Acampamento do Projeto RADAM, Rosa 668 Alto Solim?es, Mpio. Sao Paulo de Oliven?a, estrada do Bon?im, (NY).?Amazonas: logo ap?s a saida da cidade de Sao Paulo de Oliven?a, H. C. de Lima et al. 2783 (K).

da Moa,

VOLUME 64

SYSTEMATICBOTANYMONOGRAPHS

62

most similar to A. chigorodensis is morphologically Andira macrothyrsa (see note under that species, no. 7), but may also be confused with A. inermis subsp. inermis. Andira macrothyrsa differs from A. inermis in the short, appressed hairs on the abaxial leaflet sur faces and in its whitish to greenish flowers with a red-brown to carmine spot in the cen ter of the standard petal. Andira inermis has glabrous leaflets and pink flowers with a white spot on the standard. In some specimens from Brazil (Acre and Amazonas: Cid Ferreira 5247, Rosa 668, H. C. de Lima 2783) the leaflets are blunt-ended, almost oblong, matt brown abaxially, with the secondary veins curving as they approach the margin. These characters provide no basis for recognition of subspecific taxa, because states that are intermediate between those of this group and the remaining specimens of A. macrothyrsa are found in two spec imens?L. Marinho 27 (Brazil, Acre) and V?squez & Jaramillo 8745 (Peru, Loreto). 7. Andira

R. T Pennington, Colombia. sp. no v.?Type: Antioquia: 21 chigorodensis 10 Jul 1981, J. Brand & A. Cogollo 114 (holo km de Apartado hacia Chigorod?, type: HUA!; isotypes: COL! MO!).

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

fibrous. Chromosome

number unknown. Figs. 3B, 4B(ii),

13B, 21.

ANDIRA

2003

FIG. 21. Andira

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

surface.

SYSTEMATICBOTANYMONOGRAPHS

64

Phenology. vember,

Flowering

VOLUME 64

July (only one record); fruiting records from September, No

February.

January,

(Fig. 20). Colombia

Distribution

(Antioquia);

in rain forest; to 300 m.

Examined. Colombia. Turbo, 8 km de Carepa hacia Chigorod?, Antioquia: de Lomas Aisladas, Renteria et al. 3579 (HUA, JAUM, MO); Mpio. Turbo, corregimiento Renteria 4634 (JAUM, MO); Mpio. camino Malag?n-Chigorod?, (JAUM-2 sheets), MO); Mpio. Chigorod?, 13 km W of El Jardin, Finca El Amparo, Zarucchi & D. C?rdenas 4215 (COL, HUA, MO). C?ceres, Additional

Specimens

Brand & Cogollo

215

Andira chigorodensis most closely resembles leaves, 37-55 cm long, with narrowly elliptic or 5-6 (-7) mm long. The leaves of A. macrothyrsa rowly ovate, elliptic, or narrowly obovate leaflets, Andira 8. Andira

chigorodensis

is endemic

A. macrothyrsa. It differs in its larger lanceolate leaflets, and smaller flowers, are 10-35(-40) cm long and have nar and the flowers are 9-11 mm

to the Chigorod?

region of Antioquia,

long. Colombia.

galeottiana Standley, Contr. U.S. Nati. Herb. 20(6): 217. 1919.?Type: Mex ico. Veracruz: Catemaco, 26 Apr 1894, E. W. Nelson 424 (holotype: US!; iso type: NY!).

Tree to 30 m tall; buttresses absent; bark scaling; slash pinkish brown; sapwood yel lowish, heartwood reddish brown; twigs densely hairy when young, glabrescent, hairs red brown, tangled, erect. Stipules to 5 mm long, to 1 mm wide, early caducous, with red brown ? appressed hairs, glabrescent; leaf axis 10-30 cm long; rhachis hairy, glabrescent, hairs red-brown, tangled, erect; stipels 2-3 mm long, caducous; petiolules 3-6 (-7) mm long, hairy, hairs red-brown, tangled, erect; leaflets in 4-5 pairs, 4-12.5 cm long, 2.3-6 cm wide, elliptic, narrowly obovate to broadly obovate, subcoriaceous to coriaceous, base obtuse (rarely acute or rounded), apex obtuse to rounded, often with an acumen to 5 mm long, or retuse, glabrous adaxially, hairy abaxially, hairs erect, red-brown, >0.2-1.0 mm long; primary vein channelled adaxially; secondary veins (6-) 8-12, slightly sunken adax ially, raised abaxially, pattern brochidodromous, tertiary veins plane adaxially and raised abaxially. Panicles terminal or axillary, 15-50 cm long, with dense, tangled, erect pale red-brown hairs at branch tips, glabrescent towards the base; bracts 1-1.5 mm long, with ? erect, early caducous brown hairs; pedicels 2-3 mm long; bracteoles 1mm long, indu mentum like that of bracts. Flowers 13-17 mm long. Calyx 7-7.5 mm long, with ? erect, pale brown hairs; lobes 1-1.5 mm long, acute or right-angled. Petals pink to violet; stan 11-14 mm high, 9-12 mm wide, claw 4 mm; wing 8-10 mm long, 4^.5 mm wide, long, lamellate sculpturing present; keel 8-10.5 mm long, 4^.5 mm mm wide, claw 5.5-6 long. Stamens 13-13.5 mm long, filaments united for the basal 7-8 mm, free for the distal 4-5.5 mm, vexillary stamen 10-10.5 mm long. Gynoecium 13.5-15 mm long, glabrous, or with 1-3 scattered hairs on the lower ovary surface; stipe 5.5-6 mm long, ovary 4^.5 mm long, style 3.5-5 mm long; ovules (1-) 2. Fruit 6-10 cm dard blade

claw 5 mm

long, 4.3-8.5 cm high, 5-8.0 cm wide, elongated, weighing 40-125 g when dry, green be coming dark brown or black when mature, drying brown to dark brown, smooth, flattened adaxially and below with broad ribs often running diagonally from base to apex, suture a broad groove adaxially; stylar remnant minute or absent; mesocarp 5-10 mm thick, dry ing pale brown, soft, with spongelike texture and fibers extending from the endocarp; en docarp 3-5 mm thick, buff, woody, fibrous, thickened along both sutures. Chromosome number

unknown.

ANDIRA

2003

110

105

100

95

90

85

80

FIG. 22. Distribution

75

o? Andira

65

70

60

65

galeottiana

55

50

45

40

and A. verm?fuga.

Phenology. Flowering April to July. Distribution (Veracruz, Oaxaca, Tabasco, Chiapas); wet (Fig. 22). Mexico banks, seasonally flooded zones, also in non-inundated forest. Vernacular

Representative

names.

Macayo/a

(Veracruz,

Oaxaca,

35

Tabasco);

lombrisero

sites, river

(Oaxaca).

M. E. Beccara Chiapas: Tierra Colorada, Villahermosa, s.n. (MEXU); 1 km SW junction to Emiliano Zapata, road Villahermosa-Es 7606 (MEXU); Mpio. Reforma, (MEXU); La Arena, Palenque, F. Miranda Laiana (Chinantla), Galeota 19498 (MEXU).?OAXACA: 3464 (US); Dist. Ju

SPECIMENS EXAMINED. Mexico.

s.n. (MEXU); Palenque, Fuentes c?rcega, Grether & Quero 1579

Petrolero Cactus, Tenorio H. M. Hernandez & Torres 335 (MO); Mpio. Santa Maria chit?n, 6.6 km W Mago?? por camino a Guichieovi, 418 (MO); Mpio. Tuxtepec, Tuxtepec, Mart?nez Calder?n 1369 (NY, Santa Maria, H. M. Hern?ndez Cnimalapa, & Sarukh?n 9067 (NY); Acatl?n de P?rez MEXU, MO); Mpio. Tuxtepec, Ejido Benito Ju?rez, T. D. Pennington Campo

(MEXU, US); Dist. Choapain, San Juan Lalane, Schuttes & Reko 852 (F, GH); Mpio. Tux Figueroa, Salazars.n. 9451 (A, F, U); Tolosita, L. Williams 9612 (A, tepec, Las Pinas, Sousa 3667 (MEXU, US); Ubero, L. Williams R?o Rosario, carretera E Savanna Area, Barlow 30/186D F, U).?TABASCO: (MEXU); Huimanguillo de Huimangillo Ch?velas & P?rez J. ES892 hacia Francisco (NY); Km 35 de la deviaci?n Zapata-Tenonique, 3125 (A, F, NY); Mpio. Centro, Laguna Estancia Rueda, C. Cowan et al 2294 (MO, NY); Balanc?n, Matuda Novelo & Ramos Vieja, cerca de Luis Gil P?rez, S Villahermosa, La Venta, Laguna Yucateco, edge of lake, near Rio Chicozapote, on road to Balanc?n, T?llez & E. Mart?nez junction to Tenosique

(MEXU); Mpio. Huimanguillo, Novelo & Ramos 1786 (MEXU);

10 km N 6 km from

905 (MEXU); 900 m from junction Chatle-El on road to Balanc?n, T?llez & E. Mart?nez 948 (MEXU).?VERACRUZ: Mpio. San Juan Lalana, Santi 207 km S de Catemaco, 16846 Jalahui, carretera a San Juan del R?o, Calzada (MEXU); Hueyapac

Triunfo, ago

1770

SYSTEMATICBOTANYMONOGRAPHS

66

de Ocampo,

Cedillo

& Calzada

near Huaxpala, Playa Vicente, Catemaco, Faden et al. 76/125 Hanson

17 (F); San Lorenzo Tenochtitl?n, et al ES-4301 Ch?velas (MEXU);

Ch?velas

VOLUME 64

et al. ES-2396

(MEXU); Mpio.

Island, Lake Agaltepec Mpio. Catemaco, 1.5 km N Tabanca, 10 km E Catemaco, (F, K); along E edge Lago Catemaco, 121 (MO); Men?ndez Catemaco, (R?o Coscoapan), Laguna de Sontecomap?n

& Nee 7637 (F, MO); 171 (GH, MEXU-2 de Sontecomap?n, & G?mez-Pompa sheets); 2 km Adelante Nevling Panga, R?o ca. de C?rdoba, 840 (GH, MEXU, & G?mez-Pompa Chocam?n, Teschaacu, Nevling US); P. Est. Chocam?n, de los Tuxtlas, R?o Tecesapa, al E de Seteapan, Sousa 3267 (MEXU); Mpio. Salazar s.n. (MEXU); Region Barra

Catemaco,

a 2 km de Coyame

Lago Catemaco,

13407

(MEXU); Fortuno, grove, Wing 25 (GH).

Coatzacoalcos

sobre camino

river, Williams

8483

a Tebanca, (F, US);

orilla de Lago Catemaco, Sousa et al. on Tenochtitl?n village path to orange

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

cating probable dispersal by vertebrates. Andira galeottiana has hard timber, which is used in the construction of houses, bridges, and railway sleepers. Schuhes (herb, label, Schuhes 852) reports use as a ver mifuge and to treat fungal diseases of the skin, Salazar (herb, label, unnumbered collec tion) reports an extract of the bark used as a febrifuge, and Antonio and Heinrich (herb, label, Antonio & Heinrich GUI224) report "fruits" (presumably seeds) used as a rat poi son when mixed with maize flour. 9. Andira

verm?fuga (Martius) Bentham, Comm. legum. gen. 44. 1837. Geoffroea ver m?fuga Martius in Spix andMartius, Reise Bras. 2: 788. 1828. Vouacapoua ver Brazil. Minas m?fuga (Martius) Kuntze, Revis, gen. pi. 1: 212. 1891.?Type: s.n. Gerais: "Inter fr?tices ad Salgado, et in campis editis Taboleiro," Martius (holotype: M!). spinulosa Martius in Spix and Martius, Reise Bras. 2: 788. 1828. Andira spinulosa (Martius) Bentham, Comm. legum. gen. 44. 1837. Vouacapoua spinu losa (Martius) Lyons, PL nam. 396. 1900.?Type: Brazil. Minas Gerais: "in

Geoffroea

prov. Minarum campis editis siccis ad Contendas," Martius s.n. (holotype: M!). Andira paniculata Bentham, Comm. legum. gen. 45. 1837. Vouacapoua paniculata Brazil. "Barbacena et (Bentham) Kuntze, Revis, gen. pl. 1: 212. 1891.?Type: s.n. Ponte d'Erva," Pohl (holotype: K!; isotypes: K! M, PR!, photos of M iso type: G! GH!). Andira kuhlmannii N. F.Mattos,

S?o Paulo: Loefgrenia 40: 2. 1970.?TYPE: Brazil. Moji-Mirim, margem da rodovia para Campinas, pr?ximo ? entrada da Fazenda Holambra, M. Kuhlmann 3945 (holotype: HB; isotypes: K! SP).

to 12 (-25) m tall with spreading crown, occasionally a shrub, able to root or buttresses bark with absent; thick, very slight sprout; rough deep vertical fissures; slash pale brown to red-brown to orange-brown, with red ex?date; wood generally to often the hairs ? buff/cream; swollen, densely twigs sparsely hairy, glabrescent, Tree

2003

ANDIRA

67

appressed, red-brown; bark cracking on older twigs; lenticels absent. Stipules to 5 mm long, to 1mm wide, caducous; leaf axis 6-30 cm long; rhachis brown to red-brown, hairy to sparsely hairy, glabrescent, hairs pale brown, erect; stipels 1-2 mm long; peti olules 2-5 mm long; leaflets (2-) 3-5 (-6) pairs, 4-11 cm long, 1.8-6.6 cm wide, broadly elliptic, elliptic, broadly obovate (rarely ovate, broadly ovate, suborbiculate, or to coriaceous, dark green, shiny adaxially, biculate to narrowly obovate), subcoriaceous matt abaxially with red-brown indumentum, base obtuse to rounded (rarely acute), apex obtuse to retuse (rarely acute or emarginate), glabrous adaxially, hairy to sparsely hairy abaxially, hairs erect, pale red-brown, >0.2-1.0 mm long; primary vein channelled veins 5-11, slightly sunken adaxially, raised abaxially, pattern eu brochidodromous brochidodro (occasionally completely camptodromous becoming Panicles terminal veins and raised (more mous), tertiary plane adaxially abaxially. to rarely axillary), densely covered red-brown erect hairs at branch tips, glabrescent secondary

adaxially;

wards

the

base;

bracts

narrow,

caducous,

ca.

1.5-2

mm

long,

with

red-brown,

?

ap

pressed to erect hairs; pedicels 2-A mm long; bracteoles narrow, caducous, 1mm long, indumentum like that of bracts. Flowers 12.5-18 mm long. Calyx 6-7 mm long, brown to deep purple-brown, sparsely hairy, hairs red-brown to pale red-brown, ? appressed; lobes 1-2 mm long, acute to obtuse. Petals pink to purple; standard blade 9-15 mm 9-12 mm high, claw 3.5-6 mm long; wing 8-10 mm long, 3.5-6 mm wide, claw mm long, lamellate sculpturing present; keel 7-10 mm long, 3.5-5 mm wide, claw 4-6.5 mm long. Stamens 12-16 mm long, filaments united for the basal 5-9.5 mm, free for the distal 3-7 mm, vexillary stamen 8-11 mm long. Gynoecium 12-15 mm long, ovary glabrous or the upper and lower surfaces sparsely hairy, hairs ? erect, pale with

wide, 4-6.5

bases; stipe 4-6.5 mm long, ovary 3-5.5 mm long, style 2.5-4.5 mm long; (2-) 4-6. Fruits 2.4-4 cm long, 1.2-2.5 cm high, 1.7-2.5 cm wide, elongated, ca. 20 g or less when dry, green, strongly scented, drying brown (more rarely weighing red-brown or dark brown), wrinkled, suture a fine raised line or indiscernible adaxially, indiscernible abaxially; stylar remnant tiny; mesocarp 0.5-2 mm thick, fibrous, often drying with oily granules; endocarp 1-3 mm thick, pale red-brown, woody, fibrous. red-brown

ovules

Chromosome

number unknown.

Fig. 4A. Phenology. Flowering April to October, but the great majority of records from Au gust and September (occasional records throughout the year). Distribution (Fig. 22). Peru (Pasco), Bolivia (Pando, Santa Cruz), Brazil (Acre, Ama

zonas, Bahia, Cear?, Distrito Federal, Goi?s, Maranh?o, Mato Grosso, Minas Piau?, R?ndonia, Sao Paulo); in cerrado and gallery forest. Vernacular names. Angelim preto, mata barata, angelim branco (Brazil). Representative Hartshorn

et al. 2918

Specimens

Examined.

Peru.

Pasco:

PANDO: Manuripi, Bolivia. (MO). Prov. Chiquitos, Serran?a de Santiago,

Gerais,

Cerro de Pasco, central selva, Palcazu Valley, 35 km N Puerto America, A. Jardim 1050 (K, near Santiago de Chiquitos, 5 km ENE Santiago,

Cruz: MO).?Santa Serran?a de Santiago, Santiago de Chiquitos, Daly et al. 6272 (NY); Prov. Chiquitos, near mouth of Rio Macauhan mani & A. Jardim 1291 (MEXU). Brazil. Acre:

camino

a la cueva, Ma

of Rio Yaco), 150 km SE Hu (A, F, U, US).?AMAZONAS: Mpio. Humait?, Janssen 139 (K); Transamaz?nica, Sanaiotti 287 (E, INPA).?Bah?a: mait?, 22 km E Bodoc?, Mpio. Gentio do Ouro, ca. 6 km after Santo Ina cio on Xique-Xique-Gentio do Ouro road, R. T. Pennington & Brito 256, 257, 258, 259 (CEPEC, FHO, K); & Brito 265 (CEPEC, FHO, K); Mpio. tt>oti Mpio. Barreiras, ca. 2 km along road to airport, R. T. Pennington Krukoff

(tributary

5483

rama, rd. Ibotirama-Seabra

& Brito 270, 271, 272, 273 (CEPEC, FHO, K); (BR-242) Km 8, R. T. Pennington et al. 20 (?B).?Cear?: Fazenda Serra de Ibiapaba, Campo Grande, Correntina, Jatob?, Rezenda 880 (F).?DISTRITO FEDERAL: Bacia do Rio S?o Bartolomeu, et al. 5450 (US); Chapada Dahlgren Heringer da Contagem, Irwin et al. 7954 (MO, US); Fazenda Agua Limpa, near Vargem Bonita, 18 km SSW Brasilia

Mpio.

SYSTEMATICBOTANYMONOGRAPHS

68

et al 194 (UB); Chapada Alvarenga Fazenda Breij?o, (NY); Chapada dos Veadeiros, et al. 199 (E, UB); Mpio. Goi?s, S?o Joaquim, Gibbs et al. 2713 (K, NY); Mpio. Goi?s, Serra Bridgewater ca. 38 km NE Formosa, Irwin et al. 15191 (F, Dourada, Gibbs 2782 (K, NY, UB); near C?rrego Estrema, W 12 km of rd from of Presidente Presidente MEXU, NY); Mpio. Kennedy, village BR-153-Itapor?, TV

tower, Ratter

dos Veadeiros,

et al 3595 (E, F, K, UB).?Goi?s: 11 km E Cavalcante, W. R. Anderson

VOLUME 64

Mpio. 7294

Vian?polis,

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

Kennedy, arredores

Fazenda

Primavera

de Bar?o

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

Almeida

orado, Fazenda

20 km from Vilhena, Santa

221174

(UB); Mpio.

Brotas,

et al 2810 (GH, NY, US).?S?O PAULO: Araraquara, Usina Tamoio, 14 km E de S?o Manuel, 18 km N Botucatu, (US); Mpio. Botucatu, along near Est. "13 Maio" of the old railroad, /. S. Gottsberger & C. J. Campos 12 Ja?, Leit?o Filho et al. 12927 (US).

Zarucchi

Joana, Felippe to Piracicaba, road S?o Manuel

94

Andira verm?fuga ismost closely related toA. galeottiana (no. 8; see notes under that for differential characters). species Specimens of A. verm?fuga may be divided into two groups, one characterized by thick, swollen twigs, the other by thin, leptocaul twigs, which are more usual in Andira. two forms overlap somewhat geographically, though the majority of the thin are from the northeast of the range of A. verm?fuga (Bahia and twigged collections Maranh?o). No differences can be found to separate these groups in leaf, flower, or fruit in which the twigs appear intermediate (e.g., characters, and there are some collections R. T. Pennington 249, 258, C. A. Cid 6416). This indicates little basis for splitting A. verm?fuga subspecifically. Other specimens show new growth from swollen twigs to be These

slender and leptocaul, suggesting that the underlying basis for the differences may be related, e.g., repeated burning may induce formation of thick twigs. environmentally This is corroborated by personal observations of a population of A. verm?fuga (at a Fazenda Agua Limpa, Distrito Federal, Brazil), which showed gradation from tall trees with slender twigs in gallery forest to stunted trees with thick twigs in open cer rado only 50 m away. Andira kuhlmannii N. F. Mattos appears to be based upon a spec imen from a large, gallery forest individual of A. verm?fuga, and the name is here placed in synonymy. Several interesting collections from the northwestern edge of the range are assigned to A. verm?fuga. Zarucchi et al. 2810 from Rond?nia appears in all vegetative and fruit characters to be A. verm?fuga, but the notes indicate that the twigs are hollow and house ants. The information given is insufficient to determine whether this is only a casual as et al. 8719 (Rond?nia, Brazil), Krukoff 5843 (Acre, Brazil) and sociation. Prance et al. 2918 (Pasco, Peru) all represent large trees to 25 m tall, but with flow Hartshorn ers 12.5-14 mm long, which is somewhat small for A. verm?fuga. They were collected "transition forest" (sensu Ratter, 1992) on the edge of Amazonia, extending of A. verm?fuga into this ecosystem. Prado and Gibbs (1993) and Pennington the range et al. (2000) give several examples of this type of distribution of a woody species, which in seasonal

ANDIRA

2003

69

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

in the synonymy

of A.

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

10. Andira

humilis Martius

milis

Andira pauciflora Bentham, Comm. legum. gen. 45. 1837.?Type: Gerais: "Ad Barbacena et Ponte d'Erva," Pohl s.n. (lectotype,

Brazil.

Minas

here designated:

K!). Andira humilis var. cordata N. F. Mattos, Loefgrenia 40: 3. 1970.?TYPE: S?o Paulo: Itahy, 10 Dec 1929, /. /. de Lima s.n. (holotype: RB!).

Brazil.

suffrutex forming mats up to 10 m in diameter, aerial shoots to 50 cm tall Geoxylic (occasionally shrublike, to 2 m tall); bark brown, Assuring vertically; slash pale brown to pale red-brown with some clear ex?date that oxidizes red; twigs brown to buff, bark crack ing on older twigs, with short, red-brown appressed hairs, glabrescent. Stipules to 7 mm long, to 1 mm wide, with short, red-brown appressed hairs; leaf axis 9^45 cm long; rhachis sparsely to very sparsely hairy, glabrescent, hairs short, red-brown, appressed; stipels 1^1 mm long; petiolules 1-5 mm long, indumentum like that of rhachis; leaflets in 4-7 pairs, 4-12.5 cm long, 1.3-4 cm wide, elliptic, narrowly elliptic (rarely narrowly obo to coriaceous, shiny, dark green vate, oblanceolate, ovate to lanceolate), subcoriaceous matt and the venation base obtuse, rounded (occasionally adaxially, paler abaxially, paler, truncate or ? cordate), apex obtuse to rounded (occasionally acute), generally retuse, glabrous adaxially, glabrous to sparsely hairy abaxially, hairs minute (<0.2 mm long), pale with a red-brown base, appressed; primary vein channelled adaxially; secondary veins 8-11, plane to slightly sunken adaxially, ? raised abaxially, pattern eucamptodromous be coming brochidodromous, tertiary veins plane adaxially and abaxially. Panicles terminal, 10-30 cm long, hairy to very sparsely hairy at branch tips, glabrescent towards the base, hairs red-brown, short, ? appressed; bracts 1.5 mm long, with sparse, red-brown, short, appressed hairs; pedicels 1^ mm long; bracteoles 0.75 mm long, indumentum like that of bracts. Flowers 14-16 (-19) mm long. Calyx 6-8 mm long, deep purple, sparsely hairy to glabrous except on margins of lobes, hairs red-brown to golden-brown, short, ap pressed; lobes 0.5-1.7 mm long, obtuse to acute. Petals violet to purple; standard blade 10-12 mm wide, 9-12 mm high, claw 3.5-4.5 (-6) mm long; wing 7.5-10 (-12) mm long, 4-5 mm wide, claw 4-6 mm long, lamellate sculpturing present; keel 7-9 (-12) mm long, 4-5 (-6) mm wide, claw 4-6.5 (-8) mm long. Stamens 10-13 (-16) mm long, filaments united for the basal 5-9 mm, free for the distal 3-6 mm, vexillary stamen 8.5-10 (-12) mm long. Gynoecium 11.5-16 mm long, glabrous; stipe 4.5-6.5 (-7.5) mm long, ovary 3.5-5.5 (-6.5) mm long, style 3-4 mm long; ovules 4-5 (-7). Fruits 2.8-5.3 cm long, ca. 20 g or less when dry, green 2-2.8 cm high, 2-2.8 cm wide, elongated, weighing

70

SYSTEMATICBOTANYMONOGRAPHS

FIG. 23. Distribution

of Andira

VOLUME 64

humilis.

turning yellowish when ripe, long, but many fruits falling groove, less obvious below; soft when ripe, fibrous with

sweet smelling, drying wrinkled, dark brown; stipe 5-10 mm off without a stipe; suture raised adaxially with a central stylar scar tiny; mesocarp 0.5-2 mm thick, pale yellow and oily granules; endocarp 0.5-1 mm thick, brown, woody, fi

brous.

unknown.

Chromosome

number

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

Vernacular names. Mata barata, angelim h?o); manga do campo (S?o Paulo).

(Minas Gerais);

angelim

rasteiro

(Maran

ca. 100 km WSW Brazil. Specimens. Bah?a: Espig?o Mestre, Barreiras, W. R. An 26967 (K); Mpio. Morro do Chap?u, (NY); Ibiquara, ca. 25 km N Barra da Estiva, Harley Serra do Trombador, 6 km S Morro do Chap?u, Irwin et al. 32461 Feira de Santana, (MO, NY, US); Mpio. & Brito 239, 240, 241, 242 (CEPEC, FHO, K); Mpio. Angical, road Bar Campus da UEFS, R. T. Pennington Km 35, R. T. Pennington & Brito 266, 267, 268, 269 (CEPEC, FHO, K).? reiras-Ibotirama (BR-242) Representative

derson

et al. 36865

2003

71

ANDIRA

(SP); 30 km S Brasilia on road to Belo Horizonte, ca. 20 km by road N Alto Para?so, W. R. (NY).?Goi?S: Chapada Anderson 6231 (NY); Serra Geral da Paran?, 1 km E S?o Jo?o da Alian?a, W. R. Anderson 7895 (NY); Serra on road to Jatai, Irwin & Soderstrom do Caiap?, ca. 60 km S Caiaponia 7436B (NY); Serra dos Cristais, 3 km W Cristalina, Irwin et al. 9824 (NY); Chapada dos Veadeiros, Rio das Almas, 47 km from Alto Para?so de Distrito

Fazenda

Federal:

Irwin & Soderstrom

Goi?s

509.?Maranh?O:

Limpa, Heringer

Para?so

16202

dos Veadeiros,

on road to Teresina

5 km from Alto

Agua

5662

de Goi?s, de Goi?s

Mpio. Lor?to, & L. T. Eiten 5457

et al 499, 500, 501, 502, 503; Chapada dos Veadeiros, R. T. Pennington on road to the National et al. 505, 506, 507, 508, Park, R. T. Pennington 45-50 km S Lor?to on road between Santa Estev?o and Fazenda Santa Es

GROSSO: 15-120 km beyond (NY); Grajah?, Lisboa s.n. (RB).?MATO s.n. to et Anna al. road 56291 Santa das Chapadas, Malme Cuiab?, UB, (NY, US); Araguaia, Maguire do Sul: road between Aquiduana e? al 354 (E, UB); on road and Miranda, Bridgewater Grosso (F).?Mato tev?o, G. Eiten

Alto

to Porto Murtinho

from Bonito,

e? al

Gerais: 363, 364 (E, UB).?Minas e? do Cavalcan?i al s.n. Serra Cabrai, Lagoa e? al. 15968 (US); 27 km SW Fazenda Geriza, Felippe 36 (US); Mpio. Lavras, Heringer to Goveia, Irwin e? al 22146a (F,MO, NY, RB, UB, US); Morro das Pedras, ca. 40 km e? al. 25699 (MO, NY, US); between Sete Lagoas and C?rvelo, R. S. Sanios & Casiellanos Santa, B?rrelo

Serra do Cachimbo, Dombrowski

9799

5491

Bridgewaler

(F); Joaquim

road Cuiab?-Santar?m, (US); Rio

das Cinzas,

Felicio,

Km Barra

856, Prance

do Perdizes,

e? al

Haischbach

25053

Mpio.

Santa Luzia,

(SPF); Mpio. Caet?, on road Diamantina NE Patrocino,

(US).?Paran?:

24071

Irwin

(K).?Para: Jaguariahyva,

7200

(US); Mpio. Campo Mour?o, 19955 (F, NY).?PERNAMBUCO:

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

Pellado, Edwall 5650 (SP); Mpio. Moji-Gua?u, Campos das Sete Lagoas, Faz. Campin G. Eiten & L.T. Eiten 2363 (F, US); Mpio. estrada Pirassu inha, just N Rio Moji-Gua?u, Pirassununga, Forero 8278 (SP); Mpio. S?o Carlos, N of city of S?o Carlos, de Freitas Campos 31 (UB, US); nunga-Emas, 18 km N Botacatu, 14 km E S?o Manuel, road near old rail Mpio. Botacatu, along S?o Manuel-Piracicaba road station, Gottsburger et al Bicuda

Hernandes

2131

? margem do rodovia municipal, 5 km de Vitoriana, (UB); Mpio. Botacatu, rodovia Avar?-Sao /. Mattos 14530a Manoel, (SP); 37 km de Avare, (SP); Amambay: de Cerro Cora, Fern?ndez Casas & Molero Paraguay. Parque Nacional between Caaguaz? and Yh?, Fern?ndez Casas & Molero 6362 (MO); entre Yh? y 1538

Sarapui, Yano 05 (SP). 4079 (NY).?CAAGUAZ?: San Blas, Fern?ndez Casas

km N Arroyo Guaranungua, al. 1497 (MO); Dist. Lima, 734

3851 (NY); 3 km S Arroyo Taruma, Zardini & Vel?squez 25331 (K); 2 & Vel?zquez 25676 (MO).?SAN PEDRO: 4 km S de R?o Verde, P?rez et Estancia Carumbe, Pedersen 8510 (A, K); Alto Paraguay, Primavera, Woolston

& Molero Zardini

(K, U).

Andira

humilis is easy to recognize by its geoxylic suffrutex growth form, which is in the genus and was described in detail by Warming (1892). It appears that occa unique A. humilis sionally (e.g., Amorim et al. 1822, R. T. Pennington 239, 246, 247, 266) may form a shrub or tree, as occurs in other species of geoxylic suffrutices, such as Kielmey era rubriflora (Guttiferae; J. A. Ratter, pers. comm). In the case of A. humilis, this aber rant habit may be the result of the prolonged absence of fire or possibly due to hybridiza tion with shrub or tree species (see discussion under "Intraspecific cpDNA polymorphism and potential hybridization in Andira"). There is a distinct geographic partitioning of variation in the indumentum of the calyx. Specimens with glabrous calyces are restricted to western Minas Gerais, S?o Paran?, and Distrito Federal (Fig. 11), whereas specimens with sparsely are found to the northeast of these areas. I have not recognized these spec hairy calyces imens as subspecif?c taxa, because there are intermediates (particularly in Paraguay and

Paulo, Goi?s,

Distrito

Federal) and other specimens that demonstrate that the same plant can produce two inflorescences in which the flowers have calyces with widely different amounts of indumentum.

Bentham (1837) published the three names A. humilis, A. laurifolia, and A. pauciflora on the same page of the same publication. He later (1860, 1862) placed A. pauciflora in

SYSTEMATICBOTANYMONOGRAPHS

72

the synonymy of A. humilis, but maintained A. laurifolia. Handro one species, A. humilis, and therefore this name takes priority. R. T. Pennington, sp. nov.?Type: 11. Andira macrocarpa Reserva ?tnica Huaorani, Maxus road and pipeline 23 Jul 1994, N. Pitman, A. Dik & C. Aulestia 631 E! MO!).

VOLUME 64

(1969) recognized

Ecuador. construction

only

Napo: Aguarico, project, Km 98,

(holotype: QCNE!;

isotypes:

Species insignis propter foli?la parva pilis longiusculis patulis obtecta et fructus mag nos gravesque laeviusculos endocarpio lignoso fibroso. Foliis A. vermifugae & A. suri namensi similis, praecipue fructibus majoribus differt. Foliis A. galeottianae similis, prae mollibus, duris, solidis (in A. galeottiana mesocarpiis spongiosis) cipue mesocarpiis recedit.

Emergent tree more than 30 m tall, 1m in diameter; presence of buttresses unknown; bark and slash unknown; twigs dark brown, with sparse, red-brown appressed hairs when young, glabrescent, bark cracking vertically and flaking. Stipules early caducous (not seen, probably small); leaf axis 6-12 cm long; rhachis dark red-brown, hairy, hairs red brown, erect, >0.2-1.0 mm long; stipels small (ca. 0.5 mm long, but caducous, and per haps longer on younger leaves); petiolules 4-6 mm long, indumentum like that of rhachis; leaflets in 2-3 pairs, 5.5-9 cm long, 2.5^ cm wide (the lower leaflets often the smallest), narrowly obovate (rarely elliptic), subcoriaceous, base obtuse to rounded, apex rounded, often slightly retuse, glabrous adaxially, hairy abaxially, hairs long, erect, red-brown; pri mary vein channelled adaxially; secondary veins 7-10, slightly sunken adaxially, promi becoming brochidodromous, tertiary nently raised abaxially, pattern eucamptodromous veins plane to slightly impressed adaxially and raised abaxially. Inflorescences and flow ers not seen. Fruits 10-11 cm long, 7.5-8 cm high, 8 cm wide, elongated, weighing ca. 300 g when dry, flattened adaxially, drying very dark brown and slightly ridged, the sur face smooth to the touch; stipe 5-8 mm long; suture obvious, sunken adaxially, promi nently raised below; stylar remnant tiny or not apparent; mesocarp 4-18 mm thick, finely granular with small fibers, drying very hard, brown; endocarp 4-15 mm thick, brown, woody, fibrous, thickened along the sutures. Chromosome number unknown. Fig. 24. Vernacular name. Incgamuncga (Huaorani; Ecuador). is known only from the type collection gathered in rain forest in Andira macrocarpa Napo, Ecuador (Fig. 25). Although this species has never been collected in flower, its dis tinctiveness merits description. No other species have this combination of small leaflets with long, erect hairs on their undersurfaces, and a large, relatively smooth fruit with a woody, fibrous endocarp. The leaflets resemble those of A. verm?fuga and A. galeottiana in their shape, venation, and indumentum. Neither of these species is sympatric; more over, A. verm?fuga has small fruit, and the large fruit of A. galeottiana has a unique meso that is soft and somewhat spongy with numerous air gaps when dry. The carp morphology leaflets might also be confused with those of A. surinamensis, but differ in their erect in dumentum. Andira surinamensis also has a small fruit. The other sympatric species with large fruit is A. taurotesticulata, but the fruit of this species are distinctly ridged, and have an endocarp composed principally of parenchyma and stone cells, whereas the endocarp of A. macrocarpa comprises mostly woody fibers. The leaflets of A. taurotesticulata gen erally are acuminate, whereas those of A. macrocarpa are rounded or slightly retuse at the is more apex, and the venation on the undersurfaces of the leaflets of A. macrocarpa prominently

raised.

2003 ANDIRA 73

FIG. 24. Andira Fruit

in section,

A. Habit. B. Adaxial leaflet surface. C. Abaxial macrocarpa. wall structure. (Based on Pitman et al. 631.)

leaflet surface. D. Fruit. E.

showing

1994. The tree from which the type I attempted to re-collect this species inDecember, a was cut down because of concerns road and had been collection made grew alongside over must fruit about the damage that falling 500 g when fresh) might cause (which weigh to passing vehicles. surinamensis (Bondt) Splitgerber ex Amshoff, Meded. Bot. Mus. Herb. Rijks Univ. Utrecht 52: 60. 1939. Geoffroea surinamensis Bondt, Dissertatio medica surinamensis, 8, figs 1-8. 1788. Geoffrea ob inauguralis de cortice Geoffraeae 4: Bot. Mat. Med. 46. 1812, nom superfl. Andira retusa ? suri tusifolia Stokes, namensis (Bondt) DC, Prodr. 2: 476. 1825. Vouacapoua surinamensis (Bondt)

12. Andira

74

VOLUME64

SYSTEMATICBOTANYMONOGRAPHS

Kuntze,

Revis,

gen. pi. 1: 212.

1891?TYPE:

SURINAME. Bondt

s.n. (holotype:

L!). pubescens L. C. Richard, Actes Soc. Hist. Nat. Paris 1(1): 111. 1792. Ge offroea retusa Poiret in Lamarck, Encycl. 8: 182. 1808, nom. superfl. Andira re tusa (Poiret) DC, Prodr. 2(2): 475. 1825. Vouacapoua retusa (Poiret) Lyons, PL nam. 396. 1900. ?Type: French Guiana. Cayenne, L. C. Richard s.n. (lecto

Geoffroea

type, here designated: P!, isolectoype: P!). Andira retusa var. oblonga Bentham, J. Proc. Linn. Soc, Bot. 4, Suppl. ("A Synop Brazil. 121.1860.?Type: Para: Santarem, Spruce 914 sis of the Dalbergieae"): K! K! here RB!). designated: isolectotypes: (lectotype, Andira surinamensis var. ovatifoliolata N. F. Mattos, Loefgrenia 58: 3. 1973.?TYPE: Brazil.

Para: Faro, 19 Aug

1907, A. Ducke

s.n. [RB 8395]

(holotype: RB!).

Tree to 40 m tall with spreading crown in open situations; large trees buttressed; bark brown to dark brown, scaling; slash pinkish brown or reddish brown, oxidizing darker; clear, gelatinous ex?date from cuts after several days; twigs dark brown, sparsely hairy, glabrescent, hairs red-brown, appressed; lenticels not apparent. Stipules 2-3 mm long, narrow, caducous; leaf axis 10-30 cm long; rhachis sparsely hairy, glabrescent, hairs to red-brown, appressed; stipels 1-3 mm long; petiolules 3-5 mm short, golden-brown long, indumentum like that of rhachis; leaflets in 3-5 pairs, 3-15 cm long, 1.5-6 cm wide, narrowly obovate, broadly obovate, narrowly elliptic, elliptic, broadly elliptic, oblong to suborbiculate, subcoriaceous, base rounded, obtuse to acute and occasionally slightly de current

or

somewhat

cordate,

apex

retuse

to rounded,

occasionally

acute

and with

an acu

long (to 20 mm long on sapling leaflets), glabrous adaxially, sparsely hairy to red-brown; primary hairs short (<0.2 mm long), appressed, golden-brown abaxially, vein channelled adaxially; secondary veins 7-13, sunken or slightly sunken adaxially,

men

to 7 mm

raised abaxially, pattern eucamptodromous becoming brochidodromous, tertiary veins or occasion to and raised Panicles terminal plane slightly impressed adaxially abaxially. cm to at branch 11-28 with brown hairs the brown ally axillary, long, pale tips, glabres cent towards the base, hairs short, appressed; bracts 2 mm long, hairy to sparsely hairy, to red-brown; pedicels 2-4 mm long; bracteoles 1mm long, indu hairs golden-brown mentum like that of bracts. Flowers 13-18 mm long. Calyx 6-7 mm long, with red-brown hairs; lobes 0.5-1.5 mm long, acute, occasionally obtuse. Petals pink-purple to purple, the standard with a central pale marking; standard blade 10-12 mm wide, 9-12 mm high, claw 2.5-4.5 mm

long; wing 8-11 mm long, 3^4-mm wide, claw 4-5 mm long, lamellate sculpturing present; keel 7-9 mm long, 3-5 mm wide, claw 5-6 mm long. Stamens (9-) 12-13.5 (-17.5) mm long, the filaments united for the basal 7-11 mm, free for the distal 3-6.5 mm, vexillary stamen 8-12.5 mm long. Gynoecium 13-17 mm long, very sparsely hairy on the upper and/or lower surface of the ovary, hairs red-brown, erect; stipe 3-8 mm long, ovary 4.5-6 mm long, style 3-4 mm long; ovules 3-4. Fruits 4-6 cm long, 2-3.8 cm high, 2.1-4 cm wide, elongated, weighing ca. 20 g or less when dry, green, slightly glau cous, smelling very sweet, drying dark brown and distinctly wrinkled; stipe 12mm long; suture raised adaxially and below, but hard to discern because of wrinkles; stylar remnant minute; mesocarp 4-7 mm thick when fresh, thinner when dry, green, fibrous, drying with abundant air spaces; endocarp 2-8 mm thick, woody, mosome number unknown. Figs. IB, 2B, E. Phenology. Flowering year-round. Distribution (Fig. 25). Trinidad; Colombia

fibrous, brown, drying paler. Chro

(Amazonas, Meta,

Vichada),

Venezuela

75

ANDIRA

2003

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, in forest, especially along rivers and Para, Roraima, Maranh?o, Cear?, Mato Grosso); as an savanna (pers. obs. in Guyana). tree in in also isolated forest savannas, gallery arisoru (Warao), canelito negro, Vernacular names. Yatuinay (Guahibo; Colombia); blanco sob? wonka winka, (Venezuela); koraro, bat seed, (Yekuana), (Panare), palo pilon, maha, wild mango (Guyana); rode kabbes, koeraroe ibiberoe (Suriname); mangarana, manga brava, sucupira da v?rzea, angelim (Brazil); ituayuro (Peru); visguero del barbe cho, visguero (Bolivia); large jumbie bean (Trinidad). Representative serve, Gill

12621

Specimens.

Trinidad.

Toco Road,

Valencia,

N. L. Britton

et al. 1776

(NY); Arena

Re

(NY). Amazonas:

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

estaci?n medio, galer?as del Cinaruco

del R?o Orinoco y Achaguas,

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

Distritos et al. 588

Pedro (NY).?

VOLUME 64

SYSTEMATICBOTANYMONOGRAPHS

76

toward Caicura, Boom & Grillo vie. Panare, village of Corozal, 6 km from Maniapure Cede?o, of Moku-moku Mts. in NW of Kanuku Creek, A. C. Smith 3447 (COL, NY). drainage Guyana. slopes Suriname. Schulz 7321 (COL); Sectie O, Sta 433,462,463. (A, MO, U); District 8, Armai, R. T. Pennington s.n. (US). s.n. (US); Cayenne, Martin hel 57 (MEXU). French Guiana. Crique Jacques, Wachenheim

BOL?VAR: Dist. 6460

Ecuador.

Napo:

et al. 7747 (E, MO, QCNE). Peru. Cuyabeno, Laguna Cocha, Palacios a orillas del R?o Nanay, 25 km SW Iquitos, Torres 160 (MO, NY).?Madre 58003 (MO).? junction of R?os La Torre and Tambopata, Gentry & Jaramillo

Faunistica

Reserva

Prov. Maynas,

Loreto:

Nina Rimi,

DE Dios: Tambopata Reserve, von Humboldt, UCAYALl: Bosque Nacional a Puerto Pando: camino Bolivia. Cobija,

Mar?a Km 88, Gentry et al. 36390 (MO, NY). road Pucallpa-Tingo & Hartshorn 2093 (NY).?SANTA San Francisco, Meneces

Rico,

Brazil. Amap?: Rio Araguar?, El Refugio, Killeen & P?rez 6682 (K, MO). Ecol?gica Rio Negro, Santa Isabel, Ducke 145 (A)? Froes & Black 27684 (INPA).?AMAZONAS: Island of S?o Luis, Froes 11805 (A, Cear?: Fortaleza, estrada do aeroporto, Ducke 2432 (US).?Maranh?o: GROSSO: Aripua?a, Km 245 da BR-174, M. G. Silva & A. Pinheiro 4327 (F).? GH, K, NY, U, US).?MATO do Igarap? Curupira, M. Silva & R. Sousa 2284 Para: Santarem, Km 35 da estrada do Palh?o, acampamento Cruz:

Velasco,

cachoeira

Reserva

do Pared?o,

(COL, US).?Rond?NIA: Km

Manaus-Caracari,

et al. 296 F. E. Miranda Ji-Paran?, Rio Machado, Mpio. 515, bank Igarap? Dias, Steward et al. 135 (GH).

(INPA).?RORAMA:

estrada

surinamensis might be confused with A. verm?fuga, A. humilis, A. galeottiana, A. and A. galeottiana differ in their much larger and macrocarpa. Andira macrocarpa fruits and, like A. verm?fuga, by the longer, erect indumetum on the abaxial leaflet surface. Andira humilis has short, appressed hairs on its leaflet undersurfaces, like those of A. suri Andira

namensis, but it is a geoxylic suffrutex rather than a tree. Most previous workers (e.g., Mattos 1979) have cited (Bondt) Splitgerber ex Pulle; however, J. C. Lindeman Pulle (1906) made this combination in synonymy, which the ICBN (Art 34.1; Greuter et al. 2000). The correct

this species as A. surinamensis (pers. comm.) pointed out that is inadmissible under terms of attribution

is A. surinamensis

ex Amshoff.

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

13. Andira

anthelmia

(Vellozo) Kuntze, Revis, gen. pi. 1: 212. 1891.?LECTOTYPE, here designated: Vellozo's figure (Fl. flumin., icon. 7: 104. 1831). Andira stipulacea Bentham, Comm. legum. gen. 43. 1837.?TYPE: BRAZIL. Bahia: Pohl s.n. (holotype: K!). stipulacea var. bahiensis Bentham, J. Proc. Linn. Soc, Bot. 4, Suppl. ("A 119. 1860. Andira legalis var. bahiensis (Ben Synopsis of the Dalbergieae"): Brazil. Bahia: Blanchet tham) N. F. Mattos, Loefgrenia 40: 3. 1970.?Type:

Andira

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

sent; bark grey-brown

2003

77

ANDIRA

salmon pinkish, oxidizing red-brown, with clear ex?date and small amount of red ex?date; buff to yellowish, streaked; twigs often with crowded persistent stipules, stipule scars, and leaf scars, with sparse red-brown appressed hairs, glabrescent; bark pale brown to buff, Assuring vertically; lenticels few or absent. Stipules 1.5-6 cm long, generally >0.5

wood

cm wide, persistent, ovate, pale brown, with sparse, red-brown appressed hairs, glabres cent and generally glabrous at maturity; leaf axis 24-55 cm long; rhachis dark to pale brown, sparsely hairy when young, glabrescent, hairs red-brown, erect; stipels 1-9 mm 1.5^4.5 mm

long, indumentum like that of rhachis; leaflets in (3-) 4-6 4.1-12.5 (-7.5) cm wide, elliptic, nar (-7) pairs, (2.2-) (-20.5) cm long, (1-) 2.2-6.5 obovate rowly (rarely narrowly ovate, narrowly elliptic, broadly obovate to oblanceolate), thick-chartaceous (rarely chartaceous or subcoriaceous), shiny dark green adaxially, matt long; petiolules

abaxially, base obtuse or rounded, often slightly decurrent, apex obtuse or rounded (rarely acute), often slightly retuse or with a short acumen to 10mm long, glabrous adaxially ex the hairs erect, cept the groove of the primary vein very sparsely hairy, glabrescent, ? ? to to hairs red-brown to erect, very sparsely hairy abaxially, sparsely appressed mm >0.2-1.0 veins vein channelled whitish, 8-13, ? long; primary adaxially; secondary to sunken plane slightly adaxially, raised abaxially, pattern eucamptodromous becoming brochidodromous, tertiary veins plane to slightly impressed adaxially and raised abaxially. Panicles 11-35 cm long, terminal and axillary, hairy to sparsely hairy at branch tips, glabrescent towards the base, hairs red-brown, ? appressed; bracts 2-6 mm long, narrow, caducous, sparsely hairy, hairs red-brown, appressed; pedicels 2-6 mm long; bracteoles 1.5-2.5 mm

long, indumentum like that of bracts. Flowers 19-24 mm long. Calyx 8-10 mm long, purple-brown, with sparse, red-brown, appressed hairs, indumentum most dense on lobes; lobes 0.3-2 mm long, obtuse to acute. Petals rose-violet to purple; standard blade 14-16 mm wide, 13-15 mm high, claw 6-7 mm long; wing 10-13.5 mm long, 4.5-6.5 mm wide, claw 6-10 mm long, lamellate sculpturing present; keel 9-12 mm long, 5-6.5 mm wide, claw 7-10 mm long. Stamens white, 15-20 mm long, filaments united for the basal 8.5-14 mm, free for the distal 4-7 mm, vexillary stamen 12.5-16.5 mm long. 15.5-21 mm long, ovary hairy, stipe and style sparsely hairy to upper and Gynoecium lower surface of ovary hairy, stipe and style with scattered hairs, hairs red-brown, ap pressed; stipe 5-6 (-9) mm long, ovary 4.5-7 mm long, style 4.5-6 mm long; ovules 4-5. cm long, 2.5-4

cm high, 2.2-4 cm wide, elongated, weighing ca. 20 g or less when dry, strong smelling, dark brown, this surface layer thin, green beneath, drying dark brown with surface appearing smooth but minutely wrinkled (best seen with lens or mi croscope); stipe to 12mm long, but fruit usually breaking off without a stipe; suture raised Fruits 3-6.2

adaxially, ? undetectable abaxially; stylar remnant slight, but often obvious at very apex of fruit; mesocarp 1.5-3 mm thick, green to pale lime-green, drying pale brown, fibrous and granular, somewhat air-filled and soft; endocarp 1-2.5 mm thick, brown to pale brown, woody, fibrous. Chromosome number: n = 10, 11. Figs. 1A, 26. Phenology. Flowering September to December. Distribution

(Fig. 27). Brazil (Alagoas, Bahia, Espirito Santo, Rio de Janeiro); in forest and rain forest. I have also seen this species growing along river banks in restinga a in and Bahia, seasonally inundated area in Rio de Janeiro; it can clearly tolerate water seems and it logging, likely that its fruits may be secondarily dispersed by water. Vernacular Janeiro).

names. Angelim

coco,

angelim

preto

(Bahia);

angelim

coco

(Rio de

78

SYSTEMATICBOTANYMONOGRAPHS

VOLUME 64

FIG. 26. Andira

anthelmia and A. legalis. A. A. anthelmia, habit. B. A. anthelmia, abaxial leaflet surface. leaflet. D. A. legalis, abaxial leaflet surface. E. A. legalis, stipule. F. A. legalis, flower. G. A. legalis, calyx, opened to show inner surface. H. A. legalis, standard petal. I. A. legalis, outer surface of wing petal with lamellate sculpturing. J. A. legalis, keel petal, inner surface. K. A. legalis, androecium. L. A. legalis, gynoecium,

C. A. legalis,

also

shown

above

in longitudinal section. M. fruit. O. A. legalis, fruit, 281; C-F, R. T. Pennington 307; Pennington nington, 183; O, G. P Lewis & H. C. de Lima tion. N. A. anthelmia,

sec A. anthelmia, also shown above in longitudinal gynoecium, partially sectioned to show wall structure. (Based on: A-B, R. T. F-L,

A. Lima

1196.)

51-944;

M, E. Pereira

et al

4156; N, R. T. Pen

ANDIRA

2003

50

55

79

45

FIG. 27. Distribution

35

of Andira

40

anthelmia.

Examined. Brazil. Alagoas: without 27 (RB).?Bah?a: Additional Specimens locality, Uch?a Estrada Mara?-Ubaitaba, Bel?m 1879 (NY, UB); Coaraci, cocoa plantation, Bel?m & R. S. Pinheiro 2949 (US); 120 (CEPEC); Mpio. II Ilh?us, Fazenda Serra Grande, W of CEPEC cacao plantation, Cabruca, Hummel Mpio. h?us, estrada Ilh?us-Itabuna, pr?x. a CEPLAC, H. C. de Lima et al 3866 (CEPEC); Mpio. Dh?us, CEPLAC-I1 et al. 183 (CEPEC, FHO, K); Mpio. h?us road, 2 km from CEPLAC, R. T. Pennington Jussari, Fazenda Santa to Palmira, then 1 km on right hand turn, R. T. Pennington Antonio de Baixo, Km 16 on road from BR-101 205, et al. 212 (CEPEC, FHO, Km 4, R. T. Pennington 286 (CEPEC, FHO, K); Mpio. Marau, road Mara?-Ubaitaba et al 227, 238 (CEPEC, R. T. Pennington K); Mpio. Ubaitaba, banks of Rio de Contas, by road Ubaitaba-Marau, R. T. Pennington & Brito 281, 282 5 kms before CEPLAC, Ilh?us, road Ilh?us-Itabuna, K); Mpio. & F. A. de Car and road to Jussari, R. T. Pennington Jussari, Junction of BR-101 (CEPEC, FHO, K); Mpio. bank of Rio dos Frades, R. T. Pennington valho 294 (CEPEC, FHO, K); Mpio. Porto Seguro, BR-101,
360 (RB).?MINAS /. G. Kuhlmann GERAIS: Est. Exp. de Agua Limpa, Collatina, DE JANEIRO: Praia de Grumari, D. Araujo 1843 (NY); Mpio. Campos, Lagoa de (RB, UB).?R?o et al 2593 (RB); estrada dos Ban 4044 (GUA); Silva Jardim, Poco d'Anta, Carauta Cim, D. Ara?jo & Maciel da Trjuca, Machado 77 (RB); Guanabara, deirantes, Morro do Calemb?, Lanna Sobrinho 649 (US); Restinga Santo:

Gomes

Restinga

estrada do Pan?as,

2864

de Jacarepagu?, Pereira et al 291 (US).

Segadas-Vianna

et al. 4156

(NY); Goitacazes,

Lagoa

Feia, Ponta Grossa

dos Fildagos

borough,

SYSTEMATICBOTANYMONOGRAPHS

80

VOLUME 64

closely related to A. legalis (Fig. 9), which has similar The large, persistent stipules. species are clearly distinguished by their fruits: A. anthelmia has small, bat-dispersed fruits, which are never longer than 6.2 cm, whereas A. legalis has large, rodent-dispersed fruits 5.6-12 cm long. Vegetatively they may be separated by the Andira

anthelmia

is most

denser, more deeply red-brown indumentum on the stipules, leaflet undersurface, inflo rescence axis, and calyx of A. legalis. Andira anthelmia might also be confused with A. ormosioides, which has similar, large flowers, but lacks large, persistent stipules. 14. Andira

legalis (Vellozo) Toledo, Arq. Bot. Estado S?o Paulo 2(2): 29. 1946. Lumbri cidia legalis Vellozo, Fl. flumin. 306. 1829. Vouacapoua legalis (Vellozo) Kuntze, Revis, gen. pi. 1: 212. 1891.?LECTOTYPE, here designated: Vellozo's figure (Fl. flumin., icon. 7: 105. 1831.) Andira frondosa Martius ex Bentham, Comm. legum. gen. 44. 1837. Vouacapoua (Martius ex Bentham) Kuntze, Revis, gen. pi. 1: 212. 1891.?TYPE: frondosa Brazil. Brasiliae

Rio de Janeiro: "In arenosis

sylvarum ad Capo Frio et alibi circa oram orientalis maritimam," Martius s.n. (holotype: BR!; isotype: M).

Tree to 20 m tall, simply branched; buttresses absent; bark pale brown to grey-brown, rough, Assuring vertically; trunks of saplings marked with leaf and stipule scars, stipules often persistent; slash buff to pale red-brown, oxidizing darker red-brown, a little red ex ?date; wood buff to yellowish buff; twigs swollen, with crowded persistent stipules, leaf scars and stipule scars, with red-brown, erect hairs, glabrescent; lenticels not apparent. Stipules 1.5-4.5 cm long, >0.5 cm wide, persistent, ovate, with deep red-brown, appressed leaf axis 15-30 (-60) cm long; rhachis with erect, red-brown hairs, glabrescent; stipels to 9 mm long; petiolules 3-7 mm long, indumentum like that of in 4-7 (-9) pairs, 5.5-15 (-30) cm long, 2.3-7 (-11) cm wide, elliptic, leaflets rhachis; narrowly obovate, ovate (rarely narrowly elliptic to broadly elliptic), coriaceous, base ob tuse, rounded, ? truncate or very slightly cordate, apex obtuse or rounded, often slightly retuse or with an acumen to 7 mm long, glabrous adaxially except scattered hairs in the groove of the primary vein, hairy to sparsely hairy abaxially, indumentum most dense on veins, hairs red-brown, erect, >0.2-1.0 mm long; primary vein channelled adaxially; sec

hairs, glabrescent;

ondary veins 8-13, slightly sunken adaxially, raised abaxially, pattern eucamptodromous 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-) 15mm high, claw 5-7 mm long; wing 13-14 mm long, (3.7-) 6-7.2 mm wide, claw 6.5-8 mm long, lamellate sculpturing present; keel (8.2-) 11-13 mm long, (4-) 5-6 mm wide, claw 6-8.4 mm long. Stamens (12.5-) 15 mm long, filaments united for the basal 8-11 mm, free for the distal 3-6.4 mm, vexillary stamen 12.5-15 mm long. (14.5-) 17.2-18 mm long, ovary sparsely hairy, stipe and style very sparsely Gynoecium to upper (basal half only) and lower surface of ovary hairy with a few hairs extend hairy down the stipe, hairs red-brown, ? appressed; stipe 5-6 mm long, ovary (5.5-) 7 mm ing long, style (4-) 5.2-6 mm long; ovules 2-A. Fruits 5.6-12 cm long, 4.5-8.7 cm high, 4.5-9 cm wide, elongated, weighing 100-300 g when dry, rough, pale brown speckled

2003

81

ANDIRA

50

55

FIG. 28. Distribution

35

45

of Andira

40

legalis.

(both fresh and dry); stipe to 5 mm long, sutures not obvious, sunken adaxi ally, slightly raised abaxially; stylar remnant not obvious; mesocarp 4-10 mm thick, pale brown, hard, finely granular; endocarp 3-18 mm thick, brown, woody, fibrous. Chromo some number unknown. Figs. ID, 26. Phenology. Flowering August to November. dark brown

Distribution (Fig. 28). Brazil (Bahia, Espirito Santo, Minas Gerais, Pernambuco, Rio de Janeiro); in restinga and rain forest. Vernacular names. Angelim preto, angelim roxo (Bahia); angelim coco (Rio de Janeiro). Additional Specimens Examined. Brazil. Bah?a: Mara?, Bel?m 1830 (CEPEC, MEXU, NY, UB, US); 7.3 km na estrada Serra Grande-Itacar?, Fazenda Lagoa do conjunto Mpio. Uru?uca, Dist. Serra Grande, et al 3508, 3509 (CEPEC, NY); Mpio. Porto Seguro, junction on right Fazenda Santa Cruz, A. M. de Carvalho at Km 16 on road Povoada Santa Cruz, 1 km up this road, Mattos Silva et al. 340 (CEPEC, K); Area Controle da Caraiba Metais, Noblick 2299 (K, UEFS); Mpio. Porto Seguro, junction on right at Km 16 on road Povoada 1 km up this road, R. T. Pennington Santa Cruz, & Lewis 191 (CEPEC, FHO, K); Mpio. Mara?, road et al 214, 215 (CEPEC, FHO, K); Mpio. Alcoba?a, Km 6, R. T. Pennington Mara?-Ubaitaba 1 km N do centro

SYSTEMATICBOTANYMONOGRAPHS

82

VOLUME 64

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

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

Branca, D. Araujo

al. 8597

(GUA); Mpio.

et al. 8326

Saguarema,

de Marica, pr?x. a Lagoa de Marica, Cabo, L. B. Smith 6554 (US).

Lewis

& H. C de Lima

1196 (K); Mpio.

Cabo Fri?, Cabo Fr?o, Arraial

do

Andira legalis is mostly closely related to A. anthelmia and A. ormosioides. For ac counts of how to distinguish these species, see the notes under A. anthelmia (no. 13) and A. ormosioides (no. 15). The collection Duarte 4263 from Minas Gerais listed under A. ormosioides may be A. legalis (see discussion of A. ormosioides). legum. gen. 44. 1837. Andira anthelmia var. J. Proc. Linn. Soc, Bot. 4, Suppl. ("A Synop (Bentham) Bentham, Brazil. 120. I860.?Type: sis of the Dalbergieae"): Tingua, Schott s.n. (holo type: K!; isotype: F!).

15. Andira

ormosioides

Bentham,

Comm.

ormosioides

Tree (though can flower when very small) to 30 m tall, long-boled with small crown (even in open situations), with small buttresses; bark pale brown to grey-brown, Assuring vertically and flaking slightly; slash pale red-brown; wood buff/cream; twigs brown, with erect, red-brown hairs, becoming paler, glabrescent. Stipules to 16 mm long, to 1 mm wide, moderately persistent, with red-brown, ? appressed hairs; long; rhachis with red-brown erect hairs, glabrescent; stipels 2-3 2-5 mm long, indumentum like that of rhachis; leaflets in (3-) on sterile shoots) cm long, 1.8-7 cm wide, elliptic to narrowly

leaf axis 9-30 (-35) cm (-6) mm long; petiolules 4-5 pairs, 4.7-15.5 (-19 obovate, subcoriaceous, shiny, dark green adaxially, base obtuse to rounded, often very slightly decurrent, apex ob tuse to rounded, occasionally with an acumen to 7 mm long, glabrous adaxially except scattered hairs in the groove of the primary vein, hairy to sparsely hairy abaxially, indu dense on veins, hairs red-brown, erect, >0.2-1.0 mm long; primary vein adaxially; secondary veins 8-11, slightly sunken adaxially, raised abaxially,

mentum most channelled

pattern eucamptodromous becoming brochidodromous, tertiary veins plane to slightly im cm and raised Panicles 10-30 pressed adaxially long, terminal (more rarely ax abaxially. mm erect with bracts 3-7 red-brown, hairs; illary), long, with red-brown, ? appressed mm mm 3-7 bracteoles 1-2.5 hairs; pedicels long; long, indumentum like that of bracts. 18-23 mm long. Calyx 9-10 mm long, with red-brown, ? appressed to ? erect lobes 0.8-2 mm long, obtuse to acute. Petals pink to purple; standard blade 17-20 hairs; mm wide, 15-19 mm high, claw 6-7 mm long; wing 12-15 mm long, 5-6.5 mm wide, claw 8-9 mm long, lamellate sculpturing present; keel 10-12 mm long, 4-7 mm wide, claw 8-10 mm long. Stamens 15-20 mm long, filaments united for the basal 8.5-11 mm, Flowers

free for the distal 5-10 mm, vexillary stamen 11.5-15 mm long. Gynoecium 18.5-20 mm long, ovary sparsely hairy, stipe and style sparsely hairy, hairs red-brown, ? appressed; stipe 6-7 mm long, ovary 7 mm long, style 4.5-6 mm long; ovules 5-8. Fruit 4.7-5.6 cm long, 3.4-4.1 cm high, 3.4-4.1 cm wide, elongated, weighing ca. 20 g or less when dry,

83

ANDIRA

2003

50

FIG. 29. Distribution mosioides

(see discussion);

of Andira

45

55

ormosioides.

if so, the locality

35

The collection

is a considerable

40

A. P. Duarte

range extension

4263

probably for the species.

represents A. or

smooth, very dark brown with green beneath when fresh, drying very dark brown, ap tuberculate (best seen with lens or microscope); pearing smooth but minutely stipe 6-8 mm long; suture slightly raised adaxially, flanked by two shallow grooves, very slightly raised abaxially; stylar remnant raised at apex of fruit; mesocarp 3^1 mm thick, pale mm 2-3 thick, dark brown, woody, greenish white, drying pale brown, granular; endocarp fibrous. Chromosome

number unknown. Fig. 3H. Phenology. Flowering June to December. Distribution (Fig. 29). Brazil (Bahia, Espirito Santo, Minas Gerais, Rio de Janeiro, S?o Paulo); in restinga, rain forest, and semi-deciduous forest. Vernacular names. Angelim pedra (Espirito Santo); angelim tento, angelim rosa, an gelim amargoso (Rio de Janeiro); Jacaranda rajada (S?o Paulo). Additional

& F. A. de Carvalho 164/79

(RB).?Minas

Specimens 306, 309 Gerais:

Examined.

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

SYSTEMATICBOTANYMONOGRAPHS

84

VOLUME 64

de Fora, Heringer 2700 (RB); Carangola, Rio Carangola, Leoni & Leoni 856 (K); 400 km E Belo Horizonte, Caralinga, Fazenda Montes Claros, Lopes & vicinity of Rio Manhuac?, Valley of Rio Doce, Estac?o Biol?gica DE Janeiro: Serra da Estrela, de Almeida s.n. (RB); Barreira de Soberbo, Dias s.n. Andrade 850 (K, RB).?Rio s.n. (RB 81810) (RB); s.n. (RB 54881) (RB); Rio de Janeiro, Horto Florestal (R); Rio de Janeiro, Horto Florestal 528 (F, NY, RB, UB); Mpio. Mage, Cachoeiras 29 (RB); Guanabara, /. G. Kuhlmann Avellar, /. G. Kuhlmann de Macacu, Para?so, near CPRJ, banks of Rio Para?so, H. C. de Lima et al. 4253 (RB); estrada do Recreo dos s.n. (RB 76106), Machado s.n. (RB 75296) da Trjuca, Machado Lutz 1493 (R); Restinga Bandeirantes, (RB); 10503 (RB); Mpio. Mag?, & Pessoa do RJ, Martinelli (IIIo), Dist. Para?so, Centro de Primatologia Mag? s.n. (R); Ubatuba, PAULO: Aeto da Serra, de Andrade Para?so, Sampaio 6 (RB); Gavea, E. Ule LVII (R).?SAO da Cruz 13 (NY, SP). pr?ximo a Base Norte do Instituto Ocean?grafico,

Mpio.

is Field observation in Bahia of growth form and fruit indicates that A. ormosioides distinct from the three species to which it is most similar: A. fraxinifolia, A. legalis, and A. anthelmia. The fruit of A. ormosioides are small (4.7-5.6 cm long, vs 5.6-12 cm long in A. legalis) and have a pale greenish white mesocarp (in comparison to the green meso and A. anthelmia). Andira ormosioides has a long bole and small carp of A. fraxinifolia and A. anthelmia have a short bole crown, even in open situations, whereas A. fraxinifolia and a broad, spreading crown. Herbarium specimens of A. ormosioides can be difficult to separate from large-flow ered specimens o? A. fraxinifolia and particularly from specimens of A. legalis and A. an thelmia that have lost their characteristic large stipules or were collected without stipules. An example is A. P. Duarte 4263, which has the dense red-brown indumentum on the leaflet undersurface, inflorescence axis, and calyx characteristic of both A. ormosioides and A. legalis. In the absence of fruit or stipules, it is not possible to determine this spec imen with certainty. This specimen was collected in Patos de Minas, western Minas Gerais, ca. 600 km from the Atlantic coastal forests, where both A. ormosioides and A. le galis grow. Forests in Patos de Minas are semideciduous, with a few patches of dry de ciduous

forests

on

calcareous

outcrops

(A. T. Oliveira-Filho,

pers.

comm.).

Andira

ormo

forests of the Rio Doce valley of eastern Minas in the semi-deciduous are to the rain forests, whereas A. legalis grows strictly which coastal transitional Gerais, in rain forests and restingas. The Duarte specimen from the dry area of Patos de Minas is sioides

more

occurs

likely to be A. ormosioides

and is therefore included here.

fraxinifolia Bentham, Comm. legum. gen. 44. 1837. Vouacapoua fraxinifolia Brazil. Minas Gerais: (Bentham) Kuntze, Revis, gen. pi. 1: 212. 1891.?Type: prope Itamb?, Pohl s.n. (lectotype, here designated: K!; isolectotype: K!). Andira parvifolia Martius ex Bentham, Comm. legum. gen. 44. 1837. Vouacapoua (Martius ex Bentham) Kuntze, Revis, gen. pi. 1: 212. 1891.?TYPE: parvifolia Brazil. Minas Gerais: "in campis altis Serro Frio," Martius s.n. (holotype: M!).

16. Andira

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

Andira micans

1906.?Type:

Taubert

Brazil.

ANDIRA

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85

BRAZIL. Bahia: be Loefgrenia 40: 1. 1970.?Type: tween Len?ois and Itaberaba, 15 Sep 1956, E. Pereira 2064 (holotype: R!; iso type: RB!). var. latifoliolata N. F. Mattos, Loefgrenia 40: 2. 1970.?TYPE: Andira fraxinifolia s.n. 13 Aug 1949, O. Machado Brazil. Rio de Janeiro: Restinga da Gavea, (holotype: RB!). Brazil. Andira anthelmia var. gracilis N. F. Mattos, Loefgrenia 40: 3. 1970.?Type: S?o Paulo: Limeira, Navarro de Andrade 167 (holotype: R; isotype: RB!). Brazil. Bahia: Km Andira bahiensis N. F. Mattos, Loefgrenia 45: 2. 1970.?Type:

Andira handroana N. F. Mattos,

80 between Betanha

and Canavieiras,

13 Jul 1964, N. T. Silva 58414

(holotype: UB; isotype: CEPEC!). N. F. Mattos, Loefgrenia 53: 1. 1970.?TYPE: BRAZIL. Per Andira pernambucensis nambuco: Rio Formoso, Engenho S?o Manoel, 3 Sep 1954, J. Falc?o, Engler & E. Pereira 944 (holotype: RB!; isotypes: RB-2 sheets!). var. lanceata N. F. Mattos, Loefgrenia 58: 3. 1973. Brazil. S?o Andira fraxinifolia 24 Aug 1963, J. Mattos & N. F. Mattos 8383 (holo type: SP!). 58: 2. 1973.?Type: Andira pisonis var. emarginata N. F. Mattos, Loefgrenia Brazil. Bahia: Igua?u, 30 Dec 1922, P. Campos Porto s.n. (lectotype, here des Paulo: 2 km N of Atibaia,

ignated: RB!). Brazil. Andira pisonis war. puberula N. F. Mattos, Loefgrenia 58: 2. 1973.?Type: /. G. Kuhlmann do Rio Santo: 1934, Dur?o, Doce, Linhares, Apr Lagoa Espirito 163 (holotype: RB!). Shrub or small tree to 12 m tall, with broad spreading crown in open situations; but tresses absent; bark grey-brown, fissuring vertically and flaking (larger trees); slash pale brown to pale red-brown, oxidizing darker, occasionally with slight red ex?date; wood to dark brown, older twigs paler, often buff/whitish, hairy to sparsely hairy, glabrescent, hairs erect. Stipules 2-9 mm long, to 1mm wide, caducous, with sparse, red-brown ? appressed hairs; leaf axis 6-21.5 (-25) cm long; rhachis sparsely hairy (occasionally more densely hairy), glabrescent, hairs red-brown, erect; stipels 1-5 mm long; petiolules 2-3 (-5) mm long, indumentum like that of rhachis; leaflets in (2-) 3-7 pairs, 2-12 cm long, 0.7^1.2 cm wide, elliptic, narrowly elliptic, narrowly obovate, cream/buff;

twigs brown

to subcoria oblanceolate (more rarely lanceolate to broadly obovate), thick-chartaceous ceous (rarely chartaceous), dark green, shiny adaxially, matt abaxially, base obtuse to rounded (rarely acute), often very slightly decurrent, apex acute, obtuse to rounded, gener ally with an acumen to 7 mm long, glabrous adaxially except scattered hairs in the groove of the primary vein, hairy to very sparsely hairy abaxially, indumentum most dense on .0mm long; primary vein channelled adaxially; sunken adaxially, raised abaxially, pattern eu camptodromous becoming brochidodromous, tertiary veins plane adaxially and slighty raised or raised abaxially. Panicles 4-30 cm long, terminal and axillary, with red-brown, erect hairs, glabrescent towards the base; bracts 2-3.5 mm long, with appressed, red-brown hairs; pedicels 2.5-4 mm long; bracteoles 1.5-2 mm long, indumentum like that of bracts. veins, hairs pale to red-brown, erect, >0.2-l secondary veins 6-10, ? plane to slightly

Flowers 13-17 mm long. Calyx 6-7 mm long, brown to purplish, hairy to sparsely hairy, hairs ? appressed, most dense on lobes; lobes 0.25-1.75 mm long, obtuse to acute. Petals pink to purple, the standard with a white central marking; standard blade 10-14 mm wide, 10-13 mm high, claw 4-5 mm long; wing 8.5-11.5 mm long, 5-7 mm wide, claw 5-7 mm

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

Phenology. Flowering year-round. Distribution (Fig. 31). Brazil (Alagoas, Bahia, Cear?, Distrito Federal, Espirito Santo, Goi?s, Minas Gerais, Paran?, Pernambuco, Rio de Janeiro, Santa Catarina, S?o Paulo); in restinga, rain forest, campo rupestre, and secondary vegetation, often seen isolated in pas tures.

Vernacular names. Angelim branco, angelim preto (Bahia); angelim coco (Bahia, Es pirito Santo); angelim amargoso (Minas Gerais); angelim pedra (Minas Gerais, Rio de Janeiro); angelim da folha grande, pau de morcego, quaiseara, Jacaranda de morcego, fruta de c?valo (S?o Paulo); pau angelim (Santa Catarina). e? al Specimens. Brazil. Alagoas: Fazenda S?o Louren?o, Andrade-Lima Fleixeiras, Mpio. Sa?de, Cachoeira do Paulista, 6 km along road starting at Km 10 of road Sa?de-Jacobina, 1055 (CEPEC, E); Serra Jacobina, Blanchet 2723 (E, BM, F, K, NY); Mpio. Jacobina, Serra do na estrada para Morro do Chap?u, A. M. de Carvalho et al 6141 Tombador, ca. 8 km SW da sede do municipio, /. G. (E, NY); Mpio. Saude, Cachoeira do Paulista, 6 km along road starting at Km 10 of road Sa?de-Jacobina, Jardim 71 (CEPEC, FHO); Mpio. 100 m up hill from Club Tomoromba, R. T. Pennington 194 Ilh?us, Oliven?a, Representative

7 (F).?Bah?a: Amorim et al

road to Pontal, junction on left, about 300 m up this Ilh?us, ca. 3 km N Oliven?a, (CEPEC, FHO, K); Mpio. 200 (CEPEC, FHO, K); Mpio. track, R. T. Pennington Ilh?us, Agua da Oliven?a, R. T. Pennington 201, 202 & A. M. de Carvalho 204, 228 (CEPEC, (CEPEC, FHO, K); Mpio. Jussari, Fazenda Teimoso, R. T. Pennington to Palmiras, ca. Km 15, R. T. Pennington & A. M. de Carvalho 207 FHO, K); Mpio. Jussari, road from BR-101 et al. 211, 213 (CEPEC, FHO, road Mara?-Ubaitaba Km 5, R. T. Pennington (CEPEC, FHO, K); Mpio. Mara?, Km 4 (off road to Pedras from main road Oliven?a-Una), R. T. Pen Una, road to Independencia Jacobina, Km 20 along road Jacobina toMorro do Chap?u nington e? al 234, 236 (CEPEC, FHO, K); Mpio. & Brito 248,249,250; 18 km E Morro do Chap?u on BR (BR-324), R. T. Penningion Mpio. Morro do Chap?u,

K); Mpio.

& Brito 251 (CEPEC, FHO, K); Mpio. Len?ois, 5 km from Len?ois on road to BR-242, 052, R. T. Penningion ca. 2 km N Alcoba?a, R. T. Pennington & Brito 274, 275, 276, 278, 279; Mpio. Alcoba?a, R. T. Pennington & F. A. de Carvalho 302 (CEPEC, FHO, K); Mpio. Queiroz et al. Itua?u, arredores do Morro da Mangabeiras, 1610 (K, NY).?Cear?: Federal: Santana, Poranga, S. J. Filho 63 (RADAM).?Distrito Campus of Uni 10064 (UB); Fazenda Agua Limpa, near Vargem Bonita, ca. 18 km SSW Brasilia versity of Brasilia, Heringer TV Tower, Ratter 163

(NY); Mpio.

et al 3791 Linhares,

SANTO: Lagoa do Dur?o, Linha Res., Rio Doce, J. G. Kuhlmann da CVRD, H. C de Lima 1711 (RB).?GOI?S: Mpio. Santo An Cabteiro do Para?so, Elias de Paula 3147 (UB).?MINAS GERAIS: San

(UB).?ESPIRITO Reserva Florestal

tonio Descoberto, Corrego da Fazenda tana do Riacho, Serra do Cipo, pr?x. do Rio Cipo, de Barros 1356 (SP); Mpio. Jaboticatubas, Km 123 ao longo de rodovia Lagoa Santo-Concei?ao do Mato Dentro, Joly & Samir s.n. (SP); Vi?osa, Mexia 4932 (A, F, GH, U, de Mata Verde, estrada para Lamban, E. Pereira 5788 (RB); Rio Vermelho, Pedra Menina, US); Concei?ao Guaratuba, Vargem da Ang?lica, Morro da Virada do Mato Virgem, Pirani et al s.n. (SP).?Paran?: da entrada da Baia, Braga 2364 (RB); Itacar?, opp. Marungaia, Dus?n 16511 (A); Mpio. Trjucas do Sol, 15117 (NY, US); Mpio. Antonica, Rib. Lagoa Vermelha Haischbach 18102 (NY, US); Matul?o, Haischbach Fazenda

Morro

Mpio.

Morretes,

(NY).?Pernambuco: do Cabo, Graziela

Col.

Florestal, Recife,

Hatschbach Mata

20206 27144 (RB); Mpio. Senges, Rio do Funil, Hatschbach de Janeiro: Cabo Fri?, Arraial Irm?os, Lima 48-126 (RB).?Rio Reserva Biol?gica de Jacarepagu?, Lanna 124 (RB); Mpio. Maca?,

de Dois

7 (NY); Guanabara,

ANDIRA

2003

87

leaflet surface. D. A. Habit. B. Node of rhachis with stipels. C. Abaxial 30. Andira fraxinifolia. E. Calyx, opened to show inner surface. F. Standard petal, inner surface. G. Outer surface of wing petal K. Gy with lamellate sculpturing. H. Detail of lamellate sculpturing. I. Keel petal, inner surface. J. Androecium. section. L. Fruit. M. Fruit in section, showing wall structure. O. Seed. noecium, also shown above in longitudinal FIG.

Flower.

(Based on: A-K,

R. T. Pennington

228; L-M,

R. T. Pennington

202)

88

SYSTEMATICBOTANYMONOGRAPHS

60757065

FIG. 31. Distribution

55

VOLUME 64

403550 45

o? Andira fraxinifolia.

Restinga da Praia de Carapebus, H. C. de Lima 656 (RB); Mpio. Nova Igua?a, pr?x. a Lixeira, H. C. de Lima N Joinvile, J. Mattos 1173 (RB); Mpio. Parati, Parati-Mirim, H. C. de Lima 3673 (RB).?Santa Catarina: 12530 (SP); Campo Maseaiamb?, Palho?a, Reitz & Burkart 5617 (US); Mpio. Itaja?, Itaja?, Praia Braba, L. B. Smith & Klein 7288 (NY, US); Brusque, Mata do Hoffman, Veloso 98 (RB).?SAO PAULO: Canan?ia, Una do s.n. (SPF); near Moji-Mirim, M. Cardoso, Morro do Pereirinha, de Barros 2238 (SP); S?o Vicente, W. Hoehne 1477 (NY); Juqui?, Capueira, na margem do Rio Juqui?, M. Kuhlmann 4694 (SP); Ubatuba, Praia de Kuhlmann J. Mattos 13802 (SP); Mpio. de Jureia, Pirani 818 (SPF); Mpio. Perequer?-Ass?, Iguape, esta?ao ecol?gica Jundinhy, C. Smith 14 (SP); Sao Paulo, Instituto de Bot?nica, Sugiyama 888, 889 (SP).

Andira fraxinifolia is a widespread and variable species, which is most likely to be confused with A. anthelmia, A. ormosioides, and A. legalis in the rain forests and restinga forests of Atlantic coastal Brazil, and with A. verm?fuga, in the cerrado woodlands of Cen tral Brazil. Andira ormosioides, A. legalis, and A. anthelmia have flowers 18-24 mm long, are 13-17 mm long. Andira fraxinifolia whereas those o? A. fraxinifolia also lacks the large, persistent stipules that characterize A. legalis and A. anthelmia. The gynoecium of A. verm?fuga is either glabrous or at most sparsely hairy on the upper and lower surfaces it is uniformly hairy. The fruits of A. verm?fuga dry with only, whereas in A. fraxinifolia a wrinkled surface, whereas inA. fraxinifolia they dry smooth. Furthermore, A. fraxinifo lia does not grow in the cerrado woodlands, which is the main habitat for A. verm?fuga. In the cerrado biome, A. fraxinifolia

is only occasionally

found in gallery forests.

2003

ANDIRA

89

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

used.

ex Bentham, Comm. legum. gen. 45. 1837.?Type: s.n. (holotype: BR!; Rio Santo: Doce, 1827, Wied-Neuwied Espirito BM! K! M, NY!; photo of M isotype: F!).

17. Andira

n?tida Martius

Brazil. isotypes:

shrub to tree 25 m tall; buttresses absent or very small; bark brown, and Assuring vertically scaling (large trees), or pale brown, often marked with patches of lichen (shrubs and small trees in restinga); slash pale red-brown to buff, oxidizing darker, often with slight red ex?date; wood buff/cream; twigs red-brown to dark brown, or cov Multi-stemmed

ered with pale buff bark (which may be scraped away to reveal dark bark beneath), hairy, hairs red-brown, appressed, or glabrous; lenticels occasionally numerous, elongated, pale. Stipules to 10mm long, to 1mm wide, caducous, hairy to sparsely hairy, glabrescent; leaf axis 4-15 (-25) cm long; rhachis pale brown to brown, the pulvinus often darker, sparsely hairy or glabrous, hairs short, red- brown, ? appressed; stipels 1 (-2) mm long, caducous; (-5) pairs, 2.4-11.5 (-14) cm long, 1.7-4.5 (-6) cm wide, elliptic, broadly ovate ovate, elliptic, narrowly (rarely narrowly elliptic, suborbiculate to broadly ovate), coriaceous to thick-chartaceous, base obtuse, rounded to slightly cordate, often slightly decurrent, apex acute, obtuse, rounded, often slightly retuse or with a short acumen to 10 mm long, glabrous adaxially, very sparsely hairy abaxially or glabrous, hairs red-brown, short (<0.2 mm long), appressed; primary vein channelled adaxially; secondary veins 6-13, ? plane to slightly sunken adaxially, ? plane to slightly raised abaxially, pattern eu or ? completely brochidodromous, camptodromous becoming brochidodromous, tertiary leaflets in 2-4

veins plane adaxially and abaxially. Panicles 6-30 cm long, terminal and axillary, with sparse to very sparse, short, appressed, red-brown hairs at branch tips, becoming less hairy towards the base; bracts 1.5-2 mm long, narrow, caducous, sparsely to very sparsely hairy, hairs red-brown, ? appressed; pedicels to 2 mm long or flowers sessile; bracteoles 0.5-1 mm long, indumentum like that of bracts. Flowers 10-13 mm long. Calyx 4-5.5 mm long, red-brown to reddish to ? black, with sparse red-brown, appressed hairs or glabrous ex cept around the margins of the lobes; lobes 0.2-0.5 mm, obtuse. Petals pinkish white to purple, the standard pale with violet markings or pinkish with a central white marking; standard blade 8-8.5 mm wide, 7-8 mm high, claw 3-3.5 mm long; wing 5.6-8 mm long, 3-3.5 mm wide, claw 3-4 mm long, lamellate sculpturing present; keel 5-7.5 mm long, 2.6-3.5 mm wide, claw 3-4.5 mm long. Stamens 8.5-10 mm long, united for the basal 4.7-8 mm, free for the distal 2-4.5 mm, vexillary stamen 6-7.5 mm long. Gynoecium

SYSTEMATICBOTANYMONOGRAPHS

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VOLUME 64

9-13 mm

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

dry, pale green to green, fibrous, sweet-smelling, drying brown, hard, slightly granular; endocarp 2-5.5 mm thick, brown to dark brown, drying paler, fibrous. Chromosome number unknown. Figs. 3C, 4A(iii), 5B, 13B, 32. Phenology. Flowering September to January (toMay in Espirito Santo). thick when

Distribution

(Fig. 33). Brazil (Bahia, Espirito Santo, Pernambuco, Rio de Janeiro); in on white sand, closed restinga forest, and rain forest. scrub open restinga Vernacular names. Angelim branco, angelim da praia (Bahia); angelim coco (Espir ito Santo); angelim (Pernambuco). REPRESENTATIVE Specimens.

Brazil. BAHIA: Santa Cruz da Cabr?lia, Almeida & dos Santos 103 (CEPEC, Sa?da de Itaju do Colonia 356 (US); Mpio. Uru?uca, Dist. Serra Grande, para Itap?, Km 20, Almeida estrada Serra Grande-Ilh?us, 3 km from Distrito, Amorim et al. 347, 348 (CEPEC); Itacar?, Bel?m & R. S. Pin heiro 2988 (MEXU, UB, US); Mpio. et al. Ilh?us, ca. 7 km along road Oliven?a-Vila Brasil, A. M. de Carvalho

US);

3309 (CEPEC); Mpio. Porto Seguro, Esta??o Ecol?gica do Pau Brasil, Euponino 167 (US); Mpio. Nova Vi?osa, Km 9 along road Nova Vi?osa-Mucuri, Farney et al. 2615 (RB); Mpio. Mucuri, Rio Mucuri, Farney et al. 2653 on road to 16 km S Cumuruxatiba Salvador, pr?ximo ao aeroporto, M. L. Guedes 972 (RADAM); (RB); Mpio. et al. 18086 (K); Mpio. Prado, Harley Ilh?us, grounds of CEPLAC, G. P. Lewis & A. M. Prado, 12 km S Prado, estrada para Alcoba?a, Copuva,

G. Hatschbach

Ubaitaba,

G. P. Lewis

& J. M.

756 791

(CEPEC, E, NY); Mpio. Nova Vi?osa,

(K); Mpio.

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

Silva

J. G. Jardim de Carvalho

48748

de Carvalho

ca. 15 km S Itabuna on road BR-101, R. T. Pennington et al. 209 (CEPEC, FHO, Buerarema, road Porto de Campinhos-Ubaitaba, et al. 221 (CEPEC, FHO, K); Km 9, R. T. Pennington Marau, et al. 235 (CEPEC, FHO, K); Mpio. R. T. Pennington Mpio. Una, near Pedras, Km 5 on road to Independencia, et al. 288, 290, 291, 292 (CEPEC, FHO, K); Mpio. Alcoba?a, Km Salvador, dunas de Itapua, R. T. Pennington FHO, K); Mpio.

K); Mpio.

4 on road Alcoba?a-Teixeira de Freitas, R. T. Pennington & F. A. de Carvalho 300 (CEPEC, FHO, K); Mpio. ca. 2 km N Alcoba?a, 1 km from the sea, R. T. Pennington & F. A. de Carvalho 301 (CEPEC, FHO, Alcoba?a, K); Mpio. Mata de S?o Jo?o, estrada do Coco, em dire?ao a Sau?pe, G.C.P. Pinto 429/81 (RADAM); Mpio. 14 km N Serra Grande, off road to Itacar?, W. W. Thomas et al 9469 (CEPEC); Itacar?, "Campo Cheiroso," Barrol?ndia, Est. Experimental Mpio. Belmonte, "Gregorio Bondar," 48 km E BR-101 on road to Belmonte, W. W. Thomas et al. 9881 (CEPEC).?ESPIRITO SANTO: Mpio. Vila Velha, restinga de Lagoa do Milho, D. Araujo & Peixoto 324 (RB); Itaunas, G. P. Lewis et al. 1636 (K); Mpio. Linhares, Vale do Rio Doce, Reserva Florestal da Cia., H. C. de Lima 1668 (RB); Mea?pe, estrada Rodovia do Sol, 10 km depois de Guarapari, H. C. de Lima 2922 (K, MEXU); Mpio. Concei?ao do Barra, Ita?nas, ap?s a ponte para a cidade velha, H. C. de Lima 2966 (MEXU, RB); Mpio. Linhares, Reserva Florestal de Linhares, Sucre 8656 (K, NY).?Pernambuco: Recife, Mata

de Dois

limaos, Andrade

Lima 55-1988 (K); Cabo, ?rea-projecto Suape, Mata do Zumbi, Andrade-Lima 105 (F); Ilha Itamaraca, G. A. Ramage s.n. (BM).?Rio DE JANEIRO:Mpio. Quissam?, Mata do Caio, Farney et al. 3425 (E, RB).?SERGIPE: Mpio. Santo Amaro de Brotas, 8 km after "Terminal Portuario" of Aracaju, Farney 2880 (RB); Mpio. Santo Amaro de Brotas, 5 km after "Terminal Portuario" of Aracaju, Far

& Medeiros-Costa

ney 2930 Gomes

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

&

2003

ANDIRA 91

leaflet surface. C. Flower. D. Calyx, opened to show inner surface of wing petal with lamellate sculpturing and detail of sec I. Gynoecium, also shown below in longitudinal sculpturing. G. Keel petal, inner surface. H. Androecium. dried fruit. L. Fruit in section, showing wall structure. M. Seed. (Based on: A, tion. J. Fresh fruit. K. Wrinkled 288 (leaves), H C. de Lima 1668 (inflorescence); B, R. T. Pennington C-I, A. Ribeiro 62; J, L, R. T. Pennington 292; K, R. T. Pennington 291.) FIG.

32. Andira

surface. E. Standard

nitida. A. Habit.

petal,

B. Abaxial

inner surface. F. Outer

92

SYSTEMATICBOTANYMONOGRAPHS

50

FIG.

55

45

33. Distribution

35

o? Andira

VOLUME64

40

nitida.

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

is ridged from the lower to the upper suture and dries smooth. Imade

ANDIRA

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93

(mostly sterile) of both growth forms in Bahia and found them I initially considered them as possibly distinct species (A. n?tida distinct morphologically. and "A. sp. nov. 3" in Pennington 1995, 1996), and they were coded as separate terminal taxa in cladistic analyses. Subsequent examination of more herbarium specimens showed extensive

collections

that differences between the putative taxa are not consistent, especially at the northern and southern ends of the range. It is not possible to ascribe either putative taxon a unique combination of character states, and they are therefore not formally recognized. 18. Andira

Brazil. N. F. Mattos, Loefgrenia 45: 1. 1970.?Type: R. P. R. 12 Jan 1967, Belem & S. Pinheiro 3089 (holotype: UB!;

marauensis

Mara?,

Bahia: isotype:

CEPEC!). Tree to 35 m tall (though flowering from 3 m tall); buttresses absent; bark grey brown, Assuring vertically and flaking in small plates; slash pale brown, oxidizing darker; twigs brown to very dark brown, with very sparse appressed hairs, glabrescent; bark thicker and cracking on older twigs; lenticels on older twigs, elongated. Stipules ca. 2.5 mm long, to 1mm wide, caducous, sparsely hairy; leaf axis 3-10 cm long; rhachis very sparsely hairy, glabrescent, hairs short, red-brown, ? appressed; petiolules 3-7 mm long; stipels not seen (either absent or early caducous); leaflets (1-) 2 (-3) pairs, 2.5-8.5 cm terminal long, 1.1-3.7 cm wide, elliptic, narrowly obovate (rarely broadly obovate), leaflets often the most distinctly obovate, subcoriaceous, base acute to obtuse, slightly de rounded (rarely slightly retuse or with an acumen to 2 mm long), very glabrous adaxially, sparsely hairy abaxially, hairs red-brown, short (<0.2 mm long), vein channelled adaxially; secondary veins 8-10, ? plane adaxially, appressed; primary raised slightly abaxially, pattern brochidodromous, tertiary veins plane adaxially and or cm raised Panicles 5-11 plane slightly abaxially. long, terminal and axillary, with at red-brown hairs branch sparse, short, appressed tips, glabrescent towards the base; bracts narrow, caducous, ca. 2 mm long, sparsely hairy, hairs red-brown, ? appressed; 1-2 mm long; bracteoles caducous, not seen. Flowers 9-11 mm long. Calyx pedicels 4^.5 mm long, deep purple, glabrous except around the margins of the lobes; lobes 0.2-0.3 mm, obtuse, shallow. Petals pink to purple; standard blade 7.5-10 mm wide, 6-8 mm high, claw 2.5-3.5 mm long; wing 5-7 mm long, 2.5-3.5 mm wide, claw 4-4.5 mm long, lamellate sculpturing present; keel 5-7 mm long, 3-3.5 mm wide, claw 3.5-4.5 mm current, apex obtuse,

long. Stamens 8-9 mm long, united for the basal 4-6.5 mm, free for the distal 2-3 mm, vexillary stamen 5.5-6 mm long. Gynoecium 9.5-12.5 mm long, glabrous or with 1, 2, or 3 hairs; stipe 3.5-5 mm long, ovary 3-4 mm long, style 3-3.5 mm long; ovules 1-2. Fruit weighing ca. 20 g or less when dry, drying smooth (only seen as rotted fragments on the forest floor; pers. obs.). Chromosome

number unknown.

Phenology. Flowering January (3 records) and May (1 record). Distribution (Fig. 34). Brazil (Bahia); in Atlantic coastal rain forest. Vernacular

Additional

names.

Specimens

no Km 46 da Rod. BA-001 R. P. Bel?m & Pinheiro

Angelim. Examined.

Brazil.

(Ilh?us-Una),

Amorim

Bah?a: Mpio. Una, Reserva Biolog?a do Mico Le?o, estrada et al. 1411,1238,1908 (CEPEC, NY); Mpio. Mara?, Mara?, (MEXU, UB, US); Mpio. Uru?uca, Dist. de Serra Grande, 7.3 km

3091 (NY, UB), 3142 na estrada Serra Grande-Itacor?, et al. 3510 (CEPEC, NY); Mpio. Una, Reserva Biolog?a do A. M. de Carvalho Mico Le?o, estrada no Km 46 da Rod. BA001 /. G. Jardim et al. 90 (CEPEC, FHO); Mpio. Una, (Ilh?us-Una), border of Fazendas Maruim on road Oliven?a-Buerarema, and Dois de Julho, 33 km SW Oliven?a Maruim,

SYSTEMATICBOTANYMONOGRAPHS

94

50

FIG.

Mori

et al

13800

102 (CEPEC, K-2

(NY); Mpio.

34. Distribution

Ilh?us, Fazenda

35

45

55

of Andira

VOLUME 64

40

marauensis.

Barra do Manguinho,

Km

11 da Rod.

Ilh?us-Oliven?a,

Voeks

sheets).

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

2003

ANDIRA

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

long, narrow, moderately persistent, with short, appressed, red-brown mm mm long, narrow, moderately persistent, bracteoles 1.5-3 1.5-2 hairs; pedicels long; mm indumentum like that of bracts. Flowers 14-15 long. Calyx 6-7 mm long, with very sparse, appressed, pale red-brown, short hairs; lobes 0.5-1 mm long, acute to obtuse.

Petals violet, the standard with a central white marking; standard blade 10 mm high, 12 mm wide, claw 5 mm long; wing 9 mm long, 4-4.5 mm wide, claw 6 mm long, lamellate sculpturing present; keel 8 mm long, 3.5-4 mm wide, claw 6 mm long. Stamens 11mm long, filaments united for the basal 5.5-6.5 mm, free for the distal 4.5 mm, vexillary sta men 9-9.5 mm long. Gynoecium 13-13.5 mm long, with sparse short appressed hairs on the lower surface of the ovary; stipe 4.5 mm long, ovary 4.5 mm long, style 4-4.5 mm long; ovules 3. Fruits 5-10 cm long, 4.8-8 cm high, 4.5-9 cm wide, elongated, weighing 100-300 g when dry, flattened adaxially, rough, pale brown with dark brown specks (both fresh and dry); stipe 5-13 mm long; suture raised adaxially and abaxially; stylar remnant not visible; mesocarp 6-15 mm thick (fresh), pale greenish white to pale green, slowly ox idizing red-brown when cut, drying pale brown, hard, dry, finely granular; mesocarp 1.5-2 mm thick; endocarp 3-9 mm thick (fresh), brown, woody, fibrous. Chromosome number unknown. Figs. 3F, 4A(ii), 35. Phenology. Flowering in October. Distribution (Fig. 36). Brazil (Bahia); in open scrubby restinga on white

sand.

Additional Specimens Examined. Brazil. Bah?a: rod. BR-030, Porto de Mara?, Mpio. et al. 178 (CEPEC, K); Mpio. Km II, A. M. de Carvalho Ilh?us, estrada Ilh?us-Canavieiras, Campinhos-Mara?, Km 33, A. M. de Carvalho 622 (CEPEC); Mpio. Km 9, A. M. de Carvalho & Ilh?us, estrada Oliven?a-Maruim, Faria 2547 (CEPEC, UEFS); Mpio. Dh?us, 7 km on road Oliven?a-Vila et al. 3308 Brasil, A. M. de Carvalho Km 10, Gentry & Zardini 50011 Ilh?us, estrada Pontal-Oliven?a, (CEPEC); Mpio. Ilh?us, (CEPEC); Mpio. estrada Oliven?a-Maruim, Km 5-8, M. P. M. de Lima et al. 20 (CEPEC, RB-2 sheets); Mpio. Ilh?us, estrada Km 7-10, Martinelli et al. 11102 (CEPEC, RB); Mpio. Km 5, Ilh?us, estrada Oliven?a-Una, Oliven?a-Maruim, Fazenda Jairi, Mattos Silva et al. 1199 (CEPEC, K); Mpio. ramai com en Ilh?us, Fazenda Barra do Manguinho, trada no Km

10 da estrada

Km 10, Mattos Silva et al. 1393 (CEPEC, K); Mpio. Ilh?us, Pontal-Oliven?a, ? Fazenda Barra do ramai com entrada no Km 10 da estrada (junto Manguinho), Mattos Silva et al 1877 (CEPEC, K); Mpio. Km 6, R. T. Ilh?us, dirt road Oliven?a-Maruim, Ilh?us-Oliven?a, et al 181 (CEPEC, FHO, K); Mpio. R. T. Pennington Pennington Ilh?us, just S Oliven?a, 184, 185, 229 197 (CEPEC, FHO, K); Mpio. Mara?, Ilh?us, ca. 3 km N Oliven?a, R. T. Pennington (CEPEC, FHO, K); Mpio. Fazenda

Guanabara

96

FIG. opened

35. Andira

to show

carvalhoi.

inner surface.

SYSTEMATICBOTANYMONOGRAPHS

A. Habit.

B. Abaxial

F. Standard

233; C, A. M.

surface.

C.

de Carvalho

D. Flower. E. Calyx, of wing petal with lamellate also shown J. Gynoecium,

Inflorescence.

petal, inner surface. G. Outer surface H. Keel petal, inner surface. I. Androecium.

sculpturing and detail of sculpturing. section. K. Fruit. L. Fruit below in longitudinal T Pennington

leaflet

VOLUME 64

in section, showing wall structure. M. 491; D-J, M. P. M. de Lima et al. 20; K-L,

Seed. (Based on: A, B, R. R. T. Pennington 197.)

2003

ANDIRA

50

55

FIG. 36. Distribution

45

o?Andira

97

35

40

carvalhoi.

road Mara?-Porto

de Campinhos Km 20, R. T. Pennington et al 216, 217 (CEPEC, FHO, K); Mpio. Una, road Km 42, ca. 0.5 km down road to Pedras, R. T. Pennington et al 232, 233 (CEPEC, FHO, K); 8.9 km from Oliven?a, W. W. Thomas et al 8969 (CEPEC, NY); Mpio. Ilh?us, Road Oliven?a-Maruim, Mpio. then 3 km W, W. W. Thomas et al 9726 (CEPEC, NY); Mpio. Ilh?us, 10 km S Hh?us airport on road to Oliven?a, on road toMaruim, W. W. Thomas et al. 10950 (E, NY). Ilh?us, 8.9 km SW Oliven?a Oliven?a-Una,

Andira carvalhoi is only likely to be confused with A. n?tida from which it differs by itsmuch larger, brown fruits. It also has larger flowers and a longer leaf axis; the flowers of A. n?tida are 10-13 mm long and the leaf axis rarely exceeds 15 cm. Although A. carvalhoi is abundant in some sites (e.g., at Oliven?a, Bahia), itmust be considered endangered because of its restricted range. Its habitat of sandy coastal restinga is under great pressure for development and is highly prone to soil erosion if cleared. the fruits of A. carvalhoi are dispersed by large rodents (Pennington & de Lima Moreover, 1995) and may not be distributed adequately in the absence of these animals, as has been demonstrated for Hymenaea courbaril (Leguminosae-Caesalpinioideae; Asquith et al. 1999). It seems likely that many areas of restinga where A. carvalhoi grows do not

SYSTEMATICBOTANYMONOGRAPHS

98

support healthy populations of large rodents, given vegetation and proximity to centers of population.

VOLUME 64

the high levels of disturbance

of the

Brazil. parviflora Ducke in Arq. Inst. Biol. Veg. 2(1): 47. 1935.?Type: Amazonas: Manaus, estrada do Aleixo, 27 Apr 1932, A. Ducke RB 23865 (lecto type, here designated: RB!; isolectotypes: F! G! K! RB-2 sheets! U!).

20. Andira

Tree to 20 m tall, buttresses absent; bark smooth, pale brown, cracking vertically, not flaking, some horizontal markings; slash bright orange-brown; wood cream; twigs pale brown (or dark brown), with red-brown, erect, tangled hairs, glabrescent; bark Assuring vertically on older twigs. Stipules to 6 mm long, up to 5 mm wide, densely covered with erect, tangled red-brown hairs; leaf axis 2.5-17.5 cm long; rhachis with red-brown erect, tangled hairs; stipels 0.5-1.5 mm long; petiolules 0.5-1.5 mm long, indumentum like that of rhachis; leaflets in (2-) 3^1 (-5) pairs, 2-10 cm long, 1-4 cm wide, narrowly obovate, elliptic (rarely oblanceolate), coriaceous, base obtuse or rounded (rarely slightly cordate), apex obtuse, often with a short acumen to 5 mm long, glabrous adaxially, abaxially with sparse red-brown, erect hairs, >0.2-1.0 mm long, the veins more densely hairy; primary vein channelled adaxially; secondary veins 10-15, slightly sunken adaxially, raised abaxi the brochidodromous veins becoming brochidodromous, ally, pattern eucamptodromous anastamosing very close to themargin, tertiary veins impressed to slightly impressed adax ially and raised abaxially. Panicles 3-18 cm long, axillary and terminal, densely covered with red-brown, erect, tangled hairs; bracts 2.5-3 mm long, 1.5-2.5 mm wide, densely cov ered with pale brown, ? appressed hairs; bracteoles 1.5-2 mm long, 0.75-1 mm wide, in dumentum like that of bracts. Flowers 6-7.5 mm long. Calyx 3-4 mm long, lilac to purple, with

sparse, appressed, pale brown hairs. Petals sessile, whitish, the standard with purple marking; standard blade 6.2-7.5 mm wide, 5-5.5 mm high, claw 1.5mm long; wing 4.5-5 mm long, 2.8-3.2 mm wide, claw 2-2.5 mm long, sculpturing absent; keel 4-5 mm long, 2-2.5 mm wide, claw 2.5-3 mm long. Stamens 5 mm long, filaments united for the basal 1.5-2.5 mm, free for the distal 2-3 mm, vexillary stamen 3 mm long. Gynoecium 4.7-5.2 mm long, the ovary with pale brown, ? appressed hairs on upper and lower surfaces, indu mentum extending to the top of the stipe and base of the style with scattered hairs at the sides of the ovary, or ovary sparsely hairy on upper and lower surfaces only; stipe 1-1.2 mm long, ovary 2 mm long, style 1.5-2 mm long; ovules 1-2. Fruits 3-4.1 cm long, 2.4-3.2 cm high, 2.4-3.2 cm wide, ? globose to elongated, weighing ca. 20 g or less when dry, green, drying red-brown to dark brown to almost black, appearing smooth but some what tuberculate (best seen with lens or microscope); stipe insignificant (fruit ? sessile); su ture indistinct adaxially and abaxially; stylar remnant tiny; mesocarp 1.5 mm thick, pale mm 1.5-4 thick brown, granular, hard; endocarp (thickened along upper side), cream to non-fibrous. Chromosome number unknown. Fig. 37. pale brown, hard, to Phenology. Flowering February July. Distribution (Fig. 38). Brazil (Amazonas, Para); in terra firme forest on sandy and clay soil, and also in low forest on white sand. names.

Vernacular

Sucupira

vermelha,

acupu

rana,

sucupira

chorona

(Amazonas);

andira uchi (Para). Additional Rod.

BR-174,

INPA/WWF

Specimens Km

64,

1302.4500.2

Examined.

Brazil.

then 34 km E on ZF3,

Amazonas:

Fazenda

(K); estrada Manaus-ltacoatiara,

Distrito Agropecuario da Suframa, 1302 of DBFF project, Ackerly et al Km 26, Reserva Florestal Ducke, Adair s.n. (INPA);

Esteio,

Manaus, Reserve

ANDIRA

2003

99

V?rSeUAAM yC?ltf_s

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

et al 9074.)

100

FIG.

SYSTEMATICBOTANYMONOGRAPHS

38. Distribution

of Andira

parviflora,

A. cujabensis,

VOLUME 64

and A. cordata.

Manaus, Reserva Florestal Ducke, Alu?sio 115 (INRA); Manaus, Reserva Florestal Ducke, L. Co?lho s.n. (INPA); near Rio Taruma, near Cachoeira Alta, Ducke 2229 (COL, INPA); Manaus, Ducke 21348 (F); Manaus, estrada 90 km NNE Manaus, Distrito Agropecuario da S?frama, re do Taruma, Ducke RB 23866 (RB); Mpio. Manaus, serve 1501 of DBFFproject, "Km 41," Kukle 23 (K), Kukle & Boom 44 (K, MEXU); Proj. Radam, Rio Cauabari, afl. do Rio Negro, Marinho 526 (NY); Manaus, track from Km 63, road Manaus-Itacoatiara, Prance et al 9074 Km 60, Prance & Ramos 23563 (US); estrada Manaus-Itacoat (COL, F, GH, K); estrada Manaus-Caracara?, do inventario florestal, W. Rodrigues 7990 (INPA); estrada Manaus-Itacoatiara, iara, Km 130, ?rvore XIV-104 reserve Km 51, W. Rodrigues et al 11179 8484 (INPA); Manaus, WWF/INPA DBFF 85, W. Rodrigues Km 30, M. F. Silva et al. 91 (F); Rio Uaup?s, island above rapids (FHO, INPA, K); estrada Manaus-Caracara?, at Ipanor?, D. W. Stevenson et al 958 (K).?Para: do aeroporto, L. S. Co?lho et al. 270 C.PT ?as proximidades estrada da barragem do Caran?, pr?ximo ao igarap?, E. Soares 104 (INPA). (INPA); Porto Trombetas,

Km

This distinctive species is the only species o? Andira from the Amazonian region with red-brown hairs on the leaflet undersurfaces. The timber of A. parviflora is used for con struction and general carpentry (Loureiro & da Silva 1968). 21. Andira

cujabensis Bentham, J. Proc. Linn. Soc, Bot. 4, Suppl. ("A Synopsis of the tribe Dalbergieae): 120. 1860. Vouacapoua cujabensis (Bentham) Kuntze, Revis, gen. pi. 1: 212. 1891.?TYPE: BRAZIL. Goi?s: between Arragas and Navidade, Gardner 3654 (holotype: K!; isotypes: BM! E-2 sheets! F! G! GH-2 sheets! K!).

2003

ANDIRA

101

lanei N. F. Mattos, Loefgrenia 40: 1. 1970.?TYPE: BRAZIL. Mato Grosso: Tres Lagoas, Fazenda Canaan, Feb 1969, F. Lane s.n. (holotype: HB; isotype: K!).

Andira

a shrub) to 12 m; buttresses absent; bark thick, fissured; slash (occasionally wood honey-brown; pale straw-colored; twigs dark brown to pale brown, densely hairy, hairs pale brown to pale red-brown, erect, tangled, bark splitting on older twigs to reveal paler bark beneath; lenticels often numerous, pale, elongated. Stipules to 5 mm long, 3 Tree

mm wide

or narrower, early caducous, densely covered with pale brown to red-brown hairs; leaf axis 5-23 cm long; rhachis densely hairy, glabrescent, hairs buff to pale brown to red-brown, erect, tangled; stipels 1-2 mm long, caducous; petiolules 1-5 mm long, in dumentum like that of rhachis; leaflets in (2-) 3^4 (-5) pairs, (3-) 4-10.5 cm long, 2-5.5 cm wide,

broadly elliptic, elliptic, ovate, narrowly ovate (rarely broadly obovate to nar coriaceous, base rounded to cordate, apex obtuse (rarely acute or obovate), rowly a short blunt acumen to 5 mm long or slightly retuse, glabrous adax often with rounded),

ially, densely hairy abaxially and the epidermis often not visible, glabrescent, hairs gen erally buff to pale brown, occasionally red-brown, erect, tangled, >0.2-1.0 mm long; pri mary vein channelled adaxially; secondary veins 8-10, impressed to slightly impressed or ? adaxially, raised abaxially, pattern eucamptodromous becoming brochidodromous completely brochidodromous, tertiary veins plane adaxially and raised abaxially. Panicles (4.5-) 10-32 cm long, axillary and terminal, densely covered with pale red-brown, long, tangled hairs; bracts 1.5-2 mm long, 1mm wide, caducous, with long, pale red-brown hairs; bracteoles 1mm long, 0.5 mm wide, indumentum like that of bracts. Flowers 5.5-7 mm long, sessile. Calyx 3^4- mm long, black to dark purple, sparsely to very sparsely hairy, the margins of lobes hairy, hairs long, red-brown; lobes 0.2-0.5 mm long, obtuse, sometimes with the apex markings; standard blade long, 1.5-2.2 mm wide, 2.5-3 mm long. Stamens

acuminate. Petals dirty white, the standard with mauve-purple 5-7 mm wide, 4-5 mm high, claw 2 mm long; wing 3.5-4.5 mm sculpturing absent; keel 2.5-4 mm long, 1-2 mm wide, claw 4.5-5.5 mm long, filaments united for the basal (1.8-) 2.8-3.5

free for the distal 1.5-2 mm; vexillary stamen 4 mm long. Gynoecium 5-5.75 mm on or on its lower surface both the and lower surfaces, or the en upper long, ovary hairy tire distal half of the ovary hairy, hairs long, pale; stipe 1.5-2 mm long, ovary 1.5-2 mm long, style 1.75-2 mm long; ovules 1-2. Fruit 2.6-4.2 cm long, 2-3.2 cm high, 2-3.3 cm

mm,

wide, ? globose, weighing ca. 20 g or less when dry, green, drying brown to dark brown, smooth, appearing smooth but minutely tuberculate (best seen with lens or microscope); stipe \-4 mm long; suture a slight depression adaxially, not visible abaxially; stylar rem nant absent; mesocarp 1.5-2 mm thick, hard, granular, tinged slightly greenish; endocarp 1-2.5 mm thick, pale brown, woody, very hard, non-fibrous. Chromosome number un known. Fig. 2D. Phenology. Flowering January toMarch (occasional records as early as November or as late asMay). Distribution (Fig. 38). Brazil (Mato Grosso, Mato Grosso do Sul, Goi?s, Para); in cer rado and gallery forest. Oliveira Filho (1992) demonstrated that near Cuiab? (Mato Grosso), A. cujabensis can tolerate seasonal water-logging, which may be the explanation for its ability to grow in gallery forest. Vernacular names. Angelim branco, angelim do cerrado, cascudinho, Grosso); angelim amargoso (Goi?s); andira (Para).

sucupira (Mato

VOLUME64

SYSTEMATICBOTANYMONOGRAPHS

102

Additional W. R. Anderson 38339

Examined. 7150 (NY); 8 km N Terezina by road, W. R. Anderson Brazil. Goi?s: ca. 15 km S of Goi?s Velho, 7293 (NY, QCA); Serra Dourada, by road, W. R. Anderson & Kummrow (NY); Urua?u, Elias de Paula 3277 (UB); arredores de Guarai, Hatschbach

Specimens

11 km E of Cavalcante 10051

(K); Serra do Caiap?, ca. 8 km S Cavalcante,

eiros,

UB, US); Mpio. Piren?polis, Hidrel?trica da Serra da Mesa,

50 km S Caiap?nia, road to Jata?, Irwin et al 17962 (INPA); Chapada dos Vead Irwin et al. 23984 (F, G, UB); 8 km S Niquel?ndia, Irwin et al. 34872 (MO, NY, Mac?do 4352 (US); margem direita do Rio Tocantins, canteiro de obras da Usina

B. A. S. Pereira et al 1552 (NY); Rio Java?s, /. M. Pires & M. R. Santos 16257 (INPA); estrada para Formoso, a 6 km W de Cariri, J. M. Pires & M. R. Santos 16637 (US); Serra Dourada, Rizzo 4142 (RB); Mpio. Santa Isabel, Una do Bananal, Parque Nacional do Araguaia, Fazenda Bareira Branca, cam inho para Riozinho, F. C. da Silva et al 257 (SP, UB); Mpio. Goi?s, Souza Lima 208 (RADAM, RB).?Mato

GROSSO: 35 km ENE Barra do Gar?as, W. R. Anderson at western 11293 (SPF); Mpio. Chapada dos Guimar?es,

9691

(NY); Mpio. Coxim, Rio Taquar?, W. R. Anderson 11361 edge of Chapada dos Guimar?es, W. R. Anderson Geographical Society Base Camp, de Castro R2102 (E, K, U,

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

Luciara,

Santa Terezinha,

(SP); vicinity of Veu de Noiva, ena, Prance & Schaller 26266

Chapada

Santa Terezinha, perto do Lago Portugu?s, /. Mattos 15539 Prance et al 18965 (K, U, US); Fazenda Santa Filom 270 km N Xavantina, Royal Geographical Society Base Camp, Ramos

Fazenda

dos Guimar?es,

(GH, NY); & R. Souza 156 (K); Vale de Sonhos, 80 km N Barra do Gar?as on road to Xavantina, Ratter et al 2323 (E, K, U, UB); Pantanal, Rio Negro, Fazenda Santa Teresa, Schaller 160 (NY); Km 165 da rodovia Cuiab?-Santar?m entroncamento, M. G. Silva & Rosario 4939 (NY); Mpio. Luciara, lake 2 km NW Luciara, W. W. Thomas et al 4312 51

do Sul: Mpio. Nova Andradina, MS-134 a 30 km de Nova Andradina, Leite & Klein a 22 km de Nova Andradina, Nova Andradina, MS-134 Pastore & Klein 77 do Rio Araguaia, ? Santana do Araguaia, Lemes & Mileski 158 margems pr?ximo

Grosso

(K).?Mato (RADAM,

RB); Mpio.

(RADAM).?Para:

Fazenda (RADAM, RB); near Reden??o, Inaj?, N.T. Silva 4808 (NY, US).

Prof. Getulino,

Ratter

et al. 6869V

(E); Rio Araguaia,

Fazenda

do Rio

In the Brazilian cerrados, sterile specimens of A. cujabensis might be confused with A. verm?fuga-, however, the smaller (5.5-7 mm long), white flowers of A. cujabensis are completely distinct from the larger (12.5-18 mm long), pink to purple flowers of A. ver m?fuga. For a separation from A. cordata see that species (no. 22). Although Bentham (1860) spelled the specific epithet "cujabensis" in the protologue, two orthographic variants have been widely used, "cuiabensis" (Bentham 1862; I have used this incorrect spelling on many determination slips) and "cuyabensis" (Mattos 1979). 22. Andira

Arroyo ex R. T. Pennington & H. C. Lima, Kew Bull. 50: 562. Brazil. Bahia: 30 km W Barreiras, 12 Jan 1977, G. Hatschbach MBM!; (holotype: isotypes: G! K! MEXU!).

cordata

1995.?Type: 39477

a shrub; buttresses absent; bark thick (to 1 cm), Tree 6 (-15) m tall, occasionally to grey-brown, Assuring vertically; slash pale red-brown, 5-20 mm thick, a little red ex?date; wood buff streaked with cream; twigs dark brown to brown, older twigs paler, glabrous (or rarely sparsely hairy, glabrescent); lenticels numerous, elongated, pale brown. Stipules early caducous, not seen; leaf axis 4.5-15 cm long; rhachis glabrous (or rarely with sparse, pale brown, erect hairs), dark brown peeling to reveal red-brown or pale brown beneath; stipels 0.5 mm long, caducous; petiolules 1.5-4 mm long, glabrous brown

(or sparsely hairy); leaflets in 2-4 pairs, 2-7 cm long, 1.4^- cm wide

(the lower leaflets

2003

103

ANDIRA

to 5 cm wide),

broadly elliptic to ovate, (rarely suborbiculate, elliptic, narrowly ovate to ovate), broadly thinly coriaceous, base truncate, cordate (rarely rounded); apex obtuse to rounded (the terminal leaflets sometimes truncate), often slightly retuse, occasionally with an acumen to 4 mm long, glabrous (rarely very sparsely hairy abaxially, hairs erect); pri mary vein channelled adaxially; secondary veins 8-10, plane to slightly impressed adaxi ter ally, slightly raised abaxially, pattern eucamptodromous becoming brochidodromous, cm to Panicles 5-25 veins raised and abaxially. long, slightly tiary plane plane adaxially axillary and terminal, sparsely covered with pale-brown hairs at branch tips, glabrescent at base; bracts and bracteoles caducous, not seen. Flowers 6.5-7 mm long, sessile. Calyx 3-3.5 mm long, purple to black, glabrous (or with a few scattered pale appressed hairs), except themargins of the lobes sparsely hairy, hairs pale, ? erect; lobes 0.2-0.4 mm long, obtuse. Petals whitish to pale lilac, the standard with purplish markings; standard blade 5-6.5 mm wide, 4.5-5 mm high, claw 2-2.5 mm long; wing 4-4.5 mm long, 1.5-2.5 mm wide, claw 2.5-3 mm long, sculpturing absent; keel 3-3.5 mm long, 1.5-2 mm wide, claw 3 mm long. Stamens 5.5-6 mm long, filaments united for the basal 2.5-4 mm, free for the 5.5-6.5 mm long, distal 1.5-2.5 mm; vexillary stamen 3.5-4 mm long. Gynoecium mm mm mm 2-2.5 1.75-2 long; ovules 2. long, style long, ovary glabrous, stipe 1.75-2.2 cm cm ? cm Fruit 3-3.6 wide, high, 2.5-2.9 globose, weighing ca. 20 g or long, 2.5-2.9 less when dry, green with pale yellowish specks when young, drying dark brown, almost tuberculate (best seen with lens or microscope); appearing smooth but minutely mm remnant not apparent; mesocarp 1-2 mm thick, gran sutures and 3.5 stylar stipe long;

black,

ular, hard; endocarp 2^- mm

thick. Chromosome January to April.

number unknown. Figs. 3A, 4B(i),

Phenology. Flowering Distribution (Fig. 38). Brazil (Maranh?o, Bahia, Goi?s, Tocantins); Vernacular name. Gr?o de galo (Bahia). Additional Anderson

in cerrado.

ca. 100 km WSW Barreiras, W. R. Examined. Brazil. Bahia: Espig?o Mestre, (NY), 36797 (NY, UB); Mpio. Correntina, Faz. Jatob?, margem da estrada que da acesso da Silva 1352 (UB); Mpio. Barreiras, Elias de Paula 3122 (UB); Mpio. Correntina, Fazenda & Collares Rio Formoso, M. M. Fern?ndez 17 (RADAM, RB); drainage of Rio Corrente,

Specimens

et al 36607

a guarita, Aparecida Formoso do Guara, Rio Piau, ca. 150 km WSW al. 31521

39.

(UB); Mpio.

Barreiras, Irwin et al 14855 (F, G); Mpio. Barreiras, 5 km NW Barreiras, Irwin et ca. 2 km up road to airport, R. T. Pennington & Brito 261, 262, 263, 264 Cocos, Lagoa do Pratud?o, S. B. da Silva & M. G. de Lima 372 (NY); Mpio. Formoso

Barreiras,

(CEPEC, FHO, K); Mpio. do Rio Preto, proximo ao rio Riach?o, pr?ximo a vereda Olhas d'Agua, Walter et al. 228 (US).?GOI?S: Serra G?rai de Goi?s, Rio da Prata, 6 km S de Posse, Irwin et al. 14420 (F, G, NY); Rio Corda, regi?o de Xamboi?, E. Oliveira 1446 (UB); 10?45'S, 47?15'W, Orlandi 91 (RADAM, RB).?MARANH?O: 7?12'S, 47?25'W, J. A. Ferreira

& C. A. Miranda 304 (RADAM, RB); Mpio. Graja?, J. A. Ferreira & C. A. Miranda 322 (RADAM, & G. Gottsberger 15-9482 S?o Codo, Fazenda Canto do Ro?a, /. Go?tsberger RB); Mpio. (NY); Mpio. em dire?ao ? Balsas, C. A. Miranda & J. A. Ferreira Raimundo das Mangabeiras, 355 (RADAM, RB); duas Black 1630a (NY); near Carolina on road N to Estreito, transect area 2, leguas de Carolina, J. M. Pires &G.A. Ratter & V. P. de Lima 6716 olina,

Ponto

Taguatinga,

and BR-010, 35 km N of Car (E); Mpio. Carolina, Transamazonian hwy, BR-230 towards Serra de Baleia, E. L. Taylor et al E1259 (NY).?TOCANTTNS: Mpio. estrada Taguatinga-Mimoso do Oeste, Km 14, B. A. S. Pereira 1604 (NY). Tur?stico W

Andira cordata is the only tree species of Andira in the Brazilian cerrados with en tirely glabrous foliage, or with only very poorly developed indumentum on the leaflet un dersurfaces. Andira verm?fuga and A. cujabensis both have well-developed indumentum on the leaflet undersurfaces. Although many specimens of A. humilis are glabrous, this species is completely distinct in its geoxylic suffrutex growth form. This

species

closely

resembles A. cujabensis

and is parapatric

(sensu Mayr,

1982,

104

SYSTEMATICBOTANYMONOGRAPHS

VOLUME 64

cordata. A. Bark of trunk. B. Portion of twig with lenticels twig. C. Habit. D. Node of stipels. E. Flower. F Calyx, opened to show inner surface. G. Standard petal, inner surface. H. Wing K. Gynoecium, also shown at right in longitu petal, outer surface. I. Keel petal, inner surface. J. Androecium. of C and dinal section. L. Fruit. (Based on: A, R. T. Pennington 264; B-D, R. T. Pennington 262, inflorescence FIG.

39. Andira

rhachis with

E^K, G. Hatschbach

39477;

L, J. M. Pires

1630a.)

2003

ANDIRA

105

p. 275) with it. The combined phylogenetic analysis indicates that these taxa are closely from A. cujabensis by its lack of indu related (Fig. 8). Andira cordata is distinguished mentum (on foliage, gynoecium, and calyx). Andira cordata also has smaller leaflets that generally are not acuminate and secondary veins that are plane adaxially and less raised abaxially. Some specimens of A. cordata (e.g., Gottsberger 15-9482 and Taylor E-1259) have sparsely hairy foliage, but the indumentum is very poorly developed in comparison to that of A. cujabensis, and the gynoecium remains glabrous. Brazil. Ama 23. Andira micrantha Ducke, Arq. Inst. Biol. Veg. 2(1): 48.1935.?Type: zonas: Manaus, estrada do Aleixo, 10May 1932, A. Ducke KB 23864 (lectotype, here designated: RB!; isolectotypes K! U! US!). Tree to 35 m

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

along the upper suture); stylar remnant absent; exocarp thin, black in section; mesocarp 4-8 mm thick, pale brown to brown, granular, very hard; endocarp 5-9 mm thick, pale brown,

very

hard,

non-fibrous.

Chromosome

number

unknown.

Phenology. Flowering March to June. Distribution (Fig. 40). Brazil (Amazonas); in terra firme forest, on both sandy and clay soils, though some records are from river banks. Vernacular

names.

Acapu-rana,

sucupira

vermelha,

sucupira

chorona.

106

SYSTEMATICBOTANYMONOGRAPHS

FIG. 40. Distribution

50

55

60 70 65

of Andira

micrantha,

A. cori?cea,

VOLUME 64

4035 45

and A. grandistipula.

Examined. Reserva Florestal Ducke, Adair s.n. Specimens Additional Brazil. Amazonas: Manaus, estrada do Taruma, Ducke RB 23401 (G, K, RB, U); Manaus, (INPA); Manaus, Ducke RB 21365 (F); Manaus, terreno da SIDERAMA, Loureiro et al. s.n. (INPA); estrada do Aleixo, Ducke RB 35077 (G, INPA); Manaus, reserve 1501 ("Km 41") of DBFF project, A. A. 90 km NNE Manaus, Distrito Agropecuario, Mpio. Manaus, Oliveira ter Egler, (MEXU,

et al. 420

(K); Manaus, Reserva Florestal Ducke, quadra 23, O. Pires 29 (INPA); Manaus, Reserva Wal estrada Mau?, Prance el al. 11635 Km 64, Prance ei al 9049 (U); Manaus, road Manaus-Itacoatiara, ei al. 15011 (U); Prance NY, U); Basin Rio Negro, Rio Cuieras just below mouth Rio Brancinho,

1534 (US); Manaus, Reserva Florestal Ducke, ao lado da & Chagas Igarap? da Cachoeira Alta, W. Rodrigues Km 84, ?rvore no. 7929 (COL, INPA); estrada Manaus-Itacoatiara, ?rvore no. 1578, W. Rodrigues & Osmarino 7992 (INPA); Manaus, Cachoeira Baixa do Tarum?, W. Rodrigues & V-13 do inventario florestal, W. Rodrigues

L. Co?lho 8557 (INPA). is a rain forest species and ismost likely to be confused with A. co ri?cea and A. taurotesticulata, which also grow in this habitat and possess large fruit. These two species are not, however, sympatric with A. micrantha, which grows in central Amazonia in the vicinity of Manaus. Andira micrantha may be distinguished from A. co Andira micrantha

ri?cea by the persistent stipules whereas stipules of A. micrantha and presumably tiny; no stipule fruit that are ribbed from suture

15 mm long and 5 mm wide, on any specimens examined) Andira taurotesticulata has sparse hairs on the undersur

face, whereas

leaflets glabrous.

of the latter. These reach are caducous (not present scars have been observed. to suture and leaflets with the fruit of A. micrantha are smooth and the

Pulle, Rec. Trav. Bot. N?erl. 6: 267. 1909.?Type: Sectie O, tree number 61, 15May 1907, Boschbeheer (lectotype, here designated: U!).

24. Andira cori?cea Boschreserve

Suriname. (B. W.) 61

2003

ANDIRA

107

R. Benoist, Bull. Mus. Hist. Nat. (Paris) 25(4): 296. 1919.? Guiana. Maroni, Wachenheim 79 (lectotype, here designated: P!; P!).

Andira wachenheimii Type: French isolectotype:

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

number

unknown.

Phenology. Guiana). Distribution

Flowering

May

to October

(May, Suriname; August-October,

French

(Fig. 40). French Guiana and Suriname; in rain forest. names. Kobon, roode kabbes, reddie kabbes, koeraroe (Arawak), akoeli lebi (Par?maca; French Guiana). tjerere (Suriname); St. Martin rouge, lebi kiabici/kiabici Vernacular

Specimens Examined. Sectie O, B. W. 1909 (COL, U, US); Sectie O, tree 61, Suriname. I, B.W. 5450 (U); Brownsberg, 1, tree 5, B. W. 4092 (U); Zanderij (U); Forest Reserve Zanderij tree 1301, B. W. 6870 (A); Brownsberg, B. W. 6870 (U); Emmaketen, 1.5 km from main camp in direction of small Hendrik peak, Daniels in bosje op te savanna ten O. van de Brinck & Jonker 868 (U); Sabanpassie Additional

B. W. 3933

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

Charvein

Km et en

108

SYSTEMATICBOTANYMONOGRAPHS

VOLUME 64

ismost likely to be confused with A. taurotesticulata (no. 5) and A. the other rain forest species with large fruit; see those species for a (no. 23), discussion of their differences. CTFT (1989) reported that the timber of A. cori?cea is used in the construction of of items such as wooden houses and has commercial potential for the manufacture snooker cues and umbrella handles. Andira

cori?cea

micrantha

Pulle specimen,

(1909) cited two collections with his description of A. cori?cea. The flowering collected on 15May 1906, is selected here as the lectotype.

Amshoff, Bull. Torrey Bot. Club 75(4): 394. 1948.?Type: grandistipula Guyana. Kaieteur Plateau, Kaieteur Savannas, 6May 1944, B. Maguire & D. B. Fanshawe 23273 (holotype: NY!; isotypes: A! F! K! U!).

25. Andira

Small tree 2.5-6 m tall; presence of buttresses unknown; bark and slash unknown; twigs glabrous, very dark brown, almost black, this outer layer splitting and peeling to reveal red-brown beneath; lenticels pale. Stipules to 7.5 cm long, to 6 cm wide, broadly obovate, pale green drying pale brown, glabrous, persistent, crowded at shoot tips and around inflorescence bases; leaf axis 21-30 cm long; rhachis very dark brown, almost this layer thin and peeling to reveal red-brown beneath, glabrous; stipels absent (perhaps present on young leaves and caducous); petiolules 7-13 mm long, dark brown, almost black, this layer thin, peeling to reveal red-brown beneath, glabrous; leaflets in (2-) 3-4 pairs, 11-18 cm long, 4.5-10 cm wide, elliptic, broadly elliptic (rarely nar

black,

rowly obovate, some lower leaflets occasionally broadly ovate to ovate), coriaceous, base rounded, apex rounded (rarely obtuse), often with a short acumen to 3 mm long, glabrous adaxially, sparsely hairy abaxially, hairs minute (<0.1 mm long), appressed, red-brown; primary vein channelled adaxially; secondary veins 8-11, ? plane to slightly sunken adaxially, raised abaxially, pattern eucamptodromous becoming brochidodro tertiary veins plane adaxially and plane to slightly raised abaxially. Panicles cm long, terminal, dark brown, almost black, sparsely covered with short, ap pressed, red-brown hairs at branch tips, glabrescent at base; bracts not seen; pedicels 2-4 mm long, very dark brown, almost black, glabrous; bracteoles not seen. Flowers ca.

mous, 17-32

15mm

long. Calyx dark brown, almost black, ca. 6 mm long, glabrous except the mar gins of the lobes sparsely covered with red-brown hairs; lobes 0.5-1 mm long, obtuse. Petal color unknown (only old flowers seen); standard blade ca. 12 mm wide, 10 mm high, claw ca. 3.5 mm long; keel and wings not seen. Stamens ca. 10mm long, filaments united for the basal ca. 6 mm, free for the distal ca. 4 mm. Gynoecium ca. 13-15 mm long, very sparsely hairy on lower surface of the ovary, hairs red-brown, ? appressed; stipe ca. 4 mm long, ovary ca. 6 mm long, style ca. 4-5 mm long; ovules 1 (single ovary dissected). Fruit 4-7.8 cm long, 3.4-6 cm high, 3.6-6.5 cm wide, elongated, broad, flat tened adaxially, weighing 100-300 g when dry, without odor, greenish yellow when brown and dark brown, some areas somewhat yellow, appearing ripe, drying pale smooth but minutely wrinkled and tuberculate (best seen with lens or microscope); su ture a fine line adaxially, raised abaxially; exocarp thin, almost papery (fresh fruit); 5-12 mm thick, greyish brown, hard, finely granular, often splitting along mesocarp suture when dry; stylar remnant minute; endocarp 3-13 mm thick, cream-buff, upper non-fibrous,

very

Phenology. November.

hard.

Chromosome

Flowering

in May

number

(single

unknown.

record);

two fruiting

records

in June and

ANDIRA

2003

Distribution areas

of

(Fig. 40). Guyana,

restricted

109

to the Pakaraima Mountains;

in scrub in

savanna.

Specimens Examined. Region: Additional Pakaraima Mts, Imbaimadai, Guyana. Mazaruni-Potaro 7972 (U, US); 1992 (COL, K, US); Pakaraima Mts, Imbaimadai Creek W of Imbaimadai, Pipoly Hoffman Pakaraima Mts, Mt. Membaru, Maas & Westra 4299 (K, U); Upper Mazaruni River Basin, Partang Rapids, near et al 43878 (NY-2 sheets). mouth Partang River, Maguire

is distinguished by its large, persistent stipules from the other grandistipula II of Andira (defined by an endocarp of stone cells). None of the clade of eight species collections examined had fully mature flowers, and the petal color is unknown. Andira

Brazil. Arroyo ex R. T. Pennington, sp. nov.?Type: Amap?: Rio Oiapoque, 3 km E mouth Rio Mutura, 22 Sep 1960, H. S. Irwin, J. M. Pires & L. Y. Th. Westra 48442 (holotype: NY!; isotypes: F! U! ?B!).

26. Andira

praecox

tervequinata floribus majoribus et statura altiore recedit (arbor grandis m ad 30 alta, non frutex vel arbuscula usque ad 7 m alta). usque m Tree to 30 tall; presence of buttresses unknown; bark and slash unknown; twigs brown, densely covered with red-brown appressed hairs, becoming grey-brown, glabres cent. Stipules to 10 mm long, to 1mm wide, caducous, with appressed, red-brown hairs; Ab Andira

leaf axis 3-10 (-18.5) cm long; rhachis with sparse, red-brown, appressed hairs; stipels 1-2 mm long; petiolules 2-5 mm long, sparsely to densely hairy, hairs red-brown, ? ap pressed, short (<0.2 mm long); leaflets in (1-) 2 (-3) pairs, 3.5-9.5 (-14) cm long, 1.9?4.3 (-6) cm wide, narrowly obovate (rarely elliptic), terminal leaflets distinctly obovate, to subcoriaceous,

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

rent, rounded

(rarely obtuse),

base obtuse

apex slightly

plane adaxially, raised abaxially, pattern eucamptodromous 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. 3mm long, style 2.5-3 mm long; ovules 1-2. Fruits 3.1-4.1 cm long, 2.9-3.6 cm high, 2.8-3.6 cm wide, ? globose, weighing ca. 20 g or less when dry, very dark brown, almost (with brown pock-marks where the surface is broken) and slightly wrinkled tuberculate with a slight waxy bloom (best seen with lens or microscope); stipe long; sutures raised adaxially and abaxially; stylar remnant insignificant; meso carp 1-2 mm thick, pale brown, hard, granular; endocarp 2-5 mm thick, buff, woody, non

black,

minutely to 9 mm

fibrous,

very

hard.

Chromosome

number

unknown.

Fig.

41.

110

SYSTEMATICBOTANYMONOGRAPHS

VOLUME 64

FIG. 41. Andira praecox. A. Habit with developing young fruit. B. Habit with mature fruit. C. Abaxial leaflet surface. D. Calyx, opened to show inner surface. E. Standard petal, inner surface. F. Wing petal, outer sur face. G. Keel petal, inner surface. H. Androecium. I. Gynoecium, also shown at right in longitudinal section. J. Old flower with developing fruit. K. Entire fruit (at left) and in section (at right), showing wall structure. (Based on: A, C-J, H. S. Irwin et al. 48442;

B, K, D. C. Daly

et al. 3923.)

FIG. 42. Distribution

Phenology. ber

111

ANDIRA

2003

and

of Andira

praecox,

in September

Flowering

A. tervequinata,

A. trifoliolaia,

and A. unifoliolata.

(single record); two fruiting

records in Decem

January.

Distribution Vernacular Additional

(Fig. 42). Brazil (Amap?); in terra firme forest. names. Andiroba jaruba, sucupira vermelha. Brazil. Amap?: Mpio. Mazag?o, road toward BR-156, incomplete 3923 (K, NY); Mpio. 5-10 km SW Rio Preto, Daly & Cardoso Macap?, Vila Nova, 18 km SSW Cupixi, Rabelo et al 3199 (ULM, US); Serra do Navio, Ro

Specimens

Monte

75-80 Dourado, road Cupixi-Rio Macap?,

Examined.

km WSW

drigues 2917 (TNPA). Andira praecox ismost similar toA. tervequinata, especially in leaf morphology, but differs in several characteristics, especially in its smaller flowers (6.5-7 mm long, whereas those of A. tervequinata are 9 mm long). Andira praecox grows in rain forest and is a tree 15-30 m in height. In contrast, A. tervequinata is a shrub or small tree to only 7 m high, which grows in savanna and associated gallery forest habitats. The twigs of A. praecox are grey-brown, whereas those of A. tervequinata are very dark brown, generally with almost black outer bark, which peels to reveal red-brown bark beneath. Andira praecox leaves generally have five leaflets (occasionally trifoliolate, or with seven leaflets), but A. terve quinata always has five leaflets or trifoliolate leaves. Leaflet shape differs subtly. In A. leaflets are narrowly obovate (rarely elliptic), but in A. tervequinata they are praecox, broadly elliptic to broadly obovate. Further diagnostic characters may be found in the fruit, but fruiting collections of both species are few and poor. Andira praecox generally has smaller fruits (to 4.1 cm long), and its fruit surface (when dry) is dark brown to almost

112

SYSTEMATICBOTANYMONOGRAPHS

black, whereas the fruit of A. tervequinata brown with pale, raised specks.

VOLUME64

is larger (5.5 cm long) and generally more

red

R. T. Pennington, G. Aymard & N. Cuello, Novon 7: 72. tervequinata 1997.?Type: Venezuela. Bol?var: Distrito Heres, west bank of R?o Trueno km of 35 W Alto, Chiguao, high plateau, ecotone between gallery forest and sa

27. Andira

vanna, 23 Mar

1985, Huber 10345 (holotype: NY!; INPA, K, MO, MY, MYF! P, PORT, RB, US, VEN).

isotypes AAU,

E! HBG,

tree or shrub to 7 m tall; presence of buttresses unknown; bark and slash un twigs very dark brown with raised elongated lenticels, outer bark very dark brown

Small known;

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

name. Pil?n rebalsero.

Additional Specimens Examined. Venezuela. Bol?var: Distrito Heres, a lo largo del R?o El Trueno al N de la base del Guaiquinima 6147 Distrito Piar, R?o Aparam?n at Kambay NY, (MO, PORT); Aymard Tepui, mer? rapids, SE base of Amaruay-tepui, W of Aparam?n-tepui, E of Auyan-tepui, Holst & Liesner 2798 (E, MO); Distrito Piar, R?o Aparam?n, Kambay-mer? rapids, ca. 3 km SE of SSE corner of Amaruay tepui, Liesner & Hoist

20674

(E, MO).

ANDIRA

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113

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

opened

to show

SYSTEMATICBOTANYMONOGRAPHS

114

VOLUME64

Andira tervequinata might be confused with A. trifoliolata (no. 28) and A. praecox from A. trifoliolata were discussed in de (no. 26); see notes under the latter. Differences tail by Pennington et al. (1997) and are summarized here. Andira trifoliolata has trifolio late leaves and leaflets that are glabrous, with a plane primary vein on the upper surface and an acuminate

apex. Andira tervequinata has leaves with 3-5 leaflets, which have on the undersurface, a channelled primary vein on the upper sur hairs short, appressed

face,

a retuse

and

or

emarginate

apex.

Brazil. Ducke, Arq. Inst. Biol. Veg. 4(1): 22. 1938.?Type: Rio Curicuriare, 20 Feb 1936, A. Ducke RB 35079 (lectotype, here designated: RB!; isolectotypes: G! K! U!).

28. Andira

trifoliolata

Amazonas:

to 20 (-30) m tall; presence of buttresses unknown; bark dark brown, slash hard, yellow, fibrous; twigs dark brown (or almost black), sparsely hairy, occasional glabrescent; pale, elongated lenticels. Stipules to 9 mm long, to 1mm wide, with brown, caducous, appressed, red-brown hairs at tips, glabrous near base; leaves leaf axis 2.5-9 (-12) cm long. Rhachis dark brown, this layer trifoliolate, pinnately Tree

yellow; wood

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

ca. 20 g or less when dry, green, drying dark globose, weighing smooth but appearing minutely wrinkled (best seen with lens or where surface is broken; stipe 15-25 mm long; su pock-marks adaxially, not visible abaxially; stylar remnant tiny; mesocarp to dark brown (or greenish), hard, granular; endocarp 2-3 mm

thick, pale number unknown. thick, pale buff, very hard, non-fibrous. Chromosome Phenology. Flowers recorded in February, June, July, and October.

Distribution (Guainia); Venezuela (Fig. 42). Colombia (Amazonas); zonas); along rivers, but also in terra firme and secondary forest.

Fig. 4B. Brazil

(Ama

ANDIRA

2003

115

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

Acero

221

small laja along lower part 81 (K); Depto. Rio Negro, Mucuriapi, 551 (FHO); road from San Fernando (NY); Depto. Atabapo, L. Delgado km from San Fernando, Gentry & Tillett 10889 (MO); Depto. Atabapo, et al. 125856 nidades de Culebra y Huachamacari, (NY); Rio Steyermark 43368 Brazil. AMAZONAS: Barcelos, Ducke (F, NY, U, US). Adderley Coomes

of R?o Baria, Davidse & Miller 26752 to Santa Barbara, 12^0 de Atabapo entre las Comu R?o Cunucunuma, Casiquiare, RB 17293

near Solano, Wurdack

&

(RB, K); Rio Negro, Rio 764 (NY); estrada Manaus-Porto trecho entre os Rios Castanho e Tupana, Velho, Curicuari, O.C. Nascimento M. F. Silva et al. 924 (INRA); estrada Manaus-Porto Velho, Km 40-30, M. F. Silva 979 (INPA).

Andira trifoliolata is the only species in the genus that has all leaves trifoliolate. It is likely to be confused only with A. praecox, which rarely has a few trifoliolate leaves, and A. tervequinata, which has 1-2 pairs of leaflets; neither of these species is uniformly tri foliolate like A. trifoliolata. For other diagnostic characters, refer to the discussion of A. praecox (no. 26) and A. tervequinata (no. 27). The collection Gentry & Tillett 10889 may represent a new species. It is said to have been taken from a shrub, whereas all other records of A. trifoliolata are from trees. More over, the leaflets have sparse, appressed hairs on their lower surfaces, whereas all speci mens

of A. trifoliolata are glabrous. Yet, the specimen has immature fruit and lacks flow ers; more complete material is necessary to determine its true status. The fruit and bark of A. trifoliolata may be used as a contraceptive (herbarium label: Christenson et al. 1432, US). Brazil. Ducke, Arq. Inst. Biol. Veg. 4(1): 22. 1938.?Type: estrada do 3 Mar A. Ducke RB 35078 Manaus, Aleixo, 1937, (holo type: RB!; isotypes: G! K! INPA! RB-5 sheets! U!).

29. Andira

unifoliolata

Amazonas:

Tree to 35 m tall with small buttresses; bark rough, flaking; slash unknown; twigs dark brown, with very sparse, red-brown hairs when young, rapidly glabrescent, bark of older twigs splitting to reveal paler bark beneath; lenticels raised, pale, elongated. Stipules to 4 mm long, very early caducous (seen only on a seedling), with sparse, red-brown hairs; petiole 5-16 (-18) cm long, glabrous, dark brown, this layer peeling to reveal pale brown stipels to 2 mm long (seen only on seedlings); petiolule 3.5-6 mm long, unifoliolate (1-2 pairs leaflets in seedlings), leaflet 4.7-13 cm long, 1.3-5 cm glabrous; wide, elliptic (to narrowly ovate), dark green, shiny adaxially, much paler and matt abax ially, the venation pale, thick-chartaceous (seedlings) to coriaceous, base obtuse (or rarely acute), often slightly decurrent, apex acute to obtuse with an acumen to 10 mm long, beneath;

glabrous adaxially, very sparsely hairy abaxially, glabrescent, hairs short, appressed; pri mary vein plane adaxially; secondary veins 6-7, plane adaxially, raised or only slightly raised abaxially, pattern disorganized, eucamptodromous with occasional brochidodro mous linkages (particularly at the leaflet tip), the veins curving uniformly, tertiary veins plane adaxially and abaxially. Panicles 2^4.5 cm long, axillary (and terminal), with sparse, pale red-brown hairs at branch tips, glabrescent proximally; bracts and bracteoles early ca ducous (not seen); pedicels 0.8-1 mm long. Flowers 6-7 mm long. Calyx 3-4 mm long, dark brown or ? black, glabrous or with a few scattered appressed pale red-brown hairs, the margins of the lobes sparsely to very sparsely hairy; lobes obtuse with small pointed tips, 0.3-0.8 mm. Petals white, the standard marked with red; standard blade 6 mm wide,

116

SYSTEMATICBOTANYMONOGRAPHS

VOLUME 64

i\??M?^

L?\r\^

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

andira

2003

117

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

number unknown. Figs. IC, 2F, 44. Phenology. Flowering February toMarch. Distribution (Fig. 42). Brazil (Amazonas, Para, Roraima); sandy and clay soils. Vernacular

names.

zonas); andira-uchi

Sucupira

amarela,

sucupira

chorona,

in terra firme rain forest on

sucupira

vermelha

(Ama

(Para).

Additional Specimens Examined. Brazil. Amazonas: Reserva Florestal Ducke, Adair s.n. Manaus, Reserva Florestal Ducke, ?rvore do fenel?gico (INPA); Manaus, 20, Alu?sio s.n. (INPA); Mpio. Mau?s, Rio ?cima do lugar S?o Sebasti?o, Cid Ferreira 4268 (INPA); Manaus, estrada do Aleixo, Ducke 674 Apoquitaua s.n. (INPA), Mac?do s.n. (INPA), F. C. Mello (F, GH, K, NY, US); Manaus, Reserva Florestal Ducke, Lourival et al. s.n. (INPA); Manaus, Reserva Florestal Ducke; estrada Manaus-Itacoatiara, Km 133, Monteiro & F. s.n.

Mello

Reserva Florestal Ducke, W. Rodrigues & Loureiro 5953 (INPA); Manaus, (INPA); Manaus, Florestal Ducke, ?rvore do fenel?gico & D. Co?lho 7554 (INPA); estrada Man 13, W. Rodrigues Km 190, ?rvore XXVI-37 do inventario florestal, W. Rodrigues 7994 (INPA); estrada Man aus-Itacoatiara, Km 170, picada XXII, ?rvore 58 do inventario florestal, W. Rodrigues e? al. 8502 (INPA); aus-Itacoatiara,

Reserva

estrada Manaus-Porto

Velho, Rio Castanho, margem direita, estrada para Reserva Florestal Ducke, estudo de associa?ao, J. A. C. (INPA); Manaus, entre as cachoeiras e Jandi?, L. S. Co?lho 103 de Tremalhetinha Trombetas, do Caran?, entrada para a Barragem Velha, E. Soares 5 (INPA).?Roraima: reservatorio,

Maciel

o Careiro, M. F. Silva e? al. 373 Rio Souza s.n. (INPA).?Para: estrada (INPA); Porto Trombetas, UHE de Samuel, are? do futuro

104 (INPA, RB, UB).

leaves are unique to A. unifoliolata, and this species cannot be confused it is most similar to A. trifoliolata. The two any other of Andira. Morphologically are is uniformly species easily distinguished trifoliolate, vegetatively: A. trifoliolata whereas A. unifoliolata is uniformly unifoliolate. Andira unifoliolata also has flowers 6-7 mm long compared to 8-10 mm long in A. trifoliolata. Andira unifoliolata has useful Unifoliolate

with

timber.

Doubtful Andira

sect. Aristobulia

Andira

amazonum

and Excluded

Names

Bentham, Comm. legum. gen. 43. 1837.?LECTOTYPE, here des Andira amazonum Martius ex Bentham. = Watairea guianensis Aublet, Hist, ignated: t. Guiane 2: 302. 1775. 755, pi. Martius

ex Bentham,

Comm.

legum.

gen.

43.

1837.?TYPE:

Rio Negro, Martius s.n. (holotype: BR!; isotype: M). = Vatairea guianensis Hist. pi. Guiane 2: 755, t. 302. 1775; see also de Lima (1982). Andira

BRAZIL.

Aublet,

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

VOLUME 64

SYSTEMATICBOTANYMONOGRAPHS

118

Bentham, Comm. legum. gen. 44. 1837, nom americana Aublet, Hist. pi. Guiane, suppl. 9. 1775.

Andira

8: 26. 1936; see

(Aguair) Ducke, Ann. Acad. Brasil. Ciencias

Vataireopsis araroba also de Lima (1980). aubletii

superfl.

=

Vouacapoua

Brazil. Andira bracteosa Bentham, Comm. legum. gen. 43. 1837.?Type: Para, Sieber s.n. (holotype: BR!; isotypes: B-W, HAL). = Vatairea guianensis Aublet, Hist. pi. Guiane 2: 755, t. 302. 1775; see also de Lima (1982). Andira

cinerascens Martius,

in sched.; see also de Lima

(1982).

Andira gabonica B?illon, Adansonia 6: 219. 1866.?TYPE: Duparquet = Aganope gabonica (B?illon) Polhill, Kew Bull. 25: 269. 1971.

32 (holotype: P!).

Andira horsfieldii Leschenault, Ann. Mus. Nati. Hist. (Paris) 16: 481, t.12. 1810.?TYPE: = Euchresta Leschenault, Ann. Mus. Nati. Hist. (Paris) 16: 481, t.12.1810. horsfieldii rar. PL 31. 1838. 148, pi. (Leschenault) Bennett, jav. Andira

ex J. St. Hilaire, Diet. Sei. Nat. 2: 137. 1804, nom. superfl. = americana Aublet, Hist. pi. Guiane, suppl. 9. 1775.

racemosa Lamarck

Vouacapoua

Suriname. Am Andira villosa Kleinhoonte, Rec. Trav. Bot. Ne?rl. 22: 404.1925.?Type: bei dem Pulle 319 1920, Coppenameflusse Raleighstrom, Aug (holotype: U!).?The not it does not clearly does and the floral reveals that Andira, type belong morphology Itmatches a specimen from Peru (Loreto: Iquitos, fit any genus of Papilionoideae. San Juan, 19 Sep 1945, A. Ducke 1823), which Ducke described as Hymenolobium velutinum; however, it is not a species o? Hymenolobium (pers. obs.; H. C. de Lima, pers. comm.). Both specimens constituting the type collection have flowers but lack fruits; thus, it is impossible

to determine

their affinities.

Andira

Brazil. zehntneri Harms, Repert. Spec. Nov. Regni Veg. 17: 443. 1921.?Type: Bahia, Barra, Oct 1912, Zehntner 2097 (holotype: M; isotype: RB!). = Lonchocarpus sp.

Skolemora pernambucensis Arruda in H. Koster, Travels Brazil 498. 1816.?TYPE: un known.?It is not known whether Arruda's types are extant (Stafleu & Cowan 1976). The description of the anthelminthic properties of this species suggests that this name may apply to a species o? Andira.

ACKNOWLEDGMENTS This work was financially Research Council (SERC) supported initially by the Science and Engineering and the Royal Botanic Gardens, Kew, and completed with the support of the Royal Botanic Garden Edinburgh. The Gatsby Charitable Foundation funded research on intraspecific cpDNA polymorphism, carried out by Liz in Guyana was supported by The Commonwealth and Government of Guyana Iwokrama I am grateful to my Ph.D. advisers Roger Polhill, Robert Scotland, Chris Leaver, and Programme. the guidance of Brian Styles, Bennett, who supervised the start of this research. I particularly acknowledge

Caddick.

Fieldwork

Rain Forest Mike who

very sadly died before my

thesis work was

complete.

I also thank: Maureen

Warwick

(artwork), Robert Mill

ANDIRA

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119

work and cladistic analysis), Tony Cox (molecular work), (Latin diagnoses), Mark Chase (advice on molecular David Harris (numer Peter Gasson (fruit anatomy), Bente Klitgaard (pollen data), Iain Prance (biogeography), ous discussions and Barbara Richard Brummitt, Mark Watson, and reading an entire draft), David Mabberley, comments on in MacKinder (constructive (nomenclatural issues), Denis Filer (databasing), Peter Hollingsworth of Leucaena was an inspiration for this Colin Hughes (whose monograph traspecific cpDNA polymorphism), Serena Marner, Anne Sing, Alison Strugnell (help with ad and who collected Andira inermis inMexico), for invaluable help on my arrival ministrative (advice on all topics and particularly problems), Gwilym Lewis leaves and fruit of Mexican in Brazil), Lourdes Rico (who collected (who species), Clive Foster, Phil Griffiths work,

of Brazilian (discussion species), Eimear Nie Lughadha, expertly grew species of Andira), Ary Oliveira-Filho of Mex inMexico, discussion for fieldwork in Guyana), Mario Sousa (hospitality Charlie Stirton (arrangements ican species), Martin Cheek (who collected Andira inermis in Africa), Jeff Doyle (molecular systematics), Larry Jane Doyle, Paul Manos, (cladistics and critically reading portions of this manuscript), Kelly, Anne Bruneau (cladis (molecular systematics), Kevin Nixon Soreng, Stephan Maumont I particularly thank Andr? Carvalho for help and and Dada). For my fieldwork, enormous hospitality, Jim Ratter, Sam Bridgewater, and de Lima, Alvaro Cogollo, and also Haroldo Cavalcante I also Taiman dos For Fernando Herminio with fieldwork thank Ribeiro. Carvalho, Brito, ("Guga") help Felipe

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124

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APPENDIX 1 Restriction Restriction

site characters

study by Pennington

Character2 3

used

to assess

phylogenetic

Sites of species

position

not included

of Andira

Enzyme

Probe

Asel

regionb

Site Diagnostic

Mutation0

MB8+9

3.6=1.8+1.7 1.8 = 1.05+

For

Clade H (site absence)

4

Asel

MB8+9

5

Asel

MB11

2.0=1.7

6

Asel

MB11

3.7 = 2.0+1.7

MB 13

1.0 = 0.6 + 0.4

n.d.

+ n.d.

Clade

III (site presence)

Clade

I (site presence)

Clade m

(site absence) I (site absence)

Clade

7

Asel

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

5.2 = 4.0+1.2

Clade H (site presence)

MB8+9

3.6 = 3.2 + n.d.

Clade

19

HindHI

MB2

20

Hindm

34

XmnI

MB3

1.8 = 1.2 + 0.6

35

XmnI

MB11

0.96 = 0.6 + n.d.

I (site presence)

A. verm?fuga; A. humilis RTP 269 (site presence) Clade

I and Clade H

(homoplasy; 37

in the

(1995).

MB13

XmnI

3.5 = 3.4 + n.d.

Clade m

site absence) (site presence)

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

Character

striction

site, and its absence

is diagnostic

for clade H.

2003

125

ANDIRA

APPENDIX 2 Inferred

Plastome

Types

of Andira

Accessions

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

abbreviated:

= restriction site mutations characteristic of clade III; PI = restriction site mutations char of plastome site mutations characteristic of the monophyletic group I; PITH restriction group of (1995), but no muta plastome group I and clade II [these accessions were screened previously by Pennington tions diagnostic for clade II were screened]. acteristic

of clade II;m

acteristic

Species

Voucher

A. anthelmia

RTP 227

Herbarium CEPEC,

FHO, K

Population

Clade

Locality

ffl

Brazil:

Bahia;

14?19'S, A. anthelmia

RTP 281

CEPEC,

FHO, K

m

Brazil:

Bahia;

14?51'S, A. anthelmia

RTP 282

CEPEC,

FHO, K

m

39?21'W

Brazil:

39?04'W

Bahia;

14?51'S,

39?04'W

A. anthelmia

RTP 208

CEPEC,

FHO, K

m

Brazil: Bahia;

A. anthelmia

RTP 286

CEPEC,

FHO, K

m

Brazil: Bahia;

A. anthelmia

RTP 294

CEPEC,

FHO, K

15?06'S,

15?05'S,

in

RTP 217

A. carvalhoi

RTP 229

CEPEC,

FHO, K

pi/m

CEPEC,

FHO, K

pi

Brazil:

CEPEC,

FHO, K

m

Brazil:

AMC*

CEPEC

pi

Brazil:

39?06'W Bahia;

14?57'S, A. cordata

RTP 262

CEPEC,

FHO, K

pi/n

Brazil:

A. cordata

RTP 263

CEPEC,

FHO, K

ii

Brazil:

RTP 264

CEPEC,

FHO, K

n

39?01'W

Bahia; 12?07'S, 45?06'W Bahia;

12?07'S, A. cordata

39?01'W

Bahia;

15?10'S, A. carvalhoi

39?01,W

Bahia;

14?57'S,

RTP 233

39?24'W

Brazil: Bahia; 13?58'S,

A. carvalhoi

39?33'W

Brazil: Bahia; 15?06'S,

A. carvalhoi

39?33'W

45?06'W

Brazil: Bahia; 12?0rS,

45?06'W

A. cujabensis

RTP 467*

E,UB

A. cujabensis

RTP 474*

E,UB

ii

Brazil: Goi?s;

A. cujabensis

RTP 478*

E,UB

n

Brazil: Goi?s;

A. cujabensis

RTP 485*

E,UB

ii

Brazil: Goi?s;

n

Brazil:Goi?s; 16?44'S, 52^7^

16?52'S,

16?30'S,

16?44'S,

52^0^

52^3^

52?37'W

Species

VOLUME 64

SYSTEMATICBOTANYMONOGRAPHS

126

Voucher

Herbarium

Population

Clade

Locality

A. cujabensis

RTP 486*

E,UB

1

n

Brazil: Goi?s;

A. cujabensis

RTP 493*

E,UB

1

n

Brazil: Goi?s;

A. cujabensis

RTP 497*

E,UB

2

n

Brazil: Goi?s;

52?37'W

16?44,S,

16?44'S,

47*23^

13?55'S, A. cujabensis

RTP 498*

E,UB

ii

2

Brazil: Goi?s; 13?55'S,

A. cujabensis

RTP 503*

E,UB

ii

2

RTP 213

FHO, K

CEPEC,

1

in

Brazil:

RTP 236

FHO, K

CEPEC,

2

in

Brazil:

RTP 274

FHO, K

CEPEC,

3

m

39?03'W

Bahia;

15?10'S, A. fraxinifolia

47?23'W

Bahia;

14?12'S, A. fraxinifolia

47?23'W

Brazil: Goi?s; 13?55'S,

A. fraxinifolia

52?37'W

Brazil:

39?06'W

Bahia;

12?35'S,

41?16'W

A. fraxinifolia

RTP 280

CEPEC,

FHO, K

3

m

Brazil:

A. fraxinifolia

RTP 283

CEPEC,

FHO, K

4

m

Brazil:

A. fraxinifolia

RTP 318

CEPEC,

FHO, K

4

A. fraxinifolia

RTP 285

CEPEC,

FHO, K

5

m

Brazil: Bahia;

A. fraxinifolia

RTP 287

CEPEC,

FHO, K

5

m

Brazil: Bahia;

Bahia;

12?32'S,

14?56'S, ni

39?01'W

15?05'S,

15?05'S,

RTP 298

FHO, K

CEPEC,

6

in

Brazil:

RTP 302

FHO, K

CEPEC,

6

m

Brazil:

RTP 303

FHO, K

CEPEC,

6

m

Brazil:

RTP 249*

A. fraxinifolia

RTP 250

39?14'W

39?!^

Bahia;

17?30'S, A. fraxinifolia

39?33,W

Bahia;

17?30'S, A. fraxinifolia

39?33'W

Bahia;

17?13'S, A. fraxinifolia

39?01'W

Brazil: Bahia; 14?55'S,

A. fraxinifolia

41?14'W

Bahia;

39?!!^

CEPEC,

FHO, K

7

m

Brazl: Bahia;

CEPEC,

FHO, K

7

m

Brazil:

llo02'S,40?43'W Bahia;

11?02'S,40?43,W A. fraxinifolia

MS 889

K,SP

8

ni

Brazil: 23?S,

A. galeottiana

A. galeottiana

A. humilis

LR s.n.

MCh

s.n.*

RTP 239

CEPEC,

K

1

pi

Mexico:

E

2

pi

Mexico:

1

in

Brazil:

FHO, K

S?o Paulo; 38?W

Oaxaca; 17?N, 95?W

Chiapas; 17?N, 91?W Bahia; 38^7^

12?15'S, A. humilis

RTP 246

CEPEC,

FHO, K

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

2003

ANDIRA

Voucher

Species

Herbarium

127

Clade

Population

Ahumilis

RTP 267*

CEPEC,FHO,K

2

PI

Ahumilis

RTP 268*

CEPEC,

FHO, K

2

PMI

Locality

Brazil:Bahia; 12?10'S, 44^3^ Brazil:

Bahia; 44?43'W

12?10'S,

CEPEC, FHO,K

RTP 269

Ahumilis

PI

2

Brazil: Bahia; 12?10'S, 44?43'W

RTP 499*

Ahumilis

PI 3

E, UB

Brazil:

Goi?s; 47?23/W

13?55'S, RTP

A. humilis

PI 3

E, UB

500*

Brazil:

Goi?s; 47^3^

13?55'S, RTP

Ahumilis

PI3

E, UB

501*

Brazil:

Goi?s; 47?23'W

13?55'S,

RTP 502*

Ahumilis

PI 3

E, UB

Brazil:

Goi?s; 47?23'W

13?55'S, Ahumilis

RTP 506*

E, UB

Ahumilis

RTP 507*

E, UB

PI 4

Brazil:

Goi?s; 14?07,S,47?31/W

PI4

Brazil:

Goi?s; 14?07,S,47?31'W

A. humilis

RTP 508*

E, UB

PI 4

Brazil:

Goi?s; 14?07'S, 47?31'W

RTP 509*

A. humilis

PI 4

E, UB

Brazil:

Goi?s; 47?31'W

14?07'S, A.humilis MdS

1*

PI 5 Brazil:

E

D. F.; 47?50'W

15?55'S, Ahumilis

2*

MdS

E

PI 5

Brazil:

D. F; 47?5(TW

15?55'S, MdS Ahumilis

3*

E

PI 5

D. F. ;

Brazil:

47?5(TW

15?55'S, 4*

MdS Ahumilis

MdS

Ahumilis

E

PI 5

E

5*

D. F.; 15?55'S, 47?5(W

Brazil:

5 PI

D. F.;

Brazil:

47?50/W

15?55'S, TDP13558

inermis A.

1 K

I

Costa Rica: Puntarenas; 83?19/W

09?07'N, A.m^w CEH1673

FHO, K

2

3579

K 3

I Mexico:

Oaxaca; 16?22/N,94?11'W

miroi?A.

MC

A.

inermis

merm?5

RTP 512*

A.

RTP 580*

I4

E, QCNE

E, INBIO

I

4

I

Cameroon,

SW Province;

03?58'N,

09?16'E

00?33'S,

Napo; 77?50^

Ecuador:

Rica: Puntarenas; 83^0^

Costa

08?42,N, A.

?n*/row

RTP

589*

E, INBIO

5 I

Rica: Puntarenas; 83?3(TW

Costa

08?42'N, A. te^a/w

RTP

305

CEPEC,

FHO, K

1

HI

Brazil:

Bahia;

17?30'S,

Alegalis

RTP 307

CEPEC,FHO,K

1

ffl

39?!^

Brazil:Bahia; 17?30'S,

39?!!^

128

VOLUME64

SYSTEMATICBOTANYMONOGRAPHS

Species

Voucher

A.

legalis

RTP 308

legalis

HCL

Herbarium

FHO, K

CEPEC,

Population

Clade

Locality

ffl

Brazil: Bahia;

m

Brazil: Rio de Janeiro; 22?55'S, 43?10'W

17?30,S,39?11'W A.

s.n.

E

A. macrothyrsa

TDP 13550

K

PI

Peru: Loreto; 00?S, 73?W

A. macroihyrsa

RTP 523*

E,QCNE

PI

Ecuador: 00?40'S,

A. marauensis

AMC

s.n.

CEPEC

ffl

Brazil:

Bahia;

15?09'S, A. nilida

RTP 300

CEPEC,

FHO, K

ffl

Brazil:

A. niiida

RTP 304

CEPEC,

FHO, K

ffl

Brazil:

A. niiida

RTP 313

CEPEC,

FHO, K

CEPEC,

FHO, K

ffl

Brazil:

RTP 288

CEPEC,

FHO, K

ffl

Brazil:

39?14'W Bahia;

13?02'S, A. nitida

RTP 289

CEPEC,

FHO, K

ffl

Brazil:

RTP 292

CEPEC,

FHO, K

ffl

Brazil:

RTP 301

CEPEC,

FHO, K

ffl

Brazil:

A. nitida

RTP 316

CEPEC,

FHO, K

ffl

Brazil:

A. nitida

RTP 237

CEPEC,

FHO, K

ffl

Brazil:

CEPEC

ffl

Bahia;

Brazil:

RTP 433

A. surinamensis

RTP 458

FHO, K, U, US

ffl

A. surinamensis

RTP 462

FHO, K, U, US

ffl

A. surinamensis

RTP 463

FHO, K, U, US

ffl

FHO, K, U, US

ffl

39?06'W

Bahia;

15?18'S, A. surinamensis

39?13/W

Bahia; 15?06'S, 30?\1^

15?10'S,

AMC 3309

38?23"W

Bahia;

17?30'S,

A. nitida

38?23'W

Bahia;

13?02'S, A. nitida

38^3^

Bahia;

13?02'S, A. nitida

39?!^

Bahia;

17?13'S, A. nitida

39?!^

Brazil: Bahia; 17?30'S,

III

39?13'W

Bahia;

17?30'S,

RTP 311

39?05'W

Bahia;

17?30'S,

A. niiida

Napo; 77?10'W

39?04'W

Guyana:

0A?\3^, 58^8^ Guyana: 04?13'N,

58*58^

Guyana: 0A?\3f"H, 58^8^ Guyana: 04?09,N,

59^2^

A. verm?fuga

RTP 256*

CEPEC,

FHO, K

PI

Brazil: Bahia;

A. verm?fuga

RTP 257*

CEPEC,

FHO, K

PI

Brazil:

11?15/S,42?52,W Bahia;

11?15,S,42?52,W A. verm?fuga

RTP 258*

CEPEC,

FHO, K

PI

Brazil: ll015'S,

A. verm?fuga

RTP 259

CEPEC,

FHO, K

PI

Bahia; 42^2^

Brazil: Bahia; 11?15'S, 42^2^

ANDIRA

2003

Herbarium

129

Species

Voucher

A. verm?fuga

RTP 265

CEPEC, FHO,K

2

PI

A. verm?fuga

RTP 270

CEPEC,FHO,K

3

PI/H

Population

Clade

Locality Brazil: Bahia; 12?07'S, 45?06'W Brazil: 12?07'S,

A

RTP 271

verm?fuga

FHO, K

CEPEC,

PI

3

Brazil:

Bahia; 43?15'W

Bahia; 43?15'W

12?07'S, 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, RTP 465*

A. verm?fuga

E,UB

PI

4

43?15'W

Brazil: Goi?s; 16?52'S,

52?20'W

E,UB

4

PI

Brazil: Goi?s;

E,UB

4

PI

Brazil: Goi?s;

verm?fuga

RTP 471*

E,UB

4

PI

Brazil: Goi?s; 16?52'S, 52?20'W

A. verm?fuga

RTP 504*

E,UB

5

PI

Brazil: Goi?s; 14?07'S, 4713^

A. verm?fuga

JR 7232V*

E

5

PI

Brazil: Goi?s;

A. verm?fuga

RTP 466*

A. verm?fuga

RTP 469*

A

16?52'S,

16?52'S,

14?07'S,

NUMERICALLIST OF SPECIES 1. A.

inermis

la. A.

inermis

subsp.

not determined) (subspecies inermis

14. A. legalis 15. A. ormosioides

lb. A.

inermis

lc. A.

inermis

subsp. glabricalyx subsp. rooseveltii

17. A. nitida

2. 3.

A. jaliscensis A. multistipula

4.

A. cubensis

5.

A.

16. A. fraxinifolia 18. A. marauensis 19. A. carvalhoi

6.

20. A. parviflora 21. A. cujabensis 22. A. cordata

7.

23. A. micrantha

9.

A. verm?fuga 10. A. humilis

25. A. grandistipula 26. A. praecox

12. A.

11. A. macrocarpa surinamensis

28. A.

13. A. anthelmia

29. A. unifoliolata

taurotesticulata

A. macrothyrsa A. chigorodensis 8. A. galeottiana

24. A. cori?cea

27. A.

tervequinata trifoliolata

52^0^

52^0^

47?13'W

SYSTEMATICBOTANYMONOGRAPHS

130

VOLUME 64

INDEXTONUMBERED COLLECTIONSEXAMINED The numbers Species

presented

in parentheses above.

refer to the corresponding

3775 (la). 1070 (lb). P., & J. A. Cede?o 7430 (3). P., et al. 8127 (28); 8357 (28).

species

A., & W. Rodrigues F., et al. 3102(3).

Acevedo,

P., & A. Sinca

Aubreville,

Acevedo,

R., & J. Sosa

Ayala,

Acevedo-Rodr?guez,

Acevedo-Rodr?guez, Ackerly, D., et al. INPA/WWFl302.4500.2

Adams, C. D. 7843 (la). Aguilar, R. 228 (la). Alain 4404 (4). Albert de Escobar, L., & J. I. Santa 3527 Albert

de Escobar, H. A.

(la);

13885

(12).

8233 ?feL. Delgado 133 (la). C, et al. 700 (la). G.,

(12).

B.

Azuara,

(20).

Azurdia,

BAFOG 51N (24); 1183 (12); 1201 (24); 1264 (24).
L. H.,

Bailey,

149 (la);

52 (la);

321

(la); 1352 (12); 1657 (12).

(la).

L., et al. 1848 (5).

13367

681

List of

Aymard,G. 6147 (27); 7991 (12); 9263 (28). Aymard,

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

Alencar (RB 54889) (12); (RB 61016) (9). Allard,

in the text and in the Numerical

(la).

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

393 (la); 432 1090 (la). Barbosa, C, et al. 1856 (la). Barkley, F. A., & M. Hern?ndez Barlow, F. D., 30/186D (8).

D., et al. 194 (9). D. L. 145 (21). A. M. A., et al. 347 Amorim,

Barreto,M. 937 (16); 1515 (16); 1782 (16); 5485 (10); 5491 (10); 5590 (16); 5592 (16); 5608 (16); 5764 (16); 10096 (16).

J., & T. S. dos Santos

Almeida,

103 (17);

130 (17).

Alvarenga, Amaral,

(17); 348

(17);

1055

(16); 1238 (18); 1411 (18); 1896 (16); 1908 (18). C. W.

Anderson,

332

W. R., et al. 7159

(10); 36024

(16); 36607

(22); 36865 (10). Andrade-Lima

& Medeiros-Costa

105 (17);

154 (17).

et al. 7 (16). F. W. 464 (le).

Andrade-Lima Andrews,

Bartlett, H. H.,

(9);

3441 (9). Arana, A. 27 (la).

Araujo, D. 1843 (13); 3553 (16); 4612 (14); 5289 (16); 6176 (16); 6598 (16); 7390 (16); 8079 (14); 8295 (16); 9687 (16); 9763 (16). 4874 (16). D., & R. Henriques 3683 (16); 4044 D., & N. C. Maciel

Beard,

Araujo,

(13);

6031 (16). D., & A. Magnanini D,. & J.Mauro 8614 (14). D., & A. L. Peixoto 324 (17).

Araujo, D., et al. 3461 (16); 8326 (14); 8597 Archer, W. A. 2050 (la); 2563 (la). L. 2805 (la). Aristeguieta, Atchinson,

E. 276

(4).

16319

(la).

(la).

(la).

S. G.

18686 (la). S. G., et al. 19556 Becker, R. M. 42 (21). Beck,

(3).

Belem, R. P. 1830 (14); 1879 (13). R. P., ?feR. S. Pinheiro

Belshaw,

C. M.

Bena,P.

1167(12).

2445

3370

(16); 2550

(16);

(la).

A. 3658 (lb); 3796 Benitez-Paredes, Bernai, H. Y. 158 (la). Black, G. A. 54-17372 (la). Black, Black,

(lb).

G. A., & Klein 54-17197 (la); 54-17372 (la). G. A., & P. Ledoux 50-10366 (la); 50-10386

(12). Black,

4259 (16);4465 (16); 7514 (16); 8700 (14). Araujo,

40602

2591 (16); 2807 (17); 2949 (13); 2988 (17); 3089 (18); 3091 (18); 3142 (18).

L. M. 3-7 (la). N., & M. Heinrich GUI224 (8). Antonio, T. 3616 (la). da Silva, M., et al. 1352 (22); 2738 Aparecida

Araujo,

(la).

J. S. 157 (la). P. 1336 (la).

Beck,

Andrews,

Araujo,

?feT. Lasser

J. H. 6332

Beaman, Beard,

Belem,

Antonio,

Araujo,

W. N.

C.

Bautista,H. P. 533 (17); 1145 (17); 1368 (17).

(12).

Anderson,W. R. 6231 (10); 6456 (10); 6771 (21); 7150 (21); 7293 (21); 7294 (9); 7895 (10); 9560 (21); 9691 (21); 10051 (21); 11292 (la); 11293 (21); 11361 (21); 36797 (22). Anderson,

Bangham,

Barbosa,

G.A.,

et al. 54-16454

(12)

Blanchet, J. S. 607 (13); 2723 (16); 3089 (16); 3137 (10); 3672 (16). Boege, W. 1244 (8). Bohrer, C. 47 (16). Bonder, G. 2465 (16).

(14).

Bonpland 1599 (la); 3901 (la). Boom,

B. M.

Boom,

B. M., B. M.,

Boom,

6738

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VOLUME 64

SYSTEMATICBOTANYMONOGRAPHS

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2003 ANDIRA 139

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Woodworm,

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25531

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17899

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(la); 28788 (10); 25676

(5). (7).

(la). (10).

2003

ANDIRA 141

INDEXTO SCIENTIFICNAMES are in roman

names

Accepted

type; the main

(Baill.) Polhill

118

frondosa Mart, ex Benth. 80 var. longifoliolata N. F. Mattos gabonica Baill. 118

Agouti paca 14 Amerimnum affine Spreng. Juss. 3

44

Lam.

117

Benth.

sect. Euandira sect. Lumbricidia

grandistipula

(Veil.) Benth. 3, 22, 31, 36 N. F. Mattos 3, 22, 31, 36 3, 31, 36 (Veil.) N. F. Mattos

subsect. Glabratae

N. F. Mattos

acuminata

Benth.

amazonum

Mart,

anthelmia

(Veil.) Benth.

3, 31, 36

117

4, 5, 6, 11, 12, 13, 15, 16,

(Veil.) J. F. Macbr. (Veil.) Toledo 76

38, 42, 43, 69-72,76, var. cordata N. F. Mattos inermis

N. F. Mattos Benth.

carvalhoi

82

85

118 4, 9, 10, 12,

49, 50,

Perr.

?feA. Rich.)

Polhill

4, 21,

33,

43, 44,45, 49-50,129 var. riedelii Benth. 44 jaliscensis

R. T. Penn.

5, 9, 10, 12, 21, 22, 23, 24,

25, 27, 28, 32, 33, 37, 38, 39, 42, 49, 50-52,

53, 129

R. T. Penn.

5, 8, 9, 10, 21, 23, 29,

laurifolia

ex R. T. Penn. & H. C. Lima 4, 5,

Pulle 4, 5, 13, 16, 21, 27, 29, 35, 37, 40,

41, 56, 59, 60,10?-108,129 cubensis Benth. 8, 16, 21, 23, 33, 34, 37, 40, 42, 129 53, 55-56, Benth.

4, 5, 7, 12, 13, 17, 18, 21, 22,

23, 24, 25, 27, 28, 32, 33, 34, 35, 37, 39, 41,

2, 4, 5, 7, 8, 9, 10, 11, 12, 13,

14, 15, 16, 17, 18, 20, 21, 24, 25, 26, 27, 28, 30, 32, 33, 37, 39, 42, 84-S9,126,

legalis

Urban

42

N. F. Mattos

129

Benth.

66, 68

101 69, 71, 72

(Veil.) Toledo

4, 5, 6, 12,13,

14, 18, 21,

24, 25, 26, 27, 28, 29, 30, 32, 33, 36, 37, 38, 41, 78, 80-82, 84, 88, 127, 128, 129 var. bahiensis

76 (Benth.) N. F. Mattos R. T. Penn. 14, 21, 29, 33, 37, 39, 41, 72-73, 75, 76, 129 Ducke 4, 5, 8, 10, 12, 13, 21, 24, 25, macrothyrsa macrocarpa

26, 28, 29, 30, 32, 37, 40, 41, 56, 59, 60-62, 64, 128, 129 marauensis

100-102,103,105,125,126,129 Kunth 44 Benth.

jamaicensis kuhlmannii

lanei N. F. Mattos

9, 10, 18, 20, 21, 24, 25, 26, 28, 30, 32, 33, 129 34, 35, 37, 40, 41, 100,102-105,125,

fraxinifolia

51, 53, 56, 62,

(Guillem., subsp. grandiflora Gillett ex Polhill 44,48 (de Wild.) subsp. rooseveltii

37, 38, 40, 41, 61, 62-64,129 chiricana Pittier 44, 48 cinerascens Mart. 118

excelsa

12,

4*49,129

125, 129

cujabensis

1, 2, 3, 4, 5, 7, 8,10,

R. T. Penn. 21, 33,43,44,45,

subsp. glabricalyx

27, 28, 30, 32, 33, 34, 37, 40, 41, 92, 94-98,

cori?cea

127, 129

103,126,

62,129

76

R. T. Perm. & H. C. Lima

cordata Arroyo

92, 69

inermis 4, 21, 29, 33, 43, 44-48,

subsp.

13, 14, 15, 16, 17, 18, 19, 20, 21, 24, 25, 26,

chigorodensis

DC.

(W.Wright)

76

117

bracteosa

4, 5, 8, 12, 14, 16, 17, 18,

33, 34, 36, 37, 40, 42-50, 124, 127, 129

118

bahiensis

44

13, 14, 15, 16, 17, 24, 25, 26, 27, 28, 30, 32,

var. gracilis N. F. Mattos 85 var. ormosioides (Benth.) Benth.

Aguiar Benth.

?feA. Rich.

5, 15, 21, 24, 25, 26, 28,

19, 21, 23, 24, 25, 26, 28, 30, 32, 33, 35, 37,

ex Benth.

Benth.

Perr.

Amshoff

118 horsfieldii Lesch. humilis Mart, ex Benth.

44

18, 20, 21, 22, 24, 25, 26, 27, 28, 29, 30, 32,

araroba

68, 72, 76,

29, 30, 32, 34, 37, 38, 41,106,108-109,129 handroana N. F. Mattos 85

33,34,37,38,41,76-30,82,84,88,125,129 var. acuminata Benth. 76

aubletii

12, 15, 21, 23, 24, 25, 26, 28,

Guillem.,

grandiflora

3, 31, 36

sect. Paucifoliolatae subsect. Lumbricidia

anthelmintica

76

126, 129

sect. Aristobulia

anthelmia

84

30, 32, 33, 34, 37, 39, 41, 64-66,

36-37

anthelmia

Standl.

galeottiana

Andira

Andira

85

var. latifoliolata N. F. Mattos 85 var. rosea (Mart, ex Benth.) Benth.

Aganope gabonica

are in italics.

Synonyms

var. lanceata N. F. Mattos

15

Aglaia

is in boldface.

entry for each

N. F Mattos

5, 21, 23, 24, 25, 27, 28, 129

32, 37, 40, 42, 92, 93-94,128, micans Taub, ex Glaz. 64 micrantha

Ducke

5,15,

16, 21, 25, 27, 29, 35, 37, 108, 129

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

SYSTEMATICBOTANYMONOGRAPHS

142

microcarpa

Griseb.

55

Dalbergieae

multistipula

Ducke

5, 8, 21, 23, 24, 25, 27, 28, 29,

Dasyprocta

32, 37, 38, 41, 43, 51,5^-54,129 var. peruana N. F. Mattos 60 nitida Mart, ex Benth. 4, 5, 9,10, 11, 12,13,

Benth.

subsp. aurantica

18,

12, 13, 15, 20,

4, 5, 8, 9,11,

1, 2, 33 14

Diptychandra aurantica Tul. 36

21, 24, 25, 26, 27, 28, 30, 32, 33, 37, 38, 40, 42, 89-93, 94, 97, 128, 129 ormosioides

Krug & Urb. Euchresta

80, 82-S4, 88, 129 Benth. 66

Geoffroea

Ducke

4, 5, 21, 24, 25, 26, 28, 30, 32,

34, 37, 39, 41, 98-100,129 parvifolia Mart, ex Benth. 84 Benth. 69, 71 N. F. Mattos pernambucensis pisonis Mart, ex Benth. 84

pauciflora

ex R. T. Penn.

Arroyo

114, 115, 129 37, 40, 42,109-112, racemosa Lam. ex J. St. Hil. 2, 3, 118 retusa (Poir.) DC. 74

riparia Kunth 44 rosea Mart, ex Benth. sapindoides spinulosa

surinamensis

(Bondt)

73

25, 27, 28, 29, 32, 37, 39, 40, 41, 56-60,

72,

?feN. Cuello

5,

109, 111,112-114,

Ducke

3, 5, 10, 21, 22, 25, 27, 29, 31, 35, 37, 38, 40, 111, 114-115, 117, 129 unifoliolata Ducke 3, 5, 6, 7, 13, 21, 22, 24, 25, 111,

(Mart.) Benth.

4, 5, 10, 12, 13, 17, 18,

19, 21, 24, 25, 26, 28, 30, 32, 33, 37, 39, 42, 76, 88, 102, 103, 128, 129 118

65, 66-69,72, villosa Kleinhoonte wachenheimii

Benoist

Harms

Caesalpinoideae 12 Cleogonus

118

15, 97

Coursetia caribaea

Seem.

(Jacq.) Lavin var. caribaea 34

107

1

49

var. novogaliciensis var. viridiflora 49 Lonchocarpus staudtii Harms

44

Lumbricidia

Veil.

36

Veil.

76

legalis Veil. Meliaceae 15

Lavin & Sousa 49

36, 80

Millettia rooseveltii

de Wild.

Mimosoideae

115-117,129

zehntneri

viridiflora

anthelmia

129

verm?fuga

118

15, 36, 49, 97

Leguminosae Lennea

106, 108, 129

26, 28, 30, 31, 32, 34, 36, 37, 38, 40,

Ducke

J. R. Forst. & G. Forst.

2, 4, 5,

var. ovatifoliolata N. F. Mattos 74 taurotesticulata R. T. Penn. 9, 10, 16, 21, 23, 24,

trifoliolata

15, 97

Kielmeyera rubriflora71

76

21, 27, 29, 37, 40, 42,

44

21, 24, 25, 26, 28, 30, 32

velutinum Inocarpus

R. T. Penn., Aymard

Ruiz & Pav. ex G. Don

nitidum

ex Amshoff

66, 69

Benth. 1, 2, 20, 23, 33, 118 Hymenolobium flavum 21, 24, 25, 26, 28, 30, 32

84

Splitg.

74

71

Hymenaea courbaril L.

6, 7, 13, 15, 16, 18, 21, 24, 25, 26, 28, 30, 32, 34, 37, 39, 42, 72, 73-76, 128, 129

tervequinata

Glycyrrhiza undulata Guttiferae

(DC.) Benth. 44, 48 (Mart.) Benth. 66

stipulacea Benth. 76 var. bahiensis Benth.

118

acutifolia Stokes 42 inermis W. Wright 42, 44 Stokes 72 obtusifolia

verm?fuga Mart.

5, 21, 27, 29, 35,

var. oblonga Benth. 74 var. surinamensis (Bondt) DC.

(Lesen.) Benn. Jacq. 2, 4

spinulosa Mart. 66, 69 surinamensis Bondt 73

var. emarginata N. F. Mattos 85 var. puberula N. F. Mattos 85 praecox

Carvalho

ex Taub. 4

Rich. pubescens retusa Poir. 74

85

(Tul.) Lima,

Dussia

horsfieldii

paniculata

36

subsp. epunctata 36

21, 24, 25, 26, 27, 28, 30, 32, 33, 37, 39, 42,

parviflora

VOLUME 64

Myoprocta

14

Ostryoderris brownii Hoyle

49 1, 36, 49,

Papilioniodeae Parkia biglobosa multijuga Pisum

49

15

50 Benth.

15

sativum L. 29 Poeppigia procera C. Presl

69

118

& Costa

2003 ANDIRA 143

Jacq. 4 sapindoides DC. 44

anthelmia

(Veil.) Kuntze 76 (Benth.) Kuntze 55 100 (Benth.) Kuntze cujabensis (Benth.) Kuntze 84 fraxinifolia

Pterocarpus

cubensis

Pterodon emarginatus Robineae 49

35, 36

Vogel

frondosa humilis

Skolemora pernambucensis 33

Arruda

118

laurifolia

Trachypogon

guianensis

80

(Benth.) Kuntze 69 (Veil.) Kuntze 80 (Benth.) Kuntze 66 paniculata

Sophoreae

plumosus Vatairea

inermis

(Mart, ex Benth.) Kuntze (Benth.) Kuntze 69 (W.Wright) Lyons 42

legalis

112 Aubl.

117, 118

Vataireopsis araroba (Aguair) Ducke

Vigna 124 Aubl. 2 Vouacapoua americana Aubl. 2

118

(Mart, ex Benth.) Kuntze (Poir.) Kuntze 74 (DC.) Kuntze 44 sapindoides

pisonis retusa

spinulosa (Mart.) Lyons 66 surinamensis (Bondt) Kuntze verm?fuga

(Mart.) Kuntze

66

73

84