dichapetalaceae_rhabdodendraceae 1972

FLORA NEOTROPICA

Monograph No. 10

Dichapetalaceae by

Ghillean T. Prance

Monograph No. 11

Rhabdodendraceae by

Ghillean T. Prance

TROPIC Of CANCER

FLORA NEOTRO P I C

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April 20, 1972

FLORA NEOTROPICA Published for

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FLORA NEOTROPICA

Monograph No. 10

Dichapetalaceae

by Ghillean T. Prance

C^s I ?1TROPIC OP CANCER

'~--- ~ ~ ~C ---- --.--------------------

FLORA NEOTROPICA

TROPIC OF CAPRICORN //

Published for Organization for Flora Neotropica

by Hafner Publishing Company

New York April 20, 1972

Copyright ? 1972

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A MONOGRAPH OF THE NEOTROPICAL DICHAPETALACEAE GHILLEAN T. PRANCE

INTRODUCTION

The present study is confined to the 41 American species of Dichapetalaceae. Although there are comparatively few species of this family in the American tropics, they are, nevertheless, widespread. They range from Mexico and the Caribbean to eastern-central Brazil, with all 3 genera represented and one confined to America. The

Dichapetalaceae is most abundant in Africa where there are about 150 species mostly of

Dichapetalum, and 7 of Tapura. There are also 17 species of Dichapetalum widespread in Malesia.

HISTORY OF AMERICAN DICHAPETALACEAE

The first genus of Dichapetalaceae to be described was Tapura, from French Guiana, by Aublet (1775). He placed Tapura in the order Pentandria Monogynia of the Linnean sexual system, and he described and figured one species, T. guianensis. Aublet's illustration of the flower is not accurate, but since there is adequate type material of T.

guianensis, this does not present any problem in the definition or typification of the

genus. Du Petit Thouars (1806) described two genera from Madagascar, Dichapetalum and

Leucosia, which he placed in the family Terebinthaceae. Leucosia has since been regarded as a synonym of Dichapetalum by all more recent workers in the family,

Vahl (1810) published the genus Symphyllanthus which included two species, S.

rugosus and S. glaber, both from the Guianas. Vahl's two species are in fact species of

Dichapetalum. The name Symphyllanthus has not been used in the literature, except by Gleason (1924), and must be regarded as a synonym of Dichapetalum.

A.P. de Candolle (1811a, 1811b) described the new genus Chailletia based on two

species from French Guiana, C. pedunculata and C. sessiliflora. His description was based more on C. pedunculata since he examined mature flowers of only that species. Both of De Candolle's species were equivalent to species already described in other genera. Chailletia pedunculata is the same as Symphyllanthus glaber of Vahl, but when transfered to Dichapetalum the De Candolle specific epithet must be used, and Chailletia sessiliflora is synonymous with Tapura guianensis Aublet. De Candolle published his description of Chailletia in two places both in 1811. It is hard to determine which was first, but 1811a was probably the first since it was published in January; 181 lb is a much more detailed and illustrated description.

Many species of Dichapetalum were originally described in the genus Chailletia, but have since been transferred to Dichapetalum in more recent publications because

Dichapetalum is one of the two oldest names for the genus (The other, Leucosia, was described at the same time as Dichapetalum. Since Dichapetalum has been adopted by all later authors in preference to Leucosia there are no reasons for changing this situation.)

Robert Brown (1818) was the first to relate the two genera Tapura and

Dichapetalum, and to recognize the group as a family which he named Chailletieae (Chailletiaceae). Brown treated two genera in his family Chailletieae: 1. Chailletia in which he included Dichapetalum and Leucosia of Du Petit Thouars, and Mestotes of Solander; and 2. Tapura of Aublet. He stated that his reason for adopting the name

1 B. A. Krukoff Curator of Amazonian Botany, The New York Botanical Garden.

3

4 Flora Neotropica

Chailletia in preference to the earlier name Dichapetalum was because the name

Dichapetalum was derived from a character not present in the whole of the genus. Similarly he rejected the name Leucosia because it had already been applied to a genus of crustaceans. Neither of these reasons are legitimate according to the present code of botanical nomenclature, and Baillon (1874) was correct to adopt the name Dichapetalum for this genus.

A.P. de Candolle (1825) in the 'Prodromus' adopted Brown's family Chailletiaceae, and he included three genera; Chailletia with five species, one from America, one from Asia, and three from Africa and Madagascar; Leucosia with the one original species of Du Petit Thouars; and Tapura with a single species T. guianensis. Thus De Candolle did not follow Brown in treating Leucosia as a synonym of Chailletia.

The genus Patrisia was described in manuscript by Von Rohr, but it was never

published by him. The name was taken up by Steudel (1840) but was not described, and is an illegitimate name for Dichapetalum. The Von Rohr specimen at the British Museum bears Rohr's manuscript description of his genus, which was never validly published.

Endlicher (1840), included 3 genera, Moacurra, Chailletia, and Tapura, in his

synopsis of the Chailletiaceae. Moacurra was a Malesian genus described earlier by Roxburgh and later united with Dichapetalum. Endlicher cited all the synonyms of Chailletia already mentioned here, i e Symphyllanthus, Leucosia, Dichapetalum and Mestotes.

Poeppig & Endlicher (1842) proposed the new genus Stephanopodium which they placed in the Chailletiaceae. This new genus was based on the single species, S.

peruvianum. In the same work they also described a new species of Tapura, T. amazonica. Both of their species are still recognized today in this work. Endlicher (1843), in the

supplement to his 'Genera plantarum', added the genus Stephanopodium to the genera listed previously.

Bentham (1853) discussed the genus Tapura and gave descriptions of the five species known to him, two of which were new. He described the floral morphology of Tapura accurately and he corrected the mistakes of several earlier authors. He also listed the few

species of Chailletia (=Dichapetalum) from the New World known to this date. Bentham concluded that the family Chailletiaceae is related to the Celastraceae and Aquifoliaceae, but some years later Bentham & Hooker (1862), placed the Chailletiaceae in their order Geraniales next to the Meliaceae. They included three genera, Chailletia, Stephanopodium, and Tapura.

Baillon (1874), in his 'Histoire,' considered the Dichapetalaceae as part of the family Euphorbiaceae under his 'Serie des Dichapetalum'. Baillon recognized three genera, Dichapetalum, Stephanopodium and Tapura. He adopted the earlier name Dichapetalum rather than Chailletia, thus correcting those authors who had used the more recent name Chailletia. He cited the following names as synonyms of Dichapetalum: Chailletia, Leucosia, Symphyllanthus, Moacurra, Mestotes, Walhenbergia, and Plappertia. At a later

date, Baillon (1886), considered the group as the separate family Dichapetalaceae when he treated it in Martius' 'Flora Brasiliensis.' That account, which included 10 species in the 3 genera Dichapetalum, Tapura, and Stephanopodium, is an important summary of the Neotropical species and genera known to that date. Baillon established some new

synonymy, and he described 5 new species. Engler (1896), in his 'Pflanzenfamilien,' considered the Dichapetalaceae related to

the Euphorbiaceae subfamily Phyllanthoideae. He also recognized the 3 genera: Dichapetalum, Stephanopodium and Tapura, all of which he described and figured accurately. He listed 73 species of Dichapetalum from around the tropics, 4 species of

Stephanopodium from the Neotropics, and 5 species of Tapura from America and Africa. Ule (1906) described the new genus Gonypetalum for his new species G. juruanum;

later authors added three more species to this genus, but more recently it has been reduced to synonymy under Tapura, an opinion with which I concur.

Introduction 5

Engler & Krause (1931), revised Engler's earlier account of the family, and presented further details. They divided Dichapetalum into 4 sections all of which had been described in earlier publications of Engler on the African Dichapetalaceae. Since all the American species fall into their section Eudicbapetalum (which should be called section

Dichapetalum to follow the rules of nomenclature), these sections are not discussed here. The other three sections were all for African species. Engler and Krause followed Baillon (1886) in recognizing 2 sections in Stephanopodium, and they divided Tapura into 3 sections, two of which are represented in the New World. The third was to accommodate a single African species. Engler and Krause also recognized the genus Gonypetalum as

separate from Tapura. During the present century the family has not been comprehensively monographed

in the New World, and the only significant accounts of it are those in the various regional floras, each of which considered a restricted part of the Neotropical range of the family. The most significant of these accounts are discussed briefly below. Gleason (1924), treated 4 species from Central America and the Caribbean. He adopted Vahl's name

Symphyllanthus for Dichapetalum, but there are no reasons for this since the name

Dichapetalum predates Symphyllanthus by 4 years. Macbride (1950), in his account of Peruvian Dichapetalaceae recognized 4 genera: Dichapetalum (4 species), Stephanopodium (1 species), Tapura (7 species), and Gonypetalum (3 species).

Lasser (1944) described the 3 species known at that time from Venezuela. Stafleu (1951) treated 2 species of Dichapetalum and 3 of Tapura in his account for the 'Flora of Suriname.' The most useful account of the family in recent years is that by Rizzini

(1952), in which he treated all the Brazilian species of Dichapetalaceae. Rizzini was the first to reduce Gonypetalum to synonymy under Tapura. He recognized 5 species of

Dichapetalum, 5 of Stephanopodium, and 8 of Tapura. He included keys for their

identification, and gave descriptions and notes about some of the most critical species. Hutchinson (1964), in his 'Genera,' included 4 genera in the Dichapetalaceae:

Dichapetalum, Tapura, Stephanopodium, and Falya. The latter was described by Descoings (1957) for a Madagascan collection. Breteler (1969) showed that Falya is in fact synonymous with Carpolobia (Polygalaceae).

Lundell (1966) gave a summary of the Mexican and Central American species of

Dichapetalum, and Lewis (1967) treated the two species known from Panama. Also during the present century the Dichapetalaceae has been variously placed in the

phylogenetic systems. This is discussed in the section on phylogeny which follows. There have been several accounts of the Dichapetalaceae in Asian and African regional floras which are not discussed in detail here. Dichapetalum, the only genus represented in Malesia was revised by Leenhouts (1956, 1957). Some of the most important accounts of African Dichapetalaceae are those by De Wildeman (1919), Moss (1928), and Hauman

(1955). The African Dichapetalaceae are presently being revised by F. J. Breteler of

Wageningen.

SYSTEMATIC POSITION OF THE DICHAPETALACEAE

The position of the family in the early pre-Darwinian systems has been given in the

history of the family. A summary of the numerous relationships suggested for this family follows

Bentham & Hooker (1862) GERANIALES near Celastraceae and Rhamnaceae

Engler (1896) EUPHORBIALES near Euphorbiaceae subfamily Phyllanthoideae Bessey (1907) GERANIALES near Euphorbiaceae and Polygalaceae Hallier (1921) LINALES united with Chrysobalanaceae and Trigoniaceae Wettstein (1935) TRICOCCAE near Euphorbiaceae subfamily Phyllanthoideae Gundersen (1950) EUPHORBIALES near Euphorbiaceae

6 Flora Neotropica

Melchior (1964) THYMELAEALES near Geissolomataceae, Penaeaceae, Thymelaeaceae, Elaeagnaceae

Cronquist (1968) CELASTRALES near Icacinaceae, Aquifoliaceae, and Cardiopteridaceae Takhtajan (1969) EUPHORBIALES near Euphorbiaceae subfamily Phyllanthoideae Hutchinson (1969) ROSALES near Rosaceae and Calycanthaceae

From this summary it can be seen that there has been a wide divergence of opinion as to the relationship of the Dichapetalaceae. Most of the above phylogenists have placed it with reservations, for example, Hutchinson said that he placed it in the Rosales "for want of a better place."

Engler (1896), Bessey (1907), Wettstein (1935), Gundersen (1950), and Takhtajan (1969) all relate Dichapetalaceae to the Euphorbiaceae, in particular to the subfamily Phyllanthoideae. This does not seem a satisfactory relationship because the flowers of the

majority of Dichapetalaceae are very little reduced. Hutchinson (1969), placed the family in the Rosales next to Rosaceae; however, as

stated above, it was placed in the Rosales with some reservation. This again does not seem a satisfactory relationship to me, because of the anatomy, the tendency to zygomorphy and a fused corolla, the reduced disc, and the tendency to polygamous flowers.

Hallier (1908) originally placed the Dichapetalaceae in his order Polygalales together with the Trigoniaceae, Polygalaceae, and Vochysiaceae. Later (1921), he united the

Dichapetalaceae with the Trigoniaceae and Chrysobalanaceae into a single family, Neither of these positions is satisfactory. The Dichapetalaceae are not closely related to the

Chrysobalanaceae, and anatomy and floral morphology certainly do not support Hallier's united family. The Dichapetalaceae may, however, be closer to the Trigoniaceae, a small

family usually placed in the Polygalales. There are striking similarities between

Dichapetalaceae and the Polygalales, but again the Dichapetalaceae does not fit well into that order, especially since the flowers are only slightly zygomorphic.

Melchior (1964), placed the Dichapetalaceae in his order Thymelaeales together with the Geissolomataceae, Penaeaceae, Thymelaeaceae, and Elaeagnaceae. This order in itself seems an unnatural group of families which probably do not belong together, and the

Dichapetalaceae does not fit near to the Thymelaeaceae. It may be closer to the Geissolomataceae which is placed in the Celastrales by both Cronquist (1968), and

Takhtajan (1969). Cronquist (1968) placed the Dichapetalaceae in his order Celastrales together with 9

other families including the Icacinaceae, Aquifoliaceae, Cardiopteridaceae, and Celastraceae. This to me seems by far the most natural position for the Dichapetalaceae. Cronquist stated that the Dichapetalaceae does not appear to have any close relatives in the Celastrales. However, in my opinion it is quite closely related to both the Icacinaceae and Celastraceae. The Dichapetalaceae probably represents a more advanced member of the order because of its lack of endosperm, tendency to zygomorphy and polygamous flowers. Most of the morphological features of Dichapetalaceae occur in at least some

genera of Icacinaceae, which in itself is rather a diverse family. The important floral characters of the Dichapetalaceae, such as the 3 locular ovary and the pendulous ovules, are all found in the Icacinaceae. The Icacinaceae, however, differ in the absence of

stipules and a disc, and in the presence of endosperm in most genera. The disc of the

Dichapetalaceae is extremely reduced in some species and is often represented only by small nectariferous glands, especially in the genus Dichapetalum.

As already stated above the Dichapetalaceae also have affinities with the primitive members of the Polygalales such as the Malpighiaceae. While I agree with the composition of both orders Polygalales and Celastrales of Cronquist, I disagree with his derivation of the Polygalales. Cronquist derives the Celastrales from the Rosales, and the Polygalales as a simple-leaved offshoot from the Sapindales. I consider both the Celastrales and the

Polygalales to be derived from the Rosales, and consequently they are more closely related than indicated by Cronquist. Hence the Dichapetalaceae has affinities with both orders.

Introduction 7

POLLEN - Fig 1

The grains colporate, small, 10-17 u on longest axis, slightly flattened in some species of Dichapetalum, spheroidal to oblong in other species. The sexine about as thick as the nexine; the walls without distinctive patterning to distinctly reticulate when studied by a light microscope, with considerable variation in reticulation pattern between the species.

The pollen is described briefly by Erdtman (1952). When studied by the light microscope it is of a rather generalized type without distinctive features which would

help to place the family. Erdtman makes the following comment based on the pollen: "Pollen grains similar to those in Dichapetalaceae occur in several families, e g Euphorbiaceae.... The grains in Convolvulaceae, Geraniales, Rosaceae, Thymelaeaceae, Aquilariodeae etc are more or less different." From this we may conclude that pollen does not support the views of Melchior (1964) that the Dichapetalaceae belong near to the Thymelaeaceae, neither those of Hutchinson (1969) who related them to the Rosaceae, nor of Hallier (1921).

In the phylogenetic section of this work it was suggested that the two most likely positions for the Dichapetalaceae are near to the Polygalales or Celastrales. Pollen would

certainly favor the Celastrales rather than the Polygalales which have rather a distinct

pollen type. In the Celastrales there is a wide range of pollen types; for example, the Icacinaceae are extremely eurypalynous, and the inclusion of the Dichapetalaceae in the Celastrales would not increase the pollen diversity at all.

However, in a recent publication, Lobreau (1969) described and discussed the pollen of all families that have been included in the Celastrales in the widest sense, in an attempt to redefine the limits of that order. He included the pollen of the Dichapetalaceae in his

study, and said: "Par sa forme, le pollen rapelle celui de certaines Icacinaceae du type de Stemonurus.... Le pollen des Dichapetalaceae, malgre une faible resemblance avec

quelques Icacinaceae, semble plus proche de celui de certaines Euphorbiaceae Phyllan- thoideae." In spite of these conclusions of Lobreau, I do not think that pollen provides sufficient evidence for excluding the Dichapetalaceae from the Celastrales, especially if the Icacinaceae are placed there. The pollen is nearer to that of certain Icacinaceae than to Euphorbiaceae.

The pollen of Dichapetalaceae is presently under investigation by Dr. W. Punt of Utrecht in connection with Breteler's work on the African species. I have sent to Dr. Punt

pollen of all three American genera of the family. Hence the above comments are

preliminary, and it is to be hoped that Dr. Punt's detailed study will provide suggestions for the phyllogenetic position of the family and for the relationships between the genera, as well as a detailed description of the pollen morphology. Dr. Punt tells me that he has

already found differences between the pollen of Tapura and Dichapetalum. Pollen material examined: Dichapetalum flexuosum, Cansdale 3976; D. rugosum,

Silva 59726; Tapura amazonica, Prance et al 5090; T. lanceolata, Ducke 412; Stephanopodium engleri, Magalhaes 140.

ANATOMY

General descriptions of the anatomy of the Dichapetalaceae have been published by Barth (1896), Solereder (1908), Engler & Krause (1931), Metcalfe & Chalk (1950). The

following descriptions are based primarily on these works, but have been verified by preparing slides of a few species of all 3 genera. While anatomy is of some use in

suggesting the relationships of the family, it is not of much use taxonomically below the

family level. The 3 genera are not anatomically distinct. This is particulary true of Tapura and Stephanopodium where the anatomy is extremely similar.

8 Flora Neotropica

:...:

A B C :>EiNi , ̂ 9 , :: : ::" :::"; *.;: *

D E F G

J

1 ' H '~K

FIG 1. Pollen and lamina of Dichapetalaceae. A-G, pollen, X 620. A-C, Tapura amazonica (Prance 5090). D-E, Tapura lanceolata (Ducke 412). F-G, Dichapetalum rugosum (Silva 59726). H-K, lamina. H-J, Dichapetalum pedunculatum (De La Cruz 3095). H, X 32; J, X 9; K, Tapura juliani (Klug 2010), X 9.

Introduction 9

LAMINA Fig 1.

Usually dorsiventral, simple, glabrous or with unicellular hairs often with conical, wart-like papillae. The epidermis consists of one to several layers of cells on both surfaces, and mucilaginous cells are common in the upper epidermis especially when it is more than one cell thick. Hypoderm is present in some species and it is frequently mucilaginous. Stomata are confined to the lower surface, and are rubiaceous. The mesophyll is extremely varied in thickness and content; the palisade cells are usually very short, and

large spaces between the palisade and spongy cells are frequent in many species; fibrous

mesophyll cells are common in many species and are derived either from the fibers of the vascular bundle which have detached from it or from the spongy mesophyll tissue where some cells thicken and lengthen. The midrib has a simple epidermis usually covered with a

hairy cuticle; the bundle structure is similar to that of the sterile petiole. The vascular bundles of the smaller veins are accompanied by sclerenchyma in some species, and branches of the sclerenchymatous elements sometimes extend into the mesophyll.

PETIOLE Fig 2, 3.

The epidermis is always cutinized on the periclinal wall, and often partially on the anticlinal walls which also thicken all over with cellulose. Hairs are usually present on the epidermis but are absent in a few species; the epidermis consists of 2 layers in a few

species, e g Tapura latifolia. The cortex is always thickened with collenchyma or

subcollenchyma, and crystal inclusions are abundant; the membranes of certain cells often become gelatinous and frequently the lumen of the cell membrane becomes

gelatinous leaving a stone cell suspended in a gelatinous mass. As in the stem, the cork arises from the subepidermal layer. Typically the vascular bundles are horseshoe-shaped, but there is much variation owing to the epiphyllous inflorescence.

Species with an axillary inflorescence have the characteristic horseshoe-shaped petiole bundles which are less compact towards the base, and are often flanked by small lateral bundles, e g Dichapetalum axillare. The main and lateral bundles are usually surrounded by a mass of sclerotic cells.

In species with an epiphyllous inflorescence, the inflorescence usually emerges above the middle of the petiole, but in the majority of cases the floral and leaf bundles are

separate from near to the base of the petiole; they are often very close, e g in Tapura capitulifera, but are not actually fused. In a few cases, e g Stephanopodium blanchetianum and Tapura latifolia, they are fused on one side only. The sterile petioles usually show no trace of the floral bundle, but Barth (1896) observed traces of the floral bundle in Dichapetalum pedunculatum and Tapura capitulifera; this is marked by a small inverted bundle, and a small mass of sclerenchyma, but both disappear rapidly. In the sterile petioles of Tapura amazonica and Dichapetalum axillare in material examined there is no trace of a floral bundle. In Stephanopodium blanchetianum there are 2 equal bundles of about the same size which soon fuse, but the upper (floral bundle) divides in the middle. In S. engleri the floral bundle is much smaller. In Tapura cubensis there is little difference between the sterile and fertile petioles except that the floral bundle of the sterile petiole is less compact. The floral bundles definitely arise from the leaf bundle within the petiole rather than in the stem. The stem has ciruclar bundles and shows no trace of the floral bundle in the species examined.

In Tapura juliani where the flowers are borne well up the midrib, the leaf bundle of the petiole is almost cylindrical with just a short break, hence the petiole resembles the

young stem much more than in other species examined. The floral bundle of this species arises from the midrib of the lamina rather than the petiole. The structure of the midrib at this point is similar to the fertile petiole in other species.

10 Flora Neotropica

A .... '~ : ../ :,i '.-':,.z-.,

D 4 ... .A .

~ -'

Er~~ F

A, \a

FIG 2. Petiole and young stem of Dichapetalaceae. A-C, Tapura amazonica (Prance et al 5090). A, petiole below inflorescence, showing leaf and inflorescence trace; B, petiole above inflorescence, showing leaf trace only; C, sterile petiole, showing similarity to fertile petiole above inflorescence. D-G, Stephanopodium engleri (Warming s n). D, young stem with cylindrical bundle; E, petiole below inflorescence, showing leaf and inflorescence trace; F, petiole just above inflorescence, showing leaf trace only; G, leaf base, showing inflorescence bud emerging. All X 80.

Introduction 11

iA BCTj~ 7*

. ' ,': r:~~~~~~~~~~ - \;

N

E~~~~~~~ D

L14w ~ ~ ~ ~ ~ ~ ~~~~~~~~~~-

FIG 3. Petiole, midrib and peduncle of Dichapetalaceac. A-D, Dichapetalum pedunculatum (De La Cruz 3095). A, petiole at point of junction with inflorescence; B, peduncle, showing similarity with young stem; C, petiole above inflorescence, showing leaf trace only; D, midrib. E-F, Tapura juliani (Kiug 2010). E, midrib below but near junction with inflorescence; F, petiole at junction with

~~~~~~~~~~~~~lamina base. All X 40.~

Z?~~~~~~~~~~' i:, .*::*~~~~~~~~~~~~~~~~~~~~~~~~~,v . 4~~~

A:~~~~~~~,

FIG 3. Petiole, midrib and peduncle of Dichapetalaceae. A-D, Dicbapetalum pedunculatum~:-%fi~ (DeLa ru 305).A,petoleatpoit f jnctonwit inloescnc; B peunle,shoin siilrity wihyon te;C ptol bveiforsece hoigleftac ny;D irb.EF Tpr julan (lu 210. , idibbeow utner untin it iflrecene;F,peioe t untin it lamina base. All X 40

12 Flora Neotropica

The presence of floral bundles in some sterile petioles, and the separate nature of the floral bundle demonstrates that the position of the inflorescence has arisen by a gradual fusion of the peduncle with the petiole rather than being a true epiphyllous inflorescence.

Although some authors have considered the inflorescence of the Dichapetalaceae as truly epiphyllous, e g de Candolle (1890), the petiole anatomy shows otherwise. This was

clearly pointed out by Barth (1896) when he said "En effet il y a bien morphologique- ment accrescence congenitale du petiole avec le pedoncule floral. D'autre part, l'inflorescence est anatomiquement epiphylle puisque son faisceau sort du faisceau foliare, mais elle ne l'est en realite pas plus qu'un bourgeon dont le faisceau est foliare; la difference est uniquement due ai un retard plus on moins considerable dans la sortie du faisceau gemmaire."

Petiole material examined: Tapura amazonica (Prance 5090), T. juliani (Klug 2010), T. singularis (Pires et al 10474), Stephanopodium engleri (Warming s n), Dichapetalum axillare (Allen 2202), D. donnell-smithii (Gentle 2623), D. pedunculatum (De La Cruz 3095).

YOUNG STEM

The cork arises in the subepidermis, the phelloderm is frequently sclerotic, and the

primary cortex contains stone cells; mucilaginous cells are frequent in the cork, phelloderm and cortex. The pericycle contains characteristic groups of fibers which do not form a continuous mass and are not interspersed with stone cells. The phloem consists of sieve tubes with associated parenchyma cells, and the parenchyma is abundant. The xylem contains narrow vessels usually with simple and rarely scalariform perforation plates, parenchyma is extremely variable in amounts, the fibers have bordered pits, and the rays are of 2 distinct sizes, the smaller 1-4 cells wide, and the larger to 10 cells wide.

WOOD Fig 4.

Growth rings are absent or poorly defined by more abundant apotracheal parenchyma. The vessels are most frequently solitary, but often occur in multiples of 2 or 3 cells or in pore chains in some species of Tapura; the mean tangential diameter of the vessels is 50-100 p, but is sometimes larger, there are about 40 per square mm; the

perforation is simple or scalariform with up to 12 bars in some species of Tapura, but never with spiral thickening; the intervascular pitting is usually alternate and is minute, occasionally scalariform, and transitional intervascular pitting occurs; the ray-vessel pits are also usually alternate and minute, but rarely compound pits occur; tyloses have been

reported in one species of Dichapetalum but were not observed in the present study; the mean vessel length is 0.6-0.9 mm. Parenchyma is predominantly paratracheal around the vessels and with some short irregular wings; in addition there is always some diffuse

parenchyma; occasionally more abundant apotracheal parenchyma occurs in scattered cells or in narrow bands, e g in Tapura cubensis the strands are up to 8 cells high. The rays are typically up to 5 or 6 cells wide, but sometimes 8-10 cells wide in Dichapetalum; they are over 1 mm high in some species; uniseriates of square and upright cells are

frequent; there are abour 11 rays per mm, and they are heterogeneous (Kribs types I and IIA), with 2 to several marginal rows of square and upright cells; sheath cells are usually present, and solitary crystals are often present in both upright and procumbent cells. The fibers have moderately to very thick walls, and there are frequently some gelatinous fibers

especially in the young wood; they have moderately to very numerous small pits on the radial and tangential walls; the pits have small borders; the mean fiber length is about 1.5 mm.

Introduction 13

Y-34089), A, x S; B, t S; C, r s. D-F, Dichapetalum axillare (Cooper & Slater 172, SJRw Y-10525), D,

-"

X S; E, t s; F, r s; G, t s. H, Tapuraguianensis (Ducke 230, SJRw V-31957), r s. AF, all X 40; G-H, X

~.~~~~~~100. ~LL

rT=

100. 100Q.

14 Flora Neotropica

The wood is moderately hard and heavy, of a rather coarse texture, straight or fairly straight grained, and easy to cut.

Descriptions based at least partially on American Dichapetalaceae have been given by Record (1927), Tippo (1938), Heimsch (1942), Record & Hess (1943) and Metcalfe & Chalk (1950).

Heimsch (1942), stated amongst his conclusions that "it is not evident on the basis of wood structure whether or not the Dichapetalaceae belong with the families of the Gruinales (i e Malpighiaceae, Linaceae, Humiriaceae etc) or the Rosaceae or the

Euphorbiaceae." Similarly Metcalfe & Chalk (1950) do not draw any conclusions about the taxonomic position of the family based on wood anatomy. However, it appears to me that the wood anatomy of this family would fit better either with the Celastrales or with the Polygalales as defined by Cronquist, rather than with the Euphorbiaceae or Rosaceae.

Wood material examined: Dichapetalum axillare SJRw Y-10525 (Cooper & Slater 172); D. gelonoiodes SJRw Y-12495; Tapura guianensis SJRw Y-31957, FHOw 10841 (Ducke 230); T. juruana SJRw Y-36481, FHOw 20286 (Krukoff 6160); T. lanceolata SJRw Y-34089, (Ducke 325); Stephanopodium aptotum SJRw-Y-20899, Y-20920

(Espina & Giacometto A145, A124).

CHROMOSOMES

At the time of this study the chromosomes of the American Dichapetalaceae have not yet been examined. However, in connection with Breteler's studies of African

Dichapetalaceae, Dr. Th. Gadella of Utrecht is investigating the chromosomes of the

Dichapetalaceae and has already made a number of counts (Gadella 1969, 1970). The data already available indicate that the family has little variation in chromosome number as all 14 African species of Dichapetalum examined so far by Gadella have the same basic number of 2n = 24.

SYSTEMATIC TREATMENT

Dichapetalaceae Baillon, Mart. Fl. Bras. 12(1): 365. 1886 (Dichapetaleae); Engler in

Engler & Prantl, Nat. Pflanzenfam. 3(4): 345. 1896; Gleason, N. Am. Flora 25: 381-383. 1924; Hutchinson, Fam. Fl. P1. 1: 206. 1926, Gen. P1. 1: 216-219. 1964; Engler & Krause, Nat. Pflanzenfam. 19c: 1-11. 1931; Macbride, Fl. Peru, Publ. Field Mus. Bot. 13(3): 954-964. 1950; Lasser, Bol. Soc. Venez. Ci. Nat. 9: 125-128. 1944; Stafleu in Pulle, Fl. Suriname 3(2): 166-172. 1951; Rizzini, Revista Bras. Biol. 12(1): 97-108. 1952; Leon & Alain, Fl. Cuba 3: 36-38. 1953; Lewis, Fl. Panama, Ann. Missouri Bot. Gard. 54: 9-12. 1967.

Chailletiaceae R. Brown in Tuckey, Narr. Congo. Append. 5: 442-444. 1818; de Candolle, Prodr. 2: 57-58. 1825; Endlicher, Gen. P1. 2: 1104. 1840; Bentham & Hooker, Gen. PI. 1(1): 340. 1862.

Trees, shrubs, lianas, or suffruticose subshrubs. Stipules present but usually caducous. Leaves simple, alternate, entire, pinnately nerved. Inflorescence corymbose- cymose or subcapitate, or the flowers fasciculate, axillary or more frequently attached to the petiole or rarely to the midrib. Flowers small, hermaphrodite or less frequently unisexual, actinomorphic to weakly zygomorphic; pedicels often articulated. Petals 5, either free, imbricate and almost equal, or connate into a tube, with the lobes equal or

markedly unequal, the lobes usually bifid at apex and frequently bicucullate or inflexed; often clawed at base. Stamens 5, all fertile or only 3 fertile, free or adnate to the corolla tube, with filaments or rarely the anthers sessile; anthers bilocular, dehiscing longitudinally. Disc of 5 equal or unequal hypogynous glands alternating with the stamens, or united into a disc. Ovary superior, free, 2-3-locular, the ovules anatropous, pendulous, paired at the top of each loculus. Styles 2-3, free or more frequently connate nearly to

apex, often recurved, the stigma capitate or simple. Fruit a dry or rarely a fleshy drupe; epicarp most frequently pubescent; mesocarp thin; endocarp hard; 1-2 (-3) locular, the loculi usually with only one seed developing; seed pendulous, without endosperm; embryo large, erect.

A tropical family of about 240 species in three genera, distributed throughout the lowland tropical regions of both hemispheres (but absent from Polynesia and Micronesia),

extending into the subtropics in Africa and India. TYPE GENUS. Dichapetalum. The name is derived from the Greek words JlK0

(dicha-divided in two), and 1reraXov (petalon-petal), because of the petals with a bifid

apex which is frequent in many species of the genus.

Key to the Genera of Dichapetalaceae in America 2

1. Petals free and regular; stamens free; inflorescence with a long distinct peduncle. 1. Dichapetalum.

1. Petals connate or only 3 free; stamens adnate to corolla tube; inflorescence sessile or almost so. 2. Corolla with 5 equal obtuse lobes, shorter than the tube; fertile stamens 5, the anthers

sessile on tube. 2. Stephanopodium. 2. Corolla zygomorphic, the lobes bifid and bicucullate, exceeding the tube in length; fertile

stamens 3 or 5, the anthers on slender filaments. 3. Tapura.

2 This key is constructed for a quick identification of the American species, and is not satisfactory for the African species where there is much more variation in the inflorescence type.

15

16 Flora Neotropica

1. Dichapetalum Thouars, Gen. Nov. Madag. 23. 1806; Baillon, Mart. Fl. Bras. 12(1): 369. 1886; Engler in Engler & Prantl, Nat. Pflanzenfam. 3(4): 348. 1896; Engler & Krause, Nat. Pflanzenfam. 19c: 5-9. 1931; Lasser, Bol. Soc. Vanez. Cienc. Nat. 9: 126. 1944; Macbride, Publ. Field. Mus. Bot. 13(3): 955-958. 1950; Stafleu, in Pulle, Fl. Suriname 3(2): 167-169. 1951; Rizzini, Revista Bras. Biol. 12(1): 97-101. 1952; Hutchinson, Gen. Fl. PI. 1: 218. 1964; Lundell, Wrightia 3: 173-176. 1966; Lewis, Ann. Missouri Bot. Gard. 54: 9-12. 1967; Prance, Act. Bot. Venez. 3: 301-304. 1968; Ind. Nom. Gen. 10/07545.

Leucosia Thouars, Gen. Nov. Madag. 78. 1806. Symphyllanthus Vahl, Skrift. Nat. Hist. Selsk. 6: 86. 1810; Gleason, N. Amr. Flora 25: 381.

1924. Chailletia de Candolle, Nouv. Bull. Soc. Philom. Paris 40: 205-206. 1811; Ann. Mus. Paris 17:

153. 1811; Prodr. 2: 57. 1825; Endlicher, Gen. PI.: 340. 1840; Bentham & Hooker, Gen. PI. 340. 1862.

Moacurra Roxburgh (Hort. Beng. 21. 1814 nom nud) Fl. Ind. 2: 69. 1832. Mestotes Solander ex De Candolle, Prodr. 2: 57. 1825. Plappertia Reichenbach, Consp. 146. 1828. Quilesia Blanco, Fl. Filip. ed. 1: 176. 1827. Patrisia Rohr ex Steudel, Nom. Bot. ed. 2. 1: 342. 1840. Pentastira Ridley, Trans. Linn. Soc. II. 9: 27. 1916.

Small trees, shrubs or lianas. Leaves alternate, petiolate, entire. Stipules small, deciduous or persistent. Inflorescences branched cymose or corymbose panicles with long peduncles (peduncles short or absent in some African species), axillary or adnate to

petiole. Flowers small, hermaphrodite, polygamous or dioecious, actinomorphic. Bracts small. Receptacle usually convex or subplanate. Sepals 5, imbricate, free, or connate at

base, equal or subequal. Petals 5, free to the base, alternate with the sepals, usually bicucullate anc 2-lobed at apex, the margins of the lobes inflexed and sometimes

enveloping the anthers. Stamens 5, equal, free, all fertile in male and hermaphrodite flowers; filaments usually free, rarely connate at the base only; anthers broadly oblong, introrse. Disc usually consisting of 5 hypogynous glands which are opposite to the petals; glands entire, shallowly lobed, free or united. Ovary free (in American species) or rarely adnate to receptacle, globose, 2-3 locular with 2 ovules in each loculus. Styles 1-3, free or connate almost to apex, rudimentary pistil present in male flowers. Fruit a dry indehiscent drupe, 1-3 locular usually with one seed in each loculus; epicarp pubescent (in American species).

TYPE SPECIES. Dichapetalum madagascariensis Poiret, Encycl. Method. Suppl. 2: 470. 1812; Dict. Sci. Nat. 13: 178. 1819. Type. Du Petit Thouars s n, Madagascar, fl

(holotype, P; photo, NY). DISTRIBUTION. Throughout the lowland tropics in Malesia, tropical and south

Africa, and in the New World from Mexico to Peru and Amazonian Brazil, but not in the Caribbean. Most diverse and abundant in Africa.

Engler in his various publications on African Dichapetalum described four sections, which are summarized in Engler & Krause (Nat. Pflanzenfam. 19c: 5-9. 1931). All the American and Malesian species belong to his section Eudichapetalum (correctly, section

Dichapetalum). His other three sections are for groups of African species, and are not considered here. There are no grounds for the recognition of sections or subgenera in the American species.

Key to American species of Dichapetalum 1. Stipules with fimbriate margins; pedicels well developed 2.0-6.0 mm long; lamina distinctly

to slightly bullate on upper surface. 2. Leaves 15.0-32.0 cm long; densely hirsute on lower surface. 1. D. bullatum.

Syscematic Treatment 17

2. Leaves 8.0-17.5 cm long; with only a few stiff appressed hairs on lower surface. 3. Pedicels ca 5.0 mm long; inflorescence densely brown-tementose. 2. D. stipulatum. 3. Pedicels 2.0-3.0 mm long; inflorescence shortly gray-puberulous. 3. D. nervatum.

1. Stipules with entire margins; pedicels usually short, 0-2.0 mm long, rarely longer; lamina rarely slightly bullate, usually smooth on upper surface. 4. Lamina with a dense compact gray-lanate pubescence on lower surface. 4. D. latifolium. 4. Lamina glabrous, hirsute, or with a few stiff hairs on lower surface.

5. Young branches and inflorescence hispid. 6. Lamina narrowly oblong-lanceolate to lanceolate, 1.1-2.8 cm broad; inflorescence

gray-brown pubescent. 5. D. nevermannianum. 6. Lamina oblong, 3.0-5.0 cm broad; inflorescence rugose. 6. D. coelhoi.

5. Young branches and inflorescence tomentose to glabrous, never hispid. 7. Lamina thickly coriaceous, with conspicuously impressed venation on upper

surface, densely hirsute on lower surface; inflorescence usually dark brown to rufous pubescent. 7. D. rugosum.

7. Lamina coriaceous to membraneous, the venation of upper surface plane (or if slightly impressed then with only a few stiff appressed hairs on lower surface), glabrous or with a few stiff appressed hairs on lower surface; inflorescence gray to brown pubescent. 8. Lamina predominantly oblong to oblong-lanceolate (length breadth ratio

usually greater than 2.5). 9. Petioles 1.0-4.0 mm long; inflorescences terminal, the peduncles and young

stems hispidulous; lamina very unequal at base. 8. D. pauper. 9. Petioles 5.0-18.0 mm long; inflorescences predominantly petiolar or

axillary; young stem tomentellous to tomentose. 10. Young branches and inflorescences pale gray pubescent; fruit oblong-

ellipsoid, to 3.8 mm long, the epicarp pale gray-brown pubescent; leaf bases subcuneate to cuneate. 9. D. axillare.

10. Young branches and inflorescences yellow-brown pubescent; fruit ellipsoid, to 2.0 mm long, the epicarp dark yellow-brown pubescent; leaf bases usually rounded to subcordate. 11. Flowers polygamodioecious, on short pedicels to 0.3 mm long;

petiolar inflorescences spreading, 4.0-6.0 mm long. 10. D. steyermarkii. 11. Flowers hermaphrodite; pedicels 0.5-2.5 mm long; petiolar inflor-

escences compact. 11. D. donnell-smithii. 8. Lamina predominantly elliptic to ovate-elliptic (length breadth ratio usually less

than 2.3). 12. Leaves hirsute on lower surface, sometimes with slightly impressed

venation on upper surface. 13. Lamina broadly ovate-elliptic, abruptly acuminate at apex 12. D. spruceanum. 13. Lamina elliptic, the apex with long thin curved acumen.

11. D. donnell-smithii. 12. Lamina glabrous or with a few stiff appressed hairs only on lower surface,

the venation prominent to plane. 14. Lamina with mucronate or abruptly acuminate apex, the acumen

usually curved, the base markedly unequal. 12. D. spruceanum. 14. Lamina tapering to an acute or acuminate apex, the acumen erect, the

base equal or slightly unequal. 15. Leaves 12.0-28.0 cm long; petioles 8.0-32.0 mm long. 13. D. froesii. 15. Leaves 5.0-16.0 cm long; petioles 4.0-12.0 mm long.

16. Flowers with long pedicels 1.0-3.0 mm long; stipules 0.5-2.0 mm long, caducous; inflorescence branches sparsely tomentellous to glabrous. 14. D. odoratum.

16. Flowers with short pedicels 0.2-1.5 mm long; stipules 3.0-10.0 mm long, persistent or caducous; inflorescence branches tomentellous to sparsely puberulous. 15. D. pedunculatum.

1. Dichapetalum bullatum Standley & Steyermark Publ. Field Mus. Bot. 23: 169. 1944; Lundell, Wrightia 3: 174. 1966. Fig 5 A-D, 11.

Vine or shrubby vine, the young branches hispid, becoming glabrous with age. Leaves ovate-elliptic to elliptic, coriaceous, 15.0-32.0 cm long, 6.0-19.0 cm broad, acuminate at apex, the acumen 10.0-20.0 mm long, rounded to subcordate at base,

18 Flora Neotropica

bullate above, hirsute beneath; midrib slightly impressed and pubescent above; primary veins 9-12 pairs, slightly impressed above, the secondary and tertiary venation impressed above; petioles 5.0-8.0 mm long, hispid. Stipules subpersistent, hispid, the margins fimbriate. Flowers hermaphrodite; pedicels 5.0-6.0 mm long. Inflorescences short terminal or subterminal panicles, much branched at base, 2.0-3.0 cm long, hispid. Bracts and bracteoles linear, persistent, hispid. Calyx ca 3.0 mm long, gray-puberulous and

hispid on exterior, the lobes almost equal. Corolla of 5 equal deeply bifid and slightly cucullate lobes, free to base, glabrous. Fertile stamens 5, alternating with and exceeding corolla lobes. Disc of 5 free ellipsoid glands, apex of glands rounded. Ovary bilocular with 2 ovules in each loculus, lanate on exterior except for glabrous base. Style lanate at base, sparsely puberulous above, the apex shortly trifid. Fruit not seen.

TYPE. Steyermark 39874, Guatemala, Izabal, southeast of Puerto Barrios, fl

(holotype, F; isotype, US); Steyermark 38803, Guatemala, Izabal, Montana del Mico, fl

(paratypes, F, US); Standley 72813, Guatemala, Izabal, Puerto Barrios, st (paratype, F); 73053, Puerto Barrios, st (paratype, F).

DISTRIBUTION. Known only from the type collections from Guatemala. Collected in flower in April and December.

2. Dichapetalum stipulatum Macbride, Publ. Field Mus. Bot. 11: 68. 1931; Publ. Field Mus. Bot. 13(3): 957. 1950. Fig 10.

Liana, the young branches ferrugineous tomentose, soon becoming glabrous. Leaves

elliptic, coriaceous, 8.0-17.5 cm long, 4.5-8.0 cm broad, acute to acuminate at apex, subcordate to rounded at base, bullate above, with a few stiff appressed hairs only beneath; midrib slightly impressed and pubescent above; primary veins 7-9 pairs, slightly impressed above; secondary venation impressed or plane above; petioles 2.0-5.0 mm long, villous-tomentose. Stipules subpersistent, villous, fimbriate, ca 5.0 mm long. Flowers

hermaphrodite; pedicels ca 5.0 mm long. Inflorescences large terminal corymbose panicles ca 12.0 cm long, the rachis and branches villous-tomentose. Bracts and bracteoles linear, 1.0-4.0 mm long, persistent, tomentose. Calyx ca 3.0 mm long, villous-tomentose on

exterior, the lobes almost equal. Corolla of 5 equal lobes, deeply bifid and slightly cucullate, free to base, glabrous. Fertile stamens 5, alternating with and exceeding the corolla lobes. Disc of 5 ellipsoid glands, united at base into a small circular disc. Ovary bilocular or trilocular with 2 ovules in each loculus, or occasionally of 2 separate unilocular carpels, lanate on exterior. Style lanate at base, glabrescent above, the apex bifid or trifid. Fruit bilocular or trilocular; epicarp with a short compact pubescence; mesocarp very hard when dry, ca 1.5 mm thick; endocarp thin, hard, lanate within.

TYPE. Klug 1081, Peru, Loreto, Mishuyacu, fl (holotype, F; isotype, NY). DISTRIBUTION. Known only from forest in Loreto, Peru. Collected in flower in

February and in fruit in August and October. PERU. Loreto: Killip & Smith 28128 fr (F, NY, S, US); Klug 76 fl (F, NY, US).

This species and D. bullatum are closely related, and differ from the other American

species of Dichapetalaceae in their fimbriate stipules, the long hispid or villous pedicels and the bullate leaves. Dichapetalum stipulatum differs from D. bullatum in the larger branched inflorescences, with a villous rather than hispid pubescence, and the smaller leaves with almost glabrous lower surfaces.

3. Dichapetalum nervatum Cuatrecasas, Lloydia 11: 222. 1949. Fig. 10.

Tree, the young branches sparsely hirsute-tomentose, soon becoming glabrous. Leaves oblong, coriaceous, 8.0-13.0 cm long, 4.5-5.5 cm broad, acuminate at apex, with acumen 5.0-15.0 mm long, rounded and equal at base, slightly bullate above, glabrescent

Systematic Treatment 19

H~~~~~~~~~ -B

FIG 5. Species of Dichapetalum. A-D, D. bullatum (Steyermark 39874). A, habit X 0.25; B, flower X 3; C, flower section X 4; D ovary and style, X 4. E-J, D. latifolium (Pires et al 1071). E, flower X 5; F, flower section X 6; G, ovary and styles X 6; H, ovary section X 15; J, habit X 0.5.

20 Flora Neotropica

or with a few stiff appressed hairs only beneath; midrib plane, hirsutulous above; primary veins 7-9 pairs, plane above; venation plane or slightly impressed above; petioles 3.5-5.0 mm long, sparsely hirsutulous. Stipules subpersistent, hispid-hirsute, fimbriate. Flowers

hermaphrodite; pedicels 2.0-3.0 mm long. Inflorescences (part only seen) with

gray-tomentellous rachis and branches. Bracts and bracteoles lanceolate 1.5-4.0 mm long, persistent, tomentellous. Calyx 2.0 cm long in bud, tomentellous on exterior. Corolla of 5

equal bifid lobes, glabrous, free to base. Fertile stamens 5, alternating with corolla lobes. Disc of 5 separate ellipsoid glands. Ovary (one only examined) bilocular with two ovules in each loculus, lanate on exterior. Style short and with a trifid apex, lanate at base. Fruit

ellipsoid, unilocular; epicarp velutinous-tomentellous; mesocarp thin; endocarp thin, hard, bony, glabrous within.

TYPE. Cuatrecasas 15833, Colombia, Valle, Rio Yurumangui, fl fr (holotype, F;

isotype, COL). DISTRIBUTION. Known only from the type collection in flower and fruit in

January.

This species is known only from a single rather poor collection with unattached flowers and fruit. It is certainly very close to D. stipuldtum, and only further material will show whether it is in fact a distinct species. It differs from D. stipulatum in the smaller

leaves, and in the shorter gray pubescence of the inflorescence and flowers.

4. Dichapetalum latifolium Baillon, Mart. Fl. Bras. 12(1): 370. 1886; Rizzini, Revista Bras. Biol. 12(1): 100. 1952. Fig 5 E-J, 13.

Dichapetalum amazonicum Krause, Notizbl. Bot. Gart. Berlin 11: 135. 1951. Type. Tessmann 3562, Peru, Loreto, Iquitos, fl (holotype, B lost; photos, F, MO; isotype, S).

Dichapetalum vestitum (Bentham) Baillon var cinerascens Baillon, Mart. Fl. Bras. 12(1): 372. 1886. Types. Martius s n, Brazil, Amazonas, Manaus, fl (lectotype, M): Poeppig 2903, Brazil, Amazonas, Tefe, fl (paratypes, G, LE, P, W).

Dichapetalum scandens (Poiret) Johnson var cinerascens (Baillon) Macbride, Publ. Field Mus. Bot. 13(3): 957. 1950.

Liana, the young branches shortly and densely tomentellous, becoming glabrous with age. Leaves elliptic to ovate-elliptic, coriaceous, 8.0-36.0 cm long, 4.0-21.0 cm

broad, mucronate to shortly acuminate at apex, rounded to subcuneate at base, plane not bullate above, with a gray lanate tomentum beneath; midrib plane and pubescent above;

primary veins 6-8 pairs, plane to weakly impressed above; the secondary and tertiary venation usually weakly impressed above; petioles 7.0-18.0 mm long, short-tomentellous.

Stipules caducous (not seen). Flowers androdioecious; pedicels 1.5-3.0 mm long. Inflorescences spreading terminal or petiolar panicles; the rachis and branches tomentellous. Bracts and bracteoles lanceolate, 0.7-4.0 mm long, persistent, tomentose. Calyx ca 2.0 mm long, gray-tomentellous on exterior, the lobes slightly unequal. Corolla of 5 equal deeply bifid and slightly cucullate lobes, free to base, glabrous to sparsely pubescent on exterior. Fertile stamens 5, alternating and slightly exceeding corolla lobes. Disc of 5 free

glands with a weekly bifid apex. Ovary bilocular or trilocular with 2 ovules in each

loculus, lanate on exterior. Style lanate at base, glabrous above, the apex trifid. Fruit

(immature) ellipsoid; epicarp tomentose. TYPE. Spruce 2165, Brazil, Amazonas, Sao Gabriel, Rio Negro, fl (holotype, K;

isotypes, P, W). DISTRIBUTION. Forests of western Amazonia in Venezuela, Peru and western

Brazil. Flowering October to May. VENEZUELA. Apure: Steyermark et al 10515 fl (NY, VEN); 10627 fl fr (NY, VEN). PERU. Loreto: Klug 496 fl (F, NY, US); 1431 fl (F, NY, US); Schunke V. 6241 fl (F). BRAZIL. Amazonas: Black 48-2908 fl (IAN, NY, P); Ducke 881 fl (F, IAN, MG, MO, NY, RB, US); Froes 23734 fl (IAN, RB); Pires & Black 1071 fl (GH, IAN); M. Silva 2176 fl (MG). BOLIVIA. La Paz: Buchtein 1718 fl (NY).


:;~~f 6

g

A: J:>:

C\ \ ' 9 .

FIG 6. Species of Dichapetalum. A-F, D. pauper (Froes 22493, 28317). A, flower X 5; B, habit X 0.5; C, flower X 6; D, ovary and styles X 6; E, ovary section X 15; F, fruit X 0.5. G-J. D. nevermannianum (Von Wedel 1424). G, habit X 0.5; H, fruit X 0.5; J. flower section X 6.

22 Flora Neotropica

This species was placed in synonymy under D. scandens (D. rugosum) by Macbride (Publ. Field Mus. Bot. 13(3): 956. 1950). Dichapetalum latifolium is certainly distinct from that species, and differs in the gray-lanate pubescence of the leaf undersurface, the inflorescence and the longer pedicels.

5. Dichapetalum nevermannianum Standley & Valerio, Publ. Field Mus. Bot. 18: 597. 1937; Woodson & Schery, Ann. Missouri Bot. Gard. 29: 353. 1942; Lundell, Wrightia 3: 174. 1966; Lewis, Ann. Missouri Bot. Gard. 54: 10. 1967. Fig 6 G-J, 11.

Shrub, the young branches hispid. Leaves lanceolate to oblong-lanceolate, membraneous to chartaceous, 6.0-12.5 cm long, 1.1-2.8 cm broad, the apex tapered to a finely pointed acumen 10.0-16.0 cm long, subcordate and slightly unequal at base, plane not bullate above, with a sparse appressed hirsute pubescence beneath; midrib slightly prominent and hirsute above; primary veins 10-11 pairs, almost plane above; venation

plane above; petioles 2.0-4.0 mm long, hispid. Stipules lanceolate, to 5.0 mm long, persistent, hispid, the margins entire. Flowers hermaphrodite; pedicels 2.0- 3.0 mm long. Inflorescences small terminal panicles (one only seen), the rachis and branches hispid. Bracts and bracteoles lanceolate, 1.5-3.5 mm long, persistent, hispid. Calyx ca 2.5 mm

long in bud, gray-tomentose on exterior, the lobes equal. Corolla of 5 equal deeply bifid

lobes, glabrous, free to base. Fertile stamens 5, alternating with corolla lobes disc of 5

elongate glands. Ovary (one examined) bilocular, lanate on exterior. Style short, lanate at

base, glabrous above, the apex bifid. Fruit unilocular to trilocular, ellipsoid; epicarp hispid-tomentose; mesocarp thin; endocarp very thin, bony, glabrous within.

TYPE. Standley & Valerio 48497, Costa Rica, Lim6n Province, Rio Reventazon, below Cairo, fr (holotype, F; isotypes, EAP, G, US).

DISTRIBUTION. Wet forests in the Caribbean coastal region of Costa Rica and Panama. Collected in flower in October and in fruit in February. PANAMA. Bocas del Toro: Von Wedel 1424 fl bud (F, GH).

This poorly known species is most distinct amongst the American species by the narrow lanceolate leaves and the hispid branches, petioles and inflorescence. The material examined is poor and a whole inflorescence or open flowers have not yet been seen.

6. Dichapetalum coelhoi Prance, sp nov Fig 7 A-E, 10.

Frutex scandens, ramulis juvenilibus hispidis. Folia alternata, petiolata; petiolo 3.0-5.0 mm longo, hispido; laminae oblongae, coriaceae, 10.0-18.0 cm longae, 3.0-5.0 cm latae. Apice in acuminem tenue 5.0-15.0 mm longum attentuatae, basi subcordatae, inaequales, supra glabrae, planae haud bullatae, subtus sparse hispidae; costa media supra plana, sparse pubescente; costis secundariis 10-12 jugis, supra planis; venis supra planis. Stipulae lineares, circa 6.0 mm longae, hispidae, marginibus integris. Flores hermaphroditi, circa 5.0 mm longi, in paniculis terminalibus 2.0-4.0 mm longis dispositi; pedunculis circa 1.5 mm longis; rachi ramisque rugoso-hirsutis. Bracteae bracteolaeque lineari-

lanceolatae, 1.5-3.5 mm longae, persistentes, hirsutae. Calyx 3.0-3.5 mm longus, extus

tomentosus, lobis aequalibus. Petala 5, glabra, ad basim libera, aequalis, apice bifida. Stamina 5, omnis fertilis, cum calycis-lobis alternata. Discuss constans ex glandis 5, apicibus lobatis. Ovarium biloculare, ovulis in quoque loculo duobus, extus lanatum.

Stylus glabrus, apice bifidus. Fructus mihi ignoti. TYPE. Byron Albuquerque, L. Coelho & J. Lima 68-119, Brazil, Amazonas,

Manaus-Itacoatiara road, Km 104, fl (holotype, NY; isotype, INPA 21145).

This species is distinct from others on account of the hispid pubescence of the

inflorescence, leaves and young branches, and in the terminal inflorescence.


A

"S r......

FIG 7. Species of Dichapetalum. A-E, D. coelhoi (Byron et al 68-119). A, flower X 4; B, ovary and style X 5; C, flower section X 5; D, habit X 0.5; E, ovary section X 5. F-L, D. froesii (Froes 22850, 22325). F, fruit X 0.5; G, habit X 0.5; H, flower X 5; J, flower section X 7; K, ovary and styles X 7; L, ovary section X 10.

24 Flora Neotropica

It is with pleasure that I name this species for the Coelho brothers, Luiz and

Dionisio, who have done so much to help me during my field work in Amazonia.

7. Dichapetalum rugosum (Vahl) Prance, Act. Bot. Venez. 3: 303. 1968. Fig 14.

Symphyllanthus rugosus Vahl, Skr. Naturh.-Selsk. Kj0benh. 6: 86. 1810. Cordia scandens Poiret, Dict. Sci. Nat. 10: 40. 1818. Type. Martin s n, French Guiana, fl

(holotype, P). Chailletia vestita Bentham, Jour. Bot. Kew Misc. 3: 372. 1851. Type. Spruce 623 (Jan 1850),

Brazil, Para, Santarem, fl (lectotype, K; isotype, P); Spruce s n (Nov-Mar 1849-50), Brazil, Para, Santarem, fl (probable isotypes BM, NY, W); Spruce 151 (Nov-Mar 1849-50), Santarem, fl (paratype, M); Spruce 780 (s n in most sets, Apr 1850), Brazil, Para, Santarem, fl (paratypes, BM, C, CGE, G, GH, LE, M, NY, OXF, P, W).

Dichapetalum vestitum (Bentham) Baillon, Mart. Fl. Bras. 12(1): 371. 1886; Stafleu in Pulle, Fl. Suriname 3(2): 168. 1951; Rizzini, Revista Bras. Biol. 12(1): 98. 1952.

Dichapetalum vestitum (Bentham) Baillon var scandens Baillon, Mart. Fl. Bras. 12(1): 372. 1886. Type. Martius 2794, Brazil, Amazonas, Manaus, fl (lectotype, M); Spruce 1109 (Dec-Mar 1850-51, s n in most sets), Brazil, Amazonas, Manaus, fr (paratypes, BM, CGE, G, GH, GOET, K, LE, M, MG, NY, OXF, P, W).

Chailletia flavicans Klotzsch in Schomburgk, Fauna & Fl. Guiana. 1184. 1849, nom nud. Dichapetalum flavicans (Klotzsch) Engler, Bot. Jahrb. 23: 145. 1897. Type. Schomburgk 1319,

Guyana, fl (holotype B, lost; photos, F, MO). Chailletia parkeri Planchon ex Triana & Planchon, Ann. Sci. Nat. V. 15: 379. 1872 nom nud.

Type. Birschel s n, Venezuela, fl (K). Dichapetalum scandens (Poiret) Johnston, Jour. Arnold Arb. 16: 44. 1935; Macbride, Publ. Field.

Mus. Bot. 13(3): 956. 1950 pro parte.

Liana or shrub, the young branches tomentose to tomentellous, becoming glabrous with age. Leaves oblong to ovate-elliptic, thickly coriaceous, 6.0-32.0 cm long, 3.5-21.0 cm broad, most frequently acute but varying from rounded to acuminate at apex, subcuneate, rounded or subcordate at base, plane not bullate, or weakly bullate only above, densely hirsute beneath; midrib plane and pubescent above; primary veins 7-13

pairs, slightly impressed above; petioles 2.0-35.0 mm long, densely tomentose. Stipules lanceolate, 2.0-4.0 mm long, subpersistent or caducous, densely tomentose, the margins entire. Flowers hermaphrodite; pedicels 0.5-2.0 mm long. Inflorescences terminal,

axillary or petiolar corymbose panicles, 1.5-8.0 mm long, the rachis and branches

tomentose. Bracts and bracteoles triangular, 0.5-3.0 mm long, persistent, tomentose.

Calyx 3.0-3.5 mm long, densely ferrugineous-tomentose on exterior, the lobes equal. Corolla of 5 equal lobes, bifid and weakly cucullate at apex, free to base, glabrous. Fertile

stamens 5, alternating with and equalling corolla lobes. Disc of 5 separate bifid glands, united at base. Ovary bilocular or trilocular with 2 ovules in each loculus, lanate on

exterior. Style lanate at base, glabrous above, the apex shortly trifid. Fruit unilocular or

bilocular; epicarp densely velutinous tomentose; mesocarp thin; endocarp very thin, hard,

bony, glabrous within. TYPE. Herb. Vahl. s n, French Guiana, fl (holotype, C). DISTRIBUTION. Primary and secondary forests on non-flooded ground, savannas,

and stream margins in the Guianas and Amazonia, and west to the foothills of the Andes

in Peru and Colombia. Collected in flower continously in Amazonia. VENEZUELA. Barinas: Breteler 4435 fl (VEN); 4588 fl (U, VEN). Bolivar: Steyermark 89476 fl (US, VEN). SURINAME. B.W. 3219 fr (K, MO, NY, U); 6561 fr (NY, U); Lindeman 5394 st (U); Maguire 24658 fr (F, GH, IAN, NY, P, RB, U, US); Pulle 454 fl (L, NY, RB, U, US); Van Donselaar 3488 st (U). FRENCH GUIANA. Martin s n3 fl (BM, F, P); 923 fl (BM): Melinon 233 fl (P); Richard s n fl (G, P); Sagot 191 fl (P). PERU. San Martin: Klug 2626 fl (A, BM, F, G, GH, K, MO, NY, S, US). Loreto: Klug 858 fl (F, NY, US); LI. Williams 3725 fr (F, US). BRAZIL. Amazonas: Chagas INPA 3820 fl (INPA, NY); INPA 6384 fl (INPA, NY); Chagas & Coelho INPA 3704 fl (IAN, INPA, NY); D. Coelbo INPA 4011 fl (INPA, NY); Ducke 131 fl (A, F, K, MO, NY, RB, S, US); 1112 fl (IAN, K, MG, MO, NY, US); 1479 fl (A, F, IAN, MG, NY, US);MG 7656 fl (BM, MG, RB 13576);MG 11600fl (G, MG);Fr6es21604 fl (IAN, SP); 30715 fl (IAN, U, UB); Krukoff 8519 fl (A, BM, F, G, K, LE, MICH, MO, NY, P, S, U); Pires & Black 1157 fl (IAN, GH); W. Rodrigues et al 1538 fl (INPA, NY); 1541 fl (INPA, NY); 1960 fl

3 Probable isotypes of Cordia scandens.


(INPA, NY); 2235 fl (INPA, MG, NY); 2320 fr (INPA, NY); 2406 fl (INPA, NY); 2530 fl (INPA, NY); 3035 fl (INPA); 3835 fl (INPA, NY); 4107 fl (INPA, NY); 4367 fr (INPA); 4404 fr (INPA); 4457 fr (INPA, NY); 4550A fl (INPA, NY); 4845 fl (INPA, NY); 4950 fl (INPA, NY); 5060 fl (INPA, NY); INPA 443 fl (INPA, MG); INPA 2925 fl (INPA, MG); Spruce 1185 fr (BM, GH, K, NY, P, W). Para: Archer 8310 fl (IAN, K, NY, UC); Black 48-2297 fl (IAN); 48-3591 fl (IAN); 50-9067 fl (IAN); Black et al 57-19022 fl (IAN, P, UB); 57-19739 fl (IAN, U); Cavalcante 127 fl (MG); 345 fl (HB, MG); Dahlgren & Sella 375 fl (F); 504 fl (F); Ducke MG 503 fl (MG, RB 13562); MG 3457 fl (BM, MG, RB); MG 10975 fr (MG, RB 13577); MG 12124 fr (MG); Froes 30442 fl (IAN, P, SP); Guedes 126 fl (P); 131 fl (IAN, NY); 337 fl (IAN); Huber MG 4011 fl (BM, G, MG, P, RB 13578, US); J.G. Kuhlmann 1885 fl (NY, RB 139379); 2092 fl (RB 139380); M. Kuhlmann et al 365 fl (NY, SP); Martius s n fl (L, M); Monteiro da Costa 276 fl (IAN); Museu Goeldi 9686 fl (BM, MG, RB 13574, US); Oliveira 564 fl (IAN, UB); 3273 fl (IAN); 4156 fl (IAN); 4294 fl (IAN, NY); 4295 fr (IAN); Pires 8241 fl (U, UB); Pires & Black 670 fl (IAN); 1300 fl (IAN, INPA, RB); Pires & Silva 10375 fl (IAN); M. Silva 1 786a fl fr (MG); N.T. Silva 59726 fl (F, IAN, K, NY, S); Snethlage MG 8216 fl (MG). Terr. Amapa: Froes & Black 27383 fl (IAN, U). Maranhao: Ducke MG 627 fl (BM, INPA, MG, RB 13573). Without locality: Glaziou 10080 fl (C, K, P).

The species here brought together in synonymy cannot be separated from one another. Baillon (Mart. Fl. Bras. 12(1): 372. 1886) divided this species into three varieties based on small differences in leaf shape and habit. These characters are not reliable and his varieties are not maintained. One of Baillon's varieties, var cinerascens, is in fact D.

latifolium. The type of this species is the specimen in the Vahl herbarium at Copenhagen which

does not bear a collector's name. It is probably a duplicate of a Richard collection with a

Symphyllanthus manuscript name which is at Paris and Geneva. Hence: Richard s n fl (G, P) are probable isotypes. 8. Dichapetalum pauper Rizzini, Revista Bras. Biol. 12(1): 101. 1952. Fig 6 A-F, 10.

Vine, the young branches hispid-tomentose, soon becoming glabrous. Leaves oblong to oblong-lanceolate, coriaceous, 6.0-16.0 cm long, 1.7-4.0 cm broad, tapered into a

long usually curved acumen at apex, rounded to. subcordate and unequal at base, plane not bullate above, with a few stiff appressed hairs only beneath; midrib almost plane and

pubescent above; primary veins 8-11 pairs, weakly prominent above; venation more or less plane; petioles 1.0-4.0 mm long, hispid to tomentose. Stipules ca 2.0 mm long, ovate, persistent, tomentose, the margins entire. Flowers hermaphrodite; pedicels 1.5-3.0 mm

long. Inflorescences short terminal panicles, the rachis and branches hispid-tomentose. Bracts and bracteoles to 3.0 mm long, lanceolate, tomentose. Calyx ca 2.5 mm long, the lobes equal, tomentose on exterior. Corolla of 5 equal deeply bifid lobes, glabrous, free to base. Fertile stamens 5, alternating with and equalling corolla lobes in length. Disc entire, flattened. Ovary bilocular, with 2 ovules in each loculus, lanate on exterior. Style deeply divided into three, lanate at base, glabrescent above. Fruit ellipsoid, unilocular to

bilocular; epicarp densely tomentose, mesocarp thin; endocarp thin, bony, glabrous within.

TYPE. Ducke MG 15945, Brazil, Para, Gurupa, fl (holotype, RB; isotype, MG). DISTRIBUTION. This species is known from three collections in Brazilian

Amazonia. Collected in flower in October and January. BRAZIL. Amazonas: Froes 22493 fl (IAN, U); 28317 fr (IAN, U).

9. Dichapetalum axillare Woodson, Ann. Missouri Bot. Gard. 29: 353. 1942; Lundell,

Wrightia 3: 175. 1966; Lewis, Ann. Missouri Bot. Gard. 54: 10. 1967. Fig 9 G-L, 11.

Dichapetalum brenesii Standley, Publ. Field Mus. Bot. 23: 13. 1943. Type. A. Smith 20 (Herb. H.E. Stork 4120), Costa Rica, Zarcero, fl (holotype, F; isotypes, EAP, UC).

Shrub or small tree, the young branches sparsely tomentellous, becoming glabrous with age. Leaves oblong to oblong-elliptic, subcoriaceous, 6.0-16.0 cm long, 2.2-6.5 cm

broad, the apex tapered to a long slender acumen 5.0-25.0 mm long, usually subcuneate at base, plane not bullate above, glabrous or with a few stiff appressed hairs beneath; midrib impressed to slightly prominent above; primary veins 7-9 pairs, plane above,

26 Flora Neotropica

venation more or less plane above; petioles 4.0-8.0 mm long, tomentellous to tomentose.

Stipules to 2.0 mm long, deltoid, tomentose, caducous. Flowers hermaphrodite; pedicels 0.2-1.0 mm long. Inflorescences short terminal, axillary or petiolar panicles, the rachis and branches gray-brown tomentose. Bracts and bracteoles lanceolate, 0.5-2.0 mm long, tomentose, persistent. Calyx ca 2.5 mm long, the lobes almost equal, tomentose on exterior.Corolla of 5 equal, deeply bifid and slightly cucullate lobes, glabrous, free to base. Fertile stamens 5, alternating with and equalling corolla lobes. Disc of 5 tack-like, free glands. Ovary bilocular or trilocular with 2 ovules in each loculus, lanate on exterior. Style lanate at base, glabrescent above, the apex trifid. Fruit irregularly elongate-ellipsoid, to 3.8 cm long; epicarp shortly gray-brown tomentellous; mesocarp thin, endocarp thin, bony, glabrous within.

TYPE. Allen 2202, Panama, Cocle, hills north of El Valle de Anton, fl (holotype, MO; isotypes, EAP, GH, NY, US).

DISTRIBUTION. Mountain and coastal forests of Costa Rica, Panama and northern Colombia in shady places. Flowering March to July. COSTA RICA. Alajuela: Brenes 67664 fl (F, NY); A. Smith s n (Herb. H.E. Stork 4195) fl fr (MAD); s n (Herb. H.E. Stork 4120) fi (MAD); 4 (Herb. H.E. Stork 4104)4 fl (F, MAD, MICH, MO, NY, UC); 519 fl (EAP, F, MO); 746 4 fl (A, EAP, F, NY); 971 4 fl (A, F, MO, NY); 2157 fl (A, EAP); Standley 34674 st (US); 34723 fl (US). Heredia: Le6n 3705 fl (EAP); Skutch 3782 fl (A, K, NY, S, US); 3788 fl (A, K, MO, S, US). PANAMA. Chiriqui: Cooper & Slater 1724 fl (A, F, GH, MAD, NY, US). San Bias: Pittier 4349 fr (F). COLOMBIA. Choco: Romero C. 518 fr (COL, F).

10. Dichapetalum steyermarkii Prance, Act. Bot. Venez. 3: 301. 1968. Fig 15.

Liana, the young branches tomentellous, becoming glabrous with age. Leaves

oblong, coriaceous, 8.0-16.0 cm long, 3.0-5.0 cm broad, acuminate at apex, the acumen 3.0-13.0 mm long, rounded and unequal at base, plane not bullate above, glabrous beneath except for a few appressed hairs on venation; midrib more or less plane above, pubescent; primary veins 7-9 pairs, weakly impressed above; venation plane above; petioles 10.0-18.0 mm long, tomentellous when young, canaliculate. Stipules linear, ca 2.0 mm long, tomentellous, caducous. Flowers polygamo-dioecious; pedicels ca 0.3 mm

long. Inflorescences spreading petiolar panicles, 4.0-5.5 cm long, the rachis and branches

shortly tomentellous; peduncles 5.0-20.0 mm long. Bracts and bracteoles lanceolate, 0.3-1.5 mm long, persistent, tomentose. Calyx ca 2.0 mm long, tomentose on exterior, the lobes equal. Corolla of 5 equal slightly bifid lobes, free to base, glabrous. Fertile stamens 5, alternating and equalling corolla lobes. Disc of 2-3 glands, lobed at apex. Ovary bilocular, with 2 ovules in each loculus, lanate on exterior. Styles 2, short, connate at base. Fruit not seen.

TYPE. Steyermark 89875, Venezuela, Aragua, Parque Nacional Henry Pittier, fl

(holotype, VEN; isotypes, NY, US). DISTRIBUTION. Confined to the mountains of Aragua State, Venezuela. Collected

in flower in October. VENEZUELA. Aragua: Steyermark 89909 fl (paratypes, US, VEN).

This species differs from D. pedunculatum in the much longer petioles, the caducous

stipules and the petals which are only slightly divided at the apex. One of the paratypes of this species (Steyermark 62213) in fact belongs to D. pedunculatum. I associated this collection with D. steyermarkii incorrectly in the original description.

11. Dichapetalum donnell-smithii Engler, Bot. Jahrb. 23: 144. 1896; Lundell, Wrightia 3: 175. 1966. Fig 8, 11, 12.

Symphyllanthus donnell-smithii (Engler) Gleason, N. Am. Flora 25: 381. 1924.

Shrub, tree or vine, the young branches tomentose or tomentellous, becoming glabrous with age. Leaves oblong-lanceolate to ovate-elliptic, coriaceous, 6.0-28.0 cm

long, 2.5-12.5 cm broad, acuminate at apex, with acumen 2.0-12.0 mm long, cuneate to 4 Paratypes of D. brenesii


f n lero~asor sIb b

J:". o..- t I /; t?E .: " : *i

3 m :i:: :fii; :: ::: 1 ;: ! !..*....::?:%:.:.: .: ' '7 ?

FIG 8. Pictorialized scatter diagram of Dicbapetalum donnell-smitbii left of A-B, var donnell-smithbii; right of A-B var chiapasense.

- ~~.~~il>*<Kf. * pp.~? FI itnlie cte diga of Dihaeau donlszb.leto AB a

donl-mti;rgto A3-BP var cbipasnse

28 Flora Neotropica

subcordate at base, plane not bullate above, often hirsute when young above; hirsute, glabrous, or with a few appressed hairs beneath; midrib plane to weakly prominent above, pubescent; primary veins 7-12 pairs, plane to weakly prominent above; petioles 5.0-18.0 mm long, tomentose when young. Stipules linear, 1.0-25.0 mm long, tomentose. Flowers

hermaphrodite; pedicels 0.5-2.5 mm long. Inflorescences terminal and petiolar panicles, the peduncles and branches tomentose; peduncles 5.0-35.0 mm long. Bracts and bracteoles lanceolate, 0.5-2.0 mm long, tomentose. Calyx 2.0-2.5 mm long, tomentose on exterior, the lobes almost equal. Corolla of 5 equal bifid and often slightly cucullate lobes, glabrous, free to base. Fertile stamens 5, alternating and equalling the corolla lobes. Disc of 5 free, ellipsoid glands, lobed at apex. Ovary bilocular or trilocular, with 2 ovules in each loculus, lanate on exterior. Styles 3, united or free, lanate at base, glabrous at apex. Fruit unilocular to trilocular, ellipsoid; epicarp shortly tomentose; mesocarp thin; endocarp thin, hard, bony, glabrous within.

Key to the Varieties of Dichapetalum donnell-smithii

1. Leaves ovate-elliptic to oblong, 3.8-12.5 cm broad, the ratio length to breadth 1.6-2.7, usually hirsute beneath, rarely with only a few stiff appressed hairs; stipules 2.0-25.0 mm long. a. var donnell-smithii.

1. Leaves oblong to oblong lanceolate, 2.5-6.5 cm broad, the ratio length to breadth 2.5-4.1, glabrous or with a few stiff appressed hairs beneath; stipules always short, 1.0-3.0 mm long.

b. var chiapasense. la. Dichapetalum donnell-smithii var donnell-smithii. Fig 11.

TYPE. J.D. Smith 2067, Guatemala, Escuintla, fl (holotype B, lost; isotypes, G, GH, K, M, NY, P, US).

DISTRIBUTION. Scrubland, thicket and forests from southern Mexico to El Salvador and Honduras. Collected in flower around the year. MEXICO. Oaxaca: Martinez-

Calderon 17 fl (A, MEXU, UC, US). Tabasco: Matuda 6042 fl (MEXU); Rovirosa 192 fl (US); 783 fl (K, US). Chiapas: Matuda 4 fl (MICH); 630 fl (A, MICH, MO, NY, US); 5420 fl (EAP, MO); 15420 fl (EAP, F); 16535 fl (EAP, F, MICH); 16565 fl (EAP, F, MEXU); 17617 fl (EAP, F, K, MEXU, NY); 17942 fr (F); 17980 fr (F). GUATEMALA. Peten: Aguilar 243 fl (A, K, MICH, MO, NY). Alta Verapaz: Molina R. 12204 fl (NY); Standley 70549 fl (F); 91317 st (F, US). San Marcos: Steyermark 37992 st (F). Izabal: Popenoe 45 fl (F); Standley 24046 fl (GH, NY, US); 24496 fl (GH, MO, US); 24918 fl (NY, US); 25080 fl (GH, US). Quezaltenango: Steyermark 33520 fl (A, F). Retalhuleu:

Standley 87703 fl (F, S); 88436 fl (F). Escuintla: Standley 63465 st (F); 63876 fr (F, US); 64882 st (F, NY); 89569 st (F, US). Chimaltenango: Standley 62267 fl (F). Sacatepequez: Breedlove 11406 fr (F, MICH, US); J.D. Smith 2500 fr (F, K, MO, US); Roe et al 785 fr (F, WIS); 787 fl (F, WIS); Standley 63745 st (F); 65017 fl (F, US); 88939 fr (F, UC); 88968 fr (F). Santa Rosa: Heyde & Lux 4431 fl (BM, GH, K, M, US); 60659 st (F). BRITISH HONDURAS. Stann Creek: Schipp 714 fl (A, BM, F, GH, K, MICH, MO, NY, S, UC). Toledo: Gentle 6024 fr (F, NY, S, US); 6572 fr (NY); 6910 fr (S). HONDURAS. Cortes: Molina R. 5517 fr (EAP); L.O. Williams & Molina R. 14554 fl (EAP, F, GH); 17944 fl (EAP, F, GH). Atlantida: Yuncker et al 4816 fr (A, EAP, F, MICH, MO). EL SALVADOR. Ahuachapan: Standley & Padilla 3025 fl (EAP, F, NY, UC). Sonsonate: Pittier 1977 fl (BM, P, US). La Libertad: Calderon 1512 fl (NY, US). COSTA RICA. Cartago: Leon 2730 fr (EAP).

LOCAL NAMES. Mexico: Duraznito. British Honduras: Auselin.

lb. Dichapetalum donnell-smithii var chiapasense (Standley) Prance, stat nov Fig 12.

Dichapetalum chiapasense Standley, Publ. Field Mus. Bot. 17: 196. 1937.

Dichapetalum gentlei Lundell, Wrightia 3: 173. 1966, synon nov. Type. Contreras 5383, Guatemala, Alta Verapaz, Sebol, fl (holotype, LL; isotype, S).

TYPE. Matuda 679, Mexico, Chiapas, Monte Ovando, fr (holotype, MICH). DISTRIBUTION. Southern Mexico, Guatemala, British Honduras and Honduras in

scrub and forests. MEXICO. Veracruz: Sousa 3618 fr (MEXU). Chiapas: Matuda 4006 fr (A, MICH); 4208 fl (A, F, MEXU, MICH, NY, US). GUATEMALA. Peten: Contreras 6597 fr (S, US). Alta Verapaz: Contreras 4264 s fr (S, US); 429 5 fr (US). Huehuetenango: Steyermark 48618 st (F); 48800 fl (F, US). Suchitepequez: Steyermark 46613 fl (F, NY, US). HONDURAS. Atlantida: Yuncker et al 8491 fr (F, GH, K, MICH, MO, NY, S, US). Comayagua: Edwards 444 st (A, F, K, UC). BRITISH HONDURAS. Corozal: Gentle 46 fr (F); 61 fl (F); 4803 fl (F, MICH, NY); Lundell 4845 fl


FIG 9. Species of Dichapetalum. A-F, D. pedunculatum (De La Cruz 3688, 3826). A, habit X 0.5; B, flower X 7; C, flower section X 8; D, ovary and styles X 8; E, ovary section X 15; F, fruit X 1. G-L, D. axillare (Skutch 3782). G, habit X 0.5; H, flower X 4; J, flower section X 5; K, ovary and styles X 5; L, ovary section X 15.

~~~~~~~~~~~~~~ '%'.' ','.?.:. " , "g,~~~

?~~ ~-?~F.? ~S~D

..

,/

styles X 5; L, ovary section X 15.

30 Flora Neotropica

(F, MICH, MO, NY). El Cayo: Gentle 9194 fl (EAP, F, NY, S, US). Belize: Gentle 1305 fr (A, F, K, MO, NY); 1325 fl (F, K, MICH, MO, NY, S, US). Stann Creek: Gentle 2623 fl (A, F, K, MEXU, MICH, NY, US); 2864 fl (F, K, MICH, NY, US); 7999 fl (EAP, F, MICH, NY, S, US); Schipp 26 fr (F, GH, NY); 378 fl (A, BM, F, GH, MICH, MO, NY, S, UC, US). Toledo: Gentle 4219 fl (US); 5593 fl (EAP, S, US); 7206 5 fr (EAP, NY, S, US).

The following material of this species is intermediate between the two varieties, and is not placed in either here. MEXICO. Chiapas: Matuda 645 fl (MICH, MO, US); Miranda 7881 fl (MEXU). GUATEMALA. Izabal: Snedaker D-131 fl (F). BRITISH HONDURAS. Corozal: Gentle 501 fr (F, K, MEXU, MICH, US). El Cayo: Lundell 6163 fl (GH, MICH, NY, S). Stann Creek: Gentle 2644 fr (A, F, K, MEXU, MICH, NY); 2880 fl (A, F, K, MICH, NY). HONDURAS, Atlantida: Yuncker et al 8732 fr (F, GH, K, MICH, MO, NY, S, US).

Various workers have recently merged the two species D. donnell-smithii and D.

chiapasense. There is considerable overlap between these two species when all available material is studied, and it seems best to regard them as a single rather variable species. I have studied the variation in some detail, and find some correlation between the variables. While there is considerable overlap, material previously referred to as D. chiapasense tends to have a narrower more oblong leaf and glabrous or sparsely pubescent leaf undersurface, while that of typical D. donnell-smithii has broader and more elliptic leaves with a hirsute lower surface. This variation can be seen in Figure 8 which is a pictorialized scatter

diagram of the collections studied. Species lying to the left of line A-B fall into one

group which I have called variety donnell-smithii, while those to the right of the line fall into variety chiapasense. The area encircled contains specimens which are difficult to

assign to either variety, and most of them are unassigned in this work. Since the variation does not appear to be ecologically or geographically correlated,

and since there is overlap, I regard the two extremes as varieties rather than subspecies. Dichapetalum gentlei which was described recently falls well within the range of

variation in this species and is treated as a synonym of var chiapasense.

12. Dichapetalum spruceanum Baillon, Mart. Fl. Bras. 12(1): 371. 1886; Macbride, Publ. Field Mus. Bot. 13(3): 957. 1950. Fig 13.

Symphyllanthus plicatus Gleason, Phytologia 1: 26. 1933, synon nov. Type. Lawrance 414, Colombia, Boyaca, Alto Chapon, fl (holotype, NY; isotypes, A, F, MO, S).

Tree, shrub, scandent shrub or vine, the young branches tomentellous becoming glabrous with age. Leaves ovate to elliptic, coriaceous, 8.0-30.0 cm long, 4.0-16.5 cm broad, abruptly acuminate at apex, the acumen 5.0-10.0 mm long, usually curved, rounded to cuneate and unequal at base; plane not bullate above, sparsely hirsute or with a few stiff appressed hairs only beneath; midrib more or less plane above, pubescent when

young; primary veins 8-10 pairs, plane to slightly impressed above; venation plane to

slightly impressed above; petioles 7.0-15.0 mm long, sparsely short-tomentellous or

puberulous when young. Stipules to 8.0 mm long, lanceolate, tomentellous, caducous, the

margins entire. Flowers hermaphrodite; pedicels ca 0.5 mm long. Inflorescences axillary and petiolar spreading panicles, 4.0-6.5 cm long, the rachis and branches shortly puberulous; peduncles 8.0-30.0 mm long. Bracts and bracteoles triangular, 0.5-2.0 mm

long, persistent, tomentose. Calyx ca 2.0 mm long, gray-tomentose on exterior, the lobes

equal. Corolla of 5 equal lobes, the apex deeply bifid, slightly cucullate, free to base, glabrous. Fertile stamens 5, alternating and equalling the corolla lobes. Disc of 5 glands, the apices lobed. Ovary bilocular or trilocular, with 2 ovules in each loculus, lanate on exterior. Style with bifid or trifid apex, glabrous except for lanate base. Fruit ellipsoid; epicarp with short dense appressed pubescence; mesocarp thin; endocarp thin, hard, glabrous within.

TYPE. Spruce 4927, Peru, San Martin, Tarapoto, fl (holotype, K; isotypes, BM, CGE, P, W). 5 Paratype of D. gentlei.


0

b7 60 50

FIG 10. Distribution of species of Dichapetalum. D. nervatum; A D.stipulatum; o D. coelboi; A D. pauper.

32 Flora Neotropica

DISTRIBUTION. Forests of the eastern fringes of the Andes in Colombia, Ecuador and Peru. COLOMBIA. Meta: Triana s n fr (BM). Putumayo: Scbultes & Cabrera 19074 fl (A, US). Without locality: Mutis 1140 fl (US); 2957 fl (US); 2959 fl (US); 2960 fl (US); 4035 fl (US); 5095 fl (US); 5807 fl (US). ECUADOR. Napo-Pastaza: Asplund 10179 fl (G, K, US); 10311 fl (G, US);Mexia 7285 fl (F, UC, US). PERU. San Martin: Klug 2751 fl (A, BM, F, G, GH, K, NY, S, US); 4306 fl (A, BM, F, K, MO, NY, S, U, UC, US); Ll. Williams 6605 fl (F, NY).

This species is close to D. rugosum but differs in the less hirsute leaves with less

impressed venation, and the gray-tomentellous axillary inflorescence. The unicate collection, Killip 34375, from the State of Meta, Colombia probably

belongs to this species. It has much larger leaves than any other collections examined, and

only further material will show whether this is part of D. spruceanum or whether it is a new species with much larger leaves than any other American Dichapetalum.

Symphyllantus plicatus was described without comparison to South American

species of Dichapetalum. There are no differences to justify maintaining this species in

Dichapetalum since it matches D. spruceanum.

13. Dichapetalum froesii Prance, sp nov Fig 7 F-L, 15.

Frutex scandens, ramulis juvenilibus sparse tomentellis. Folia alternata petiolata; petiolo 8.0-32.0 cm longo, sparse tomentello; laminae ellipticae, chartaceae, 12.0-28.0 cm longae, 5.0-15.0 cm latae, apice in acuminem 8.0-15.0 mm longum contractae, basi rotundatae vel subcuneatae, aequales, supra planac haud bullatae, subtus cum pilis rigidis sparse munitae; costa media supra plana vel leviter impressa; costis secundariis 7-10 jugis, supra planis; venis supra planis leviter impressis. Stipulae lanceolatae, ad 7.0 mm longae, caducae, tomentellae. Flores hermaphroditi, in paniculis 3.0-8.0 mm longis ad petiolos affixi, rachi ramisque sparse tomentellis vel puberulis, pedunculis 1.5-4.5 mm longis, pedicellis 1.5-2.0 mm longis. Bracteae bracteolacque 0.5-2.0 mm longae, lanceolatae, persistentes, puberulae. Calyx circa 2.5 mm longus, extus griseo-puberulus, lobis

aequalibus. Petala 5, glabra, aequalis, ad basim libera, apice leviter bifida. Stamina 5, omnis fertilis, cum calycis-lobis alternata aequantes. Discus constans ex glandulis 5, ellipsoideus, integer. Ovarium biloculare, ovulis in quoque loculo duobus, extus lanatum.

Styli 2, versus basim lanati, supra glabri. Drupa ellipsoidea; epicarpio breviter appresso- tomentello; mesocarpio tenue; endocarpio tenuissimo, oseo, intus glabro.

TYPE. Black 48-2884, Brazil, Amazonas, Rio Igana, Tunui, fl (holotype, NY;

isotypes, COL, IAN, P). DISTRIBUTION. Western Amazonia and the foothills of the Colombian Andes.

Flowering April to June. COLOMBIA. Caqueta: Cuatrecasas 8975 fl (COL, F, NY, US). Putumayo: Garcia-Barriga et al 18677 fl (COL, NY). Brazil. Amazonas: Froes 22325 fl (IAN, U, UB, WIS); 22850 fr (IAN).

This species is close to D. pedunculatum but differs in the much larger leaves, and the longer petioles and pedicels.

14. Dichapetalum odoratum Baillon, Mart. Fl. Bras. 12(1): 371. 1886; Lasser, Bol. Soc. Venez. Cienc. Nat. 9: 127. 1944; Macbr., Publ. Field Mus. Bot. 13(3): 955. 1950.

Fig 15. Chailletia odorata Spruce ex Baillon, Mart. Fl. Bras. 12(1): 371. 1886, in synon.

Liana, the young branches puberulous to glabrous. Leaves elliptic to oblong, chartaceous to coriaceous, 8.0-16.0 cm long, 3.5-9.0 cm broad, acuminate at apex, with acumen 5.0-15.0 mm long, rounded to cuneate, and equal or slightly unequal at base, plane not bullate above, glabrous or with a few stiff appressed hairs on primary venation beneath; midrib more or less plane, sparsely puberulous to glabrous above; primary veins 7-9 pairs, slightly prominent above; venation plane to prominent above; petioles 4.(-8.0


. - ----4-20

?_____ _ _.

95 90 85 80

FIG 11. Distribution of species of Dichapetalum. * D. donnell-smithii var donnell-smithii; A D. bullatum; * D. nevermannianum; o D. axillare. bullatum; ? D. nevermannianum; o D. axillare.

34 Flora Neotropica

mm long, puberulous to glabrous. Stipules linear-lanceolate, to 5.0 mm long, caducous, the margins entire. Flowers hermaphrodite; pedicels 1.0-3.0 mm long. Inflorescences

spreading axillary or petiolar panicles, the rachis and branches tomentellous to glabrous. Bracts and bracteoles 0.5-1.0 mm long, lanceolate, persistent, puberulous. Calyx 1.5-2.0 mm long, gray-puberulous on exterior, the lobes equal. Corolla of 5 equal lobes, deeply bifid, glabrous, free to base. Fertile stamens 5, alternating and equalling corolla lobes. Disc of 5 short united glands, the apices bifid. Ovary bilocular with 2 ovules in each loculus, lanate on exterior. Style lanate on lower portion, the apex trifid. Fruit not seen.

TYPE. Spruce 2864, Brazil, Amazonas, Panure, fl (holotype, P; isotypes, BM, BR, C, CGE, F, G, GH, GOET, K, LE, NY, OXF, P, RB, W).

DISTRIBUTION. Forests and caatingas of western Amazonia in Venezuela, Peru and Brazil. VENEZUELA. Terr. Amazonas: Ll . Williams 14110 fl (G). PERU. Loreto: Asplund 14097 fl (G, US); Killip & Smith 29834 fl (US); Klug 122 fl (F, NY, US); 182 fl (F, NY, US); 226 fl (F, NY, US); 551 fi (F, NY, US); 3070 fl (A, BM, F, G, GH, K, MO, NY, S, US); Sandeman 2254 fl (K, OXF); 2265 fl (K, OXF); Tessmann 4514 fl (G); 5377 fl (S); LI. Williams 3640 fl (F, NY); 3783 fl (F, US); BRAZIL. Amazonas: Krukoff 8077 fl (A, BR, F, G, K, LE, MICH, MO, NY, P, S, U); 8351 fl (A, BM, BR, F, G, K, LE, MICH, MO, NY, P, S, U); 8368 fl (A, BM, BR, F, G, K, LE, MICH, MO, NY, P, S, U); J.G. Kuhlmann RB 20714 fl (S, U, US).

This species is close to D. pedunculatum, and earlier I suggested that it was

synonymous. However, it is distinct by a number of differential characters, and should be maintained as a separate species. It differs from D. pedunculatum in the tendency to

longer pedicels, the smaller caducous stipules, and the less pubescent inflorescence and leaves.

15. Dichapetalum pedunculatum (De Candolle) Baillon, Adansonia 11, t. 9, fig. 7, 1874; Hist. P1. 5: 140. 1874; Mart. Fl. Bras. 12(1): 369. 1886; Stafleu in Pulle, Fl. Suriname 3(2): 167. 1951; Rizzini, Revista Bras. Biol. 12(1): 100. 1952. Fig 9 A-F, 15.

Chailletia pedunculata De Candolle, Nouv. Bull. Soc. Philom. Paris 40: 205. 1811; Prodr. 2: 57. 1825.

Symphyllanthus glaber Vahl, Skr. Naturh.-Selsk. Kj0benk 6: 86. 1810. Type. Richard s n, French Guiana, fl (holotype C, herb. Vahl; isotypes, G, P).

Dichapetalum glabrum (Vahl) Prance, Act. Bot. Venez. 3: 304. 1969, nom illegit.

Liana, the young branches tomentellous to glabrous. Leaves ovate-elliptic to oblong, coriaceous, 5.0-16.0 cm long, 2.0-7.5 cm broad, acuminate at apex, subcordate to subcuneate and slightly unequal at base, plane not bullate above, glabrous or with a few stiff hairs on venation only beneath; midrib more or less plane above, pubescent to

glabrous; primary veins 7-9 pairs, more or less plane above, the venation plane above; petioles 4.0-12.0 mm long, shortly tomentellous at least when young. Stipules persistent, linear, to 10.0 mm long, puberulous to tomentellous, the margins entire. Flowers

hermaphrodite; pedicels 0.2-1.5 mm long. Inflorescences spreading panicles, usually petiolar, sometimes terminal, the rachis and branches tomentellous to glabrescent; peduncles 1.0-3.5 cm long. Bracts and bracteoles 0.5-2.0 mm long, lanceolate, persistent, tomentose. Calyx 1.5-2.0 mm long, tomentose on exterior, the lobes almost equal. Corolla of 5 equal lobes, deeply bifid, glabrous, free to base. Fertile stamens 5, alternating and equalling or shorter than the corolla lobes. Disc of 5 large glands with lobed apices. Ovary bilocular or trilocular with two ovules in each loculus, lanate on exterior. Styles lanate at base, glabrous above, divided into 2 or 3 or united with a trifid apex. Fruit

ellipsoid, most frequently unilocular, but often bilocular; epicarp short-tomentellous; mesocarp thin; endocarp thin, hard, bony, glabrous within.

TYPE. Patris s n, French Guiana, fl (holotype, G-DC). DISTRIBUTION. Primary and secondary forest, and river margins from Trinidad

and northern Venezuela through the Guianas to the western part of Brazilian Amazonia.


20

I' S . L' ------------"-- - -----, ._: L -. : .2 ( ] -2 0

?1 a .

95 90 85 FIG 12. Distribution of Dichapetalum donnell-smitbii; o var chiapasense; A var uncertain. 2~IG 12. Distribution of Dichapetalum donnell-smithii; o var chiapasense;

x var uncertain.

36 Flora Neotropica

Flowering around the year. TRINIDAD. Bot. Garden 487 fl (NY, US); 2134 fl (US); Broadway s n fl (NY); Crueger sn fl (GOET, K); Purdie sn fl (K); Simmonds 364 fl (K, TRIN); 395 fl (K). VENEZUELA. Dist. Federal: Karsten s n fl (LE, W); Steyermark 94763 fl (NY). Monagas: Steyermark 62213 fl (F, NY, VEN). Terr. Delta Amacuro: Rusby & Squires 131 fl (BM, F, G, GH, K, LE, M, MO, NY); Steyermark 87305 fr (NY, US, VEN). Bolivar: Lasser 1927 fl (F, NY, US, VEN); Maguire, Steyermark, & Maguire 46878 fl (NY); Steyermark 60427 fr (F, NY, VEN); 86347 fl (NY, VEN); 86499 fl (NY, US, VEN); 89121 fl (NY, VEN); 90708 fl (NY, US, VEN). GUYANA. Aitkin 40 fl (K); Archer 2397 fl (K, US); 2468 fl (K, US); Beckelt & Kortright 8534 fl (K, U); De La Cruz 1108 fl (NY); 1193 fr (GH, NY, US); 1596 fl (F, GH, MO, NY, UC, US); 1895 fl (F, GH, MO, NY, UC, US); 2625 fl (F, GH, MO, NY, UC, US); 3095 fl (F, GH, MO, NY, UC, US); 3543 fl (F, GH, MO, NY, US); 3688 fr (F, GH, MO, NY, UC, US); 3826 fl (F, GH, K, MO, NY, UC, US); 3905 fl (F, GH, MO, NY, UC, US); 4041 fl (GH, NY, US); 4337 fr (F, NY, US); 4517 fl (F, GH, MO, NY, US); Fanshawe 205(F.D.2941) fl (K); 230(F.D.2966) fl (K, S); 1079(F.D.3815) fr (K); Gleason 205 fl (NY, US); Irwin 198 fl (US); Jenman 1718 fl (K); 2355 fl (K, NY); 4326 fl (K); Parker s n fl (K); Persaud 238 fl (F); Sandwith 121 fr fl (K, LE, NY, P, S, U, US); Schomburgk 477 fl (BM, CGE, F, G, GH, K, L, P, US, W); A.C. Smith 2800 fr (A, F, K, NY, P, U, US); Stockdale 8808 fl (K); Tillett & Tillett 45762 fl (NY)); Tutin 163 fl (BM, K, RB, U, US); 315 fi (BM, K, U, US). SURINAME. Gonggrijp & Stahel BW 6541 fl (BR, K, MO, NY, U); Herb. Dept. Lanbouw 286 fl (U); Heyligers 470 st (U); Lanjouw 1275 fr (K, NY, U); Lindeman 4652 fl (U); Maguire 24669 fr (MICH, MO, NY, U, UC, W);Maguire & Cowan 39003 fr (NY); Pulle 305 fl (L, RB, U); Samuels s n fl (US); Van Donselaar 3085 st (U). FRENCH GUIANA. Broadway 659 fr (GH, NY); 831 fl (GH, NY, US); Crevaux s n fl (P); Leblond 347 fl (P); 454 fl (P); s n fl (P); Lemee s n fl (P); Leprieur s n fl (G); s n (1838) fl (P); s n (1840) fl (P); s n (1849) fl (P); sn (1850) fl (P); Martin s n fl (BM, K);Melinon s n (1842) fl (P); Perrottet s n fl (P); Poiteau s n fl (K, P); Rothery 117 fl (K); Sagot 191 fl (BM, BR, G, GH, GOET, K, L, NY, P, S, U, W); Wachenheim s n fl (BM, K, P); 250 fl (P); 254 fl (P). BRAZIL. Amazonas: Ducke RB 23656 fl (RB); Krukoff 7072 fl (BM, BR, F, G, IAN, K, LE, MICH, MO, NY, S, W); Martius s n fl (M); Prance et al 8116 fl (INPA, NY). Terr. Roraima: Prance et al 9849 fl (INPA, NY). Para: Black 47-2030 fl (IAN); Black & Ledoux 50-10252 fl (IAN); Burchell 10107 fl (BR, GH, K, P);Ducke MG 3285 fl (MG);MG 15991 fl (BM, MG, RB 13567); RB 20713 fl (RB, S, U, US); RB 20715 fr (RB, S, US); Froes 30871 fr (IAN); T. Guedes 90 fr (IAN); Killip & Smith 30538 fl (NY, US); Pires 3106 fl (IAN, U, UB, US); Pires & Black 1287 fl (IAN, MICH, RB); 1652 fl (COL, IAN, P, U, UC); Pires & Silva 11653 fr (IAN); A. Silva 421 fl (IAN, NY, RB). Without locality: Ferreira s n fl (P).

LOCAL NAMES. Guyana: Kahakudiballi rope, Massari (Arawak)

I originally referred Steyermark 62213 to D. steyermarkii, but this collection is in

fact D. pedunculatum and is cited here. In making the new combination D. glabrum (Vahl) Prance, I inadvertently

overlooked the name D. glabrum Elmer; hence my new name is illegitimate and this

species must be referred to as D. pedunculatum by which it has generally been known.

De Candolle, in his original description of Chailletia pedu;culata, did not give any details about this type specimen except to say that it was from Cayenne. Baillon in 'Flora

Brasiliensis' cited one of the Paris sheets as the type: Herb Olyssip s n, from Para, Brazil.

This sheet came from the herbarium Lusitania in Lisbon, and was presumably collected

by A.R. Ferreira in Brazil between 1783 and 1792. It could have been at Paris prior to De

Candolle's description of C. pedunculata, and was possibly seen by him. However, it

disagrees with de Candolle's description in two features, firstly in the locality, Brazil not

Cayenne, and secondly in the mature flowers. De Candolle was unable to describe mature

flowers of C. pedunculata, yet the Herb. Olyssip. material has many open flowers. Baillon

was certainly incorrect in his designation of the type collection of D. pedunculatum. De

Candolle in the 'Prodromus' gave a synoptic description of D. pedunculatum which

included the citation of a single collection by Patris. Thus the type is certainly the Patris

collection. The De Candolle herbarium at Geneva has three sheets of D. pedunculatum all without field notes or collectors' names. One of these sheets is, however, marked Cayen- ne. This is almost certainly the type collection, and at least one of the three sheets in the De Candolle herbarium, possibly all three, is the Patris collection.

2. Stephanopodium Poeppig & Endlicher, Nov. Gen. et Sp. 3: 40, t. 246. 1842;

Endlicher, Gen. Suppl. 3: 97. 1843; Bentham & Hooker, Gen. PI.: 341. 1862;


. - ~ ~( '

.. ~~~-"10

I O

80 70 60

FIG 13. Distribution of species of Dichaperalum. A D. spruceanum; * D. latifolium.

FIG 13. Distribution of species of Dichapetalum. a D. spruceanum; ? D. latifolium.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.

38 Flora Neotropica

Baillon, Mart. Fl. Bras. 12(1): 376. 1886; Engler in Engler & Prantl, Nat. Pflanzenfam. 3(4): 350. 1896; Engler & Krause, Nat. Pflanzenfam. 19c: 9-10. 1931;

Macbride, Publ. Field Mus. Bot. 13(3): 958. 1950; Rizzini, Revista Bras. Biol. 12(1): 101-104. 1952; Hutchinson, Gen. PI. 1: 218. 1964; Prance, Act. Bot. Venez. 2(2): 50. 1966; Ind. Nom. Gen. 10/03000.

Slender trees or shrubs. Leaves alternate, petiolate, entire. Stipules small, deciduous. Inflorescences of small sessile or shortly pedunculate glomerules adnate to the petioles. Flowers hermaphrodite or polygamo-dioecious, actinomorphic. Bracts small. Receptacle campanulate to cylindrical. Sepals 5, imbricate, slightly to very unequal. Petals united into a long obconical or rarely cylindrical, tube, with 5 equal broadly ovate lobes which are shorter than the tube, the lobes entire at the apex. Stamens 5, all fertile, alternate with corolla lobes and adnate to mouth of corolla tube; anthers sessile or subsessile. Disc

FIG 14. Distribution 50

FIG 14. Distribution of Dichapetalum rugosum.


. 10~

~~~~~10

70 60 50

FIG 15. Distribution of species of Dichapetalum. o D. froesii; A D. odoratum; + D. steyermarkii; * D. pedunculatum.

40 Flora Neotropica

of 5 hypogynous glands which are opposite the corolla lobes, the glands entire, equal to

unequal, free or connate. Ovary globose, free, 2-3-locular with 2 ovules in each loculus.

Styles 2-3, free or connate to apex. Fruit a dry coriaceous drupe, usually bilocular but sometimes with only one loculus developing.

TYPE SPECIES. Stephanopodium peruvianum Poepp. & Endl. The name Stephano- podium is derived from the Greek words aTreyoc0yo (stephanos = a crown or wreath), and irobLov (podion = a foot), referring to the flowers which 'crown' the petioles at the base 'foot' of the leaf.

DISTRIBUTION. Colombia, Peru, Venezuela, and in eastern central Brazil, but not in the intervening area.

Baillon separated S. engleri into a separate section, Isorthosiphon Baillon, Mart. Fl. Bras. 12(1): 378. 1886. This was followed by Engler and Krause (1931), and was based on the shape of the corolla tube. Since the tube varies in shape and since S. engleri does not seem to be more isolated in any way from the other species of this small genus, no sections are adopted in this treatment.

Key to species of Stephanopodium 1. Corolla tube exserted beyond calyx lobes, a long narrow cylinder; calyx lobes extremely

uneual. 2. Leaves oblong to lanceolate, 5.0-9.5 cm long, 1.7-2.8 cm broad; petioles canaliculate;

corolla tube lanate within. 1. S. engleri. 2. Leaves elliptic, 10.0-16.0 cm long, 3.5-6.0 cm broad; petioles terete; corolla tube

glabrous within. 2. S. aptotum. 1. Corolla tube and usually the lobes included, broad and obconical; calyx lobes slightly

unequal. 3. Leaf undersurface hirsutulous pubescent. 3. S. estrellense. 3. Leaf undersurface glabrous or with a few stiff appressed hairs only.

4. Midrib extremely prominent above. 5. Calyx 2.0-2.5 mm long; pedicels 1.5-2.0 mm long; corolla tube glabrous within;

leaves 9.0-18.0 mm long; primary veins 7-10 pairs. 4. S. peruvianum. 5. Calyx 5.0-6.0 mm long; pedicels 3.0-5.0 mm long; corolla tube lanate within;

leaves 12.0-22.0 mm long; primary veins 10-12 pairs. 5. S. angulatum. 4. Midrib impressed or plane above.

6. Petioles 9.0-15.0 mm long; tomentellous when young; leaf apex rounded to acute. 8. S. blanchetianum.

6. Petioles 4.0-7.0 mm long (to 10.0 mm in S. venezuelanum); leaf apex usually distinctly acuminate. 7. Flowers sessile to subsessile; midrib usually impressed above, rarely plane.

8. Leaves ovate-elliptic, with a long acumen 3.0-20.0 mm long; inflorescences often borne near to stem; flowers polygamous. 6. S. venezuelanum.

8. Leaves oblong, with a short blunt acumen 0-3.0 mm long; inflorescences always borne on middle of petioles; flowers hermaphrodite. 7. S. sessile.

7. Flowers distinctly pedicellate, pedicels 1.0-2.0 mm long; midrib plane above. 9. S. organense.

1. Stephanopodium engleri Baillon, Adansonia 11: 109, t.9. 1873; Baillon, Mart. Fl. Bras. 12(1): 378. 1886; Rizzini, Revista Bras. Biol. 12(1): 102. 1952. Fig 16 A-G, 19.

Small tree, the young branches sparsely puberulous, soon becoming glabrous. Leaves

oblong to lanceolate, subcoriaceous, 5.0-9.5 cm long, 1.7-2.8 cm broad, gradually tapering to an acuminate apex, the acumen 6.0-15.0 mm long, cuneate and equal to

slightly unequal at base, glabrous beneath; midrib slightly impressed above, prominent and with a sparse appressed pubescence beneath; primary veins 7-10 pairs, arcuate,

anastomosing; petioles 5.0-12.0 mm long, canaliculate, sparsely pubescent. Stipules lanceolate, to 3.0 mm long, tomentellous, subpersistent. Flowers hermaphrodite, borne in dense-flowered sessile glomerules inserted on petioles, with very short pedicels; bracteoles


^SH J I

IN

or .- i

FIG 16. Species of Stepanopodium A S engleri (Magales s IAN 14709, 14564). A,

habit X 0.5; B, flower X 5; C, flower section X 5; D, ovary section X 15; E, fruit X 0.5; F, corolla and stamens X 5; G, ovary and style X 5. H-O, S. estrellense (Glaziou 13795, Weddell 769). H, flower X 5; J, flower section X 2.5; K, young fruit X 0.5; L, corolla and stamens X 5; M, ovary and styles X 5; N, ovary section X 15; 0, habit X 0.5.

42 Flora Neotropica

ca 0.5 mm long, persistent, tomentose. Calyx 3.0-4.0 mm long, tomentose on exterior, the lobes unequal. Corolla exserted well beyond calyx lobes, of a long exserted tube and 5 small bifid lobes which are not cucullate, the tube glabrous on exterior, lanate near mouth and glabrous at base within. Fertile stamens 5, alternating with corolla lobes, the anthers sessile, inserted at mouth of corolla tube. Ovary bilocular with 2 ovules in each loculus, tomentose on exterior. Style deeply divided into two, sparsely pubescent. Fruit

irregularly ellipsoid, ca 1.5 cm long; epicarp velutinous-ferrugineous; mesocarp thin; endocarp thin, bony, rather soft, glabrous within.

TYPE. Warming 1841, Brazil, Minas Gerais, Lagoa Santa, fl (holotype, P; isotype, C).

DISTRIBUTION. Known only from woodland and forest in the State of Minas Gerais, Brazil. Flowering December to April. BRAZIL. Minas Gerais: Barreto 10584 fl (RB); 10585 fl (RB); 10750 fr (RB); Damazio 13386 fl (RB 112331); Herb. Schwacke 13386 fl (P); Magalhaes s n (Mar. 1940) fr (IAN, US); s n (Apr. 1940) fl (IAN, US); 140 fl fr (RB); Warming s n fl (C, F, P). 6

The typification of this species presents a problem since, in the original description, Baillon cited, "Lagoa Santa inter sylvas legit Engler (exs., n. 1091), aprili et decembre floriferum (Herb. Warming)." There is no Engler collection in the Warming herbarium, and Engler never collected at Lagoa Santa. The actual material examined by Baillon is

undoubtedly the material cited above, Warming 1841, which was collected at Lagoa Santa, "in sylvis." There are three gatherings of this number one of which was collected in April and another in December 1864. Furthermore the collectors name is abbreviated to Eug. W. which could easily be mistaken for Engler since the script is of a flowing nature. In 'Flora Brasiliensis' some years later, Baillon cited Warming as the collector of this species, and did not mention Engler. Both the Paris and Copenhagen sheets of

Warming 1841 are annotated in Baillon's handwriting.

2. Stephanopodium aptotum Wheeler, Proc. Biol. Soc. Wash. 53: 7. 1940. Fig 18.

Small tree, the young branches glabrous. Leaves elliptic, coriaceous, 10.0-16.0 cm

long, 3.5-6.0 cm broad, acute to bluntly acuminate at apex, cuneate and equal at base,

glabrous beneath; midrib plane to weakly impressed above, prominent and glabrous beneath; primary veins 9-10 pairs, arcuate, anastomosing; petioles 6.0-10.0 mm long, glabrescent, terete, rugose. Stipules lanceolate, glabrescent, to 4.0 mm long, caducous. Flowers hermaphrodite, borne in dense-flowered sessile glomerules on petiole, with long pedicels to 5.0 mm long; bracteoles 0.5-1.0 mm long, persistent, sparsely tomentellous.

Calyx 5.0-6.0 mm long, sparsely puberulous on exterior, the lobes unequal, puberulous near apex on exterior. Corolla exserted well beyond calyx lobes, of a long exserted tube and 5 small entire equal lobes, the tube glabrous on both surfaces. Fertile stamens 5,

alternating with corolla lobes, the anthers sessile, inserted at mouth of corolla tube. Ovary trilocular with 2 ovules in each loculus, shortly tomentellous on exterior. Style single but

deeply divided into three for upper portion, shortly tomentellous. Fruit not seen. TYPE. H.H. Smith 1701, Colombia, Magdalena, Santa Marta, fl (holotype, GH;

isotypes, BM, G, K, MO, NY, US). DISTRIBUTION. Endemic to the highland forest of the Santa Marta mountains,

Colombia. COLOMBIA. Magdalena: Espina & Giacometto A23 st (MAD); A59 st (MAD); A 124 st (MAD);A145 st (MAD); Kernan 102 fl bud (NY, P, US).

3. Stephanopodium estrellense Baillon, Mart. Fl. Bras. 12(1): 377. 1886; Rizzini, Revista Bras. Biol. 12(1): 102. 1952. Fig 16 H-0, 19.

Stephanopodium sessiliflorum Kuhlmann ex Rizzini, Revista Bras. Biol. 12(1): 104. 1952, synon nov. Type. Kuhlmann RB 73047, Brazil, Guanabara, Horto Florestal, Rio de Janeiro, fl

(holotype, RB; isotype, NY).

6 Probable isotypes


Small tree, the young branches tomentellous, becoming glabrous with age. Leaves

oblong to oblong-lanceolate, coriaceous, 9.0-12.5 cm long, 2.5-4.2 cm broad, acuminate at apex, the acumen 5.0-15.0 mm long, cuneate and equal to slightly unequal at base, hirsutulous beneath; midrib plane to weakly impressed beneath; primary veins 8-10 pairs, arcuate, anastomosing; pedicels 6.0-10.0 mm long, canaliculate, tomentose when young. Stipules lanceolate, to 4.0 mm long, tomentose, caducous. Flowers hermaphrodite, borne in dense-flowered sessile glomerules inserted on the petiole, with short pedicels; bracteoles ca 0.5 mm long, tomentose, persistent. Calyx 4.0-5.0 mm long, tomentose on exterior, the lobes almost equal. Corolla with 5 small equal, slightly bifid lobes and a broad included tube, the tube pubescent above and glabrous below on exterior, lanate above and glabrous below within. Fertile stamens 5, alternating with corolla lobes, the anthers sessile, inserted at mouth of corolla tube, equalling corolla lobes in length. Ovary 2- 3-locular with 2 ovules in each loculus, villous on exterior. Style 1, but divided into 2 or 3 for half of length, pubescent almost to apex. Young fruit rotund, bilocular; epicarp velutinous tomentose; pericarp thin, bony, glabrous within.

TYPE. Weddell 769, Brazil, Guanabara, Rio de Janeiro, fl (holotype, P; isotypes, F, G).

DISTRIBUTION. Known only from forests around Rio de Janeiro, Brazil. BRAZIL, Gaunabara: Glaziou 13795 fr (C, G, K, LE, P).

This species is easily recognized within the genus by the hirsutulous undersurfaces of the leaf. The only significant difference mentioned by Kuhlmann between S. sessiliflorum and S. estrellense was the sessile flowers of the former. This difference does not hold, and these two names are conspecific.

4. Stephanopodium peruvianum Poeppig & Endlicher, Nov. Gen. et Sp. 3: 40, t. 246. 1842; Macbride, Publ. Field Mus. Bot. 13(3): 958. 1950. Fig. 17 A-F, 18.

Tapura costata Cuatrecasas, Lloydia 11: 221. 1949. Type. Cuatrecasas 16746, Colombia, El Valle, R;o Calima, fl bud (holotype, F; isotype, COL).

Small to medium sized tree, the young branches sparsely puberulous, soon

becoming glabrous. Leaves elliptic, coriaceous, 9.0-18.0 cm long, 3.0-7.0 cm broad, acuminate at apex, the acumen abrupt, 5.0-13.0 mm long, rounded to cuneate at base, glabrous beneath; midrib distinctly prominent above, prominent and glabrous or with a few stiff appressed hairs beneath; primary veins 7-10 pairs, arcuate, anastomosing; petioles 5.0-9.0 mm long, glabrous or sparsely puberulent when young, shallowly canaliculate, rugose. Stipules deltoid, to 3.5 mm long, sparsely pubescent, caducous. Flowers androdioecious, borne in dense-flowered sessile glomerules inserted on petioles, pedicels 1.5-2.0 mm long; bracteoles ca 0.5 mm long, ovate, tomentellous, persistent. Calyx ca 2.5 mm long, gray puberulous on exterior, the lobes unequal. Corolla equalling calyx in length, with 5 equal lobes inserted on long tube, the tube glabrous on exterior,

glabrous except at base of anthers within. Fertile stamens 5, alternating with corolla

lobes, the anthers sessile, inserted at mouth of corolla tube. Ovary bilocular, tomentose on exterior. Style single, the apex trifid. Disc of 5 large free glands surrounding the ovary. Fruit ellipsoid; epicarp with short dense appressed pubescence; mesocarp thin, fleshy; endocarp thin, very hard, bony, glabrous within.

TYPE. Poeppig 1986, Peru, San Martin, Tocache, fl (holotype, W; isotypes, G, L, LE, P).

DISTRIBUTION. Primary forests of Pacific Colombia, Amazonian Colombia and Peru, and upland in Peru to Huianuco. Flowering April to July. COLOMBIA. Amazonas: Gar- cia-Barriga 13846 fr (US); Schultes & Cabrera 14157 fl (A, US). PERU. Loreto: Klug 1985 fl (A, BM, F, GH, K, MICH, MO, NY, US). Hutanuco: Schunke V. 858 fr (f).

Material of this species has been much confused with species of Tapura in the past.

44 Flora Neotropica

0 H

FIG 17. Species of Stephanopodium. A-F, S. peruvianum (Klug 1985, Schunke 858). A, flower X 6; B, flower section X 6; C, habit X 0.5; D, ovary section X 10; E, fruit X 0.5; F, corolla and stamens X 6. G-L, S. organense (RizziniRB 132051). G, flowerX 6;H,flowersectionX 6; J,corolla and stamens X 6; K, ovary and style X 10; L, habit X 0.5.

X 6; , floer setion 6; habi .. ..:.5; ,oayscinX1;EfrtX0.";Fool n

stamens X 6~~~~~~~~~~~~. j-,S raes Rzn B125) ,fowrX6 ,foe eto ;J ool

and stamens X 6; K, ovary and style X 10; L, habit X 0~~~~~~~~~~~~~.5.


Sterile and fruiting material is quite similar to T. guianensis and T. peruviana, but is easily distinguished by the extremely prominent leaf midrib.

The species Tapura costata was described from poor material with young flower buds only. Although difficult to assess from this material the anthers are certainly sessile and the corolla lobes are equal, making this a species of Stephanopodium. Although the

type of T. costata was collected far from any other known collections of Stephano- podium peruvianum, it matches the latter species in all respects, and is hence placed in

synonymy. Stephanopodium peruvianum is most closely related to S. angulatum; for differences

see under that species.

5. Stephanopodium angulatum (Little) Prance, comb nov Fig 18.

Tapura angulata Little, Phytologia 18: 412. 1969.

Tree ca 10.0 m tall, the young branches sparsely puberulous-glabrescent. Leaves

elliptic, coriaceous, 12.0-22.0 cm long, 3.8-7.8 cm broad, acuminate at apex, the acumen 7.0-18.0 mm long, cuneate at base, papillose on both surfaces, glabrous beneath; midrib prominent above, prominent and glabrous or with a few stiff appressed hairs beneath; primary veins 10-12 pairs, arcuate, anastomosing; petioles 7.0-12.0 mm long, shallowly canaliculate, puberulous-glabrescent, rugose. Stipules 1.0-1.5 cm long, lanceolate, glabrous, caducous. Flowers hermaphrodite, borne in dense-flowered sessile

glomerules inserted on petioles, pedicels 3.0-5.0 mm long; bracteoles ovate, 0.5-1.0 mm

long, persistent, sparsely puberulous on exterior. Calyx ca 5.0 mm long, very sparsely appressed puberulous on exterior, the lobes unequal. Corolla equalling calyx in length, with 5 equal lobes with rounded apices, and a broad cylindrical tube, the tube sparsely puberulous on exterior near apex, lanate within. Fertile stamens 5, alternating with corolla lobes, the anthers sessile, inserted at mouth of corolla tube. Ovary trilocular with 2 ovules in each loculus, tomentose on exterior. Style hirsute, with a trifid apex. Disc of 5 free large glands surrounding the ovary, the apices rounded. Fruit (immature) ellipsoid, ca 2.0 cm long; epicarp velutinous tomentose; mesocarp thin, fleshy; endocarp thin, bony, glabrous within.

TYPE. Little & Dixon 20451, Ecuador, Esmeraldas, Camino Pacto, NW of Quito, fl fr (holotype, US; isotype, NY).

DISTRIBUTION. Known only from the type, collected in lower montane forest of Ecuador. Collected in flower and fruit in February.

Stephanopodium angulatum was described in Tapura, but the sessile anthers and

equal corolla lobes show that it is a species of Stephanopodium. Its affinities are with S.

peruvianum another poorly known species. Since so few collections of S. peruvianum have been made, and since they are from geographically widely separated areas, the material is obviously inadequate for a detailed study. Only further material will show the exact relationship between S. peruvianum and S. angulatum. Judging from the present material, S. angulatum is distinct from S. peruvianum and differs in the large'r leaves with a greater number of primary veins, the larger flowers, the longer petioles, and the more

pubescent interior of the corolla tube.

6. Stephanopodium venezuelenum Prance, Acta. Bot. Venez. 2(2): 50. 1966. Fig 18.

Small tree or shrub, the young branches puberulous-glabrescent. Leaves oblong to

elliptic, membranceous, 5.5-10.5 cm long, 2.0-5.5 cm broad, acuminate at apex, the acumen 3.0-20.0 mm long, subcuneate and markedly unequal at base, glabrous or with a few stiff appressed hairs only beneath; midrib plane or slightly impressed above,

46 Flora Neotropica

t10

*.8

N . .

' .:" .; . .. .*

'

. . .

FIG 18. Distribution of species of Stephanopodium. o S. aptotum; * S. venezuelanum; * S. peruvianum; A S. angulatum.

,I~~ ~~Ss~T ??!1

lllrl[' [l ~ I ll ~ ~ I I [ ~l i I . ...~~~~~~~~?L O~~~ ~ --r I i. . i I ~

/j ~M.~ ~~~

~~~~~,;r~~~~~~~~~~~~~ 1~~~~~~ I~~~~!

~' .. ?~~~~

F~ ~IG 1 . isrbto ofseiso tpaooim . pou;?S eeulnr;?

peruvanur ; .agltm


prominent and with a sparse appressed pubescence beneath; primary veins 6-8 pairs, arcuate, anastomosing; petioles 4.0-10.0 mm long, canaliculate, puberulous when young. Stipules lanceolate, to 3.0 mm long, tomentellous, caducous. Flowers polygamous (male flowers only seen), borne in few-flowered almost sessile glomerules inserted at base or middle of petiole, sessile; bracteoles ovate, persistent, tomentose. Corolla with 5 small

equal lobes inserted on long tube, the tube sparsely pubescent on both surfaces. Fertile stamens 4-5, alternating with corolla lobes, the anthers sessile, inserted at mouth of corolla tube. Ovary not seen. Fruit not seen.

TYPE. Steyermark & Agostini 91198, Venezuela, Sucre, Peninsula de Paria, Cerro

Patao, fl (holotype, VEN; isotypes, K, NY, US). DISTRIBUTION. Endemic to the forests of the Paria Peninsula of Venezuela.

Collected in early flower in July and August. VENEZUELA. Sucre: Steyermark & Rabe 96107 fl (F, NY, US, VEN).

7. Stephanopodium sessile Rizzini, Revista Bras. Biol. 12(1): 103. 1952. Fig 19.

Small tree, the young branches glabrescent. Leaves elliptic to oblong, coriaceous, 4.0-12.0 cm long, 1.5-5.5 cm broad, acute or bluntly acuminate at apex, cuneate and

unequal at base, glabrous beneath; midrib slightly impressed to plane above, prominent and glabrous or with a few stiff appressed hairs beneath; primary veins 8-11 pairs, arcuate, anastomosing; petioles 4.0-7.0 mm long, canaliculate, sparsely puberulous- -glabrescent. Stipules lanceolate, to 4.0 mm long, persistent, sparsely puberulous. Flowers

hermaphrodite, borne in dense-flowered sessile glomerules inserted on petioles, sessile.

Calyx 4.0-5.0 mm long, sparsely tomentellous on exterior, the lobes slightly unequal. Corolla of 5 equal slightly bifid lobes inserted on a long tube, the tube glabrous on both

surfaces. Fertile stamens 5, alternating with corolla lobes, the anthers sessile, inserted at

mouth of corolla tube. Ovary bilocular with 2 ovules in each loculus, tomentellous on exterior. Styles 2, glabrescent. Fruit not seen.

TYPE. Pessoal do Horto Florestal R 20641, Guanabara, Rio de Janeiro, Horto

Florestal, fl fr (holotype, RB 139381; isotype, R); Rizzini RB 66163, Guanabara, Rio de

Janeiro, Horto Florestal, st (paratypes, NY, RB). DISTRIBUTION. Known only from forest around Rio de Janeiro. BRAZIL. Guana-

bara: Glaziou 18910 fl (C, K, MO, P).

One sheet of this species at New York is labelled Curran RB 10586 which would

make it an isotype of Tapura obovata (=S. blanchetianum). However, this sheet does not

match the Curran material of the same number at Rio de Janeiro, and the New York sheet is certainly mis-labelled. The New York sheet matches the type of S. sessile

perfectly, and is almost certainly a duplicate of it. Since it is fertile it is probably a

duplicate of Museu Nacional 20641, cited above.

Stephanopodium sessile is quite distinct from S. blanchetianum by its shorter

petioles, more acuminate thinner leaves, and its sessile flowers.

8. Stephanopodium blanchetianum Baillon, Mart. Fl. Bras. 12(1): 377. 1886. Fig 19.

Tapura obovata Rizzini, Revista Bras. Biol. 12(1): 107. 1952, synon nov, nom illegit. Type. Curran 67, Brazil, Bahia, Rio Grongogy basin, fl bud (syntype RB 10585; isosyntype, US); 83 fl bud (syntype, RB 10585).

Tree to 15.0 m tall, the young branches shortly tomentellous, becoming glabrous with age. Leaves oblong-elliptic, chartaceous, 6.5-11.5 cm long, 2.8-5.7 cm broad, rounded to acute or bluntly acuminate at apex, subcuneate and equal at base, glabrous beneath; midrib slightly impressed above, prominent and tomentellous beneath; primary veins 8-10 pairs, arcuate, anastomosing; petioles 9-14 mm long, canaliculate, tomentel-

48 Flora Neotropica

? " ': ' " '

'O 400

FIG 19. Distribution of species of Stephanopodium. o S. blanchetianum; + S. engleri; A S. organense; A S. estrellense; o S. sessile.


lous when young. Stipules ca 2.0 mm long, triangular, pubescent, caducous. Flowers

hermaphrodite, borne in dense-flowered sessile glomerules inserted on petioles, with distinct pedicels 0.5-1.5 mm long; bracteoles ca 0.5 mm long, persistent, tomentose.

Calyx 3.5-4.0 mm long, shortly tomentose on exterior, the lobes slightly unequal. Corolla with 5 equal lobes inserted on short broad tube, the tube tomentose on exterior

except at base, densely lanate within. Fertile stamens 5, alternating with corolla lobes, the anthers sessile, inserted at mouth of corolla tube, equalling corolla lobes in length. Ovary bilocular with 2 ovules in each loculus, pilose on exterior. Styles 2, sparsely pubescent to

apex or glabrous. Disc of 5 large separate glands. Fruit not seen. TYPE. Blanchet 2338, Brazil, Bahia, Ilheus, fl (lectotype, P; isotypes, BR, F, G, K,

LE); Blanchet 72 fl (paratype, P). DISTRIBUTION. Confined to the forests of the coastal region of Bahia, Brazil.

BRAZIL. Bahia: Belem & Pinheiro 2917 fl (UB); Bondar 2285 st (NY, SP);2459 fl (NY, SP); Riedel 735 fl (NY).

Rizzini described Tapura obovata based on material with young flower buds. This material is certainly Stephanopodium rather than Tapura as the 5 sessile anthers, and the

equal corolla lobes show. It matches S. blanchetianum, one of the most distinct species from the same region. Consequently T. obovata is reduced to synonymy.

Stephanopodium blanchetianum can be distinguished from the other related eastern Brazilian species by the blunt leaf apices, and the much longer tomentose petioles.

9. Stephanopodium organense (Rizzini) Prance, stat nov Fig 17 G-L, 19. S. estrellense var organense Rizzini, Revista Bras. Biol. 12(1): 102. 1952.

Small tree, the young branches sparsely tomentellous, becoming glabrous with age. Leaves oblong to oblong-lanceolate, membraneous, 3.5-8.5 cm long, 1.5-3.7 cm broad, the apex tapered to a long thin acumen 5.0-15.0 mm long, cuneate and equal or slightly unequal at base, with a few stiff appressed hairs especially near the base and midrib beneath; midrib plane above; prominent and with an appressed pubescence beneath; primary veins 8-10 pairs, arcuate, anastomosing; petioles 5.0-7.0 mm long, shallowly canaliculate, appressed pubescent. Stipules lanceolate, subpersistent, to 4.0 mm long, tomentellous. Flowers hermaphrodite, borne in small sessile glomerules inserted on

petiole; pedicels 1.0-2.0 mm long; bracteoles minute, ovate, tomentellous, persistent. Calyx 3.0-3.5 mm long, gray puberulous on exterior, the lobes slightly unequal. Corolla included, of 5 small equal, entire lobes inserted on a long tube, the tube glabrous on both surfaces except for a few hairs beneath anthers. Fertile stamens 5, alternating with corolla lobes, the anthers sessile, inserted at mouth of corolla tube, exceeding corolla lobes.

Ovary bilocular with 2 ovules in each loculus, tomentose on exterior. Style single, the

apex trifid, glabrous. Disc of 5 large glabrous glands, inserted at base of corolla tube. Fruit not seen.

TYPE. A. Barbosa 62, Brazil, Rio de Janeiro, Serra dos Orgaos, fl (holotype, RB;

isotype, NY). DISTRIBUTION. Known only from the forests near to Rio de Janeiro, Brazil.

Collected in flower in March. BRAZIL. Minas Gerais: Glaziou 19612 fl (C, K, P). Rio de Janeiro: Rizzini RB 132051 fl (RB).

This species is quite distinct from S. estrellense with which Rizzini associated it. It differs in the glabrous and smaller leaves, and in the flower structure. I have therefore elevated Rizzini's variety to specific rank.

3. Tapura Aublet, P1. Guiane 1: 126, t. 48. 1775; Jussieu, Gen. P1. 419. 1789; de Candolle, Prodr. 2: 58. 1825; Endlicher, Gen. P1. 1104. 1840; Bentham & Hooker,

50 Flora Neotropica

Gen. P1. 1: 341. 1862; Baillon, Mart. Fl. Bras. 12(1): 373-376. 1886; Engler in

Engler & PrantI, Nat. Pflanzenfam. 3(4): 351. 1896; Gleason, N. Am. Fl. 25: 382-383. 1924; Engler & Krause, Nat. Pflanzenfam 19c: 10. 1931; Lasser, Bol. Soc. Venez. Ci. Nat. 9: 127-128. 1944; Macbride, Publ. Field Mus. Bot. 13(3): 958-963. 1950; Stafleu in Pulle, Fl. Suriname 3(2): 169-172. 1951; Rizzini, Revista Bras. Biol. 12(1): 104-108. 1952; Leon & Alain, Fl. Cuba 3: 37-38. 1953; Hutchinson, Gen. Fl. P1. 1: 219. 1964; Ind. Nom. Gen. 10/03104.

Rohria Schreber, Gen. PI. 30. 1789, non Rohria Vahl. Gonypetalum Ule, Verh. Bot. Ver. Brandenb. 48: 174. 1906; Macbride, Publ. Field Mus. Bot.

13(3): 963-964. 1950.

Trees or shrubs. Leaves alternate, petiolate, entire, usually coriaceous. Stipules small, deciduous or persistent. Inflorescences of densely crowded glomerules adnate to and sessile or subsessile on the petioles or midrib (except for small axillary cymules in T.

singularis, and long pedunculate in some African species). Flowers small, hermaphrodite or polygamous, weakly zygomorphic. Bracts small and often scale-like. Sepals 5, imbricate, connate at base, usually unequal rarely equal. Petals 5, connate at base to form a long distinct tube, or free almost to base, the lobes imbricate, usually with 2 much

larger broad lobes which are bifid and bicucullate at apex, and 3 smaller linear-lanceolate entire lobes, rarely with 3 equal or nearly equal bicucullate lobes only. Stamens 5, all fertile or more frequently 3 fertile and 2 reduced to sterile staminodes, the filaments adnate to inside of corolla tube or to base of corolla in species without a distinct tube; anthers introrse. Disc semiannular or 2- 3-partite. Ovary free, globose, 2-3-locular with 2 ovules in each loculus. Style single, 2-3-lobed at apex or divided for much of length. Fruit a dry coriaceous drupe, 1- 3-locular with one seed in each loculus.

TYPE SPECIES. Tapura guianensis Aublet. The name Tapura is derived from the local name in French Guiana.

DISTRIBUTION. Greater and Lesser Antilles and South America from Pacific coastal Colombia through the Guianas and common in Amazonia. Also 5 species in Africa.

Species numbers 15-17 would fall into the section Dischizolaeana Baillon

(Adansonia 11: 112. 1873). However, I do not adopt any sections in Tapura since there are no clear cut divisions. Baillon created the section Dischizolaeana for the species with 5 fertile stamens. The other species which he examined had 3 fertile stamens and 2 sterile staminodes and were placed in his section, Eutapura. This study of all the species described to date shows that in addition to the two stamen types seen by Baillon, there are 2 species with 3 fertile stamens and no staminodes (T. coriacea & T. juliani), and one

species with 4 fertile stamens and no staminodes (T. colombiana). In light of this it is

obviously not possible to maintain the sections based on the staminal character which is not correlated with any other character.

Key to American species of Tapura 1. Inflorescences borne on midrib of leaf.

2. Flowers ca 2.0 mm long; midrib prominent above; inflorescence of many flowers; corolla lobes almost equal; staminodes absent. 1. T. juliani.

2. Flowers 8.0-12.0 mm long; midrib slightly impressed above; inflorescence of few flowers; corolla lobes markedly unequal; 2 staminodes present. 3. T. peruviana.

1. Inflorescences borne on petioles or rarely axillary. 3. Inflorescences with a distinct peduncle, axillary or petiolar.

4. Inflorescences axillary; primary veins 6-8 pairs. 4. T. singularis. 4. Inflorescences borne on petioles; primary veins 12-17 pairs. 5. T. juruana.

3. Inflorescences sessile on petioles or almost so. 5. Leaf undersurface densely hirsutulous. 6. T. amazonica. 5. Leaf undersurface glabrous or with a few stiff appressed hairs only.


6. Leaf apex retuse, rounded or acute or rarely bluntly acuminate; exterior of calyx glabrescent; plants of Cuba and Hispaniola. 7. Leaves 1.0-2.4 cm long, orbicular; midrib plane above; inflorescences 1-3

flowered. 7. T. orbicularis. 7. Leaves 3.0-12.0 cm long, rarely orbicular usually elliptic; midrib slightly

impressed above; inflorescences many flowered. 8. Leaves orbicular. 8. T. haitiensis. 8. Leaves elliptic to obovate. 9. T. cubensis.

6. Leaf apex acuminate, usually with a fine acumen rarely only bluntly acuminate; exterior of calyx pubescent; plants of South America or Lesser Antilles. 9. Upper surface of midrib impressed.

10. Flowers with distinct pedicels 1.5-2.5 mm long. 11. Fertile stamens 5. 15. T. lanceolata. 11. Fertile stamens 3.

12. Leaves bluntly acuminate flowers 5.0-6.0 mm long; corolla lobes 5; 2 staminodes present. 10. T. latifolia.

12. Leaves with long thin acumen; flowers ca 3.0 mm long; corolla lobes 3; staminodes absent. 2. T. coriacea.

10. Flowers sessile or almost so. 13. Leaves bullate above. 11. T. bullata. 13. Leaves plane not bullate above.

14. Corolla lobes free almost to base, only slightly connate. 15. Leaves thinly membraneous; petioles tomentose. 12. T. acreana. 15. Leaves coriaceous; petioles sparsely appressed pubescent.

13. T. tessmannii. 14. Corolla lobes united into a long distinct tube with the filaments

inserted on it. 16. Fertile stamens 3, staminodes 2.

17. Petioles and inflorescences densely tomentellous; leaves usually hirsute beneath. 6. T. amazonica.

17. Petioles glabrescent; inflorescences and flowers gray- puberulous; leaves glabrous or with a few appressed hairs beneath. 18. Petioles 3.0-5.0 mm long; inflorescences few-

flowered, inserted at junction of petiole with lamina. 3. T. peruviana.

18. Petioles 5.0-15.0 mm long; inflorescences many- flowered, inserted below junction of lamina with petiole. 14. T. guianensis.

16. Fertile stamens 5, staminodes absent. 19. Calyx lobes unequal; corolla lobes unequal, 2 much

larger and cucullate. 15. T. lanceolata. 19. Calyx lobes almost equal; corolla lobes equal.

16. T. capitulifera. 9. Upper surface of midrib prominent. 17. T. colombiana.

1. Tapura juliani Macbride, Publ. Field Mus. Bot. 13(3): 961. 1950. Fig 20 A-C, 25.

Small tree, the young branches glabrous. Leaves oblong to oblong-elliptic, membraneous, 8.5-12.0 cm long, 2.7-4.0 cm broad, caudate-acuminate at apex, the acumen 10.0-15.0 mm long, cuneate to subcuneate and slightly unequal at base, glabrous on both surfaces except for a few stiff appressed hairs beneath; midrib prominent above,

prominent and glabrous beneath; primary veins 5-6 pairs, arcuate, anastomosing; petioles 1.5-4.5 mm long, rugose, glabrous when mature, terete to shallowly canaliculate. Stipules deltoid, ca 1.5 mm long, glabrous, subcaducous. Flowers polygamous, sessile, borne in dense sessile glomerules on midrib of lamina; bracts ca 0.4 mm long, ovate, pubescent, persistent. Calyx 1.5-2.0 mm long, shortly pubescent on exterior, the lobes almost equal. Corolla of 3 equal cucullate lobes united at base into a short tube, the tube glabrous on

exterior, lanate within around base of lobes and filaments, the lobes and base of tube

glabrous, the lobes shortly exserted beyond the calyx. Stamens 3, alternating with corolla

52 Flora Neotropica

lobes, the filaments short and broad, inserted at mouth of corolla tube, no staminodes

present. Ovary not seen, vestigial in the d flowers examined. Fruit not seen. TYPE. Klug 2010, Peru, Loreto, mouth of Rio Zubineta, fl (holotype, F; isotypes,

A, K, NY, S, US). DISTRIBUTION. Known only from the type collection; flowering in March-April.

LOCAL NAME. Hibina.

This species is easily recognized by the flowers borne on the leaves. The flowers are much smaller than those of T. peruviana, the only other species of Tapura with flowers on the leaves.

Tapura juliani together with the next species, T. coriacea, forms a link between the

genera Tapura and Stephanopodium, and demonstrates their proximity. Tapura juliani has very short filaments, almost equal corolla lobes and unisexual flowers. I keep it in

Tapura because of the staminal filaments which, although, short, are quite distinct. All

species of Stephanopodium have entirely sessile anthers. Stephanopodium has 5 fertile stamens and T. juliani has only three which is common in Tapura. Tapura juliani remains in Tapura as a rather isolated species.

2. Tapura coriacea Macbride, Publ. Field Mus. Bot. 11: 68. 1931; Publ. Field Mus. Bot. 13(3): 960. 1950. Fig 20 D-F, 25.

Small to medium sized tree, the young branches sparsely puberulous-glabrescent. Leaves oblong to oblong-elliptic, coriaceous, 5.5-11.5 cm long, 1.8-4.2 cm broad, acuminate at apex, the acumen 5.0-10.0 mm long, cuneate to subcuneate and equal or

slightly unequal at base, glabrous beneath; midrib impressed above, prominent and

glabrous beneath; primary veins 6-8 pairs, arcuate, anastomosing; petioles 5.0-10.0 mm

long, sparsely puberulous becoming glabrous with age, shallowly canaliculate. Stipules triangular, 1.5-2.0 mm long, tomentose, caducous. Flowers androdioecious, borne in dense-flowered sessile glomerules inserted on upper portion of petioles; pedicels 1.5-2.0 mm long, tomentellous; bracteoles ca 0.5 mm long, persistent, pubescent. Calyx 3.0-4.0 mm long, gray-tomentellous on exterior, the lobes slightly unequal. Corolla exserted well

beyond calyx lobes, of 3 equal lobes two of which are markedly bicucullate, and the third

only slightly bifid, united at base into a long tube, the tube puberulous-glabrescent on

exterior, tomentose within. Fertile stamens 3, alternating with corolla lobes, the filaments short and broad, staminodes absent. Ovary reduced to a residual swelling in d flowers examined. Fruit not seen.

TYPE. Klug 602, Peru, Loreto, near Iquitos, fl (holotype, F; isotypes, G, NY, US). DISTRIBUTION. Known only from the vicinity of Mishuyacu, Loreto Department,

Peru, and from one collection from Huanuco. Collected in flower in October and November. PERU. Loreto: Klug 302 fl (F, NY, US); 586 fl (F, NY, US). Huanuco: Weberbauer 3693 fl bud (G).

This species is distinct from all other species of Tapura except T. juliani by its trimerous corolla and stamens. Like T. juliani, T. coriacea has distinct filaments and only 3 fertile stamens which places it in Tapura. Tapura coriacea differs from T. juliani in the inflorescence on the petioles, the pedicellate flowers, the leaf shape and size, and the

strap-like filaments.

3. Tapura peruviana Krause, Notizbl. Bot. Gart. Berlin 11: 135. 1931; Macbride, Publ. Field Mus. Bot. 13(3): 962. 1950. Fig 21 F-J, 25.

Tapura leucantha Krause, Notizbl. Bot. Gart. Berlin 11: 136. 1931; Macbride, Publ. Field Mus. Bot. 13(3): 962. 1950, synon nov. Type. Tessmann 4545, Peru, Loreto, fl (holotype B, lost;

photos, F, MO; isotypes, G, NY).


FIG 20. Species of Tapura. A-C, T. juliani (Klug 2010). A, habit X 0.5; B, flower X 8; C, opened flower X 8. D-F, T. coriacea (Klug 602). D, opened flower X 8; E, flower X 8; F, habit X 0.5.

54 Flora Neotropica

Small tree, the young branches puberulous, soon becoming glabrous. Leaves oblong, membraneous, 9.0-13.0 cm long, 2.8-5.0 cm broad, cuspidate-acuminate at apex, the acumen 10.0-18.0 mm long, rounded to subcuneate and unequal at base, glabrous beneath except for a few stiff appressed hairs; midrib impressed above, prominent and with a stiff appressed pubescence beneath; primary veins 7-16 pairs, arcuate, inconspicuous, anastomosing near margin; petioles 1.0-4.0 mm long, appressed pubescent, shallowly canaliculate to terete. Stipules ca 2.0 mm long, oblong, caducous. Flowers

hermaphrodite, borne in few-flowered sessile glomerules on lower portion of lamina midrib; pedicels ca 1.0 mm long; bracts ca 0.5 mm long, broadly ovate, pubescent, persistent. Calyx 5.5-7.0 mm long, shortly pubescent on exterior, the lobes unequal. Corolla far exceeding calyx lobes, with 2 large bicucullate lobes and three small narrow lobes, the large lobes free to near base, the other lobes joined in a long strap, the lobes

slightly fused at base only, densely pubescent at base of lobes and filaments within, with a few sparse hairs only on exterior. Fertile stamens 3, alternating with corolla lobes, inserted on base of corolla, 2 staminodes present. Ovary 3-locular with 2 ovules in each loculus, tomentose on exterior. Style with trifid apex, pubescent throughout. Fruit

ellipsoid, unilocular or bilocular; epicarp sparsely tomentellous; mesocarp thin; endocarp hard, bony, thin, glabrous within. This species is divided into two varieties.

Key to the Varieties of Tapura peruviana 1. Inflorescences borne on the midrib well above lamina base. a. var peruviana. 1. Inflorescences borne on upper portion of the petiole. b.var petioliflora.

3a. Tapura peruviana Krause var peruviana

Inflorescence borne on midrib well above lamina base. TYPE. Tessmann 4054, Peru, Loreto, near mouth of Rio Santiago, fl (holotype B,

lost; photos, F, MO; isotype, NY). DISTRIBUTION. Known only from the Maranron River region of Loreto, Peru.

Collected in flower September to November. PERU. Loreto: Mexia 6188 fl (BM, F, G, GH, K, MICH, MO, NY, S, U, UC, US).

There is no significant difference between the types of T. peruviana and T. leucantha, which were said to differ mainly in size by Krause.

3b. Tapura peruviana Krause var petioliflora Prance, var nov.

Flores in glomerulis ad petiolam adnati dispositi. TYPE. Mexia 6349, Peru, Loreto, Rio Marafion above Pongo de Manseriche, fl

(holotype, NY; isotypes, BM, F, G, GH, K, MICH, MO, S, U, UC, US). DISTRIBUTION. Forest in Amazonian Peru and Ecuador ECUADOR. Santiago-Zamora:

Taisha, Cazalet & Pennington 7636B fl (K). PERU. Ayacucho, Rio Apurimac Valley, Killip & Smith 22857 fr (F, NY, US); 22900 fr (F, NY, US).

This variety differs solely in the position of the inflorescence. Since this seems a constant character in most species it is recognized formally here.

4. Tapura singularis Ducke, Trop. Woods 90: 21. 1947; Rizzini, Revista Bras. Biol. 12(1): 106. 1952. Fig 21 A-E, 25.

Tree to 25.0 m tall, the young branches shortly tomentellous, becoming glabrous with age. Leaves oblong to oblong-lanceolate, coriaceous, 5.0-15.0 cm long, 1.4-4.5 cm broad, acuminate at apex, the acumen 2.0-12.0 mm long, subcuneate and equal to

slightly unequal at base, glabrous beneath except for a few stiff appressed hairs; midrib


\A. '<--^.-cc^ 5

A'- E

fA

G ;

FIG 21. Species of Tapura A-E, T. singularis (Ducke 1930 Pires et al 10972) A, habit X 0.5; B, flower X 3; C, fruit X 0.5; D, ovary section X 5; E, opened flower X 4. F-J, T. peruviana (Mexia 6188). F, opened flower X 4; G, ovary section X 5; H, flower X 3; J, habit X 0.5.

56 Flora Neotropica

slightly impressed above, prominent beneath and with a stiff appressed pubescence, becoming glabrous with age; primary veins 6-8 pairs, arcuate, anastomosing near margin; petioles 3.0-10.0 mm long, shortly tomentellous, becoming almost glabrous with age, canaliculate to terete, rugose. Stipules lanceolate, to 4.0 mm long, persistent. Flowers

hermaphrodite, borne in short axillary cymes, the peduncles 1.0-12.0 mm long, tomentellous; pedicels 1.5-3.0 mm long; bracteoles 0.3-1.5 mm long, triangular, persistent, pubescent. Calyx 4.0-5.0 mm long, shortly pubescent on exterior, the lobes

slightly unequal. Corolla far exceeding calyx lobes, with 2 large bicucullate lobes and 3 small simple lobes, all lobes united at base to form a long tube; the tube sparsely pubescent-glabrescent on exterior, filled with a lanate mass of hair within. Fertile stamens 3, alternating with corolla lobes, inserted on corolla tube at base of the lobes, 2 staminodes present. Ovary 3 locular with 2 ovules in each loculus, tomentose on exterior.

Style with a trifid apex, pubescent throughout. Fruit globose to ellipsoid, ca 2.0 cm long, with 1 or 2 loculi developing; epicarp with a dense compact velutinous pubescence; mesocarp thin; endocarp very thin, hard, bony, glabrous within.

TYPE. Ducke 1930, Brazil, Para, Belem, fl (holotype, MG; isotypes, A, F, GH, IAN, K, NY).

DISTRIBUTION. Confined to forests of eastern Amazonian Brazil on periodically flooded or non-flooded ground. Flowering April to July, fruiting May to January.BRAZIL. Para: Black 815 fr (IAN); DuckeMG 17032 fr (BM, G, MG, US); RB 20610 fl (RB, S, US); T. Guedes 201 fl (IAN, UB); Huber MG 6983 fr (MG); Pires 6854 fr (IAN); 7036 fr (IAN, SP); 10137 st (IAN); 51814 st (NY, US); Pires & Silva 10312 fr (IAN); 10474 fr (IAN); 10972 fr (IAN); 11172 fr (IAN); R.S. Rodrigues MG 3670 fr (MG); N.T. Silva 12 fr (COL, IAN, INPA, K, NY, UC, US); 19 fl (IAN). Terr. Amapa: Black 49-8335 fr (IAN, U).

This species differs from all species of Tapura in the axillary inflorescence. The flower structure shows that it belongs to Tapura. It is also the largest species of American

Dichapetalaceae, the trunk growing up to one meter in circumference.

5. Tapura juruana (Ule) Rizzini, Revista Bras. Biol. 12(1): 106. 1952. Fig 25.

Gonypetalum juruanum Ule, Verh. Bot. Ver. Brandenb. 38: 174. 1907; Macbride, Publ. Field Mus. Bot. 13(3): 964. 1950.

Tree to 20.0 m tall, the young branches shortly tomentellous, soon becoming glabrous. Leaves oblong, chartaceous, 6.0-16.0 cm long, 2.2-4.5 cm broad, acuminate at

apex, the acumen 8.0-15.0 mm long, subcuneate and slightly unequal at base, sparsely appressed hirsutulous beneath; midrib impressed above, prominent and appressed pubescent-glabrescent beneath; primary veins 12-17 pairs, arcuate, anastomosing; petioles 5.0-12.0 mm long, shortly tomentellous when young, canaliculate. Stipules lanceolate, ca 2.0 mm long, pubescent, subpersistent. Flowers hermaphrodite, sessile, borne in 2 short clustered cymules inserted on upper portion of petioles, the peduncles 1.0-4.0 mm long, tomentellous; bracteoles ca 0.5 mm long, ovate, persistent, pubescent. Calyx 3.0-4.0 mm

long, gray-puberulous on exterior, the lobes unequal. Corolla exceeding calyx lobes, with

2 large bicucullate lobes and 2 smaller simple lobes, the larger lobes free almost to base, the other lobes and filaments united into a small tube, the tube glabrescent on exterior, filled by a dense lanate mass of hair within. Fertile stamens 3, alternating with corolla lobes and inserted on short corolla tube, 2 staminodes present. Ovary 3-locular, tomentose on exterior. Style with a trifid apex, sparsely pubescent. Fruit ellipsoid, ca 1.5 cm long, unilocular; epicarp with a dense compact pubescence; endocarp very thin, hard,

bony, glabrous within. TYPE. Ule 5172, Brazil, Amazonas, Rio Jurua, fl (holotype B, lost; photos, F, MO,

NY; isotypes, G, HBG, K, L, MG, RB 20716). DISTRIBUTION. Periodically flooded forest and beside rivers in western Amazonia.


G H

t<0VJ~~~~~~t

FIG 22. Species of Tapura. A-D, T. cubensis (Jack 5668). A, habit X 0.5; B, flower X 4; C, ovary section X 15; D, opened flower X 5. E-K, T. amazonica (Irwin et al 5691, 9939). E, fruit X 0.5; F, opened flower X 5; G, ovary section X 15; H, flower X 2; J, habit X 0.5; K, leaf margin X 5.

58 Flora Neotropica

Flowering February to September. COLOMBIA. Amazonas: Black & Schultes 46-297 fl (IAN). BRAZIL. Amazonas: Ducke 1551 fr (A, F, MG, NY, RB, UC, US); RB 35409 fl (G, K, INPA, NY, P, RB, S, U, US); Froes 23828 fl (IAN, U); Krukoff 5126 fl (BM, F, G, K, LE, M, MICH, NY, RB, S, SP, U, UC, US); 6160 fl (A, BM, BR, F, G, K, MAD, MICH, MO, NY, S, U, US); 6214 fl (A, BM, BR, F, G, IAN, K, MAD, MICH, MO, NY, S, U, US); Traill 113 fr (K, P). Acre: Krukoff 5381 fl (A, BM, F, G, K, LE, M, MICH, MO, NY, RB, S, SP, U, UC, US); 5625 fl (A, BM, F, G, K, LE, M, MICH, MO, NY, S, SP, U, US). BOLIVIA. Pando: Prance et al 6609 fl (INPA, NY).

This species is easily recognized by the pedunculate inflorescence inserted on the petiole. Like the preceeding species it is a much larger tree than other members of the genus.

6. Tapura amazonica Poeppig & Endlicher, Nov. Gen. et Sp. 3: 41, t. 246. 1842; Baillon, Mart. Fl. Bras. 12(1): 374. 1886; Macbride, Publ. Field Mus. Bot. 13(3): 959. 1950; Stafleu in Pulle, Fl. Suriname 3(2): 172. 1951; Rizzini, Revista Bras. Biol. 12: 105. 1952. Fig 22 E-K, 26.

Tapura ciliata Gardner, Hook, Ic. II. 1, t. 466. 1842. Type. Gardner 3087, Brazil, Goias, fl (holotype, K; isotypes, BM, CGE, F, G, GH, NY, OXF, P, W).

Tapura amazonica var cuspidata Baillon, Mart. Fl. Bras. 12(1): 375. 1886. Tapura amazonica var dasyphylla Baillon, Mart. Fl. Bras. 12(1): 375. 1886. Tapura amazonica var sublanceolata Baillon, Mart. Fl. Bras. 12(1): 375. 1886. Tapura amazonica var ciliata (Gardner) Baillon, Mart. Fl. Bras. 12(1): 375. 1886.

Tree to 30.0 m tall, usually much smaller, the young branches fulvous-tomentose, becoming glabrous with age. Leaves elliptic to obovate-oblong or oblong, thickly coriaceous, 3.0-25.0 cm long, 3.0-9.0 cm broad, obtuse to shortly acuminate at apex, the acumen 0-10.0 mm long, rounded to cuneate and often slightly unequal at base, usually plane rarely slightly bullate above, sparse to dense-hirsutulous beneath; midrib

impressed above, prominent and pubescent when young beneath; primary veins 8-22

pairs, arcuate, anastomosing near margins; petioles 6.0-16.0 mm long, tomentose, canaliculate. Stipules triangular, 2.0-4.0 mm long, pubescent, subpersistent. Flowers

hermaphrodite, sessile or with short pedicels 0.25-2.0 mm long; borne in dense

glomerules on upper portion of petioles; bracteoles 0.5-1.0 mm long, persistent, tomentose. Calyx 3.5-5.5 mm long, tomentose on exterior, the lobes unequal. Corolla

exceeding calyx lobes, with 2 larger bicucullate lobes and 3 smaller simple lobes, united at base into a very short tube, the tube glabrous on exterior, filled by a lanate mass of hair within. Fertile stamens 3, alternating with corolla lobes, and inserted at mouth of short corolla tube, 2 staminodes present. Ovary 3 locular with 2 ovules in each loculus, pilose on exterior. Style with a trifid apex, pubescent throughout. Fruit oblong-ellipsoid, to 3.0 cm long, unilocular or bilocular; epicarp shortly appressed velutinous pubescent; mesocarp 1.0-4.0 mm thick; endocarp thin, hard, bony, glabrous within.

This species is divided into two varieties.

Key to the Varieties of Tapura amazonica 1. Small tree to 8.0 m tall; leaves 6.0-25.0 cm long, obtuse to acuminate at apex. 6a. var amazonica. 1. Large tree to 30.0 m tall; leaves 3.0-7.5 (-9.0) cm long, retuse to acute at apex.

6b. var manausensis.

6a. Tapura amazonica Poeppig & Endlicher var amazonica.

Small tree to 8.0 m tall; leaves 6.0-25.0 cm long, obtuse to acuminate at apex. TYPE. Poeppig 2673, Brazil, Amazonas, Tefe, fr (holotype, W; isotype, F). DISTRIBUTION. Forest on non-flooded ground in the Guianas, Amazonia, and the

Planalto of Central Brazil. Flowering and fruiting throughout the year. COLOMBIA. Caque- ta: Romero C. 4200 st (COL). Amazonas: Schultes & Black 8308 fr (COL, GH, K, U, US); Schultes & Cabrera 13565 fl (BM, COL, GH, U, US). VENEZUELA. Terr. Amazonas: Breteler 4763 fl (VEN).


SURINAME. Daniels & Jonker 1088 st (U); Lanjouw & Lindeman 2409 st (K, NY, U). FRENCH GUIANA. Melinon 171 fl (P); 332 st (F, P); 393 fl (P); s n fl (K, P, US); Sagot 1275 fl (BM, K, P, S, U, W); s n fl (P); Wachenheim 221 fl (P); 281 fl (P); 292 fl (P); 408 fl (P). PERU. Loreto: Klug 1003 fl (F, NY, US); 1529 fl (F, NY, US); Killip & Smith 27330 fr (NY, US); 29868 fr (F, NY, US); 29929 fl (F, NY); Ule 6258 fl (G, HBG, K, L, MG); LI. Williams 3651 fl (F). BRAZIL. Amazonas: Ducke 1864 fr (F, IAN, K, MG, NY, US): MG 7349 fl (MG);MG 7665 fl (BM, MG, RB, US); Froes 20656 fr (COL, F, IAN, K, NY, SP, US); 22650 fl (IAN, SP, US); 23721 (A, IAN); 23852 fr (IAN, U, UB); HuberMG 4659 fl (BM, G, MG, RB 13570, US); Krukoff4625 fl (A, F, G, K, M, MICH, MO, NY, S, U, UC, US); 5035A fl (A, NY); 5147 fl (A, G, K, NY, U); 8528 fr (A, BM, F, G, K, LE, MO, NY, S, US); Pires 1307 fr (IAN); Pires & Black 940 fr (IAN); Spruce 2808 fr (CGE, K, P, W). Para: Archer 8301 fl (IAN, K, NY, S, UC, US); Black 55-18129 fl (IAN, U, UB); Burchell 9453 fr (K); Ducke RB 20617 fl (K, RB, S, US); RB 20618 fl (RB); Froes 27746 fl (IAN, UC); Martius s n fl (M); Oliveira 4703 fl (IAN, NY); Pires & Black 193L fl (IAN); 288L fl (IAN); 1109 fl (GH, IAN, P); Pires & Silva 10970 fr (IAN); Prance, Pennington & Silva 1392 fr (NY); R.S. Rodrigues MG 3658 fr (MG, RB 13567); N.T. Silva 1293 fr (IAN, NY). Maranhao: Pires & Black 1735a fl (IAN). Terr. Rondania: Prance & Ramos 6866 fr (INPA, NY); J.F. Silva 27 fr (IAN). Mato Grosso: Irwin et al 17359 fl (F, NY, S, UB, US); J.G. Kublmann 2079 fr (RB 20711); Moore 411 fr (BM). Goias: Burchell 7131 fr (K, P); 8216 fl (BR, GH, K, P); Glaziou 20820 fl (A, BR, C, G, LE, P); Irwin et al 9939 fl (F, NY, S, UB, US); 21600 fr (NY); Macedo 3908 fr (IAN); Prance & Silva 58448 fr (F, K, NY, RB, S, U, UB, UC, US). Dist. Federal: Duarte 8348 fr (HB, RB 12348, UB); Heringer 8891/1085 fr (HB, RB); Irwin et al 5323 fr (MO, NY, RB, SP, UB, US); 5691 fr (GH, LE, M, NY, RB, SP, UB, UC, US); 8759 fl (F, NY, S, UB, UC, US); 9612 fl (F, MO, NY, S, UB, US); 15875 fr (F, NY, S, UB, US); 18154 fr (F, NY, S, UB, US);Maguire et al 57115 fl (NY); Pereira 9044 fr (HB, RB);Pires 57115 fl (UB);Pires, Silva, & Souza 9518 fr (RB, SP, UB). BOLIVIA. Pando: Prance et al 5788 fl bud (COL, INPA, NY); 6504 fl (COL, INPA, NY).

LOCAL NAMES. Suriname: Tassi-tihi. Brazil: Abiu, Mao de gato.

6b. Tapura amazonica var manausensis Prance, var nov

Arbor magna. Folia alternata, petiolata; laminae oblongae vel ellipticae, 3.0-7.5(-9.0) cm longae, apice retusae vel acutae.

TYPE. Ducke RB 23654, Brazil, Amazonas, Manaus, Igarape Mindu, fl (holotype, US: isotypes, G, INPA, K, RB, S, U).

DISTRIBUTION. Confined to forest on non-flooded ground in the region of

Manaus, Brazil. Flowering May to August. BRAZIL. Amazonas: Vicinity of Manaus, L. Coelho INPA 6385 fl (INPA, NY); Rodrigues et al 5994 fr (INPA 15043, NY); 6921 fr (INPA 15474, NY); 6929 fr (INPA 15482, NY); 7087 fr (INPA 15749, NY); Rio Urubu, Prance et al 5090 fl (INPA, NY).

This variety is recognized by the smaller leaves and the size of the tree. Since it is of restricted geographical range, and is so distinct in the field, I have treated it as a separate

variety. Tapura amazonica is the most widely distributed species of the genus. It is usually

easily recognized by the hirsutulous leaf undersurface (throughout its range with the

exception of Amazonian Peru), and by the tomentose inflorescence and petioles. Tapura amazonica is more variable in leaf shape than most other species of the genus. With the

exception of the one new variety described here, the variation is continuous, and is not

geographically correlated. Hence no other subspecific taxa are adopted here. Baillon

(1886), described the four varieties listed above, based on minor variations in leaf shape and size. With the abundant material now available, it is not possible to maintain any of

Baillon's varieties.

7. Tapura orbicularis Ekman ex Urban, Fedde Repert. Nov. Sp. 20: 305. 1924; Leon &

Alain, Fl. Cuba 3: 38. 1953. Fig 27.

Small tree, the young branches sparsely puberulous, soon becoming glabrous. Leaves orbicular to broadly obovate, thickly coriaceous, 1.3-2.8 cm long, 1.0-2.0 cm broad,

60 Flora Neotropica

?~~~~~~~~~~~~~~~

(l.. r?r 1 'V. ''

A'. A

~'t..t tI~~~.]

+:~~~~~~~~~~~~J

FIG 23. Distribution and leaf shape of Tapura guanensis. A, Rushy et al 287; B, De La Cruz 3304; C, Forest Dept. 2752; D, FlorscbUltz 699; E, Irwin et a! 55846; F, Cowan 39033; G, Irwin 48676; H, Irwin 47934;- J, Pires et a! 51516; K, Rodrigues et a! 3440; L, Krukoff 6979; M, Cavalcante 683. All leaves)X 0.25.

.... ~: . :'..:':!, ~ ' ~ ?? ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~''

i;~~~~~~~~~i

s. . .~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ r 2:~~~~~~~~~~~~~~~l ..,

: , 'i'"',-

?~~~~~~~~~~~~t

~~~~~~~~~~~~~~~~~~~~~~~ '

?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~.

~~~~~~~~~~~~~~~~~~~~~~?c...-.t

?~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ .../:.; -

'""~~~~~~~~~~~~~~~~~t ...f . :~

:' :. : ......./.. ~~~~~~~? ~,: , ,

:::': ?"~?":! ' ' : "~"

? ":'.:',." ", . . . .?.i":-,'' ,,i X,' ..

FIG 25. Distribution and leaf shape of Tapura guianensis. A, Rusby et al 287; B, De La Cruz 3304; C, Forest Dept. 2752; D, Florscbiitz 699; E, Ir~in et al 55846; F, Cowan 39033; G, Irwin 48676; H, Irwin 47934;? J, Pires et al $1516; K, Rodrigues et a1 3440; L, Krukoff 6979; M, Caualcante 683. All leaves X 0.25.


B

~ .. ........................'

..............................................

C e

G H

H ^^

FIG 24. Species of Tapura. A-E, T. capitulifera (B.W. 4525, 6878). A, habit X 0.5; B, young fruit X 0.5; C, flower X 4; D, opened flower X 5; E, ovary section X 20. F-J, T. acreana (Schunke 142). F, opened flower X 5; G, flower X 4; H, ovary section X 20; J, habit X 0.5.

62 Flora Neotropica

rounded at apex, rounded and equal at base, glabrous beneath; midrib plane above,

prominent and glabrous beneath; primary veins 3-5 pairs, arcuate, anastomosing; petioles 1.5-4.0 mm long, glabrous, rugose, terete. Stipules to 1.0 mm long, deltoid, subpersistent. Flowers hermaphrodite, with short pedicels ca 0.5 mm long, borne in pairs or solitary on the upper portion of petioles; bracts to 0.4 mm long, persistent, puberulous, Calyx 2.5-3.0 mm long, puberulous on exterior, the lobes slightly unequal, Corolla included, with 2 large bicucullate lobes, and 3 smaller simple lobes, the lobes shortly connate at base, but not forming a tube, glabrous on exterior, with a few sparsely distributed hairs only within. Fertile stamens 3, alternating with corolla lobes, the anthers borne on very short thick filaments, 2 staminodes present. Ovary trilocular with 2 ovules in each loculus, tomentose on exterior. Styles 3, glabrous. Fruit narrowly oblong, 11.0-20.0 mm

long, unilocular or bilocular; epicarp with a short velutinous pubescence; mesocarp thin; endocarp thin, hard, bony, glabrous within.

TYPE. Ekman 15303, Cuba, Oriente, Sierra Nipe, fl (holotype, S; isotypes, G, NY, S).

DISTRIBUTION. Confined to the arid scrublands of the Serpentine Barrens in Cuba. Collected in flower January to September.CUBA. Oriente: Alain et al 4857 st (GH); 4936 fl (GH); Ekman 9590 fl (S).

This species is quite distinct from all others by the very small orbicular leaves and the two-flowered inflorescence.

8. Tapura haitiensis Urban & Ekman, Ark. Bot. 20A(15): 44. 1926. Fig 27.

Medium sized tree, the young branches puberulous, soon becoming glabrous. Leaves

broadly obovate, 4.5-8.5 cm long, 3.2-6.5 cm broad, retuse to rounded at apex, rounded to subcuneate and unequal at base, glabrous beneath; midrib slightly impressed at least on lower portion above, prominent beneath and with a few stiff 'appressed hairs only; primary veins 6-8 pairs, arcuate, anastomosing; petioles 2.0-8.0 mm long, sparsely appressed pubescent when young, becoming glabrous, rugose, terete. Stipules triangular, ca 1.0 mm long, caducous. Flowers borne on upper portion of petiole. Inflorescence, flower and fruit not seen.

TYPE. Ekman 5241, Haiti, Massif de la Hotte, st (holotype, S). DISTRIBUTION. Apparently endemic to Massif de la Hotte in Haiti. HAITI. Ekman

7434 st (S, US).

This species was described from sterile material, and is probably distinct. The

existing material shows that it is closest to T. cubensis, but apparently differs in the much broader leaves. Further material is needed to decide whether T. haitiensis is distinct from T. cubensis.

9. Tapura cubensis (Poeppig & Endlicher) Grisebach, Cat. P1. Cub.: 56. 1866; Baillon, Mart. Fl. Bras. 12(1): 375. 1886; Gleason, N. Am. Fl. 25: 382. 1924; Leon & Alain, Fl. Cuba 3: 37. 1953. Fig 22 A-D, 27.

Chailletia cubensis Poeppig & Endlicher, Nov. Gen. et Sp. 3: 41, t. 246. 1842. Tapura cubensis var wrightiana Baillon, Mart. Fl. Bras. 12(1): 375. 1886. Type. Wright 1299 fr

(syntypes, BR, G, GH, GOET, K, LE, MO, P): 2220 fl (syntypes, BM, G, GH, GOET, K, LE, MO, NY, P, S, UC, US, W).

Tapura obovata Britton & Wilson, Bull. Torrey Club 53: 465. 1916; Gleason, N. Am. Fl. 25: 382. 1924; Le6n & Alain, Fl. Cuba 3: 38. 1953. Type. Britton & Wilson 15607, Cuba, Isle of Pines, fl (holotype, NY; isotypes, F, MO, P, S); 15524 fl (paratypes, F, GH, NY, US).

Small to medium sized tree or shrub, the young branches sparsely pubescent, soon

becoming glabrous. Leaves oblong, orbicular, elliptic or lanceolate, thickly coriaceous,


2.0-10.0 cm long, 1.0-5.8 cm broad, rounded to acute or rarely bluntly acuminate at

apex, rounded to subcuneate and equal at base, glabrous beneath; midrib slightly impressed above especially on lower portion, prominent and glabrous beneath or with a few stiff appressed hairs; primary veins 5-8 pairs, slightly arcuate, anastomosing; petioles 2.0-7.0 mm long, puberulous to glabrous, rugose, shallowly canaliculate. Stipules deltoid, to 1.0 mm long, caducous. Flowers hermaphrodite, sessile, borne in many-flowered sessile

glomerules on upper portion of petioles; bracteoles ca 0.5 mm long, ovate, pubescent, persistent. Calyx 2.0-4.5 mm long, puberulous on exterior, the lobes unequal. Corolla

exceeding calyx lobes, with 2 large bicucullate lobes and 3 smaller simple lobes, united at base to form a distinct tube, the tube glabrous on exterior, sparsely pubescent at base of lobes within. Fertile stamens 3, alternating with corolla lobes, 2 staminodes present. Ovary trilocular with 2 ovules in each loculus, pilose on exterior. Style with trifid apex, pubescent throughout. Fruit roundish, 1.0-1.5 cm diameter, unilocular to trilocular; epicarp with a short appressed pubescence; mesocarp thin; endocarp very thin, hard, bony, glabrous within.

This species is divided into two subspecies.

Key to the Subspecies of Tapura cubensis 1. Leaves 4.0-10.0 cm long, elliptic, orbicular to oblong; calyx ca 5.0 mm long; inflorescences

many-flowered; corolla tube short. 9a. subsp cubensis. 1. Leaves 2.6-5.0 cm long, elliptic to lanceolate; calyx ca 3.0 mm long; inflorescences

few-flowered (4-8 flowers); corolla tube long. 9b. subsp minor.

9a. Tapura cubensis subsp cubensis

Tree. Leaves 3.0-10.0 cm long, 1.7-5.8 cm broad, oblong, obovate, elliptic or orbicular. Calyx 3.0-4.5 mm long. Corolla tube short. Inflorescences many-flowered.

TYPE. Peoppig s n, Cuba, Las Piedras, fl (holotype, W). DISTRIBUTION. Growing on a wide range of habitats on acid soil. Collected from

Serpentine Barrens, dry woods, wet woodland, coastal thicket and pinelands in Cuba and Isle of Pines. Collected in flower between February and August.CUBA. Isla de Pinos: Ekman 12456 st (S); 12601 st (S). Pinar del Rio: Alain 2414 st (GH); 6104 st (GH, US); Britton & Cowell 9986 fl bud (NY); Ekman 18642 fl (S); Leon 12781 st (NY). Habana: Ekman 13648 fl (S); Leon 11563 st (NY). Las Villas: Jack 5668 fl (A, BM, F, G, K, LE, MAD, NY, P, S, US); 5743 st (A, NY, US); 8555 st (A, F, NY, S, US); Luna 659 fl (NY); Salvoza 723 st (A). Oriente: Acuna 12476 fr (US); Alain 3210 fl (GH); Clemente & Alain 3834 fr (GH, US); Clemente 4552 st (GH); Ekman 4199 st (G, NY, S, US); 4704 st (F, MICH, NY, S); 4844 st (S); 6169 st (G, K, S, US); 6678 st (F, S); 9002 st (G, NY, S); 9285 st (S, US); 9297 st (F, S, US); Le6n et al 17188 fl (GH); 18226 fl (GH); Morton & Alain 9087 st (US); Roig 119 st (NY); Sbafer 4209 fr (NY, US); 8287 st (GH, NY, US). Without precise locality: Gill & Whitford 135 st (MAD).

LOCAL NAMES. Vigueta naranjo, Cayada de aura, Palo de aura.

9b. Tapura cubensis subsp minor Prance, subsp nov

Frutex parva. Folia alternata, petiolata; laminae lanceolatae vel anguste oblongae, 2.0-6.0 cm longae, 0.8-2.8 cm latae. Calyx 2.0-3.0 mm longa. Corollae-tubus longus. Flores in glomerulis paucifloribus dispositi.

TYPE. Clemente 4367, Cuba, Oriente, Centeno, Moa, fl (holotype, GH; isotype, US).

DISTRIBUTION. Confined to pinelands and Serpentine Barrens in Oriente Province, Cuba. CUBA. Oriente: Acuna 12474 st (US); Alain 3245 st (GH); Alain & Morton 5163 fr (GH); Clemente 4375 fl (GH, US); Shafer 4028 fr (NY); 4159 st (K, NY); 8044 fr (K, NY); 8289 fr (GH, K, NY, U).

64 Flora Neotropica

F8 I 70 Ditibto60 5ospisfTpa+TbuaaATprva pu

2 Distritio T. rblaa; 0 T. eruvana var p FIG 25. Distribution of spe cies of T apura. + T. bullata; * T. corruvia na var peruviana; * T.

peruviana var petioliflora; & T. juliani; a T. acreana; * T. coriacea; o T. juruana; ? T. singularis.


Previous workers have divided the Cuban material of Tapura into three species, T. cubensis, T. obovata, and T. orbicularis. The latter species is quite distinct, but the other two do not differ significantly. There is a complete gradation between T. cubensis and T. obovata which was separated mainly by leaf shape. However, within T. cubensis there is a large variation in leaf shape and size. Some collections from Oriente Province are much more distinct than is T. obovata. This material is separated here into a subspecies. It may ultimately prove specifically distinct, but I hesitate to describe another new species from Cuba when most of the existing herbarium material is sterile.

The following sterile material is intermediate between T. cubensis subsp minor and T. orbicularis. CUBA. Oriente: Alain & L6pez 4878 st (GH). Isla de Pinos: Roig & Clemente 1818 st (NY). Pinar del Rio: Lors 4789 st (NY).

Baillon (1886) described a variety of T. cubensis, var wrightiana. This variety does not differ significantly from T. cubensis, and is considered here as part of subspecies cubensis.

10. Tapura latifolia Bentham, Jour. Bot. Kew Misc. 5: 291. 1853. Fig 27.

Tapura pedicellaris Chodat, Bull. Herb. Boiss. 4: 498. 1896, nom nud. Type. Hahn 1478, Martinique, fl (BM, BR, K, W).

Tapura antillana Gleason, N. Am. Fl. 25: 382. 1924. Type. Fishlock 13, Lesser Antilles, Dominica, fl (holotype, NY; isotypes, F, GH, K, MO, NY).

Tree to 18.0 m tall, the young branches glabrous. Leaves oblong, oblong-ovate, or

elliptic, thickly coriaceous, 6.5-15.0 cm long, 2.5-7.5 cm broad, acute or bluntly acuminate at apex, rounded to subcuneate and markedly unequal at base, glabrous or with a few stiff appressed hairs towards base beneath; midrib impressed at least on lower

portion above, sometimes plane on upper portion, prominent and glabrous or with a few

appressed hairs beneath; primary veins 7-12 pairs, arcuate, anastomosing; petioles 7.0-13.0 mm long, glabrous, rugose, terete or shallowly canaliculate. Stipules triangular, to 2.0 mm long, puberulous, caducous. Flowers hermaphrodite, borne in dense-flowered sessile glomerules inserted on upper portion of petioles; pedicels 2.5-4.5 mm long, pubescent; bracteoles ca 0.4 mm long, ovate, persistent, pubescent. Calyx 4.5-5.5 mm

long, gray-tomentellous on exterior, the lobes unequal. Corolla exserted beyond calyx lobes, with 2 large bicucullate lobes, and 3 smaller simple lobes, united at base to form a distinct tube, the tube sparsely puberulous to glabrous, filled by a lanate mass within. Fertile stamens 3, alternating with corolla lobes, inserted on corolla tube at base of lobes, 2 sterile staminodes present. Ovary 3 locular, with 2 ovules in each loculus,

tomentose-pilose on exterior. Style with a trifid apex, pubescent throughout. Fruit

ellipsoid, ca 2.0 cm long, ca 1.7 cm broad, usually unilocular; epicarp short-dense-

tomentose; mesocarp thin; endocarp thin, hard, bony, glabrous within. TYPE. Herb. Forsyth s n, without locality (certainly from Lesser Antilles) fl

(holotype, K). DISTRIBUTION. Forest of the Lesser Antilles from Guadeloupe to St. Lucia.

Flowering mostly from November to April, but collected in flower throughout the year. LESSER ANTILLES. Guadeloupe: Duss 973 fl (P); 3462 fl (F, GH, MO, NY, US); 3622 fl (F, NY, US); L'Herminier s n fl (G); Proctor 20137 fl (BM, US); 20174 fr (BM, US); Questel 5141 fr (US); s n fr (P); Richard s n fl (P); Stehle 7003 fl (US); 7012 fr (US). Dominica: Eggers 397b fl (HBG, M, P, W); 1037 fl (K); Fishlock D13 fl (K); H388-16 fl (K); Hodge 1322 st (GH); 2976 st (GH); 2980 fl (GH);

Ramage s n fl (BM, K). Martinique: Duss 183 fl (F, NY, S, US); Hahn s n fr (GH, K); 528 fl (G, K, L, M, P, U); 975 fl (BR, P); 1308 fl (BM, P); 1478 fl fr (G, P); Stehle 5362 fl (F, G, U, US). St. Lucia: Proctor 21622 fl (A, BM, US); Ramage s n st (BM, K).

LOCAL NAMES. Rosillet, Bois cote, Bois cotelette, Bois cote gebois.

66 Flora Neotropica

The name T. antillana has more generally been applied to this species because T. latifolia was described from a type specimen without locality. There is no doubt that this material from the Forsyth Herbarium comes from the Antilles and is the same as T. antillana, since it matches perfectly. T. latifolia was overlooked for many years because of the lack of locality, but it must take preference over T. antillana which is now reduced to synonymy.

Tapura latifolia is closely related to T. guianensis but is distinct by the long pedicels and the blunter leaf apices.

11. Tapura bullata Standley, Publ. Field Mus. Bot. 22: 86, 1940. Fig 25. Small tree, the young branches sparsely pubescent, becoming glabrous with age.

Leaves oblong-lanceolate, coriaceous, 15.0-23.0 cm long, 4.8-6.5 cm broad, acuminate at

apex, the acumen 10.0-25.0 mm long, subcuneate and markedly unequal at base, glabrous except for a few stiff appressed hairs beneath; midrib slightly impressed to plane above, prominent and sparsely appressed pubescent beneath; petioles 11.0-16.0 mm long, sparsely pubescent, becoming glabrous, shallowly canaliculate, rugose. Stipules lanceolate, to 7.0 mm long, persistent, tomentellous. Flowers hermaphrodite, sessile, borne in dense-flowered sessile glomerules on upper portion of petioles; bracteoles to 2.0 mm long, ovate to lanceolate, tomentellous, persistent. Calyx ca 5.0 mm long, tomentellous on exterior, the lobes unequal. Corolla exserted beyond calyx lobes, with 3 large lobes (the apex not cucullate) and 2 small almost filamentous lobes, united at base into a very short tube, the tube densely lanate within. Fertile stamens 3, alternating with corolla lobes, inserted on corolla tube at base of the lobes, 2 staminodes present. Ovary 3 locular with 2 ovules in each loculus, pilose on exterior. Style with a trifid apex, hirsute throughout. Fruit not seen.

TYPE. Haught 1510, Colombia, Sur de Santander, vicinity of Barranca Bermeja, fl

(holotype, F; isotypes, BM, COL, NY, US). DISTRIBUTION. Known only from the type collection.

This species is distinct by its bullate leaves, and the corolla lobes which are not hooded.

12. Tapura acreana (Ule) Rizzini, Revista Bras. Biol. 12: 107. 1952. Fig 24 F-J, 25.

Gonypetalum acreanum Ule, Notizbl. Bot. Gart. Berlin 6: 312. 1915; Macbride, Publ. Field. Mus. Bot. 13(3): 963. 1950.

Medium sized tree, the young branches tomentellous, becoming glabrous with age. Leaves oblong, membraneous, 6.5-10.0 cm long, 2.2-4.0 cm broad, acuminate at apex, the acumen 7.0-10.0 mm long, subcuneate and equal at base; with a few stiff appressed hairs especially near to midrib beneath; midrib impressed above, prominent and with an

appressed pubescence beneath; primary veins 9-10 pairs, arcuate, anastomosing; petioles 8.0-16.0 mm long, tomentellous, canaliculate. Stipules triangular, to 5.0 mm long, pubescent, caducous. Flowers hermaphrodite, sessile, borne in many-flowered sessile

glomerules on upper portion of petioles; bracteoles ca 0.5 mm long, persistent, tomentose. Calyx ca 3.0 mm long, gray-tomentose on exterior, the lobes unequal. Corolla exserted beyond calyx lobes, with 2 large cucullate lobes and 3 smaller simple lobes, very briefly connate at base, sparsely pubescent to glabrous on exterior, filled by a lanate mass within. Fertile stamens 3, alternating with corolla lobes, inserted on base of corolla lobes, the filaments long, lanate, 2 staminodes present. Ovary 3 locular with 2 ovules in each

loculus, sparesely pilose on exterior. Style with a deeply trifid apex. Fruit not seen. TYPE. Ule 9524, Brazil, Acre, Sao Francisco, fl (holotype B, lost; photos, F, MO,

NY; isotypes, G, K, L, MG, RB 20717, UC).


~f A ?7

10

70 60 ' 50

FIG 26. Distribution of Tapura amazonica. A var amazonica; * var manausensis.

68 Flora Neotropica

DISTRIBUTION. River banks in western Amazonia, known from only 2 collections. Collected in flower in January and October. PERU. Loreto: Schunke 142 fl (A, F, IAN, NY, S, UC, US); 14241 fl (US, field notes the same as for 142).

13. Tapura tessmannii (Krause) Prance, comb nov Fig 28. Gonypetalum tessmannii Krause, Notizbl. Bot. Gart. Berlin 11: 137. 1931; Macbride, Publ. Field

Mus. Bot. 13(3): 964. 1950.

Large tree, the young branches puberulous, becoming glabrous with age. Leaves

oblong-elliptic, coriaceous, 5.5-6.8 cm long, 2.0-3.5 cm broad, acuminate at apex, the acumen 3.0-7.0 mm long, subcuneate and equal at base; with a few sparse appressed hairs

especially near to midrib beneath; midrib impressed above; prominent and appressed pubescent beneath; primary veins 9-12 pairs, arcuate, anastomosing; petioles 8.0-15.0 mm long, sparsely tomentellous, shallowly canaliculate. Stipules triangular, to 3.0 mm

long, pubescent, caducous. Flowers hermaphrodite, sessile, borne in many-flowered sessile

glomerules on upper portion of petioles; bracteoles ca 0.5 mm long, persistent, pubescent. Calyx ca 3.0 mm long, gray-tomentose on exterior, the lobes unequal. Corolla exserted

beyond calyx lobes, with 2 large bicucullate lobes and 3 smaller simple lobes, the lobes united only at extreme base, glabrous on exterior, the lobes and staminal filaments with a dense lanate mass. Fertile stamens 3, alternating with corolla lobes, inserted on extreme base of corolla, filaments long, lanate, 2 staminodes present. Ovary 3 locular with 2 ovules in each loculus, tomentose on exterior. Style with trifid apex. Fruit not seen.

TYPE. Tessmann 4987, Peru, Loreto, Isidro, fl (holotype B, lost; photos, F, MO;

isotypes, G, NY, S). DISTRIBUTION. Known only from the type collection.

This species is close to T. acreana, but differs in the smaller coriaceous leaves with blunter apices.

14. Tapura guianensis Aublet, P1. Guiane 1: 126, t. 48, 1775; de Candolle, Prodr. 2: 58. 1825; Bentham, Jour. Bot. Kew Misc. 5: 291. 1853; Baillon, Mart. Fl. Bras. 12(1): 371. 1886; Lasser, Bol. Soc. Venez. Cienc. Nat. 9: 128. 1944; Macbride, Publ. Field Mus. Bot. 13(3): 961. 1950; Stafleu in Pulle, Fl. Suriname 3(2): 170. 1951; Rizzini, Revista Bras. Biol. 12(1): 107. 1952. Fig 23 A-M.

Rohria schreberi J.F. Gmelin, Syst. 2(1): 93. 1791. Rohria tapura J.F. Gmelin, Syst. 2(1): 93. 1791. Rohria petioliflora Willdenow, Sp. PI. 1(1): 186. 1797. Chailletia sessiliflora DC., Nouv. Bull. Soc. Philom. Paris 40: 205. 1811; Ann. Mus. Hist. Nat. Paris

17: 153. 1811. Type. Col incog s n, French Guiana, fl (G-DC). Tapura cucullata Bentham, Jour. Bot. Kew Misc. 5: 291. 1853. Type. Spruce 2226, Brazil,

Amazonas, fl (holotype, K; isotypes, BM, M, P, W). Tapura negrensis Suessenguth, Fedde Repert. Nov. Sp. 51: 199. 1942. Type. Luetzelburg 22231,

Brazil, Amazonas, Rio Negro, S. Felipe, fl (holotype, M); 22225 (paratype, M).

Tree to 9.0 m tall or shrub, the young branches glabrous or sparsely puberulous, soon becoming glabrous. Leaves most frequently oblong to ovate-elliptic rarely oblong-lanceolate or lanceolate, coriaceous, 6.0-23.0 cm long, 2.1-9.0 cm broad, acuminate at apex, the acumen 4.0-18.0 mm long, rounded to cuneate and unequal at

base, glabrous or with a few stiff appressed hairs beneath; midrib impressed above, prominent and glabrous or with a few stiff appressed hairs only beneath; primary veins 7-14 pairs, arcuate, anastomosing; petioles 5.0-14.0 mm long, sparsely puberulous to

appressed pubescent when young becoming less pubescent with age, rugose, terete to

shallowly canaliculate. Stipules lanceolate, to 2.0 mm long, caducous. Flowers hermaphrodite, sessile or on short pedicels, borne in dense sessile glomerules inserted on upper


portion of petioles; bracteoles 0.5-1.0 mm long, persistent, pubescent. Calyx 3.5-5.5 mm

long, tomentellous to sparsely puberulous on exterior, the lobes unequal. Corolla exserted

beyond calyx lobes, consisting of 2 larger bicucullate and 3 smaller simple lobes, united at base to form a long tube, the tube glabrescent on exterior, filled by a dense lanate mass within. Fertile stamens 3, alternating with corolla lobes, the filaments inserted on corolla tube at base of the lobes, the bases densely pubescent, 2 staminodes present. Ovary 3 locular with 2 ovules in each loculus, pilose-tomentose on exterior. Style with trifid apex, pubescent throughout. Fruit ellipsoid to narrowly oblong, most frequently unilocular, but often bilocular; epicarp with a short compact velutinous pubescence; mesocarp very thin; endocarp very thin, hard, bony, glabrous within.

TYPE. Aublet s n, French Guiana, fl (BM, P). DISTRIBUTION. Primary forest on flooded and non-flooded ground in the Guianas

and Amazonia. Flowering throughout the year in the Guianas, especially November to

April; most commonly flowering August to January in Amazonia. VENEZUELA. Terr. Delta Amacuro: Rusby & Squires 287 fl (A, BM, F, G, GH, K, LE, M, MICH, MO, NY, US, W); 291 fr (BM, F, G, GH, K, M, MICH, MO, NY, US). Bolivar: Agostini 337 fl bud (NY, US); Bernardi 1497 fl (NY); 2115 fl (NY); Blanco 45 fr (US); 96 fr (US); 613 fr (US); Koyama & Agostini 7395 fl (NY); Steyermark 86696 fl (US, VEN); Wurdack 34335 fl (NY). GUYANA. Abraham 249 st (NY); Anderson s n fr (BM); Davis 85 (F.D. 2076) fl (K); 595 (F.D. 2715) fl (K); De La Cruz 1917 fr (F, GH, MO, NY, US); 3304 fl (F, K, MO, NY, UC, US); Fanshawe 143 (F.D. 2752) fl (K, NY, U); 182 (F.D. 2918) fl (K, S); 585 (F.D. 3321) fr (K, NY); 1322 (F.D. 4068) fr (K, NY); 2744 (F.D. 5543) fl (K, NY); For. Dept. 935 fr (K); Hitchcock 17084 fr (NY, S, US); Hohenkerk 792 fr (K); Im Thurn s n fl (K, P); Jenman 32 fl fr (NY, P); 3955 fl (K); 4263 fl (K); 5270 fl (K); 5077 fl (K); Persaud 180 fr (F, K, NY); Sandwith 178 fr (K, RB, U); 677 fr (K, NY, RB); A.C. Smith 2120 fl (A, F, G, K, MO, NY, P, S, U, US); 2530 fl (A, F, G, K, MAD, MO, NY, P, S, U, US); 2926 fl fr (A, F, G, K, MAD, MO, NY, P, S, U, US); 3573 fl (A, F, G, IAN, K, LE, MAD, MO, NY, P, S, U, US, W); 3613 fr (A, F, G, IAN, K, LE, NY, P, U, US, W); Tutin 144 fl (BM, K, US). SURINAME. B.W. 699 st (K, NY, U); 1181 st (K, MO, NY, U); 1875 st (U); 2314 st (K, L, NY, U); 3218 st (NY, U); 3627 fr (BR, L, MO, NY, RB, U); 4148 fl (L, NY, RB, U, US); 4184 fl (BR); 4379 fr (C, K, MO, NY, U); 4486 fr (NY, U); 4844 fl (BR, C, GH, K, MO, NY, U); 5068 fr (K, NY, U, US); 6694 fl (BR, C, K, NY, U); Cowan 38986 fr (MICH, NY, S, U, UC); 39033 fl fr (GH, MO, NY, P, U, W); Elburg LBB 9883 fl (U); Florschutz 699 fl

(BR, C, NY, U);Heyligers 171 fl (U);406 st (U);Irwin, Prance, Soderstrom, & Holmgren 55524 fl bud

(NY, U, US); 55846 fr (F, MO, NY, S, SP, U, UB, US); Kappler s n fl (S); 135 fl (L); 2114 fl (GOET, L, LE, M, P, W); Lanjouw 370 fl (C, NY, U); 819 st (NY, U); 1284 fl (K, MO, NY, U); Lanjouw & Lindeman 2143 fr (NY, U); 2232 st (U); 2862 fr (IAN, K, NY, U); Lindeman 3756 fl (U); 4801 fl (U); 4542 st (U); 5935 st (U); 6143 fr (U); LBB 12062 fl (NY, U); LBB 12096 st (NY); LBB 12118 fl (U); Maguire & Stahel 23621 fr (K, NY, U, US); Maguire 40793 fl (NY, U); 40798 fl (NY, U); Pulle 95 st (U); Stahel 326 st (A, IAN, K, MAD, NY, U, WIS); Van Donselaar 1763 fr (U); 2665 fr (U); 2708 fl (U); 2820 fl (NY, U); 3069 fr (U); Versteeg 187 fl (U); 488 fl (U); Wessels Boer 1567 fl (U). FRENCH GUIANA. Benoist 130 fl (P); 235 st (P); 833 fl (P); 1579 fl (P); Black et al 54-1766 fl (IAN); 57-17523 fl (IAN); Cowan 38908 fl (F, NY, P, US); For. Serv. 242M fl (NY, P, U); Gabriel s n st (G);

Gandoger 146 fl (P); Geay s n fl (P); Irwin, Pires, & Westra 48431 fl (IAN, MG, NY, UB); Leblond s n fl (G); Le Moult 14 fl (P); Leprieur 256 fl (P); 278 fl (G);Martin s n fl (BM, K, MO, P 7); 104 fl (P 7); 289 fl (BM); Melinon 113 fl (P); 137 fl (P); 139 fl (P); 182 fl (P); 218 fl (P); 225 fl (P); 229 fl (P); 284 fl (P); 319 fl (P); 326 fl (P); 406 fl (P); 433 fl (P); s n (1862) fl (A, BM, BR, F, G, K, L, LE, NY, P, US); s n (1865) fl (F, P, US); Perrottet s n fl (G, P); Poiteau s n fl (P); Richard s n st (G, P);Sagot 192 fl (BM, BR, G, GOET, K, NY, P, U, W); Wachenheim 15 fl (BM, GH, K, NY, P, US); 210 fl (P); 356 fl (P); 438 fl (K, P); 498 fl (P); s n fl (A, BM, NY, P, US). PERU. Loreto: Killip & Smith 27089 fr (F, NY, US); 28245 fl (US); 29447 fl (F, NY, US); 29512 fl bud (F, NY, US); 29722 fl (F, NY, US); 29757 (NY, US). BRAZIL. Amazonas: Albuquerque 67-88 fl (INPA, NY); Black 48-2787 fl (IAN, P); Cavalcante 683 fr (MG); Chagas INPA 72 fl (INPA, NY, UB); L. Coelho & Chagas INPA 4294 fl (INPA, NY); Ducke 50 fl (A, F, K, MO, NY, S, US); 230 fl (MAD); 737 fl (F, IAN, MG, MO, US); 1409 fl (A, F, IAN, MG, NY, UC, US);MG 11152 fl (MG, RB 20613); RB 23655 fl (RB, S, US);Fr6es 21242 fl (IAN, NY, US); 22171 fl (IAN, U); 33664 fl (IAN); 33809 fl (IAN, U); 33823 fl (IAN); Killip & Smith 30147 fl (NY, S, US); Krukoff 6944 fl (A, BM, BR, F, G, K, LE, MICH, MO, NY, RB, S, U, US); 6979 fl (A, NY);Mello INPA 2846 fl (INPA, NY); Pires 72 fl (IAN); Prance et al 2197 fl (F, INPA, MG, NY, P, S, US); 2243 fl (INPA, MG, NY, S, US); 3044 fl (INPA, NY, S, US);J.S. Rodrigues

7Probable isotypes of Chailletia sessiliflora; Kew sheet labelled H.F. Talbot but probably one of the

pirated Martin collections.

70 Flora Neotropica

108 fr (IAN, U); W.A. Rodrigues et al 519 fl (INPA, NY); 910 fr (INPA, NY); 1377 fl (INPA, NY); 1385 fr (INPA, NY); 3166 fl (INPA, NY); 3420 fl (INPA, NY); 3440 fl (INPA, NY); 3467 fl (INPA, NY); 4712 fl (INPA, NY); 5060 fr (INPA, NY); 7377 fl (INPA, NY). Terr. Roraima: Black 51-13906 fl (IAN). Para: Dahlgren & Sella 89 fl (F, G);DuckeMG 8876 fl (MG);MG 9066 fl (MG, RB 20615); MG 9167 fl (MG, P, RB 13571, S, US);MG 10874 fl (MG);MG 11767 fl (MG, RB 20614); RB 20611 st (RB, S, US); RB 20612 fl (RB, US); Egler & Irwin 46567 fl (IAN, MG, NY, US); Martius 3175 fl (M); Pires 3884 fl (IAN, US). Terr. Amapa: Bastos 19 fl (IAN, NY, RB); 2019 fl (NY, RB); Black 49-8515 fl (IAN); Cowan 38658 fr (IAN, K, NY, P, RB); Cowan & Maguire 38087 fl (F, NY, RB, US); 38119 fl (IAN, K, NY); Frdes 25865 fr (IAN, U, UB); 25996 fr (IAN); 26008 fr (IAN); Irwin 48676 fl

(GH, IAN, K, MG, MO, U, UB, US); Irwin & Westra 47722 fl (IAN, MG, NY, U, US); Irwin, Pires, & Westra 47934 fl (IAN, K, MG, MICH, NY, SP, U, UC, US); 47975 fl (IAN, NY, SP); 48134 fl (IAN, MG, NY, U, UB, UC, US); Pires & Cavalcante 52681 fl (IAN, MG, NY, US); Pires, Rodrigues, & Irvine 51516 fl (NY). Guanabara: Glaziou 9922 fl (C, K, P), cultivated.

LOCAL NAMES. Venezuela: Areka-yek (Arekuna), Hierrito, Jabon. Guyana: Lukuchi-danni, Muribania, Waiaballi, Waidan. Suriname: Boesi Kofi, Pakiratiki, Wassakao.

Several authors have already commented on Aublet's interpretation of the flower of T. guianensis. His figure of the flower is quite inaccurate since it completely misinterprets the corolla and the stamens. I have examined the Aublet type to confirm the identity of this species, and later authors, such as Baillon (1886), were correct in their interpretation of T. guianensis.

Tapura guianensis is the commonest and most widespread species of Tapura, and

consequently it is also the most variable morphologically. The flower structure is quite uniform throughout the range except for minor variations in size. The leaf size and shape however, is quite variable. This variation was examined carefully and it was found to be continuous and not well correlated geographically. Consequently it is not recognized formally here. The variation in leaf shape is illustrated in Figure 23. This figure shows the

tendency to a large narrow leaf on the upper Rio Negro (23M), which was described as both T. negrensis and T. cucullata. These two names represent the same variation of the

upper Rio Negro which cannot be maintained as a separate taxon. There is also a

tendency to narrower and thinner leaves in Terr. Amapa and Para, Brazil. This variation is also too inconsistent to recognize taxonomically. The same individual can show several of the different leaf types, as is shown well by collections from the forest reserves of Suriname made over a number of years from the same numbered trees.

15. Tapura lanceolata (Ducke) Rizzini, Revista Bras. Biol. 12: 105. 1952. Fig 28.

Gonypetalum lanceolatum Ducke, Bull. Mus. Hist. Nat. Paris II. 4: 737. 1932; Arq. Jard. Bot. Rio de Janeiro 6: 43. 1933.

Small to medium sized tree, the young branches sparsely puberulous, soon

becoming glabrous. Leaves oblong-elliptic, chartaceous, 6.0-14.0 cm long, 2.0-5.5 cm

broad, acuminate at apex, the acumen 6.0-20.0 cm long, subcuneate and slightly unequal at base, glabrous except for a few stiff appressed hairs towards base beneath; midrib

impressed above, prominent and glabrous beneath; primary veins 9-13 pairs, arcuate,

anastomosing; petioles 5.0-12.0 mm long, sparsely appressed pubescent, rugose, canaliculate. Stipules triangular, ca 1.0 mm long, pubescent, caducous. Flowers

hermaphrodite, borne in many-flowered sessile glomerules inserted on upper portion of

petioles; pedicels 1.0-2.0 mm long, pubescent; bracteoles ca 0.4 mm long, pubescent, persistent. Calyx 2.5-3.0 mm long, gray-puberulous on exterior, the lobes unequal. Corolla exserted beyond calyx lobes, with 2 large bicucullate lobes and 3 smaller simple lobes, the lobes united at base to form a distinct tube, the tube sparsely pubescent- glabrescent on exterior, lanate within. Fertile stamens 5(-4), alternating with corolla

lobes, the filaments inserted at base of corolla lobes, lanate pubescent at base, staminodes absent. Ovary 3 locular with 2 ovules in each loculus, pilose-tomentose on exterior. Style with trifid apex, pubescent throughout. Fruit not seen.


25

.~ _---220

FIG 27. Distribution of species of Tapura. *T. cubensis subsp cubensis; T. cubensis subsp minor; + T orbicularis; o T. baitiensis;o ; . .atifolia.

72 Flora Neotropica

TYPE. Ducke RB 23657, Brazil, Amazonas, Manaus, fl (lectotype, RB; isolecto-

types, G, INPA, K, NY, P, S, U); Ducke RB 23658 fl (paratypes, G, K, P, RB, S, U, US). DISTRIBUTION. Known only from forest on non-flooded ground around Manaus,

Brazil. Flowering December to March. BRAZIL. Amazonas: Chagas INPA 3283 fl (INPA, NY); D. Coelho INPA 3279 fl (IAN, INPA, NY); INPA 3294 fl (INPA, NY); L. Coelho INPA 2910 fl (MG); L. Coelho & Mello INPA 3229 fl (IAN, INPA, NY); Ducke 325 fl (MAD, NY); 412 fl (A, F, K, MO, NY, S, US).

This species differs from T. capitulifera in the distinctly pedicellate flowers. Tapura lanceolata differs from the other pedicellate species of the genus in the 5 fertile stamens.

16. Tapura capitulifera Baillon, Adansonia 2: 112. 1873; Baillon, Mart. Fl. Bras. 12(1): 376. 1886; Lasser, Bol. Soc. Venez. Ci. Nat. 9: 127. 1944; Stafleu in Pulle, Fl. Suriname 3(2): 170. 1951; Rizzini, Revista Bras. Biol. 12(1): 105. 1952. Fig 24 A-E, 28.

Chailletia capitulifera Spruce ex Baillon, Compt. Rend. Assoc. Fr. Lyon. 1874. 479. 1874, nom nud.

Tree to 15.0 m tall, the young branches tomentellous, soon becoming glabrous. Leaves oblong to obovate-lanceolate, coriaceous, 5.0-12.0 cm long, 1.8-5.0 cm broad, acuminate at apex, the acumen 3.0-10.0 mm long, subcuneate and slightly unequal at base, glabrous or with a few stiff appressed hairs only beneath, papillose; midrib slightly impressed above, prominent and with a sparse appressed pubescence beneath; primary veins 9-15 pairs, arcuate, anastomosing; petioles 5.0-15.0 mm long, tomentellous when

young, becoming less so with age, canaliculate. Stipules triangular-lanceolate, ca 1.5 mm

long, pubescent, caducous. Flowers hermaphrodite, sessile or subsessile, borne in dense-flowered glomerules inserted on upper portion of petioles; bracteoles ovate, ca 0.5 mm long, persistent, pubescent. Calyx 2.0-3.0 mm long, gray-tomentellous on exterior, the lobes unequal. Corolla slightly exceeding the calyx lobes, with 2 large bicucullate lobes and 3 slightly smaller simple lobes, united at base to form a short tube, the tube

glabrous on exterior, filled by a dense lanate mass within. Fertile stamens 5, alternating with corolla lobes, the filaments inserted on corolla tube at base of lobes, lanate at base, staminodes absent. Ovary 3 locular with 2 ovules in each loculus, pilose on exterior. Style with trifid apex. Fruit ellipsoid, usually unilocular, 12.0-18.0 mm long; epicarp short-dense-velutinous-tomentose; mesocarp thin; endocarp thin, hard, bony, glabrous within.

TYPE. Spruce 3188, Venezuela, Terr. Amazonas, Rio Casiquiari, fl (holotype, P; isotypes, BM, BR, C, CGE, G, GH, GOET, K, LE, NY, OXF, P, RB, W).

DISTRIBUTION. Forests of Venezuela and the Guianas. Flowering December to March. VENEZUELA. Bolivar: Steyermark & Nilsson 475 st (NY, VEN). Terr. Delta Amacuro: Berti 341 fr (F, NY, VEN); 342 fr (F, NY, VEN); 380 fr (F, NY, VEN). GUYANA. J. Boyan 52 (F.D. 7736) fr (K, NY, US). SURINAME. B.W. 1683a st (K, L, RB, U); Stahel 133 fl (A, K, MAD, NY, U, WIS); Tree No. 1014, Brownsberg Reserve, B.W. 2402 st (K, NY, U), 2741 st (K, NY, U, US), 4525 fl (BR, GH, K, L, MO, NY, U), 4692 st (C, U), 6137 fr (K, MO, NY, U), 6763 st (C, K, NY, U), 6878 fr (NY, RB, U, US); Tree No. 616, Sectie 0 Reserve, B. W. 1838 st (C, K, MO, NY, U), 2588 fl (BR, C, K, MO, NY, U), 2625 fl (K, MO, NY, U, US), 4515 fl (BR, C, GH, K, L, MO, NY, RB, U, US), 5830 st (C, U), 6072 fl (BR, C, K, NY, U).

Tapura capitulifera is clearly distinct from T. guianensis by the floral structure, but sterile herbarium material is often impossible to distinguish. It is possible that some of the

fruiting and sterile material here placed in this species belongs to T. guianensis. There are some small, but not consistent, vegetative differences such as the papillose under surface of the leaf, and the tendency to smaller narrower leaves in T. capitulifera. However, in both species there is much variation in leaf shape and size. This is well demonstrated in various Suriname collections where a large number of gatherings have been made from the same numbered tree over several years. Flowering herbarium material is needed to


10

FIG 28. Distribution of species of Tapura. o T. c; . Ttessmannii; T. capitulifera; A

T. lanceolata.

74 Flora Neotropica

distinguish between T. guianensis and T. capitulifera. However, Dr. Jan Lindeman of Utrecht informs me that these two species are quite easy to distinguish in the field.

Tapura capitulifera has a distinct pale dull grey leaf undersurface while both sides of the leaves of T. guianensis are green. Also T. capitulifera usually develops into a much larger tree.

17. Tapura colombiana Cuatrecasas, Lloydia 11:220. 1949. Fig 28.

Tree 8.0 m tall, the young branches sparsely pubescent, becoming glabrous with age. Leaves obovate-oblong, coriaceous, 9.0-16.0 cm long, 3.0-6.5 cm broad, abruptly acuminate at apex, the acumen 5.0-10.0 mm long, cuneate and slightly unequal at base, glabrous beneath; midrib prominent above, glabrous and prominent beneath; primary veins 6-7 pairs, arcuate, anastomosing; petioles 7.0-10.0 mm long, sparsely pubescent, becoming glabrous with age, rugose, terete. Stipules caducous (not seen). Flowers

hermaphrodite, sessile, borne in dense sessile glomerules inserted on upper portion of

petioles; bracteoles ca 0.5 mm long, persistent, pubescent. Calyx 5.0-6.0 mm long, yellow-brown-velutinous-tomentellous on exterior, the lobes slightly unequal. Corolla exserted beyond calyx lobes, with 4 equal short bicucullate lobes united at base into a

long tube, the tube glabrous on exterior, densely lanate within. Fertile stamens 4, alternating with corolla lobes, the filaments inserted on corolla tube at base of the lobes, lanate pubescent, staminodes absent. Ovary 3 locular. Styles 2 or 3, if 2 then one deeply divided near to apex, hirsute throughout. Fruit not seen.

TYPE. Cuatrecasas 22470, Colombia, El Valle, between Tabor and Carrizales, fl

(holotype, F; isotypes, COL, P, US). DISTRIBUTION. Known only from the type collection from high woodland at

2100-2350 meters altitude in eastern Colombia.

This species approaches the genus Stephanopodium on account of its equal corolla. However, it belongs to Tapura because of the distinct staminal filaments, the

hermaphrodite flowers and the separate styles.

18. Tapura sp 1.

Some collections from Santander, Colombia are very close to T. guianensis, but

apparently differ in a pedunculate inflorescence. This has been observed in a single damaged inflorescence, and a second sheet in early bud where the peduncles are not yet developed. This material is probably a new species but cannot be adequately determined without further collection from the same region. The collections are: COLOMBIA. Santander: southeast of Barranca Bermeja, Romero C. 4790 fl bud (COL, US); 4824 st (COL); 4857 fl (COL, US); 4972 st (COL); between Rios Horta & Guayabito, Romero C. 4309 fr (COL).

ACKNOWLEDGMENTS

I am especially grateful to Mr. David Woolcott who prepared all the line drawings of the species, and to Mr. C. C. Clare who prepared the distribution maps.

I should like to thank the directors and curators of the numerous herbaria and the wood collections consulted for making their material available for study. All the herbaria concerned are cited in the systematic part of this work, and the wood collections are cited in the anatomical part. I am also grateful to the director and other staff members of The New York Botanical Garden for much help and advice, especially to Dr. Howard


Irwin, Jr. and Dr. Bassett Maguire. I am grateful to Mr. E. Forero for much help with

many aspects of this study while he was my graduate assistant at New York. This study was greatly assisted by grants from the National Science Foundation, GB

4641 and GB 7356 which enabled me to study the family in the field and also provided much of the assistance involved in the production of this monograph.

Throughout this study it has been extremely useful to have someone studying the African Dichapetalaceae with whom I could discuss the family. I am indebted to Dr. F. J. Breteler of Wageningen for the continual exchange of material and information which has taken place.

LITERATURE CITED

Aublet, M. F. 1775. Histoire des plantes de la Guiane francaise 1: 126; 3: t. 48. Baillon, H. 1874. Euphorbiaceae-Dichapetaleae. Histoires des Plantes 5: 139-142, 233-234.

1886. Dichapetaleae. In: Martius, Flora brasiliensis 12(1): 365-378. Barth, F. 1896. Anatomie comparee de la tige et de la feuille des Trigoniacees et des Chailletiacees

(Dichapetalees). Bull. Herb. Boiss. 4: 497-520. Bentham, G. 1853. Notes on two little-known genera connected with the South American flora. Jour.

Bot. Kew Misc. 5: 289-296. & J. D. Hooker. 1862. Chailletiaceae. Genera Plantarum 1(1): 340-342.

Bessey, C. E. 1907. A synopsis of plant phyla. University Studies 7: 336. Lincoln, Nebraska. Breteler, F. J. 1969. The African Dichapetalaceae 1. Acta Bot. Neerl. 18: 375. Brown, R. 1818. Chailletiaceae. In: Appendix 5 to Tuckey, Capt. J. K. Narrative of an expedition to

explore the River Zaire. 442. Cronquist, A. 1968. The evolution and classification of flowering plants. 254-257. de Candolle, A. P. 1811a. Description du Chailletia, nouveau genere de plantes. Nouv. Bull. Soc.

Philom. Paris 40: 205-206. 181 b. Description du Chailletia, nouveau genre de plantes. Ann. Mus. Paris 17: 153. 1825. Chailletiaceae. Prodromus Systematis Naturalis regni vegetabilis 2: 57-58.

de Candolle, C. 1890. Recherches sur les inflorescences Cpiphylles. Geneva. Descoings, B. 1957. Note sur un genre nouveau de Dichapetalacees. Natural. Malgache 9(2): 171, fig.

1. De Wildeman, E. 1919. Notes sur les especes africaines du genre "Dichapetalum" Thou. Rev. Zool.

Africaine 6, Suppl. Bot. 1-75. Du Petit Thouars, A. 1806. Genera nova madagascariensia. 23. Endlicher, I. L. 1840. Chailletiaceae. Genera plantarum secundum ordines naturales disposita 2: 1104;

Suppl. 3: 97. 1843. Engler, A. 1896. Dichapetalaceae. In: Engler & Prantl, Die natiirlichen Pflanzenfamilien 3(4):

345-351. & K. Krause. 1931. Dichapetalaceae. Die natiirlichen Pflanzenfamilien 19c: 1-11.

Erdtman, G. 1952. Dichapetalaceae. In: Pollen morphology and plant taxonomy. Angiosperms. 145-146.

Gadella, Th. W. J. 1969. Chromosome numbers of some Angiospermae collected in Cameroun and the

Ivory Coast. Proc. Koninkl. Ned. Akad. Van Wetensch. Amsterdam C. 72: 306-310. 1970. Chromosome numbers of some Angiosperms collected in Cameroun and the Ivory

Coast II. Acta. Bot. Neerl. 19: 431-435. Gleason, H. A. 1924. Dichapetalaceae. North America Flora 25: 281-283. Gundersen, A. 1950. Dichapetalaceae. In: Families of Dicotyledons. 151. Hallier, H. 1908. Uber Juliana eine Terebinthaceen Gattung mit Cupula and die wahren Stammelten

der Katzchen bliitler. Dresden. 1921. Beitrage zur Kentnis der Linaceen (DC. 1819) Dumort. Bot. Centralbl. Biehefte 21:

1-178. Hauman, L. 1955. Notes sur le genre Dichapetalum Thou en Afrique Centrale. Bull. Jard. Bot. Brux.

25: 339-351. Heimsch, C. 1942. The 'Gruinales' and 'Terebinthales' of Wettstein with reference to taxonomic

grouping. Lilloa 8: 83-198. Hutchinson, J. 1964. Dichapetalaceae. In: Genera of flowering plants 1: 216-219.

1969. Evolution and phylogeny of flowering plants. 63-66. Lasser, T. 1944. Las especies venezolanas de las Dichapetalaceae. Bol. Soc. Venez. Cienc. Nat. 9:

125-128.

76 Flora Neotropica

Leenhouts, P. W. 1956. Some notes on the genus Dichapetalum (Dichapetalaceae) in Asia, Australia and Melanesia. Reinwardtia 4: 75-87.

1957. Dichapetalaceae. Flora Malesiana I. 5: 305-316. Lewis, W. H. 1967. Dichapetalaceae. In: Woodson, R. E. Jr. & Schery, R. W. Flora of Panama. Ann.

Missouri Bot. Gard. 54: 9-12. Lobreau, D. 1969. Les limites de 1' "ordre" des Celastrales d'apres le pollen. Pollen et Spores 11:

499-555. Lundell, C. L. 1966. The Mexican and Central American species of Dichapetalum. Wrightia 3:

173-176. Macbride, F. 1950. Dichapetalaceae. In: Flora of Peru. Publ. Field Mus. Bot. 13(3): 954-964. Melchior, H. 1964. Dichapetalaceae. Engler's Syllabus der Pflanzenfamilien ed. 12. 2: 317. Metcalfe, C. R. & L. Chalk. 1950. The anatomy of the Dicotyledons 1: 358-361. Moss, M. B. 1928. The genus Dichapetalum in east, south tropical and subtropical Africa. Kew Bull.

1928:115-130. Poeppig, E. F. & S. Endlicher. 1842. Stephanopodium. Nova genera et species plantarum 3: 4, t. 246. Record, S. J. 1927. The wood of Tapura cubensis (Poepp. & Endl.) Grisebach. Trop. Woods 9: 18-19.

& R. W. Hess. 1943. Timbers of the New World. 140-141. Rizzini, C. T. 1952. Dichapetalaceae Brasiliensis. Revista Bras. Biol. 12(1): 97-108. Solereder, H. 1908. Systematic anatomy of the Dicotyledons 1: 198-200. Stafleu, F. A. 1951. Dichapetalaceae. In: Pulle, Fl. Suriname 3(2): 166-172. Steudel, E. G. 1840. Nomenclator botanicus ed. 2: 342. Takhtajan, A. 1969. Flowering plants, origin and dispersal. 221. Tippo, 0. 1938. Comparative anatomy of the Moraceae and their presumed allies. Bot. Gaz. 100:

1-97. Ule, E. 1906. Dichapetalaceae. In: II. Beitrage zur Flora der Hylaea. Verh. Bot. Ver. Prov. Brandenb.

48: 174. Vahl, M. H. 1810. Symphyllanthus. In: IV. Beskrivelse over nye Planteslaegter. Skrift. Nat. Hist. Selsk.

6: 86. Wettstein, R. 1935. Dichapetalaceae. In: Handbuch der Systematische Botanik. 682-683.

List of Taxa 77

NUMERICAL LIST OF TAXA

1. Dichapetalum 1. D. bullatum Standl. & Steyerm. 2. D. stipulatum Macbride 3. D. nervatum Cuatr. 4. D. latifolium Baill. 5. D. nevermannianum Standl. & Valerio 6. D. coelhoi Prance 7. D. rugosum (Vahl) Prance 8. D. pauper Rizzini 9. D. axillare Woodson

10. D. steyermarkii Prance 11. a. D. donnell-smithii Engl. var donnell-

smithii 11. b. D. donnell-smithii Engl. var chiapasense

(Standl.) Prance 12. D. spruceanum Baill. 13. D. froesii Prance 14. D. odoratum Baill. 15. D. pedunculatum (DC.) Baill.

2. Stephanopodium 1. S. engleri Baill. 2. S. aptotum Wheeler 3. S. estrellense Baill. 4. S. peruvianum Poepp. & Endl. 5. S. angulatum (Little) Prince 6. S. venezuelanum Prance 7. S. sessile Rizzini

8. S. blanchetianum Baill. 9. S. organense (Rizzini) Prance

3. Tapura 1. T. juliani Macbride 2. T. coriacea Macbride 3. a. T. peruviana Krause var peruviana 3. b. T. peruviana Krause var petioliflora

Prance 4. T. singularis Ducke 5. T. juruana (Ule) Rizzini 6. a. T. amazonica Poepp. & Endl.

var amazonica 6. b. T. amazonica Poepp. & Endl.

var manausensis Prance 7. T. orbicularis Ekman ex Urban 8. T. haitiensis Urban & Ekman 9. a. T. cubensis (Poepp. & Endl.) Griseb.

subsp cubensis 9. b. T. cubensis (Poepp. & Endl.) Griseb.

subsp minor Prance 10. T. latifolia Benth. 11. T. bullata Standl. 12. T. acreana (Ule) Rizzini 13. T. tessmannii (Krause) Prance 14. T. guianensis Aubl. 15. T. lanceolata (Ducke) Rizzini 16. T. capitulifera Baill. 17. T. colombiana Cuatr.

78 Flora Neotropica

LIST OF EXSICCATAE

Abraham, A. A. 249 (3-14)

Acufia, J. 12474 (3-9b); 12476 (3-9a)

Agostini, G. 337 (3-14)

Aguilar H., M. 243 (1-lla)

Aitkin, J. B. 40 (1-15)

Alain, Bro. et al 2414 (3-9a); 3210 (3-9a); 3245 (3-9b); 4857 (3-7); 4936 (3-7); 5163 (3-9b); 6104 (3-9a)

Allen, P. H. 2202 (1-9)

Anderson, A. s n (3-14)

Archer, W. A. 2397 (1-15); 2468 (1-15); 8301 (3-6a); 8310 (1-7)

Asplund, E. 10179 (1-12); 10311 (1-12); 13192 (3-14); 14097 (1-14)

Aublet, J. B. C. F. s n (3-14)

Barbosa, A. 62 (2-9)

Barreto, M. 10584 (2-1); 10585 (2-1); 10750 (2-1)

Bastos, M. 19 (3-14); 2019 (3-14)

Beckelt, J. E. et al 8534 (1-15)

Belem, R. P. et al 2917 (2-8)

Benoist, R. 130 (3-14); 235 (3-14); 833 (3-14); 1579 (3-14)

Bernardi, A. L. 1497 (3-14); 2115 (3-14)

Bernoulli, C. G. et al 2582 (1-lla)

Berti, L. M. 341 (3-16); 342 (3-16); 380 (3-16)

Birschel, J. W. sn (1-7)

Black, G. A. et al 46-297 (3-5); 47-815 (3-4); 47-2030 (1-15); 48-2297 (1-7); 48-2787 (3-14); 48-2884 (1-13); 48-2908 (1-4); 48-3591 (1-7); 49-8335 (3-4); 49-8518 (3-14); 50-9067 (1-7); 50-10252 (1-15); 51-13906 (3-14); 54-17666 (3-14); 55-18129 (3-6a); 57-17523 (3-14); 57-19022 (1-7); 57-19739 (1-7)

Blanchet, J. S. 72 (2-8); 2338 (2-8)

Blanco, C. 45 (3-14); 96 (3-14); 613 (3-14)

Bondar, G. 2285 (2-8); 2459 (2-8)

Boyan, J. 52 (3-16)

Breedlove, D. E. 11406 (1-1 la)

Brenes, A. M. 6766 (1-9)

Breteler, F. J. 4435 (1-7); 4588 (1-7); 4763 (3-6a)

Britton, N. L. et al 9986 (3-9a); 15524 (3-9a); 15607 (3-9a)

Broadway, W. E. 659 (1-15); 831 (1-15); s n (1-15)

Buchtein, 0. 1718 (1-4)

Burchell, W. J. 7131 (3-6a); 8216 (3-6a); 9453 (3-6a); 10107 (1-15)

B. W. (Boschwezen Suriname) 699 (3-14); 1181 (3-14); 1683a (3-16); 1838 (3-16); 1875 (3-14); 2314 (3-14); 2402 (3-16); 2588 (3-16); 2625 (3-16); 2741 (3-16); 3218 (3-14); 3219 (1-7); 3627 (3-14); 4148 (3-14); 4379 (3-14); 4486 (3-14); 4515 (3-16); 4525 (3-16); 4692 (3-16); 4844 (3-14); 5068 (3-16); 5830 (3-16); 6072 (3-16); 6137 (3-16); 6561 (1-7); 6694 (3-14); 6763 (3-16); 6878 (3-16)

Bryon (W. de Albuquerque) 67-88 (3-14); 68-119 (1-6)

Calder6n, S. 1512 (1-lla)

Cavalcante, P. B. 127 (1-7); 345 (1-7); 683 (3-14)

Cazalet, P. C. D. et al 7636B (3-3b)

Chagas, J. et al INPA 72 (3-14); INPA 3283 (3-15); INPA 3704 (1-7); INPA 3820 (1-7); INPA 6384 (1-7)

Clemente, Bro. et al 3834 (3-9a); 4367 (3-9b); 4375 (3-9b); 4553 (3-9a)

Coelho, D. INPA 3279 (3-15); INPA 3294 (3-15); INPA 4011 (1-7)

Coelho, L. et al INPA 2910 (3-15); INPA 3229 (3-15); INPA 4294 (3-14); INPA 6385 (3-6b)

Contreras, E. 4264 (1-llb); 4295 (1-llb); 5383 (1-llb); 6597 (1-llb)

Cooper, G. P. et al 172 (1-9)

Cowan, R. S. et al 38087 (3-14); 38119 (3-14); 38658 (3-14); 38908 (3-14); 38986 (3-14); 39033 (3-14)

List of Exsiccatae 79

Crevaux, A. s n (1-15)

Cuatrecasas, J. 8975 (1-13); 15833 (1-3); 16746 (2-4); 22470 (3-17)

Curran, H. M. 67 (2-8); 83 (2-8)

Dahlgren, B. E. et al 89 (3-14); 375 (1-7); 504 (1-7)

Damazio, L. 13386 (2-1)

Daniels, A. G. H. et al 1088 (3-6a)

Davis, T. A. W. 85 (3-14); 595 (3-14)

De La Cruz, J. S. 1108 (1-15); 1193 (1-15); 1596 (1-15); 1895 (1-15); 1917 (3-14); 2625 (1-15); 3095 (1-15); 3304 (3-14); 3543 (1-15); 3688 (1-15); 3826 (1-15); 3905 (1-15); 4041 (1-15); 4337 (1-15); 4517 (1-15)

Duarte, A. P. 8348 (3-6a)

Ducke, A. 50 (3-14); 131 (1-7); 230 (3-14); 325 (3-15); 412 (3-15); 503 (1-7); 737 (3-14); 881 (1-4); 1112 (1-7); 1409 (3-14); 1479 (1-7); 1551 (3-5); 1864 (3-6a); 1930 (3-4); MG 627 (1-7); MG 3285 (1-15); MG 3457 (1-7); MG 7349 (3-6a); MG 7656 (1-7); MG 7665 (3-6a); MG 8876 (3-14); MG 9066 (3-14); MG 9167 (3-14); MG 10874 (3-14); MG 10975 (1-7); MG 11152 (3-14); MG 11600 (1-7); MG 11767 (3-14); MG 12124 (1-7); MG 15945 (1-8); MG 15991 (1-15); MG 17032 (3-4); RB 13567 (1-15); RB 20610 (3-4); RB 20611 (3-14); RB 20612 (3-14); RB 206717 (3-6a); RB 20618 (3-6a); RB 20713 (1-15); RB 20715 (1-15); RB 23654 (3-6b); RB 23655 (3-14); RB 23656 (1-15); RB 23657 (3-15); RB 23658 (3-15); RB 35409 (3-5)

Duss, A. 183 (3-10); 973 (3-10); 3462 (3-10); 3622 (3-10)

Edwards, J. B. 444 (1- lb)

Eggers, H. von 397b (3-10); 1037 (3-10)

Egler, W. A. et al 46567 (3-14)

Ekman, E. L. 4199 (3-9a); 4704 (3-9a); 4844 (3-9a); 5241 (3-8); 6169 (3-9a); 6678 (3-9a); 7434 (3-8); 9002 (3-9a); 9285 (3-9a); 9297 (3-9a); 9590 (3-7); 12456 (3-9a); 12601 (3-9a); 13648 (3-9a); 15303 (3-7); 18642 (3-9a)

Elburg, J. LBB 9883 (3-14)

Espina, R. et al A23 (2-2); A59 (2-2); A124 (2-2); A145 (2-2)

Fanshawe, D. B. 143 (3-14); 182 (3-14); 205 (1-15); 230 (1-15); 585 (3-14); 1079 (1-15); 1322 (3-14); 1565 (3-14); 2744 (3-14); 2752 (3-14)

Ferreira, A. R. s n (1-15)

Fishlock, W. C. 13 (3-10)

Florschutz, P. A. 699 (3-14)

Forest Dept. Br. Guiana (Guyana) 935 (3-14)

Forest Serv. French Guiana 242M (3-14)

Fr6es, R. L. et al 20656 (3-6a); 21242 (3-14); 21604 (1-7); 22171 (3-14); 22325 (1-13); 22493 (1-8); 22650 (3-6a); 22850 (1-13); 23721 (3-6a); 23734 (1-4); 23828 (3-5); 23852 (3-6a); 25865 (3-14); 25996 (3-14); 26008 (3-14); 27383 (1-7); 27746 (3-6a); 28317 (1-8); 30175 (1-7); 30442 (1-7); 30891 (1-15); 33664 (3-14); 33809 (3-14); 33823 (3-14)

Gabriel sn (3-14)

Gandoger, A. M. 146 (3-14)

Garcia-Barriga, H. et al 13846 (2-4); 18677 (1-13)

Gardner, G. 3087 (3-6a)

Geay, F. sn(3-14)

Gentle, P. M. 46 (1-llb); 61 (1-llb); 501 (1-11); 1305 (1-llb); 1325 (1-llb); 1332 (1-llb); 2623 (1-llb); 2625 (1-llb); 2644 (1-11); 2864 (1-llb); 2880 (1-11); 4219 (1-llb); 4803 (1-llb); 5593 (1-llb); 6024 (1-lla); 6572 (1-lla); 6910 (1-lla); 7206 (1-llb); 7999 (1-llb); 9194 (1-llb)

Gill, et al 135 (3-9a)

Glaziou, A. F. M. 9922 (3-14); 10080 (1-7); 13795 (2-3); 18910 (2-7); 19612 (2-9); 20820 (306a)

Gleason, H. A. 205 (1-15)

Gonggrijp, J. W. et al BW 6531 (1-15)

Guedes, T. 90 (1-15); 126 (1-7); 131 (1-7); 201 (3-4); 337 (1-7)

Hahn, L. 528 (3-10); 975 (3-10); 1308 (3-10); 1478 (3-10); s n (3-10)

Haught, 0. 1510(3-11)

Herb. Dept. Landbouw (Suriname) 286 (1-15)

Herb. Forsyth s n (3-10)

80 Flora Neotropica

Herb. Schwacke 13386 (2-1)

Herb. Vahl s n (1-7)

Heringer, E. P. 8891/1085 (3-6a)

Herminier, F. L. L' s n (3-10)

Heyde, E. T. et al 4431 (1- la); 60650 (1-lla)

Heyligers, P. C. et al 171 (3-14); 406 (3-14); 470 (1-15)

Hitchcock, A. S. 17084 (3-14)

Hodge, W. H. et al 1322 (3-10); 2976 (3-10); 2980 (3-10)

Hohenkerk, L. S. 792 (3-14)

Horto Florestal, Rio de Janeiro R 20641 (2-7)

Huber, J. E. MG 4011 (1-7); MG 4659 (3-6a); MG 6983 (3-4)

Im Thurn, E. F. s n (3-14)

Irwin, H. S. et al 5323 (3-6a); 5691 (3-6a); 8759 (3-6a); 9612 (3-6a); 9939 (3-6a); 15875 (3-6a); 17359 (3-6a); 18154 (3-6a); 21600 (3-6a); 47722 (3-14); 47934 (3-14); 47975 (3-14); 48134 (3-14); 48431 (3-14); 48676 (3-14); 55524 (3-14); 55846 (3-14)

Jack, J. G. 5668 (3-9a); 5743 (3-9a); 8555 (3-9a)

Jenman, G. S. 32 (3-14); 1718 (1-15); 2355 (1-15); 3955 (3-14); 4263 (3-14); 4326 (1-15); 5077 (3-14); 5270 (3-14)

Kappler, A. 135 (3-14); 2114 (3-14); s n (3-14)

Karsten sn (1-15)

Kernan, H. S. 102 (2-2)

Killip, E. P. et al 22857 (3-3b); 22900 (3-3b); 27089 (3-14); 27330 (3-6a); 28128 (1-2); 28245 (3-14); 29447 (3-14); 29512 (3-14); 29722 (3-14); 29757 (3-14); 29834 (1-14); 29868 (3-6a); 29929 (3-6a); 30147 (3-14); 30538 (1-15); 34375 (see 1-12)

Klug, G. 76 (1-2); 122 (1-14); 182 (1-14); 226 (1-14); 302 (3-2); 496 (1-4); 551 (1-14); 586 (3-2); 602 (3-2); 858 (1-7); 1003 (3-6a); 1081 (1-2); 1431 (1-4); 1529 (3- 6a); 1985 (2-4); 2010 (3-1); 2626 (1-7); 2751 (1-12); 3070 (1-14); 4306 (1-12)

Koyama, T. et al 7395 (3-14)

Krukoff, B. A. 4625 (3-6a); 5035a (3-6a); 5126 (3-5); 5147 (3-6a); 5381 (3-5); 5625 (3-5); 6160 (3-5); 6214 (3-5); 6944 (3-14); 6979 (3-14); 7072 (1-15); 8077 (1-14); 8351 (1-14); 8368 (1-14); 8519 (1-7); 8528 (3-6a)

Kuhlmann, J. G. 1885 (1-7); 2079 (3-6a); 2092 (1-7); RB 20714 (1-14); RB 73047 (2-3)

Kuhlmann, M. et al 365 (1-7)

Lanjouw, J. et al 370 (3-14); 819 (3-14); 1275 (1-15); 1284 (3-14); 2143 (3-14); 2232 (3-14); 2409 (3-6a); 2862 (3-14)

Lasser, T. 1927 (1-15)

Lawrance, A. E. 414 (1-12)

Leblond, J. B. 347 (1-15); 454 (1-15); sn (1-15); sn (3-14)

Lemee, A. s n (1-15)

Le Moult, E. 14 (3-14)

Leon, Bro. H. 11563 (3-9a); 12781 (3-9a); 17188 (3-9a); 18226 (3-9a)

Leprieur, F. R. 256 (3-14); 278 (3-14); s n (1-15); s n (1838) (1-15); sn (1840) (1-15); sn (1849) (1-15); s n (1850) (1-15)

Lindeman, J. C. 3756 (3-14); 4542 (3-14); 4801 (3-14); 4652 (1-15); 5394 (1-7); 6143 (3-14); LBB 12062 (3-14); LBB 12096 (3-14); LBB 12118 (3-14)

Little, E. L. et al 20451 (2-5)

Luetzelburg, P. von 22225 (3-14); 22231 (3-14)

Luna, A. 659 (3-9a)

Lundell, C. L. 4845 (1-llb); 6163 (1-11)

Macedo, A. 3908 (3-6a)

Magalhaes, M. 140 (2-1); sn (2-1); sn (2-1)

Maguire, B. et al 23621 (3-14); 24658 (1-7); 24669 (1-15); 39003 (1-15); 40793 (3-14); 40798 (3-14); 46878 (1-15); 57115 (3-6)

Martin, J. 92 (1-7); sn (1-7); sn (1-15); sn (1842) (3-14); s n (1845) (3-14)

Martinez-Calderon, G. 17 (1-lla)

Martius, K. F. P. von 2794 (1-7); 3175 (3-14); 3290 (3-6a); sn

List of Exsiccatae 81

(1-4); s n (1-7); s n (1-15); s n (3-6a) Matuda, E.

4 (1-lla); 630 (1-lla); 645 (1-11); 679 (1-llb); 4006 (1- lb); 4208 (1- lb); 5420 (1-lla); 6042 (1-lla); 15420 (1-l1a); 16535 (1-lla); 16565 (1-lla); 17617 (1-lla); 17942 (1-lla); 17980 (1-lla)

Melinon, M. 113 (3-14); 137 (3-14); 139 (3-14); 171 (3-6a); 182 (3-14); 218 (3-14); 225 (3-14); 229 (3-14); 233 (1-7); 284 (3-14); 319 (3-14); 326 (3-14); 332 (3-6a); 393 (3-6a); 406 (3-14); 433 (3-14); sn (1-15); sn (3-6a)

Mello, F. INPA 2846 (3-14)

Mexia, Y. 6188 (3-3a); 6349 (3-3b); 7285 (1-12)

Miranda, F. 7881 (1-llb)

Molina R., A. 12204 (1-lla)

Monnier, L. s n (3-14)

Monteiro da Costa, R. C. 276 (1-7)

Moore, S. 411 (3-6a)

Morton, C. V. et al 9087 (3-9a)

Museu Goeldi 9686 (1-7)

Mutis, J. C. 1140 (1-12); 2957 (1-12); 2959 (1-12); 2960 (1-12); 4035 (1-12); 5095 (1-12); 5807 (1-12)

Oliveira, E. 564 (1-7); 3273 (1-7); 4156 (1-7); 4294 (1-7); 4295 (1-7); 4703 (3-6a)

Parker, C. S. s n (1-15)

Patris, J. B. s n (1-15)

Pereira, E. 9044 (3-6a)

Perrotet, G. S. s n (1-15); sn (3-14)

Persaud, A. C. 180 (3-14); 238 (1-15)

Pires, J. M. et al 72 (3-14); 193L (3-6a); 288L (3-6a); 670 (1-7); 940 (3-6a); 1071 (1-4); 1109 (3-6a); 1157 (1-7); 1287 (1-15); 1300 (1-7); 1307 (3-6a); 1652 (1-15); 1735a (3-6a); 3106 (1-15); 3884 (3-14); 6854 (3-4); 7036 (3-4); 8241 (1-7); 9518 (3-6a); 10137 (3-4); 10302 (3-4); 10375 (1-7); 10474 (3-4); 10970 (3-6a); 10972 (3-4); 11172 (3-4); 11653 (1-15); 51516 (3-14); 51814 (3-4); 52681 (3-14); 57115 (3-6a)

Pittier, H.

1977 (1-lla); 4349 (1-9) Poeppig, E.

1986 (2-4); 2673 (3-6a); 2903 (1-4); s n (3-9a)

Poiteau, P. A. s n (1-15); sn (3-14)

Popenoe, H. 45 (1-lla)

Prance, G. T. et al 1392 (3-6a); 2197 (3-14); 2243 (3-14); 3044 (3-14); 5090 (3-6b); 5788 (3-6a); 6504 (3-6a); 6866 (3-6a); 6609 (3-5); 8116 (1-15); 9849 (1-15); 58448 (3-6a)

Proctor, G. R. 20137 (3-10); 20174 (3-10); 21622 (3-10)

Pulle, A. 95 (3-14); 305 (1-15); 454 (1-7)

Purdie, W. s n (1-15)

Questel, A. 5141 (3-10); s n (3-10)

Ramage, G. A. s n (3-10); s n (3-10)

Richard, L. C. s n (1-7); s n (1-15); s n (3-10); s n (3-14)

Riedel, L. 735 (2-8)

Rizzini, C. RB 61663 (2-7); RB 132051 (2-9)

Rodrigues, J. S. 108 (3-14)

Rodrigues, R. S. MG 3658 (3-6a); MG 3670 (3-4)

Rodrigues, W. A. et al 519 (3-14); 1377 (3-14); 1385 (3-14); 1538 (1-7); 1541 (1-7); 1960 (1-7); 2235 (1-7); 2320 (1-7); 2406 (1-7); 2530 (1-7); 3035 (1-7); 3166 (3-14); 3420 (3-14); 3440 (3-14); 3467 (3-14); 3835 (1-7); 4107 (1-7); 4367 (1-7); 4404 (1-7); 4457 (1-7); 4550 (1-7); 4712 (3-14); 4845 (1-7); 4950 (1-7); 5060 (1-7, 3-14); 5994 (3-6b); 6921 (3-6b); 7087 (3-6b); 7377 (3-14); INPA 443 (1-7); INPA 2925 (1-7)

Roe, K. et al 785 (1-1la); 787 (1-lla)

Roig, J. T. 119 (3-9a)

Romero C., R. 518 (1-9); 4200 (3-6a); 4309 (3-18); 4790 (3-18); 4824 (3-18); 4857 (3-18)

Rothery 117 (1-15)

Rovirosa, J. N. 192 (1-1la); 783 (1-1 la)

Rusby, H. H. et al 131 (1-15); 287 (3-14); 291 (3-14)

Sagot, P. 191 (1-7 and 1-15); 192 (3-14); 1275 (3-6a); s n (3-6a)

Salvoza, F. M. 723 (3-9a)

82 Flora Neotropica

Samuels, J. A. sn(1-15)

Sandeman, C. 2254 (1-14); 2265 (1-14)

Sandwith, N. Y. 121 (1-15); 178 (3-14); 677 (3-14)

Schipp, W. A. 26 (1-llb); 378 (1-llb); 714 (1-lla)

Schomburgk, Rob. 477 (1-15); 1319 (1-7)

Schultes, R. E. et al 8308 (3-6a); 13565 (3-6a); 14157 (2-4); 19074 (1-12)

Schunke, J. 142 (3-12); 14241 (3-12)

Schunke V., J. 858 (2-4); 624 (1-4)

Shafer, J. A. 4028 (3-9b); 4159 (3-9b); 4209 (3-9a); 8044 (3-9b); 8287 (3-9a); 8289 (3-9b)

Silva, A. 421 (1-15)

Silva, J. F. 27 (3-6a)

Silva, M. B. 1786a (1-7); 2176 (1-4)

Silva, N. T. 12 (3-4); 19 (3-4); 1293 (3-6a); 59726 (1-7)

Simmonds, N. W. 364 (1-15); 395 (1-15)

Skutch, A. F. 3782 (1-9); 3788 (1-9)

Smith, Au. 4 (1-9); 20 (1-9); 519 (1-9); 746 (1-9); 971 (1-9); 2151 (1-9); sn, H.E.S. 4120 (1-9); s n, H.E.S. 4195 (1-9)

Smith, A. C. 2120 (3-14); 2530 (3-14); 2800 (1-15); 2926 (3-14); 3573 (3-14); 3613 (3-14)

Smith, H. H. 1701 (2-2)

Smith, J. D. 2067 (1-lla); 2500 (1-lla)

Snedaker, S. C. D-131 (1-11)

Snethlage, E. H. MG 8216 (1-7)

Spruce, R. 623 (1-7); 780 (1-7); 1109 (1-7); 1185 (1-7); 2165 (1-4); 2226 (3-14); 2808 (3- 6a); 2864 (1-14); 3188 (3-16); 4927 (1-12)

Sousa, M. 3618 (1-llb)

Stahel, G. 133 (3-16); 326 (3-14)

Standley, P. C. et al 3025 (1-lla); 24046 (1-lla); 24496 (1-lla); 24918 (1-lla); 25080 (1-lla); 34674 (1-9); 34723 (1-9); 48497 (1-5); 62267 (1-lla); 63745 (1-lla); 63876 (1-lla); 64882 (1-1la); 65017 (1-11a); 70549 (1-lla); 72813 (1-1); 73053 (1-1); 87703 (1-lla); 88939 (1-lla); 88968 (1-lla); 89569 (1-lla); 91317 (1-1 la)

Stehle, H. 5362 (3-10); 7003 (3-10); 7012 (1-10)

Steyermark, J. A. et al 475 (3-16); 33520 (1-lla); 37992 (1-lla); 38803 (1-1); 39874 (1-1); 46613 (1-llb); 48618 (1-llb); 48800 (1-1lb); 60427 (1-15); 62213 (1-15); 86347 (1-15); 86599 (1-15); 86696 (3-14); 87305 (1-15); 89121 (1-15); 89476 (1-7); 89875 (1-10); 89909 (1-10); 90708 (1-15); 91198 (2-6); 94763 (1-15); 96107 (2-6)

Stockdale, J. A. 8808 (1-15)

Tessmann, G. 3562 (1-4); 4054 (3-3a); 4514 (1-14); 4545 (3-3a); 4987 (3-13); 5377 (1-14)

Tillett, S. S. et al 45762 (1-15)

Traill, J. W. H. 113 (3-5)

Triana, J. J. s n (1-12)

Trinidad Bot. Gard. 487 (1-15); 2134 (1-15)

Tutin, T. G. 144 (3-14); 163 (1-15); 315 (1-15)

Ule, E. 5172 (3-5); 6258 (3-6a); 9524 (3-12)

Van Donselaar, J. 1763 (3-14); 2665 (3-14); 2708 (3-14); 2820 (3-14); 3069 (3-14); 3085 (1-15); 3488 (1-7)

Versteeg, G. N. 187 (3-14); 448 (3-14)

Von Wedel, H. 1424 (1-5)

Wachenheim, H. 15 (3-14); 210 (3-14); 221 (3-6a); 250 (1-15); 254 (1-15); 281 (3-6a); 292 (3-6a); 356 (3-14); 408 (3-6a); 438 (3-14); 498 (3-14); s n (1-15); s n (3-14)

Warming, J. E. B. 1841 (2-1); sn (2-1)

Weberbauer, A. 3693 (3-2)

Weddell, H. A. 769 (2-3)

Wessels Boer, J. G. 1567 (3-6a)

Williams, Ll. 3640 (1-14); 3651 (3-6a); 3725 (1-7); 3783 (1-14); 6605 (1-12); 14110 (1-14)

Williams, L. 0. et al. 14554 (1-lla); 17944 (1-lla)

Wright, C. 1299 (3-9a); 2220 (3-9a); s n (3-9b)

Wurdack, J. J. 34335 (3-14)

Yuncker, T. G. et al 4816 (1-lla); 8491 (1-llb); 8732 (1-11)

Collector unknown s n (3-14)

Index 83

INDEX TO LOCAL NAMES

Abiui 59 Areka-yek 70 Auselin 28 Boesi kofi 70 Bois cote 65 Bois cote gebois 65 Bois cotelette 65 Cayada de aura 63 Duraznito 28 Hibina 52 Hierrito 70 Jabon 70

Kahakudiballi rope 36 Lukuchi-danni 70 Mao de gato 59 Massari 36 Muribania 70 Pakiratiki 70 Palo de aura 63 Rosillet 65 Tassi-tihi 59 Vigueta naranjo 63 Waidan 70 Waiaballi 70 Wassakao 70

INDEX OF SCIENTIFIC NAMES

Chailletia 16 capitulifera 72 cubensis 62 flavicans 24 odorata 32 parkeri 24 pedunculata 34 sessiliflora 68 vestita 24

Cordia scandens 24 Dichapetalum

amazonicum 20 axillare 25 brenesii 25 bullatum 17 chiapasense 28 coelhoi 22 donnell-smithii 26

var chiapasense 28 var donnell-smithii 28

flavicans 24 froesii 32 gentlei 28 glabrum 34 latifolium 20 madagascariensis 16 nervatum 18 nevermannianum 22 odoratum 32 pauper 25 pedunculatum 34 rugosum 24 scandens 24

var cinerascens 20 spruceanum 30 steyermarkii 26 stipulatum 18 vestitum 24

var cinerascens 20 var scandens 24

Gonypetalum acreanum 66 juruanum 56 lanceolatum 70 tessmannii 68

Leucosia 16 Mestotes 16 Moacurra 16 Patrisia 16 Pentastira 16 Plappertia 16 Quilesia 16 Stephanopodium

angulatum 45 aptotum 42 blanchetianum 47 engleri 40

estrellense 42 var organense 49

organense 49 peruvianum 43 sessile 47 sessiliflorum 42 venezuelanum 45

Rohria petioliflora 68 schreberi 68 tapura 68

Symphyllanthus 16 donnell-smithii 26 glaber 34 plicatus 30 rugosus 24

Tapura acreana 66 amazonica 58 var amazonica 58 var ciliata 58 var cuspidata 58 var dasyphylla 58 var manausensis 59 var sublanceolata 58

angulata 45 antillana 65 bullata 66 capitulifera 72 ciliata 58 colombiana 74 coriacea 52 costata 43 cubensis 62

subsp cubensis 63 subsp minor 63 var wrightiana 62

cucullata 68 guianensis 68 haitiensis 62 juliani 51 juruana 56

lanceolata 70 latifolia 65 leucantha 52 negrensis 68 obovata Britt & Wilson 62 obovata Rizzini 47 orbicularis 59 pedicellaris 65 peruviana 52

var peruviana 54 var petioliflora 54

singularis 54 sp 1 74 tessmannii 68

84

FLORA NEOTROPICA

Monograph No. 11

Rhabdodendraceae

by Ghillean T. Prance

TROPIC

OF

CANCER --// 'A

" " " - ~^ ^ ̂ - -----------------------------

FLORA NEOTROPICA

TROPIC OF CAPRICORN

Published for Organization for Flora Neotropica

by Hafner Publishing Company

New York April 20, 1972

A MONOGRAPH OF THE RHABDODENDRACEAE GHILLEAN T. PRANCE1

INTRODUCTION

I first became interested in the genus Rhabdodendron during a study of the status and circumscription of the family Chrysobalanaceae, because it was included in that

family under the name Lecostomion (a misspelling of Lecostemon) by Focke (1891) and

by the earlier workers. This led to a detailed study of Rhabdodendron, a genus which has been associated with 3 widely separated families, Chrysobalanaceae, Phytolaccaceae and

Rutaceae, and has been confused with the genus Sloanea of the Tiliaceae. My original study of Rhabdodendron (Prance 1968) led me to conclude that is is better considered as a separate family. At that time I was more concerned with the status and position of Rhabdodendron than with the monographic details of the species. Since then I have had the opportunity to study 2 of the 3 species of Rhabdodendron in the field, and to study a

greater amount of herbarium material. Hence the aim of this account is to supplement my earlier work with the details of the species.

Rhabdodendron is confined to South America and has 3 known species. One species is widespread in the Guianas and central and eastern Amazonian Brazil. The 2 other

species are of much more restricted range, one in central Amazonia and the other in northern Bahia.

TAXONOMIC HISTORY

The history of Rhabdodendron is rather complicated because of its confusion with Lecostemon Sesse & Moc. ex DC. Lecostemon was described by de Candolle (1825) based on one of the original drawings of Mexican plants by Mociiio & Sesse. Since Lecostemon was based on a drawing only, it was difficult to interpret. Bentham (1853) described 4 new species from South America which he considered to be additional species of Lecostemon. In fact Bentham's 4 species were not at all closely related to Lecostemon of Mocino & Sesse, but belong to what was later to be called Rhabdodendron. It was not until 1944 that Standley, who studied material in Mocino and Sesse herbarium, showed that Lecostemon terniflorum Moc. & Sesse ex DC. is a species of Sloanea in the Tiliaceae. Further details about the Mocino and Sesse drawings and about the identity of Lecostemon were given in Prance (1968); and since Lecostemon Moc. & Sesse ex DC. has

nothing to do with Rhabdodendron, it is not discussed further here. Rhabdodendron was not described until 1905 by Gilg and Pilger, and Bentham's 4

species of Lecostemon were transferred to Rbabdodendron by Huber (1909). Since Lecostemon sensu Bentham was based entirely on 4 species of Rhabdodendron, a taxonomic history of Rhabdodendron includes the history of Lecostemon sensu Bentham from 1844 until it was placed in Rhabdodendron in 1909.

Bentham (1853) placed Lecostemon in the Chrysobalanaceae with the following

1B. A. Krukoff Curator of Amazonian Botany, The New York Botanical Garden.

3

4 Flora Neotropica

comment: "The characters .... place it clearly among Chrysobalanaceae, though as a

very distinct genus." Later authors until 1909 followed Bentham's positioning of the

genus, for example, C. Mueller (1858) and J.D. Hooker (1867) who both listed only the 4 species of Bentham. Lecostemon was placed in Rosaceae tribe Chrysobalaneae by Bentham and Hooker (1865) in their Genera plantarum.

Baillon (1869) included Lecostemon sensu Bentham in his first group of

Chrysobalanoideae, but he pointed out its differences from other genera in the group. Fritsch (1888) treated Lecostemon in a separate tribe of the Chrysobalanaceae. Focke (1891) included it as a member of his subtribe Chrysobalaninae mentioning its anomalies and suggesting an affinity with the Phytolaccaceae. Hallier (1903) placed Lecostemon in the Chrysobalanoideae, a separate and rather isolated tribe.

When Gilg and Pilger (1905) described the new genus Rhabdodendron, they placed it in the Rutaceae and suggested affinities with the tribe Cusparieae. They based their new genus on the single species, R. columnare. Huber (1909) realized that Rhabdodendron and Lecostemon sensu Bentham were congeneric, and he transferred 3 of Bentham's 4 species of Lecostemon to Rhabdodendron, rightly considering that L. terniflorum Moc. & Sesse ex DC. belonged to a different genus though he had not seen the Mocifio herbarium specimen. Huber reduced R. columnare into synonymy since he observed that it was the same as Bentham's L. macrophyllum. He gave a parallel summary of the characters of Lecostemon Sesse & Moc. ex DC. and Rhabdodendron Gilg & Pilg. to establish the fact that they are different genera. Unfortunately he made no reference to Bentham's paper on Lecostemon but apparently took his facts from J. D. Hooker's account in the Flora Brasiliensis which he cites. Huber made no reference to L. gardneranum of which Hooker (1867) said that it seemed to come nearest to the Sesse & Mocinio specimen, nor does Huber include it in his key. Lecostemon gardneranum is known solely from the type gathering and is certainly congeneric with the other 2 species as is shown by floral structure and the wood anatomy, although it differs strikingly in vegetative characters. Huber's assumption that Lecostemon Sesse & Moc. ex DC. was different from Lecostemon sensu Benth. was correct though not the result of a full and critical examination. His transfer of three Lecostemon species to Rhabdodendron stands and he described four new species in the latter genus: R. duckei, R. paniculatum, R. longifolium and R. arirambae all of which have later been reduced to synonymy under the previously published species. Furthermore, he created a new tribe: Rhabdodendreae in the Rutaceae instead of placing it in the tribe Cusparieae of the same family where Gilg & Pilger suggested it might belong. The genus Rhabdodendron was included in Engler's, Pflanzenfamilien, by Krause (1914) where it was placed in the Rutaceae with the observation that Huber had created a separate tribe for it. Engler (1931) later elevated the Rhabdodendron to subfamily rank as the Rhabdodendroideae.

Most later authors except Gleason have accepted Huber's reasoning and used Rhabdodendron as the correct name. Gleason (1927), however, described Lecostemon

sylvestre from British Guiana although he was evidently familiar with Huber's work since he says that "the genus Lecostemon is one of the most poorly known members of (the Chrysobalaneae) ... it has been placed in the Rutaceae."

Ducke (1922) reduced all 4 of Huber's species as well as Bentham's L. crassipes into

synonymy under R. amazonicum (Spruce ex Benth.) Huber. He commented on the

variability within this species and showed that there is a complete gradation between the material described as new by Huber.

Sandwith (1943) further reduced Lecostemon sylvestre Gleason into synonymy under R. amazonicum, and he transferred Lecostemon gardneranum into Rhabdoden- dron, thus leaving only three species in the genus; R. amazonicum, R. macrophyllum, and R. gardneranum.

Maguire (1948) recognized 5 species of Rhabdodendron when he reinstated R.

Introduction 5

crassipes and R. sylvestre and made the necessary new combination in Rhabdodendron for the latter species.

I presented an account of Rhabdodendron (Prance 1968) and discussed the

systematic position of the genus. I concluded that it is better regarded as a separate family near to the Phytolaccaceae.

POLLEN (Fig 1A)

The grains have three furrows and three pores: the ora are circular or slightly lalongate. There is a large polar area and the exine is of medium thickness. The grains are

finely reticulate. Grain size is 18-25 p. Polar length X 100 / equatorial length = 100. There is no significant pollen difference between R. amazonicum and R.

macrophyllum. The pollen does not fit well in either the Rutaceae or the Phytolaccaceae.

LEAF ANATOMY (Fig 1B)

The leaf has a distinctively bifacial mesophyll which contains abundant fatty bodies. It is traversed by fiber-like "spicular cells," both simple and branched, which begin at the vein endings. Silica, in the form of silica bodies, is abundant in the mesophyll, but silica in other forms does not occur. There is a true hypoderm. The palisade tissue is on the upper side of the leaf only and is from 1-4 cells thick. The cells are very short. Characteristic

secretory cavities occur on both sides of the leaf and are filled with resinous contents. The stomata have no accessory cells. The only type of hairs are peltate hairs: their shield has a jagged margin owing to the projecting ray cells, which have thin walls and narrow lamina; each one contains a silica body; the ray cells do not extend to the central part of the shield. The nerves are enclosed in a continuous ring of sclerenchyma. In addition to this they are encircled by a ring of silica-rich parenchyma.

WOOD ANATOMY

1. Mature wood (Fig 1 C-F)

Since the woods of R. amazonicum and R. macrophyllum are so different they are described separately. Details of materials examined are given in Prance (1968).

Rhabdodendron amazonicum (Fig 1 E-F)

The wood shows anomalous secondary thickening in that successive bundles of

xylem and phloem repeat the structure of the young stem and are separated by bands of

conjunctive parenchyma. Vessels abundant, particularly near the anomalous parenchyma, usually solitary,

showing radial and oblique arrangement: size variable, mean diameter 68 ,u; vessels up to 100 p in length, frequent, but larger vessels (up to 135 p) infrequent; perforation porous, semi-circular in shape; the area of the perforation generally oblique to almost vertical; inter-vascular pits minute, alternate, often coalescent, 3.5-4.5 pu in diameter; vessel-to-

parenchyma pits larger, alternate; some vessels filled with gum. Fibers have thick to moderately thick walls; they are thick walled next to the

6 Flora Neotropica

A B

c D

E

FIG 1. Pollen and anatomy of Rhabdodendron. A, Pollen of R. amazonicum (Franshawe &

Maguire 32141), X 425. B, transverse section of lamina of R. amazonicum (Ducke RB 25466), X 38. C-F, The wood of Rhabdodendron, X 27: C-D, R. macrophyllum (INPA-X1396); C, transverse section; D, tangential section; E-F, R. amazonicum (INPA-X1406); E, transverse section; F, tangential section.

Introduction 7

anomalous phloem and thinner away from it; the pits are simple or bordered, few on the

tangential walls, few to numerous on the radial walls; bordering is most marked near the rays [the bordering caused Heimsch (1942) to describe the fibers as tracheids].

Parenchyma: apotracheal parenchyma is very sparse, there being a few cells scattered between the fibers; there is also some paratracheal parenchyma which does not

completely surround the vessels, but is often confluent where the vessels are more crowded; in addition to this there are masses of parenchyma between the successive

phloem and xylem groups. Rays: there is great variation in size between individuals studied; Milanez (1943)

pointed out that there is great variation in ray size according to height above ground level of the specimens studied; the rays are up to 10 cells wide and 55-75 cells high; heterogeneous rays of type IIA occur and have gum inclusions in many cells; pits are scattered and moderately large.

Rhabdodendron macrophyllum (Fig 1 C-D)

The two wood specimens examined of this species differ greatly from those of the

species described above in that they do not possess anomalous structure. This species has been described by Heimsch (1943) and Chalk & Chattaway (1937) and Metcalfe & Chalk (1950). There are further important differences from the other species. The parenchyma is abundant and in apotracheal bands which are continuous or broken; the paratracheal parenchyma is aliform or confluent. The rays are all narrow, 1-2 cells wide, and are short, heterogeneous IIB to homogeneous I. The vessels have simple perforations. The one

similarity with the wood of the other Rhabdodendron species is that the fibers have bordered pits and so are "fiber-tracheids."

2. Twig specimens (Fig 2 A-B)

Sections of twigs for all three species were examined and they have a similar structure to the mature wood. Only one species, R. gardneranum, has anomalous structure in the young wood. In this species a distinct new cambium has developed. The other samples are too young to show it. The 3 species examined show great similarity in the young wood and the bark. Inside, the bark has a layer of thin walled cells and then a

layer of stone cells. Furthermore the leaf trace patterns are all similar. There are definite

spirals in the vessels. On vessel characters they would be hard to separate at all. Rhabdodendron macrophyllum only differs in having more parenchyma and fewer fibers, which is also a feature of its mature wood.

BLASTOGENY

Seedlings have been grown of one species only, R. macrophyllum. In R. macrophyllum the endocarp of the fruit fractures in an indefinite manner. The

germination is hypogeal. The minute (3.0-5.0 mm long) first leaves are alternate and have a rounded apex. The following leaves are progressively larger and have a more pointed apex and are thus more similar to the adult leaves in their shape. The prominent marginal nerve, characteristic of this species, is present even in the youngest leaves. The epicotyl is 3.0-5.0 mm long, and there is a single tap root.

8 Flora Neotropica

A B

D

C

FIG 2. The twig anatomy and ovary of Rhabdodendron. A-B, The twig anatomy, transverse section, X 11. A, R. amazonicum (Tutin 243); B, R. gardneranum (Gardner 2814). C-D, Ovary of R. amazonicum, X 11; C, Pires et al 51199; D, Prance et al 1610.

Introduction 9

OVARY (Fig 2 C-D)

Since the nature of the ovule is important in determining the position of the family, sections were made of the ovary of R. amazonicum (material used: Pires et al 51199, Prance et al 1610).

The ovary is sessile, globose, unilocular and contains a single campylotropous ovule (see Fig 2).

RANK AND POSITION OF RHABDODENDRON

In the taxonomic history it was shown that in the past Rhabdodendron has been linked to 3 families, the Chrysobalanaceae, Phytolaccaceae and Rutaceae. The morphological and anatomical descriptions given here are enough to show that it is completely different from the Chrysobalanaceae. It was placed in the Chrysobalanaceae on the basis of a single character, the basal style. Prance (1968) listed the major anatomical, pollen and morphological differences from the Chrysobalanaceae and it is clearly not closely related to that family. Thus, the Chrysobalanaceae is not considered further here as a

possible position for Rhabdodendron. There are, however, striking similarities and differences between Rhabdodendron

and both the Rutaceae and Phytolaccaceae. In Prance (1968) I have listed the most

important similarities and differences of Rhabdodendron with both families. Rhabdodendron was placed in Rutaceae by Gilg and Pilger (1905) when they first

described the genus, and it was kept there by Engler (1931). Record (1934), on the basis of wood structure, first suggested a definite relationship to Phytolaccaceae, although Focke (1891) had hinted at it earlier on purely morphological grounds. Metcalfe and Chalk (1950) also preferred the Phytolaccaceae on anatomical grounds. Heimsch (1942) pointed out that on the basis of wood structure Rhabdodendron was anomalous in the Rutaceae, a family which is relatively uniform in wood structure. Heimsch did not suggest another position for Rhabdodendron. The only wood anatomist to keep Rhabdodendron definitely in the Rutaceae, largely on the basis of the pit characters which he considered basic, was Milanez (1943).

The most recent phylogenetic systems, with the exception of Takhtajan (1969), have largely overlooked Rhabdodendron. Takhtajan included it in the Rutaceae rather than the Phytolaccaceae.

Rhabdodendron differs from the Rutaceae in its reduced sepals, the numerous stamens, the long linear anthers, the long linear stigma, the absence of a disk, the basal

style, the solitary ovule, the presence of minute stipules, and of peltate hairs, in the

pollen structure, the anomalous secondary phloem of the wood in some species and in the fiber-tracheids.

Rhabdodendron differs from the Phytolaccaceae in its reduced sepals, the presence of petals, the long basal style, the linear stigma, the absence of a disk, the presence of

peltate hairs, the pollen, and the small distinctly bordered pits of the fibers. Rhabdodendron differs from Phytolaccaceae and Rutaceae both in pollen grain

structure and some important morphological characters, but shares some characters of wood anatomy with both, especially the former, although there are important differences as well. On balance the differences far outweigh the resemblances and the affinity with Rutaceae is slight.

Rhabdodendron has a combination of characters which is sufficiently distinct to exclude it from all known families. The most important features are: the minute calyx lobes, the absence of a disk, the long linear anthers with short persistent filaments, the basal style, the single unilocular, uniovulate carpel, the gland-dotted leaves, the peltate hairs, the anomalous secondary phloem and the conspicuously bordered fiber pits.

10 Flora Neotropica

Since many of its features occur in the Centrospermae (sensu Engler & Prantl), though scattered through several families, it seems appropriate to place it there, in a

family of its own, the Rhabdodendraceae (Prance 1968).

ACKNOWLEDGMENTS

I am grateful to the directors and curators of all the herbaria cited in the systematic part for making material available for this study. I am especially grateful to the director and other staff members of The New York Botanical Garden for much help and advice, particularly to Dr. Bassett Maguire, also to the director and staff members of the Instituto Nacional de Pesquisas da Amazonia, Manaus, for much assistance with field studies. I am

especially grateful to Dr. William A. Rodrigues for sending me correlated wood material and for assistance in locating 2 species in the field.

This study was greatly assisted by grants from the National Science Foundation, GB

4641, GB 7356 and GB 18655 which enabled me to study Rhabdodendron in the field and provided much of the assistance involved in the preparation of this account.

I should like to thank Mr. C. C. Clare for preparing the line drawings and distribution maps.

SYSTEMATIC TREATMENT

Rhabdodendraceae Prance, Bull. Jard. Bot. Brux. 38: 141-142. 1968. Rutaceae tribus Rhabdodendron Huber, Bol. Mus. Emilio Goeldi 5: 425. 1909. Rutaceae subfam Rbabdodendroideae Engler in Engler & Prantl, Nat. Pflanzenfam. ed. 2, 19a:

213,357. 1931.

Shrubs or small trees. Leaves entire, alternate, gland-dotted, coriaceous, with small

peltate hairs on the undersurface; stipules small, subulate or obscure. Inflorescence of

supra-axillary racemose panicles or racemes; bracts and bracteoles small and reduced to scales. Flowers hermaphrodite; receptacle broad, slightly concave; calyx very short, the lobes 5 or indistinct; petals 5, caducous, sepaloid, oblong or oblong-elliptic, the apex rounded or minutely apiculate, minutely punctate, aestivation imbricate; disk absent; stamens numerous (about 45), the filaments short, flattened, persisting after the anthers fall, and then recurved; anthers linear, erect, basifixed, caducous, 4-locular, dehiscing longitudinally; ovary sessile, globose, glabrous, unilocular, inserted at the base of the concave receptacle; ovule 1, basally attached, campylotropous; style arising from the base of the ovary to one side of it, fairly thick, elongated; stigmatic surface on the outermost side ascending from the base or the middle. Fruit a small drupe, globose, terminating a short stipe in the cup-shaped receptacle; exocarp thin, crustaceous when dry; endocarp slightly woody. Seed 1, reniform-globose, exalbuminous, with a thin testa; cotyledons thickly fleshy, conferruminate; radicle small and bent inward towards the hilum.

TYPE GENUS. Rhabdodendron Gilg & Pilger. Rhabdodendron Gilg & Pilger, Verh. Bot. Ver. Prov. Brand. 47: 152. 1905

Key to Species of Rhabdodendron 1. Leaves 2.0-4.5 cm long; stem with thick corky bark. 1. R. gardneranum. 1. Leaves 9.0-40.0 cm long; stem with thin bark.

2. Leaves chartaceous; primary leaf veins strongly anastomosing to form a marginal vein; wood without secondary phloem; leaves subsessile. 2. R. macrophyllum.

2. Leaves coriaceous; primary leaf veins not anastomosing and marginal vein absent; wood with secondary phloem present; leaves distinctly petiolate. 3. R. amazonicum.

1. Rhabdodendron gardneranum (Bentham) Sandwith, Jour. Arnold Arb. 24: 224. 1943; Prance, Bull. Jard. Bot. Brux. 38: 143. 1968. Fig 4, 6 A-B.

Lecostemon gardneranum Bentham, Jour. Bot. Kew Misc. 5: 295. 1853; Hooker f., Mart. Fl. Bras. 14(2): 54. 1867.

Small tree to 5.0 m tall, the young branches densely peltate pubescent, soon

glabrous, with a thick corky bark, the wood with anomalous secondary phloem. Leaves

narrowly oblong, coriaceous, the laminas 1.2-4.0 cm long, 0.5-1.6 cm broad, the apex

11

12 Flora Neotropica

shortly and abruptly acuminate or apiculate, the acumen to 1.5 mm long, cuneate to subcuneate at base, glabrous and slightly rugose above, rugose and with sparse peltate pubescence beneath; midrib plane above, prominulous beneath; primary veins plane and

inconspicuous on both surfaces; petioles 4.0-8.0 mm long, terete, peltate pubescent. Stipules lanceolate, caducous, to 2.5 mm long. Inflorescences of axillary racemes to 3.0 cm long, with sparse peltate pubescence. Bracteoles lanceolate, 1.5-2.5 mm long, persistent, with peltate pubescence; pedicels 2.0-3.0 mm long, with sparse peltate pubescence. Calyx-tube turbinate-campanulate, ca 2.0 mm long, the lobes represented by 5 small teeth only, the exterior with scattered peltate hairs. Petals 5, oblong-elliptic, ca 5.0 mm long, sepaloid, minutely punctate. Stamens ca 45, the filaments minute, flattened, persisting after anthers fall and then recurved; anthers linear, erect, basifixed, caducous. Ovary globose. Style arising from base of ovary to one side of it, elongate, the

stigmatic surface long and linear. Fruit not seen. TYPE. Gardner 2814, Brazil, Bahia, Rio Preto, fl (holotype, K; isotypes, BM, CGE,

GH, NY, OXF, P). DISTRIBUTION. Rhabdodendron gardneranum is known only from the type

gathering from northwest Bahia which lacks details of the habitat.

The type of this species is labelled as Piaul, but the locality according to Gardner's

itinerary is in fact in the northwest corner of the present state of Bahia. Rhabdodendron

gardneranum is the most distinct species of the genus differing in its small leaves and thick corky bark. The flower structure is identical with the other species. That this

species is known only by the type indicates the paucity of collecting in northeastern Brazil, and the need for further field work in the region.

2. Rhabdodendron macrophyllum (Spruce ex Bentham) Huber, Bol. Mus. Emilio Goeldi 5: 428. 1909. Prance, Bull. Jard. Bot. Brux. 38: 143. 1968. Fig 3, 4, 5.

Lecostemon macrophyllum Spruce ex Bentham, Jour. Bot. Kew Misc. 5: 296. 1853; Hooker f., Mart. Fl. Bras. 14(2): 55-56. 1867.

Rhabdodendron columnare Gilg & Pilger, Verh. Bot. Ver. Brandenburg 47: 152. 1905. Type. Ule 6165, Brazil, Amazonas, Manaus, fl (holotype, B lost; lectotype, K; isotype, L; isotype fragment, F).

Shrub to 6.0 m tall with numerous thin trunks arising from below ground, the young branches glabrous, with a thin hard bark, the wood without secondary phloem. Leaves oblanceolate, lanceolate or obovate-oblong, gradually narrowing from above middle to

base, with much variation in size and shape within the same individual, chartaceous, (7.0-)9.0-32.0(-38.0) cm long, (1.5-)2.0-7.5(-8.6) cm broad, the apex rounded, acute or minutely acuminate or mucronate, gradually narrowed to a cuneate base, glabrous above, with scattered peltate hairs beneath, not rugose on surfaces; midrib plane to

prominulous above, prominent beneath; primary veins 25-40 pairs, prominulous on both

surfaces, arcuate and anastomosing 1.0-6.0 mm from margin and forming a prominulous

marginal nerve; petioles 2.0-6.0 mm long, glabrous, usually minutely winged and

shallowly canaliculate. Stipules small, subulate. Inflorescences of axillary racemes 3.0-9.0 cm long, with sparse peltate pubescence. Bracts and bracteoles ovate, persistent, 0.5-1.0 mm long, membraneous, glabrous; pedicles 5.0-12.0 mm long, glabrous and frequently with up to 3 small bracteoles. Calyx-tube turbinate-campanulate, ca 3.0 mm long, the lobes represented by 5 obscure teeth only, the exterior glabrous. Petals 5, oblong. 6.0-8.0 mm long, membraneous, glabrous; pedicels 5.0-12.0 mm long, glabrous and frequently with up to 3 small bracteoles. Calyx-tube turbinate-campanulate, ca 3.0 mm long, the

basifixed, caducous. Ovary globose, glabrous. Style arising from base of ovary to one side of it, elongate, the stigmatic surface long and linear. Fruit subglobose, 5.0-6.0 mm

diameter; exocarp glabrous, smooth, but wrinkled when dry; mesocarp very thin, fleshy;


::i~~~~~~~~~~~~~~i

8' frl,J'~~~~~~~~~~~~~~~~~~~~~~~~~~~~~: . s~~~~~~~~~~~~~i~~~~~~ ~~~~~~s . . . . :

A ~

FIG 3. The habit of species of Rbabdodendron. A, R. amazonicum (Prance 58955); B, R.

macropbyllum (Prance 2219).

14 Flora Neotropica

endocarp thin, bony, fragile, with median line of fracture, glabrous within. Germination

hypogeal, first leaves alternate, minute, 2.0-5.0 mm long, with a single long tap root. TYPE. Spruce 1408, Dec-Mar 1850-51, s n in some sets), Brazil, Amazonas,

Manaus, fl (holotype, K; isotypes, CGE, F, LD, NY, OXF, P). DISTRIBUTION. Characteristic of and common in sandy savannas and secondary

savanna in the lower Rio Negro region of Amazonian Brazil and eastward to western Para. Collected in flower around the year but more in December to June. BRAZIL. Amazo- nas: Chagas INPA 5556 fl (IAN, INPA, MG); Coblho INPA 1173 fl (F, IAN, INPA); Corner 106 fr (IAN); Ducke 163 fl (A, F, IAN, K, MG 18296, NY, R, US); 177 fl fr (A, F, NY);MG 11173 fr (MG); MG 12198 fr (BM, MG); RB 13623 fl (IAN, K, RB); Ferreira 77 fr (INPA 5825, IAN, MG); Froes 25050 fr (IAN, IPA, R); J.G. Kuhlmann 1997 fl (RB 15925); Mello & Collho INPA 3488 fl (INPA, MG); Pires & Silva 8074 fl (IAN); Prance 2080 fr (NY, U); Prance et al 2219 fr (INPA, NY); 2691 fr (INPA, NY); 3114, fr (INPA, NY); 4742 fl (INPA, NY); 4842 fl (FHO, INPA, K, MICH, MO, NY, R, S, US, VEN); 5044 fr (A, F, INPA, K, MG, NY, R, S, U, US); 5078 fr (A, F, FHO, INPA, K, MG, NY, R, S, U, US); 5084 fr (INPA); Riedel 1436 fl (NY); Rodrigues 474 fl (FHO, INPA 5648, MG); INPA 1173 fl (INPA, MG); Rodrigues &Chagas 1731 fl (INPA 8094, MG); 2787 fl (FHO, INPA 9330); 4484 fl (INPA 12975); 4494 fl (INPA 12994); 4617 fr (INPA 13126); 4692 fr (INPA 13206); 5187 fl (INPA 13854); 5241 fr (INPA 13909); Rodrigues & Collho 1190 fl (FHO, INPA 7503); Rodrigues, Coelho & Lima 2774 fl (INPA 9317); Rodrigues & Lima 2647 fr (FHO, INPA 9190); Spruce 1181 fl (M); 5752 fl (MG); Ule 8986 fl (K, L). Para: Ducke MG 8595 fr (BM, MG, RB 13622, US). Martius s n st (M).

Rhabdodendron macrophyllum is closest to R. amazonicum but differs in the leaf

shape, the strongly anastomosing primary veins which form a marginal nerve, the shorter

petioles, and in the absence of anomalous secondary phloem in the wood. This species is

extremely polymorphic in leaf shape and size, even within the same individual. I have observed and collected it several times around Manaus and Prance 2080 is a series showing the range of leaf shape on the same bush, varying from narrowly lanceolate leaves 9.0 X 1.5 cm to oblong leaves 25.0 X 8.0 cm.

3. Rhabdodendron amazonicum (Spruce ex Bentham) Huber, Bol. Mus. Emilio Goeldi 5: 427. 1909; Sandwith, Jour. Arnold Arb. 24: 223. 1943; Prance, Bull. Jard. Bot. Brux. 38: 142. 1968. Fig 3, 4, 6 C-J.

Lecostemon amazonicum Spruce ex Bentham, Jour. Bot. Kew. Misc. 5: 295. 1853; Hooker f., Mart. Fl. Bras. 14(2): 54. 1867.

Lecostemon crassipes Spruce ex Bentham, Jour. Bot. Kew Misc. 5: 295. 1853; Hooker f., Mart. Fl. Bras. 14(2): 55. 1867. Type. Spruce 1497, Brazil, Amazonas, Manaus, fl (holotype, K; isotypes, BM, CGE, M, NY, OXF, P).

Rhabdodendron crassipes (Spruce ex Bentham) Huber, Bol. Mus. Emilio Goeldi 5: 428. 1909; Maguire, Bull. Torrey Club 75: 396. 1948; Prance, Bull. Jard. Bot. Brux. 38: 143. 1968.

Rhaoaoaendron crassipes var cayenense Bentham, Jour. Bot. Kew Misc. 5: 295. 1853. Type. Martin s n, French Guiana, fl (holotype, K; isotype, BM).

Rhabdodendron duckei Huber, Bol. Mus. Emilio Goeldi 5: 428. 1909. Type. Ducke MG 8546,

Brazil, Para, Faro, fl (holotype, MG, isotypes, RB 8559, US). Rhabdodendron paniculatum Huber, Bol. Mus. Emilio Goeldi 5: 429. 1909. Type. Ducke 8W54,

Brazil, Para, Obidos, fl (holotype, MG; isotypes, BM, US). Rhabdodendron longifolium Huber, Bol. Mus. Emllio Goeldi 5: 429. 1909. Types. Ducke MG

8504, Brazil, Para, Faro, fl (syntypes, MG, RB 8559); Ducke MG 8989, Brazil, Para, Rio

Mapuera, Morro do Taboleirinho (not seen). Rhabdodendron arirambae Huber, Bol. Mus. Emilio Goeldi 5: 430. 1909. Type. Ducke MG 8000,

Brazil, Para, Alto Ariramba (not seen). Lecostemon sylvestre Gleason, Bull. Torrey Club 54: 68. 1927. Type. Gleason 211, Guyana,

between Kangaruma and Potaro Landing, fl (holotype, NY; isotype, GH). Rhabdodendron sylvestre (Gleason) Maguire, Bull. Torrey Club 75: 397. 1948; Prance, Bull. Jard.

Bot. Brux. 38: 144. 1968.

Trees to 15.0 m X 20.0 cm diameter, usually smaller, the young branches with scattered peltate hairs, with a thin hard bark, the wood with anomalous secondary

phloem. Leaves oblanceolate, oblong to oblong-obovate, gradually narrowing from above


so '.

. , . ,

ae

FG 4. Distribution of species of Rbabdodendron. R. macropbyllum; o R. gardnerau; * R.

amaIG 4 Distribution of species of

Rabdcmndron. R. macropylum; R. garneranum; R.

amazonicum.

16 Flora Neotropica

middle to base, coriaceous, 20.0-39.0 cm long, 3.0-10.0 cm broad, the apex acute, acuminate or mucronate, most frequently with acumen 2.0-9.0 mm long, gradually narrowed to a cuneate base, glabrous above, with few scattered peltate hairs beneath, not

rugose on surfaces; midrib plane to prominulous above, prominent beneath; primary veins 30-45 pairs, plane to prominulous above, prominulous beneath, anastomosing but not

forming a conspicuous marginal nerve; petioles 1.5-3.5 cm long, with scattered peltate hairs, not winged, terete. Stipules absent. Inflorescences of axillary and sometimes terminal panicles or occasionally reduced to racemes, 9.0-17.0 cm long, sparsely peltate pubescent becoming glabrous with age. Bracts and bracteoles ovate to lanceolate, persistent, 1.0-2.0 mm long, chartaceous; pedicels 6.0-15.0 mm long, glabrescent, frequently recurved, often with 2 lanceolate bracteoles. Calyx-tube turbinate- campanulate, 2.0-4.0 mm long, the exterior glabrescent, the lobes small but distinct and apparent in young flowers only. Petals 5, oblong, 7.0-8.0 mm long, sepaloid, minutely punctate. Stamens ca 45, the filaments short and flattened, persisting after flowering and then recurved; anthers linear, ca 7.0 mm long, basifixed, caducous. Ovary globose, glabrous. Style arising from base of ovary to one side of it, elongate, the stigmatic surface

long and linear. Fruit subglobose, 6.0-10.0 mm diameter; exocarp glabrous, smooth but wrinkled when dry; mesocarp very thin, fleshy; endocarp thin, bony; fragile, with median line of fracture, glabrous within.

TYPE. Spruce 377, Brazil, Para, Santarem, fl (holotype, K; isotypes, LD, MG, OXF, P).

DISTRIBUTION. Forest on non-flooded sandy ground in the Guianas and in central to eastern Amazonian Brazil. Collected in flower June to October in the Guianas, August to December in Amazonia. GUYANA. Fanshawe 138 (F.D. 2747) fl (F, FHO, K, U); 2497 (F.D. 5233) fr (F, K, NY, U); Hohenkerk 450 fr (K); 450A fl (K); Maguire & Fanshawe 23558, fr (F, GH, K, NY, U); 32141 fl (K, NY); Sandwith 1237 fl (K, NY, U); Tutin 243 fl (BM, K, RB). SURINAME. L.B.B. 10694 fr (NY, U); Schulz 8322a fl (U); Wessels-Boer 334 fr (U). FRENCH GUIANA. For. Serv. 7406 fr (NY, P); Leprieur s n (1838) fr (F, P); s n (1840) fl (P). BRAZIL. Amazonas: Chagas INPA 389 fr (FHO, INPA); Ducke 220 fr (A, F, K, NY, R, US); 794 fl (F, IAN, NY, R, US);MG 11218 fl (INPA); MG 11606 fr (BM, MG); RB 25466 fl (K, RB); Froes 21340 fl (K, IAN); 33110 fl (IAN); 33549 fr (IAN); 33796 fr (IAN); Oliveira 2930 fr (IAN); Prance et al 2079 fr (GH, NY); 3652 fr (INPA, NY); Riedel 1355 st (NY); Rodrigues 3401 fr (FHO, INPA 10768); 4202 fr (INPA 10768); 5499 fl (INPA 14172, RB); Rodrigues & CoNlho 5627 fr (INPA 14319); Ule 8882 fl (INPA, K, L, MG 13904, NY, US). Para: Black 55-18744?2 fr (IAN); Burchell 9279 fl (K); Ducke MG 4856 (RB 13625); MG 9950 fl fr (BM, MG, US); MG 9964 fl (BM, MG);MG 10216 fr (BM, MG);MG 10339 fr (BM, MG, US); MG 10525 fr (BM, MG); MG 10826 fr (BM, MG, US); MG 10938 fl (BM, MG, P, RB 8560, US); MG 10968 fl (BM, MG, R); MG 11370 fr (BM, MG, RB 13624, US); MG 11371 fr (BM, MG, P, R, RB 13624, US); MG 14986 fl (BM, MG, US); MG 15618 fl (BM, MG);MG 15933 fl (BM, MG, RB 13627); Egler 374 fl (MG 22313, INPA); 1314 fl (MG 24361, NY);Fr6es 20461 fr (IAN, K, NY); 31234 fl (IAN, SP); 31601 fr (IAN, SP); Froes & Pires 24132 fr (IAN); Huber MG 10448 fl (BM, MG, RB 13626); Killip & Smith 30588 fl (NY, US);J.G. Kuhlmann 1738 fl (RB 21022); Lima 1592 fr (IAN, IPA); Monteiro da Costa 17 fl (R); 213 fr (F, IAN); Oliveira 963 fl (IAN); 2480 fr (IAN); 3892 fr (IAN, NY); 3937 fr (IAN, NY); 4160 fr (IAN, NY); Pires 1444 fr (IAN, US); Pires & Silva s n fl (IAN); Pires et al 6490 fl fr (IAN); Prance et al 1610 fr (F, IAN, NY, U); 58955 fl (F, IAN, NY, RB); N.T. Silva 1108 fl (IAN, NY); 1134 fl (IAN, NY); 1470 fl (IAN, NY); Spruce 473 fl (M); s n fl (CGE). Terr. Amapa: Black 49-8174 fl (IAN, NY); 49-8542 fl (IAN); Cowan 38643 fl fr (IAN, NY, US); 38656 fl (NY, RB); Froes 25704 fr (IAN); 26600 fl (IAN, SP); Froes & Black 27566 fr (IAN); Irwin 47524 fr (IAN, MG, NY); 48635 fl (IAN, MG, NY);Maguire 47095 fr (IAN, MG, NY);Mattos 10061 fl (SP); Pires & Cavalcante 52208 fl fr (MG, NY, R, RB); 52251 fr (GH, MG, NY); Pires, Rodrigues & Irvine 50561 fr (IAN, MG, NY, R, RB, U); 51059 fl (IAN, MG, NY); 51125 fl fr (IAN, NY); 51199 fl (IAN, INPA, MG, NY, SP); 51336 fl (GH, IAN, INPA, MG, NY).

LOCAL NAMES. Guyana: Mabua (Arawak). Suriname: Powisitere. Brazil: Batiputa.

Much of the synonymy given here for this species has already been suggested, e g by Ducke (1922), Sandwith (1943) and Prance (1968). However, Maguire (1948) and myself (Prance 1968) both recognized R. sylvestre and R. crassipes as distinct from R. amazonicum. I now have seen a larger amount of material, have studied the problem in


F E

H E F G

FIG 5. Rbabdodendron macrophyllum (Ducke 177, Prance et al 2080, 2691, 5098). A, habit X 0.5; B, seedling X 0.5; C, flower X 5; D, flower section X 5; E, anther and filament, ventral view X 10; F, anther and filament, dorsal view X 10; G, style, ventral view X 10; H, fruit X 2.

18 Flora Neotropica

the field, and have concluded that R. sylvestre and R. crassipes are not distinct from R. amazonicum. R. crassipes was distinguished from R. amazonicum by the broader more

elliptic leaves with a less abrupt acumen, and the larger flowers. However, there is much variation in leaf shape and flower size, and there is a complete gradation in those characters and even within the same individual. It is possible to collect leaves similar to typical Lecostemon amazonicum at one height of the tree and those that have been called L. crassipes at another. Similarly L. sylvestre cannot be maintained nor can Bentham's L. amazonicum variety cayenense. Thus for this single rather polymorphic species there are 6 synonyms.

LITERATURE CITED

Baillon, H. 1869. Rosaceae. Histoire des plantes 1: 345-483. Bentham, G. 1853. Notes on two little-known genera connected with the South American flora. Jour.

Bot. Kew Misc. 5: 289-296. Bentham, G. &J. D. Hooker 1865. Rosaceae tribus Chrysobalaneae. Genera plantarum 1(2): 600-609. Chalk, L. & M. M. Chattaway 1937. Identification of woods with included phloem. Trop. Woods 50:

1-31. De Candolle, A. P. 1825. Rosaceae. Prodromus systematis naturalis regni vegetabilis 2: 525-639. Ducke, A. 1922. Rutaceae. Plantes nouvelles ou peu connues de la region amazonienner. Arch. Jard.

Bot. Rio de Janeiro 3: 181-182. Engler, A. 1931. Rutaceae. Engler, A. & K. Prantl, Die natiirlichen Pflanzenfamilien, ed 2, 19a:

187-359. Focke, W. 0. 1888, 1891. Rosaceae. Engler, A. & K. Prantl, Die natiirlichen Pflanzenfamilien, ed. 1.

3(3): 1-48 (1888). 49-61 (1891). Fritsch, C. 1888. Uber die Gattungen der Chrysobalanaceen. Verh. Zool-Bot. Ges. Wien 38: 93-95. Gilg, E. & R. Pilger 1905. Rutaceae. Pilger, R., Beitrage zur Flora der Hylea nach deu Sammlungen von

E. Ule. Verh. Bot. Ver. Prov. Brandenburg 47: 152-154. von E. Ule. Verh. Bot. Ver. Prov. Brandenburg 47: 152-154.

Gleason, H. A. 1927. Studies on the flora of northern South American -X. Bull. Torrey Club 54: 603-618

Hallier, H. 1903. Uber die Verwandschaftverhaltnisse bei Engler's Rosalen, Parietalen, Myrtifloren und in anderen Ordnungen der Dikotylen. Abh. Naturw. Ver. Hamburg 18: 98.

Heimsch, C. Jr. 1942. Comparative anatomy of the secondary xylem in the Gruinales and Terebinthales of Wettstein, with reference to taxonomic grouping. Lilloa 8: 83-198.

Hooker, J. D. 1867. Rosaceae. Martius, K.F.P., Flora Brasiliensis 14(2): 1-76. Huber, J. 1909. Rhabdodendron. Materiaes para a flora amazonica, VII, Plantae Duckeanae austro-

guyanenses. Bol. Mus. Emilio Goeldi 5: 424-431. Krause, K. 1914. Rutaceae. Engler, A. & K. Prantl, Die natiirlichen Pflanzenfamilien, Ergazungshaft 3,

Nachtr. 4: 157. Maguire, B. 1948. Rutaceae. Plant explorations in Guiana in 1944, chiefly to the Tafelberg and the

Kaieteur Plateau-IV. Bull. Torrey Club 75: 396-397. Metcalfe, C. R. & L. Chalk 1950. Phytolaccaceae. Anatomy of the Dicotyledons 2: 1086-1091. Milanez, F. R. 1943. Anatomia das principais madeiras brasileiras das Rutaceae. Rodriguesia 7: 5-22. Mueller, C. 1858. Chrysobalaneae. Synopsis plantarum phanerogamicarum etc, Walp. Ann. Bot. Syst.

4: 640-649. Prance, G. T. 1968. The systematic position of Rhabdodendron Gilg & Pilg. Bull. Jard. Bot. Brux. 38:

127-146. Record, S. J. 1934. The woods of Rhabdodendron and Duckeodendron. Trop. Woods 33: 6-10. Sandwith, N. Y. 1943. Rutaceae. New and Noteworthy Polypetalae from British Guiana. Jour. Arnold

Arb. 24: 223-224. Standley, P. C. 1944. The status of the genus Lecostemon Moc. & Sesse. Trop. Woods 79: 9-10. Takhtajan, A. 1969. Flowering plants, origin and dispersal. 213, 225.


1:t~~~~~~~~

:i~?::

'."-!f D I_JE F G H

FIG 6. Species of Rhabdodendron. A-B, R. gardneranum (Gardner 2814). A, habit X 0.5; B, flower bud X 4. C-J, R. amazonicum (Ducke 794, Pires et al 52251). C, habit X 0.5; D, flower X 5; E, flower section X 5; F, anther and filament, ventral view X 5; G, anther and filament, dorsal view X 5; H, style X 5;J,fruitX 2.

20 Flora Neotropica

NUMERICAL LIST OF TAXA

1. Rhabdodendron gardneranum (Bentham) Sandwith 2. R. macrophyllum (Spruce ex Bentham) Huber 3. R. amazonicum (Spruce ex Bentham) Huber

LIST OF EXSICCATAE

The figures in parentheses refer to the species number in the list above.

Black, G. A. 49-8174 (3); 49-8542 (3); 55-18744 (3)

Burchell, W. J. 9279 (3)

Chagas, J. INPA 389 (3); INPA 5556 (2)

Coelho, L. INPA 1173 (2)

Corner, E. J. H. 106 (2)

Cowan, R. S. 38643 (3); 38656 (3)

Ducke, A. 163 (2); 177 (2); 220 (3); 794 (3); MG 4856 (3); MG 8000 (3); MG 8504 (3); MG 8546 (3); MG 8595 (2); MG 8854 (3); MG 8989 (3); MG 9950 (3); MG 9964 (3); MG 10216 (3); MG 10339 (3); MG 10525 (3); MG 10826 (3); MG 10938 (3); MG 10968 (3); MG 11173 (2); MG 11218 (3); MG 11370 (3); MG 11371 (3); MG 11606 (3); MG 12198 (2); MG 14986 (3); MG 15618 (3); MG 15933 (3); RB 13623 (2); RB 25466 (3)

Egler, W. A. 374 (3); 1314 (3)

Fanshawe, D. B. 138 (3); 2497 (3)

Ferreira, E. 77 (2)

Forest Serv. French Guiana 7406 (3)

Fr6es, R. L. et al 20461 (3); 21340 (3); 24132 (3); 25050 (2); 25704 (3); 26600 (3); 27566 (3); 31234 (3); 31601 (3); 33110 (3); 33549 (3); 33796 (3)

Gardner, G. 2814 (1)

Gleason, H. A. 211 (3)

Hohenkerk, L. S. 450 (3); 450A (3)

Huber, J. MG 10448 (3)

Irwin, H. S. 47524 (3); 48635 (3)

Killip, E. P. et al 30588 (3)

Kuhlmann, J. G. 1738 (3); 1997 (2)

L. B. B. (Lands Bosbeheer, Suriname) 10694 (3)

Leprieur, F. R. s n (1838) (3); s n (1840) (3)

Lima, D. A. 1592 (3)

Maguire, B. et al 23558 (3); 32141 (3); 47095 (3)

Martin, J. sn(3)

Martius, K. F. P. von sn(2)

Mattos, J. R. 10061 (3)

Mello, F. et al INPA 3488 (2)

Monteiro da Costa, R. C. 17 (3); 213 (3)

Oliveira, E. 963 (3); 2480 (3); 2930 (3); 3892 (3); 3937 (3); 4160 (3)

Pires, J. M. et al 1444 (3); 6490 (3); 8074 (2); 50561 (3); 51059 (3); 51125 (3); 51199 (3); 51336 (3); 52208 (3); 52251 (3); s n (3)

Prance, G. T. et al 1610 (3); 2079 (3); 2080 (2); 2219 (2); 2691 (2); 3114 (2); 3652 (3); 4742 (2); 4842 (2); 5044 (2); 5084 (2); 58955 (3)

Riedel, L. 1355 (3); 1436 (2)

Rodrigues, W. A. et al 474 (2); 1190 (2); 1731 (2); 2647 (2); 2774 (2); 2787 (2); 3401 (3); 4202 (3); 4484 (2); 4494 (2); 4617 (2); 4692 (2); 5187 (2); 5241 (2); 5499 (3); 5627 (3); INPA 1173 (2)

Sandwith, N. Y. 1237 (3)


Schulz, J. P. 8322a (3)

Silva, N. T. 1108 (3); 1134 (3); 1470 (3)

Spruce, R. 377 (3); 473 (3); 1181 (2); 1408 (2); 1497 (3); 5752 (2); sn (2); sn (3)

Tutin, T. G. 243 (3)

Ule, E. 6165 (2); 8882 (3); 8986 (2)

Wessels-Boer, J.G. 334 (3)

INDEX OF LOCAL NAMES

Batiputa 16 Mabua 16 Powisitere 16

INDEX OF SCIENTIFIC NAMES

Lecostemon amazonicum 14 crassipes 14 gardneranum 11 macrophyllum 12 terniflorum 3

Rhabdodendron 11-18 amazonicum 5, 14 arirambae 14

columnare 12 crassipes 14 duckei 14 gardneranum 11 longifolium 14 macrophyllum 7, 12 paniculatum 14 sylvestre 14

22

BIOGRAPHICAL SKETCH

Dr. Ghillean T. Prance was born July 13, 1937 in Brandeston, England and was educated at Malvern College, Worcestershire, England, 1952-1956, and Keble College, Oxford, 1957-1963, where he received the Bachelor of Arts degree with honors in botany in 1960. In 1963 Dr. Prance received the degree of Doctor of Philosophy in forest botany at Oxford University on the completion of his thesis "A Taxonomic Study of the

Chrysobalanaceae." Since 1963 he has been on the staff of The New York Botanical Garden from 1963-1966 as Research Assistant, and from 1966-1968 as Associate Curator. In 1968 he became the first holder of the B. A. Krukoff Curator of Amazonian Botany. He has carried out extensive field work in Brazilian Amazonia since 1964. His principal research interests are taxonomy of the Chrysobalanaceae, Caryocaraceae, Dichapetalaceae and Lecythidaceae, the flora of the Amazon basin, and numerical systematics.

Flora Neotropica is designed to present in monographic form taxonomic accounts of all plants growing spontaneously within the Western Hemisphere tropics. Geographic, ecologic, cytologic, anatomic, morphologic, chemical and economic data will provide complementary information for each contribution. Bibliography, citation of specimens and indexes are intended to facilitate consultation.

Monographs of Flora Neotropica will be issued separately without regard to taxonomic sequence. Each author will be wholly responsible for his own contribution, being restricted only by the general style and form of the work.

Those interested in preparing a monograph for Flora Neotropica must consult the Staff Committee.

For further information write to Bassett Maguire, Executive Director, Organization for Flora Neotropica, The New York Botanical Garden, Bronx, N. Y. 10458, U. S. A.

dichapetalaceae_rhabdodendraceae 1972

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