AGRICULTURAL UNIVERSITY WAGENINGEN PAPERS 86-3 (1986) THE AFRICAN DICHAPETALACEAE IX. A taxonomical revision F.J. BRETELER

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1 AGRICULTURAL UNIVERSITY WAGENINGEN PAPERS 86-3 (1986) THE AFRICAN DICHAPETALACEAE IX A taxonomical revision F.J. BRETELER Department of Plant Taxonomy Wageningen, Agricultural University, The Netherlands Received 29-V-1986 Date of publication 3 nov«bter 19Ö& (Lift) Agricultural University Wageningen The Netherlands 1986

2 CIP ISBN x ISSN x No part ofthis publication, apart from bibliographic data andbrief quotations embodiedincriticalreviews,maybe reproduced, re-corded orpublished inany form including print, photocopy, microform, elektronic or elektromagneticrecord without written permission from the publisher Agricultural University, P.O.Box 9101,6700 HBWageningen,the Netherlands. Printedinthe Netherlandsby DrukkerijVeenmanb.v.,Wageningen

3 CONTENTS INTRODUCTION AND ACKNOWLEDGEMENT 1 GENERAL PART 1 Cytology additions and survey by J. C. Arends & F. M. van der Laan 1 Pollen morphology: additions by W. Punt 5 TAXONOMIC PART 8 Description of the family 8 Delimitation of the genera 9 Key to the genera in Africa 9 Revision of Dichapetalum 10 Subdivision of the genus 10 Keys to the African species 18 Additions, corrections and emendations to the previous parts III- VIII 31 Unidentified material, possibly new taxa 42 Revision of Tapura 43 History of the genus 43 Description of the genus 44 Sectionnai arrangement 45 Key to the species 46 Alphabetical treatment of the species 46 REFERENCES 70 INDEX OFNAMES 72

4 INTRODUCTION AND ACKNOWLEDGEMENTS This publication, the ninth in the series The African Dichapetalaceae, appears much later than announced in part VIII. It is intended that part IX will be the last part dealing with the revision of the African Dichapetalaceae, although the additions to the previous parts presented in here indicate that it is likely that further botanical exploration, especially of Western Central Africa, will yield important additional information to be published. The author is grateful to the directors and curators of the herbaria cited for their continued loan of material. For the field work in Gabon, which yielded the first record for Dichapetalum rudatisii, a new variety of D. choristilum and a new species, a grant was received from the -Foundation for the Advancement of Tropical Research (WOTRO). The illustrations have been made by Miss Y. F. TAN (fig. 4,5), Mrs W. WESSEL- BRAND (fig. 7) and Miss H. G. D. ZEWALD (fig. 1, 2, 3, 6, 9, 10, 11, 12). The drawings of N. HALLE (fig. 8, 13), first published in 1967, are reproduced with the kind permission of the editors of Adansonia. Thanks are due to the late Miss G. J. H. AMSHOFF and Mr. D. O. WIJNANDS for their help with the Latin descriptions, to Mrs J. M. VAN MEDENBACH DE ROOY-RONKEL for the correct typing of the manuscript, and to Mr. J. J. Bos for polishing the English text. CYTOLOGY J. C. Arends & F. M. van der Laan Sofar only African members of the Dichapetalaceae have been analyzed for their chromosome numbers. Table 1 presents the 2n numbers of 21 species of Dichapetalum and of Tapura africana. They represent approximately 25% of the African species. The karyotypes have been analyzed in root tip cells of seedlings grown from seeds collected in the field. The vouchers of the mother plants of these seedlings are mentioned in the second column of Table 1. Most of the chromosome number counting in Dichapetalum has been done by GADELLA, whereas some additional counts in this genus and the one for Tapura africana have been done by the present authors (for references see the last column of Table 1). There is one erroneous record of 2n = 20 for D. pallidum (as D. liberiae by MANGENOT & MANGENOT (1962). GADELLA analyzed slides produced by microtome sectioning and haematoxylin staining; we employed the orcein squash staining method as outlined in VAN DER LAAN & ARENDS (1985). A photograph of one of the cells analyzed (Photog. 1) of Dichapetalum affine shows that the length of the somatic chromosomes Agric. Univ. Wageningen Papers 86-3 (1986) 1

5 TABLE 1. Somatic chromosome numbers in Dichapetalaceae Taxa Voucher Provenance Reference species with 2n = 24 Dichapetalum acuminatum De Wild. affine (Planch, ex. Bth.) Bret. altescandens Engl. choristilum Engl. choristilum Engl. congoense Engl. & Ruhl. - published as: D. mekametane Engl. congoense Engl. & Ruhl. - published as: D. mekametane Engl congoense Engl. & Ruhl. cymulosum (Oliv.) Engl. cymulosum (Oliv.) Engl. cymulosum (Oliv.) Engl. cymulosum (Oliv.) Engl. dewildei Bret. dictyospermum Bret. filicaule Bret. heudelotii (Planch, ex Oliv.) Baill. var. heudelotii heudelotii(planch, ex Oliv.) Baill. var. heudelotii heudelotii (Planch, ex Oliv.) Baill. var. longitubulosum (Engl.) Bret. - published as: D. longitubulosum Engl. heudelotii (Planch, ex Oliv.) Baill. var. longitubulosum (Engl.) Bret. - published as: D. longitubulosum Engl. heudelotii (Planch, ex Oliv.) Baill. var. longitubulosum (Engl.) Bret. - published as: D. longitubulosum Engl. heudelotii (Planch, ex Oliv.) Baill. var. longitubulosum (Engl.) Bret. heudelotii (Planch, ex Oliv.) Baill. var. ndongense (Engl.) Bret. - published as: D. martineaui Aubr. & Pellegr. resp. D. ndongense Engl. madagascariense Poir. var. madagascariense - published as: D. brevitubulosum Engl. madagascariense Poir. var. madagascariense - published as: D. guineense (DC.) Keay madagascariense Poir. var. madagascariense madagascariense Poir. var. madagascariense mombuttense Engl. mombuttense Engl. oblongum (Hook.f. ex Bth.) Engl. oblongum (Hook. f.ex Bth.) Engl. oblongum (Hook.f. ex Bth.) Engl. Breteler6445 Gabon Breteler& de Wilde 822 Gabon Bos 3631 Cameroun Bos 4542 Cameroun Bos 4637 Cameroun Bos 5029 Cameroun Bos 5258 Breteler6747 Bos & Breteler 3059 Bos & Breteler 3066 Bos & Breteler 3102 Bos 3249 de Wilde 8269-A de Koning 3690 Breteler 5503 Leeuwenberg Leeuwenberg 7906 Bos & Breteler 3092 Bos 4185 Bos 5002 Bos 5006 Breteler 5216 Bos 5431 Breteler 7294 Leeuwenberg Leeuwenberg Bokdam 3097* Breteler 2113 Breteler 5330 Breteler 5946 Versteeg & Den Outer 713 Cameroun Gabon Cameroun Cameroun Cameroun Cameroun Cameroun Ivory Coast Ivory Coast Gabon Ivory Coast Cameroun Cameroun Cameroun Cameroun Ivory Coast Cameroun Togo Ghana Gabon Zaire Cameroun Ivory Coast Ivory Coast Ivory Coast GADELLA 1977 new record, nob. GADELLA 1970 GADELLA 1970 GADELLA 1970,197 GADELLA 1970 GADELLA 1970 GADELLA 1972 GADELLA 1969 GADELLA 1970 GADELLA 1969 GADELLA 1970 BRETELER 1978 (p. 1 BRETELER 1978 (p. t GADELLA 1972 nob. GADELLA 1977 GADELLA 1970 GADELLA 1970 GADELLA 1972 GADELLA (pers. con GADELLA 1969, BRETELER 1973 GADELLA 1970 GADELLA 1977 verified nob. nob. nob. GADELLA 1977*, verified nob. GADELLA 1969 GADELLA 1969 GADELLA 1970 GADELLA 1977 Agric. Univ. Wageningen Papers 86-3 (1986)

6 TABLE 1. (continued) Taxa Voucher Provenance Reference species with 2n =24 oliganthum Bret. pallidum (Oliv.) Engl. - published as: D. liberiae Engl. & Dinkl. pallidum (Oliv.) Engl. - published as: D. liberiae Engl. & Dinkl. pallidum (Oliv.) Engl. pierrei Pellegr. rudatisii Engl. rudatisii Engl. rudatisii Engl. rudatisii Engl. rudatisii Engl. rudatisii Engl. rudatisii Engl. rudatisii Engl. thollonii Pellegr. toxicarium (G.Don) Baill. toxicarium (G.Don) Baill. toxicarium (G.Don) Baill. zenkeri Engl. zenkeri Engl. species with 2n = 48 Bos 5028 Breteler5282 Breteler5327 Breteler 6032** Breteler 7660 Bos & Breteler 3115 Bos 3180 Bos 3255 Bos 3375 Bos 3412 Bos 3464 Bos 3560 Bos 3645 Breteler 6473 Breteler 5325 Breteler 5342 Breteler 5347 Bokdam 3092 Bos 6943 Cameroun Ivory Coast GADELLA (pers. comm.) GADELLA 1972 Ivory Coast GADELLA 1972 Ivory Coast Gabon Cameroun Cameroun Cameroun Cameroun Cameroun Cameroun Cameroun Cameroun Gabon Ivory Coast Ivory Coast Ivory Coast Zaire Cameroun GADELLA 1972 new record, nob. GADELLA 1970 GADELLA 1970 GADELLA 1970 GADELLA 1970 GADELLA 1970 GADELLA 1970 GADELLA 1970 GADELLA 1970 GADELLA 1972 GADELLA 1969 GADELLA 1970 GADELLA 1970 GADELLA 1972 GADELLA (pers. comm.) Tapura africana Oliv. Leeuwenberg 9910 Cameroun ARENDS1979 species with 2n = 96 Dichapetalum crassifolium Chod. var. crassifolium Oldeman 816 Ghana new record, nob. *: erroneously published with voucher Bokdam 3092 **: published with voucher de Bruijn 1901, a seedling of Breteler 6032 falls within a range of approximately 1 to 2/mi. All chromosomes have primary constrictions in the median region. The size of the chromosomes of all species in the Dichapetalaceae is similar to those shown in Photog. 1. It is obvious that the Dichapetalaceae are characterized by a basic number of x = 12, as 2n numbers of 24, 48 and 96 have been found. Dichapetalum is almost homogeneous according to chromosome number; the octoploid number (2n = 96) of D. crassifolium places this species apart from the remainder of the species, which is partly supported by BRETELER (see p. 17). It remains tobe seen whether Tapurais a polyploid taxon within the Dichapetalaceae as only one species of the genus has been investigated. Regarding the taxonomie position of the Dichapetalaceae there are conflicting Agric. Univ. Wageningen Papers 86-3 (1986)

7 ** i* # f PHOTOG. 1. Somatic metaphase plate in root tip cell of Dichapetalum affine (collection Breteler & de Wilde 822, Gabon), showing 24 metacentric chromosomes. The arrows indicate two, probably homologous chromosomes with an elongated centromere region (8-hydroxyquinolin pretreatment, orcein staining, magn x ). opinions as to whether the family should be accommodated within the Geraniales or within orders such as the Celastrales and Rosales. BRETELER (1973) has pointed out that most evidence supports a place in the Geraniales as conceived by ENGLER. He further suggests that on the basis of comparitive morphology the Dichapetalaceae are more related to the Trigoniaceae than to other families of this order. Earlier BARTH (1896) proposed a close affinity between these two families on anatomical grounds. The (unfortunately approximate) figure of 2n = 20 for Trigonia virens is the only record for the Trigoniaceae (GOLDBLATT 1979). It is clear that members of this family should be investigated further for their chromosome numbers in order to establish whether Trigoniaceae are possibly characterized by x = 12. Some other families of the Geraniales (i.e. the Vochysiaceae, Meliaceae, Malpighiaceae and Polygalaceae) have some morphological characteristics in common with the Dichapetalaceae. From the compilative works in respect of chromosome numbers by FEDOROV (1969), MOORE (1973, 1974, 1977) and GOLDB- LATT (1981, 1984) it can be seen that there is no record for the Vochysiaceae and that none of the remaining families is characterized by a single basic number. However, a very minor fraction of the species of these families have the chromosome numbers n = 12 and/or 2n = 24. In the Meliaceae the number pertains to the genera Heynea (a synonym of Trichilia) and Cedrela, in the Malphighiaceae to Byrsonima, Galphimia and Tryallis and in the Polygalaceae to Polygala. According to BRETELER none of these taxa has a particular affinity with the Dichapetalaceae. 4 Agric. Univ. Wageningen Papers 86-3 (1986)

8 The occurrence of similar chromosome numbers in these different though to some extent related plant groups may be the result of convergent evolution. However the present cytological evidence does not support this or another opinion. As reliable and sufficient information on chromosome numbers of Trigoniaceae and Vochysiaceae in particular is lacking, the basic number of x = 12 of the Dichapetalaceae does not provide cogent support for or against the inclusion of the latter family within the Geraniales. The other orders which have been suggested to include the Dichapetalaceae have also been considered for their chromosome numbers. In the Celastrales there is only a record of 2n = 24 for one species each of'gymnospora and Tripterygium (both Celastraceae). There is conflicting evindence for n = 12 or 13 in Salvador a (Salvador aceaé). For the Rosales there are incidental records of 2n = 24 (or n = 12) for a species of Sedum (Crassulaceae), some species of Chrysosplenium and Castilleja (Saxifragaceae), some species of Lupinus, Vicia, Tamarindus and Xylia (Leguminosae). Within the same order all species of the genera Pittosporum and Sollya ( Pittosporaceae) have 2n = 24. None of the taxa mentioned above has an obvious relationship with the Dichapetalaceae. In conclusion: cytological data do not support a transfer of the Dichapetalaceae from the Geraniales to Celastrales or Rosales, nor do they firmly back the position given to it by BRETELER, as in the Engler system. COMMENTS ON SOME POLLEN GRAINS OF DICHAPETALUM AND TAPURA by W. Punt, Laboratory of Palaeobotany and Palynology Dichapetalumchoristilum Engl. var.louisii Bret. Pollen of this variety certainly belongs to the Dichapetalum choristilum type because of the characteristic straight to slightly concave sides and the slightly decreasing lumina of the reticulum towards the colpi. The colpi, however, are shorter than in the specimens of D. choristilum var. choristilum and, moreover, the size of the grains is distinctly less (Material: /. /. de Wilde c.s. 519). Dichapetalum gassitae Bret. Pollen grains of this species have to beclassified in the Dichapetalum choristilum type because of their coarse reticulum and long colpi. Also other characters like the straight sides, slightly decreasing size of the lumina towards the colpi and the rather large size are identical (Material: Louis c.s. 761). Dichapetalumpotamophilum Bret. Thisspecies is represented by pollen samples of two specimens. Unfortunately the pollen grainsof the two samplesdiffer slightly in their morphological characters. Agric. Univ. Wageningen Papers 86-3 (1986) 5

9 A. Letouzey9806. Pollen of this specimen resembles remarkably that of D. gassitae in ornamentation, shape, and size. The colpi, however, are short. B. Breteler & J. J. de Wilde 604. Pollen of this specimen fits also in the D. choristilum type, butin this specimen the colpi are very long. It differs moreover, from Letouzey 9806 and pollen of D. gassitae by the less obtuse angles and a finer reticulum, a character often found in pollen of the D. angolense type. Although differences occur in the pollen of both specimens they both should beclassified as representatives of the D. choristilum type. PHOTOG. 2. Pollen of Tapura carinata with detail of bi-reticulate ornamentation. For explanation see text. Agric. Univ. Wageningen Papers 86-3 (1986)

10 Tapura carinata Bret. Description of the pollen grains Pollen class: 3-zonocolporate. P/E ratio : Varying from suboblate to oblate. Apertures: Ectoaperture - corpus, long, narrow but broadening towards the equatorial plane, not sunken; margins distinct, straight; ends acute; no costae; distinct fastigium present. Endoaperture - indistinct aperture, probably a large poms of rectangular outline; no costae; ends diffuse. Exine: Sexine about as thick as nexine. Columellae short and indistinct; capita forming a semi-tectate layer. Ornamentation : Bi-reticulate. A coarse outer reticulum with narrow muri and large lumina, angular and varying in outline; lumina uniform in size, not or only scarcely decreasing in size towards the colpi; inner, infra-reticulum very fine. Outlines: Equatorial view - elliptic. Polar view - triangular; sides convex, angles gaping. Measurements: P \i; E \i; P/E ratio Exine ca 2 u; lumina at poles and in mesocolpium up to 2.5 i wide, but usually varying from 1 2 jj. (Material: Pobeguin s.n.). Comments: Pollen of this species is different from all other Tapura pollen. It most resembles that of T. ivorensis, but differs by its size and remarkable ornamentation, absence of costae around the endoaperture and more convex sides in polar view. The most remarkable feature of this pollen typeis the bi-reticulate ornamentation, clearly visible in SEM-micrographs (Photog. 2). This character is certainly not common in pollen of the Angiosperms, but appears regularly in completely different families. It is described from Phyllanthus species (Euphorbiaceae) (BOR, 1979), various Labiatae taxa (NABLI, 1976), and from the Tiliaceous genera Grewia and Sparmannia (PRESTING, STRAKA and FRIEDRICH, 1983). Agric. Univ. Wageningen Papers 86-3 (1986)

11 TAXONOMIC PART DESCRIPTION OF THEFAMILY Dichapetalaceae Bâillon, 1886: 365 (as Dichapetaleae) nom. cons.; Engler, 1896: 345; Engler & Krause, 1931:1; Lawrence, 1951: 549; Wagenitz, 1964:317; Hutchinson, 1973:187; Mabberly, 1978:182. Chailletiaceae Brown, 1818: 23 (as Chailleteae); De Candolle, 1825: 57; Endlicher, 1840: 1104; Bentham & Hooker, 1862:340; Hutchinson 1926: 206; 1959: 150. Trees, shrubs, subshrubs or lianas. Stipules present, caducous or persistent, simple, entire, or variously lobed or divided. Leaves alternate, simple, entire, pinnately nerved, often glandular. Inflorescences axillary, sometimes arranged on leafless axillary or terminal shoots, cymose, distinctly branched to subglobose,sessile to pedunculate, the peduncle free or adnate to the petiole and sometimes to the midrib. Bracts and bracteoles usually small. Flowers small, actinomorphic to zygomorphic, (4-)5-merous, bisexual (Africa) to unisexual; pedicel usually articulate, the upper part (the true pedicel) absent to very distinct. Sepals 5(4), subequal to strongly unequal, imbricate,free or shortly united, rarely forming a tube. Petals 5(4), equal to very unequal, alternating with the sepals, free or nearly so, or, more often, united with the alternating stamens into a very short to distinct tube composed of (8-)10(-ll) elements, entire or bilobed to bicuculate apically. Androecium of 2-5 stamens and 3-0 staminodes, opposite the sepals; anthers bilocular, introrse, opening by longitudinal slits, usually with a distinct connective. Basal staminodes (disc glands, disc lobes, hypogynous glands) 1-5, epipetalous, free or united, variously shaped. Pis?z72-3(4)-merous; ovary superior with two collateral, pendulous anatropous ovules in each locule, raphe ventral, obturator distinct or not; styles 2-3(4), free or nearly so, more often almost completely united with free apical parts. Fruit a drupe with 1-3(4) one-seeded, free pyrenes, deeply lobed or not; exocarp dehiscent or indéhiscent; mesocarp more or less fleshy; endocarp indéhiscent, usually with a distinct apical, and usually partly ventral, suture, pergamentaceous to woody or bony, hairy inside or not. Seeds exalbuminous, rarely with some albumen; testa usually thin, glabrous, rarely hairy; cotyledons usually planoconvex. Germination hypogeal, rarely epigeal, first pair of leaves opposite or alternate. Type genus: Dichapetalum Thouars (syn. Chailletia DC.) An almost circumtropical family of ca 160 species in three genera with a few extensions into the subtropics. Africa (including Madagascar) is most species diverse with 86species in Dichapetalum and 7 species in Tapura. The neotropics are second with 19 species in Dichapetalum, 21 in Tapura, and 9 species in the 8 Agric. Univ. Wageningen Papers 86-3 (1986)

12 endemic genus Stephanopodium, 49 species in all (PRANCE, ). In the Asian tropics and adjacent areas only Dichapetalum is present with 19 species in all (LEENHOUTS ; KOSTERMANS, 1980). Note. The Dichapetalaceae are a very natural group with uncertain affinities (see BRETELER, 1973: 5). The differing and often rather extreme opinions held on the taxonomie position of the family will be maintained when, for instance, the flower tube is either interpreted as a gamopetalous corolla with the stamens adnate to it (PRANCE ; HUTCHINSON, 1969: 65) or as a staminal tube like in Meliaceae, a view expressed by the present author in 1973 and, after studying the genus Tapura even more strongly advocated now. DELIMITATION OFTHE GENERA The three genera of Dichapetalaceae are closely related. BÂILLON (1873: 113) already questioned the proposed genera within Dichapetalaceae and asked himself if it was not possible to consider the family as having but a single genus with 4 subgenera notably Chailletia, Stephanopodium, Dischizolaena, and Tapura. Segregation of Dichapetalum and Stephanopodium particularly would be difficult if the latter occurred in Africa as well. The American species of Dichapetalum have distinctly bilobed and free or nearly free petals, which makes delimitation against Stephanopodium quite easy. In Africa, however, entire petals as well as a distinct staminal tube with sessile or nearly sessile anthers occur, that are common features in Stephanopodium. In species like the African D. insigne Engl., D. melanocladum Bret., and D. montanum Bret, only the bilobed petals distinguish them from Stephanopodium and even this character is not completely reliable as S. estrellense Baill. has slightly bifid petals. As regards the segregation of Tapura and Dichapetalum this is very easy in Africa, but much more difficult when American species of Tapura have to be distinguished on a generic level from African Dichapetalum. The zygomorphy of the Tapura flower may be so slight in American species that in fact only the reduced number of basal staminodes (or disc lobes) establishes a flower to be Tapura and not Dichapetalum. In conclusion it can be stated that disregarding continental separation, viz. Stephanopodium and American Tapura separated from African Dichapetalum, the generic delimitation in Dichapetalaceae is very problematic. KEY TOTHE GENERA IN AFRICA Petals equal; stamens all fertile; basal staminodes (or disc lobes) 5, free or forming a complete ring Dichapetalum Agric. Univ. Wageningen Papers 86-3 (1986) 9

13 Petals unequal, 1-2 distinctly larger; fertile stamens 2-3; basal staminodes (or disc lobes), either free or united, wanting with the large petals... Tapura REVISION OF DICHAPETALUM Short history with some critical remarks. SUBDIVISION OFTHE GENUS All sections that have been distinguished in Dichapetalum are first described by ENGLER. Before ENGLER, DE CANDOLLE, in his Prodromus (1825), also classified the known Dichapetalum species, but he did soin series not in sections. In Engler's first classification (1896: 348) the genus is divided in two sections namely Eudichapetalum (= section Dichapetalum) and Brachystephanium. The second section only contains 2 species, the syntypes D. adnatiflorum and D. kamerunense, the type section all other 72species known at that time. It is interesting to note that D. adnatiflorum and D. kamerunense are synonyms of respectively D. mombul tense and D. oblongum, both species mentioned in Engler's Eudichapetalum under number 30 and 54 respectively. This fact gives an indication how well these sections are separated. VAN TIEGHEM (1903: 230) elevated the section Brachystephanium to generic level choosing Stephanella as its name. As I have lectotypified (1973: 37) this genus with D. kamerunense Engl., the same specific name is chosen here to lectotypify the section Brachystephanium Engler. In 1902 ENGLER introduced a third section with a Latin description but without a name, mentioning D. integripetalum as the only species. In the same publication a fourth section named Tapurina is described with D. longitubulosum as the only species. In 1912 (1912-a) ENGLER gives a complete classification of all the species and presents a key to the 4 sections and the 33 series that are distinguished. The type section contains most species, accommodated in 25 series. The unnamed section of 1902 is now named Rhopalocarpus and remains monotypic. The section Brachystephanium of 1896, now deprived of its two syntypes, which are classified in the next section, has 11 species in 3 series. Of these species 3 were classified in Eudichapetalum in 1896, the other have been described as new since then. The last section, now named Tapurinia instead of Tapurina as in 1902, has 5 series. Two of these accommodate the two syntypes of section Brachystephanium under the names Adnatiflora and Kamerunensia respectively. In doing so ENGLER in fact united the sections Brachystephanium and Tapurina, so 3 sections remain. I think that taking type species out and bringing species from other sections in is once more an illustration of the weakness of the characters that define these sections. 10 Agric. Univ. Wageningen Papers 86-3 (1986)

14 In 1914 ENGLER and KRAUSE enriched the type section by a 26th series named Brauniana based on the new species D. braunii. Engler's classification is more or less adopted by some authors (FRIES 1916, DE WILDEMAN 1919, EXELL 1927, HAUMAN 1955, 1958-b), but has in fact never been accepted completely. Objections against Engler's system. ENGLER'S classification is based on a very superficial investigation of characters as observed on a single or a few specimens per species. Characters most frequently used are the size of the inflorescence in relation to petiole length of the supportingleaf, the peduncle beingfree or adnate to the periole, the adnation between petals and stamens, the petal length in relation to sepal length, the petal shape, the incision of petals and its ratio to the entire petal, and last but not least, the position of the ovary: superior, half inferior, or inferior. Inflorescences and petals vary most times too much to be of great value in specific segregation and can hardly be used for species groupings. The same can be said of the other characters used by ENGLER. As regards the position of the ovary it has already been stipulated (BRETELER 1979: 26) that all species have superior ovaries, only the receptacle may be more or less concave! In conclusion it can be stated that ENGLER'S system is most probably not accepted because it is too artificial, more a classification of specimens than a classification of species. Review and evaluation of main characters and their supposed evolutionary trends. Before presenting any new classification of the genus a review and evaluation of important characters is given. This review will also illustrate the difficulties that have been met with in the attempt to structure the subdivision of the genus. Most vegetative characters vary to such an extent and/or may occur so inpredictably, i.e. not in relation with any other character, that their use in the classification of the Dichapetalum species has been very little. The production of a reddish, slimy exudate in wounded phloem, for example,is onlyseen in D. crassifolium and D. librevillense, two species which do not show any relation as other characters are concerned. The leaf glands, another example, often vary in number and position and within a single specimen, although they seem, in general, to be more numerous and more obvious in the advanced species than in the primitive ones. As regards the inflorescence some evolutionary tendencies may be mentioned. The widely branched, distinctly peduncled, many flowered inflorescence (e.g. D. angolense) is considered to be original and the few flowered, indistinctly branched, sessile or nearly sessile inflorescence derived, with a glomerule or fascicle as the most advanced type. The development of the dichasial branching system into a scorpioid one, as seen in many species, is also considered as an evolutionary trend. These trends are very general and it is difficult to use them Agric. Univ. Wageningen Papers 86-3 (1986) 11

15 in characterizing a species. Therefore inflorescence characters have been of little value in species grouping. The absence of the joint in the pedicel considered to be an advanced character, occurs in three species, twoof them, D. glomeratum and D. pulchrum are certainly closely related, the third, D. trichocephalum stands far apart. The position of the sepals whether being erect or reflexed is of importance in some groups of species but is completely unreliable in others. It is the author's opinon, however, that in general reflexed sepals represent the more primitive condition, notwithstanding the fact that some primitive species as D. bangii and D. lujae have erect sepals. The length of the adnation between petals and stamens, sometimes resulting in a distinct staminal tube which characterizes for example the genus Stephanopodium, varieseven within a single species (e.g. D. heudelotii) and seems to occur indepedently from any other character. The type of indumentum of the ovary is certainly a character of importance when the original or the more advanced status of a species has to be established. An arachnoid indumentum is considered to be primitive and preceeds the glabrous or velutinous condition of the ovary. Glabrous ovaries may result when the arachnoid indumentum disappears (compare D. mundense with D. bellum) and glabrous fruits may be produced as happens in these species. The glabrous condition of the ovary may, however, also lead to velutinous*) fruits as in D. umbellatum. It is likely that in this case the appearance of the velutinous indumentum is initiated by the fertilisation of the ovary. In several species from Madagascar, however, it seems that the change from glabrous to velutinous may be initiated independently from fertilisation (see BRETELER 1979: 59, note under D. leucosid). As both the arachnoid and the glabrous condition may be observed in the closely related D. mundense and D. bellum, so the arachnoid and the velutinous condition can be seen in the two closely related D. pallidum and D. albidum. In the former the ovary becomes velutinous immediately after fertilisation shedding its arachnoid hairs, in the second the ovary is velutinous from the very start, sometimes mixed with a trace of an arachnoid indumentum. Glabrous fruits are not only obtained from species with arachnoid or glabrous ovaries but also from species with velutinous ovaries as happens, for example, in D. integripetalum, D. minutiflorum and D. montanum. So in fact three different stages in the evolution of the ovary indumentum may lead to glabrous fruits: arachnoid, glabrous, and velutinous. It is expected that velutinous ovaries leading to glabrous fruits may evolve into glabrous ovaries and it is asked whether in D. braunii such an evolution has already happened or not. In fig. 1 the evolution of the ovary indumentum, including the hypothetical last stage, and the two fruit conditions which may *) Forconvenience,allindumentumtypes notbeing arachnoidarecomprisedunder the term velutinous, although some ovaries and fruits have in fact an indumentum consisting of weaker and/or curly hairs. 12 Agric. Univ. Wageningen Papers 86-3 (1986)

16 *»«iu.it,,cnotes frxtiis ois-ar<y sèxxius rfatziroie.3,0. xzzu7m-ntxèu.77i 2). Zf-o-7-e-rt- Sö. -fictivx dum. / Arxxcnnvtd \ i a '. xif-fîn-e 25. <Ztzlyosp*.rmum 2. munz/ense. %). n-eu-u-e^oitt. Y Vcûciinvus \ Z. /%. i-cxxu. e» ). tnïexjriptitxliltn.25.77ttnuit/ t>rum, # hxort)xis (fyfn>i-&ei%*ctc)y& Z?; yrtxtcnzz. FIG. I.Relationbetweenovaryandfruit indumentum.forexplanationseetext. result are schematically reproduced, with examples of species. If the hypothetical last stage in the ovary indumentum does not occur, D. braunii becomes an example next to D. bellum. Whether the species have a 2-locular or a 3-locular ovary has not proven to be of any value in delimitating groups of species. Although the bilocular condition is considered to be more advanced than the 3-locular one, it must be observed that the advanced status not only occurs in advanced species like D. montanum and D. oliganthum, but also in more primitive ones like D. bocageanum and D. zenkeri. As regards the fruit shape it is not clear how evolution is directed. The fruits with very distinct lobes sometimes even with almost separate mericarps, a distinct beak, and a thin layer of mesocarp as in D. arachnoideum, D. bangii and D. lujae, are considered to be primitive. Globose fruits or fruits which are distinctly lobed but apically impressed and not or hardly apiculate and have a juicy mesocarp as for example in D. congoense, D. crassifolium, and D. cymulosum are considered to be advanced. However, the fruit shape may be very variable within a single species (e.g. D. madagascariense, D. mundense) and is unknown for quite a few species. Therefore fruit shape has been of little importance in classifying the Dichapetalum species. Agric. Univ. Wageningen Papers 86-3 (1986) 13

17 The combination dehiscent exocarp and Mucuna-like irritating hairs bounds the species D. acuminatum, D. albidum, D. altescandens, D. gilletii, D. pallidum, D. pedicellatum, and probably also D. beilschmiedioides. In D. pallidum, however, specimens occur with indéhiscent fruits that have no irritating hairs and fruits of D. staudtii, a completely different species, often have irritating hairs but are never dehiscent. Dehiscent fruits are considered as primitive. The nature of the endocarp and its inside being glabrous or hairy, is, partly for lack of information, partly for its variability within one species, considered to be of little value in species classification. The same holds for the seedcoat, in which respect D. mossambicense, with densely hairy seeds, has a rather isolated position. The seedling type, which has so far been established for 20 African species, seems to be constant in some groups ofspecies (e.g. 'Adnatae ', p. 16) and variable in others. The type with the first pair of leaves opposite (see BRETELER 1973: 29) is considered to be the primitive one. In Table 2 a summary is given of the parts of the plant that have been discussed and their supposed primitive and advanced character state. The new 'classification '. When publishing the first part of the revision (1973) the present author stated that a new subdivision of the genus would be presented at the completion of it. At present, 13 years later and the revision being completed, a new or a revised subdivision of the genus into sections will not be presented. The review and evaluation of the main characters revealed that clear cut divisions cannot be made because the characters vary mostly independently from one another. A TABLE 2. For explanation see text. Part of plant Character state leaf glands inflorescence pedicel sepals ovary fruit exocarp seedling Primitive few, obscure stalked, many flowered, widely branched, dichasial branching jointed reflexed 3-locular apiculate, distinctly lobed, little mesocarp dehiscent first pair of leaves opposite Advanced many, obvious sessile,few flowered, indistinctly branched, scorpioid branching not jointed erect 2-locular rounded, subglobose, juicy indéhiscent first pair of leaves alternate 14 Agric. Univ. Wageningen Papers 86-3 (1986)

18 FIG. 2. 'Classification' of the 124 species of the genus Dichapetalum. For explanation see text. reticulate pattern of relation between the species is the result. I think that in thisgenusthe evolution isstillin 'full swing'and that therefore most evolutionary trendshave notyetresultedindistinctsections. Some clustersofspecies cancertainlybeachieved, buththe charactersofsuch clusters are weak and not constant which means that they are not acceptable as distinctsections. Nevertheless a few of such clusters will be mentioned and briefly discussed as it isfelt that those clusters may become sections in future development. In relation to the other speciesof the genus these clusters may at present beseen asbulgeson agloberepresenting theentiregenus (see fig. 2). When,eventually, theseclustersbecome more independenti.e.maybe considered assections,they maythenbeseen assatellitesof thelarge centralelement,the typesection. The following 5clusters of species are figured as bulges and indicated by a Agric. Univ. Wageningen Papers 86-3 (1986) 15

19 namebetween ' ',which means thatthis name should not have any status under the Code. - 'Adnatae' characterizedby the peduncle adnate to the petioleof the supporting leaf, with flowers and fruits which resemble each other very much, and all with the sametypeofseedling namelywith oppositefirstleaves. Thefollowing 5speciesbelong to thisentity: D. affine, D. mombuttense, D. pierrei, D. rudatisii, D. toxicarium. Except for the latter which is confined to West Africa, allspecies have a CentralAfrican distribution. - 'Arachnoideae'. The four species of this cluster, D. arachnoideum, D. bangii, D. lujae, D. nyangense,are characterizedby an arachnoiddeciduous indumentum on the vegatative parts. This element together with their primitive fruit type are the main characters of this cluster. The species are almost entirely confined tocentralafrica, only D. bangiioccurssomewhat outside thisarea aswell. - 'Bicornes''characterizedby the sametypeof hairy petals and two-hornedbasal staminodes. Also the inflorescences and fruits (as far as known) resemble. Theonly non-central African species (D. lofense) ofthis clusterissomewhat aberrant. Species: D. lofense, D. reticulatum, D. ruficeps, D. trichocephalum and D. umbellatum. - 'Integrae' consisting of 5speciesfrom Madagascar, allwith entirepetals: D. bojeri, D. leucosia, D. rufum, D. virchowiiand D. vondrozanum. Onthe continententire petalsonlyoccurin D. crassifoliumwhichspecies does notbelong tothis clusteraccordingto other characters. - ' Venenatae'is a clusterofspecieswhich are, with the exceptionof D. integripetalum, known for their poisonous properties. Apart from this its species are relativelylarge-flowered with rather flat and onlyshallowlylobed petals. The following sixspeciesbelongin thiscluster: D. barterifrom West and Central Africa, D. integripetalum from CentralAfrica, D. cymosum, thegifblaar, the mostsouthernspeciesofafrica, and D. braunii, D. ruhlandii and D. stuhlmannii mainlyfrom East Africa. The last cluster isin my opinion the most advanced, the 'Arachnoideae' the most primitive one. The other 3clusters deserve a more or less intermediate position. Theglobecontains themajority of theafrican speciesand all the American and Asiatic ones. Also within this large entity, but on weaker grounds, some speciesgroupings can bemade and some rather isolated speciesmay bedistinguished.theamerican speciesfor instance, are ratheruniform andveryclosely relatedbetween each other.they maybeconsideredassuch a group. The Asiatic species are more diverse and not soeasily placed in a single group.as regards 16 Agric. Univ. Wageningen Papers 86-3 (1986)

20 the remaining African members the following isolated and (some) twins of strongly related species may be distinguished: Isolated species Twins D, chalotii ( D. arenarium D. chlorinum k { D. barbosae D. crassifolium ƒ D. bellum D. dewildei \ D. mundense D. edule f D. choristilum D. eickii I { D. potamophilum D.gassitae f D.gabonense D. parvifolium \ D. thollonii A group of four species with juicy, glabrous fruits is formed by D. bodyi, D. cymulosum, D. dictyospermum, and D.filicaule. Another group with dehiscent exocarp has already been mentioned in the review of characters. Any further distinction of groups seems to me of very little value. Comparison with Punt's classification. Punt's classification of the pollen (1975) was published before the revision of the genus had been completed. This is why some differences are present in the nomenclature between his work and mine. Moreover some new species published after 1975 are not investigated by PUNT. The pollen of the Dichapetalum species is classified in 23 pollen types, 17 for the African species, 5 for the Asiatic ones, and only onefor the American species. In some cases different pollen samples of the same species are clasified in different pollen types. Pollen of D. heudelotii var. ndongense is classified in the D. zenkeri type under its synonym D. sankuruense, and in the D. heudelotii type under D. martineaui and D. ndongense. D. zenkeri pollen is found in three different types namely in the D. angolense type, the D. heudolotii type and the D. zenkeri type. Most clusters of species that are distinguished and figured as bulges in Fig. 2, are found in Punt's classification as pollentypes with more or less the same species. My 'Adnatae', 'Integrae', and ' Venenatae' respectively fit Punt's D. mombuttense type, the D. leucosia type, and the D. barteri type. The 'Arachnoideae' are more narrowly circumscribed than in Punt's D. bangii type, as they contain only 4 of the 6 species of this pollen type. The species of the 'Bicornes', however, are not found in a single pollen type. D. reticulatum and D. umbellatum are classified under the Reticulatum group of the D. gilletii type and D. lofense is found under the Pallidum group of the D. angolense type. The other 2 species of this cluster, namely D. ruficeps and D. trichocephalum, have been published after Punt's publication. Concerning the remainder of the African species the differences between Punt's classification and mine are obvious. The ca 60 species that are left are Agric. Univ. Wageningen Papers 86-3 (1986) 17

21 classified by PUNT in 13different pollen types whileinmy 'classification' they arekeptas an entity. There are,however,alsosome correspondencies. Somespeciesas D. chlorinum and D. parvifoliumwhichhavebeenmentioned as isolated species, arein Punt's work also isolated in a pollen type of their own. Thesame isobserved for some twins ofspecies like D. arenarium + D. barbosae and D. bellum + D. mundense. Moreover when Punt's commentson eachpollen type are taken into consideration, thedifferences appear tobe not so fundamental as mightbeexpected at first sight. PUNTalsodescribed the evolutionary trends in the pollen characters andbased an evolutionary scheme on it.the evolutionarytrendsin the macromorphology that Ihaveindicated areinsufficient inquantity andquality tobuild an evolutionary schemeupon it. Ithink, however, that Punt's schemeis generally quite acceptablefrom my pointofview. KEYSTOTHEAFRICANSPECIES The distribution of the species over thecontinent and Madagascar has led to thedecision not to attempt tomake a singlekey totheafrican species but to produce 4keys, each onefor a separate geographic entity or area, i.e. (see map1) WestAfrica (I), CentralAfrica (II), East andsouth (Sub)Tropical Africa (III), andmadagascarandthe Comores (IV). Thefirst entitycomprises the area ofthe Flora ofwesttropicalafrica excludingthe southern part of the former British CameroonandFernando Po: these territories are added to CentralAfrica, following state borders between Nigeria on theonehand and Cameroun and Equatorial Guinea on the other hand. Theborder between areas IIand III follows almostcompletely the state bordersof the countriesconcernedwith the exception of Angola that hasbeen divided along latitude 13 South into a northern part belonging tocentral Africa and a southern part belonging to area III.Along thislatitudein Angola, in azone of at least afew hundred kilometerswide, not asingle Dichapetalum species hasbeen recorded. Moreover, there are nospecies occurring onboth sidesofthis zone.the border between the entities IIIandIVisobvious. It is undeniable that the borders between these geographic entities are often artificial asregardsspecies distribution, but itisof greatadvantage not to have todeal with a large numberof,for example, CentralAfrican specieswhen trying to identify a specimen from Ghana. Ofthe26 West African species only 6 do not occur in the area of key II, butof the 64species of the latter area only 23occur inthewest African entity, which means that 41 speciesdo not. The overlap between thecentral African area and area III iseven less: 11of the 20speciesof area III do not occurelsewhere.areaiv is almost completely differentfrom the continent, only havingonespecies incommon. Only when itwas inevitable andin order to distinguish afewspecies only, characters ofthe fruits have been used inthekeys, otherwise only characters of flowering material areused. Akey based onfruiting material has notbeen 18 Agric. Univ. Wageningen Papers 86-3 (1986)

22 MAP 1. Key areas: I - West Africa; II - Central Africa; III - East and South (Sub)Tropical Africa; IV - Madagascar and the Comores. attempted for two reasons: there are too many species with fruits still unknown, and in many instances the fruits offer too few distinguishing characters. KEY I: WEST AFRICAN SPECIES 1 a Leaves beneath with a felty, long persistent, white or dirty brown cobweb-like indumentum 2 b Leaves glabrous or glabrescent beneath, if hairy the indumentum of different nature 3 2 a Indumentum of ovary stiff, erect-hairy. Liberia-western Ghana.... D. albidum Agric. Univ. Wageningen Papers 86-3 (1986) 19

23 b Indumentum of ovary like cotton-wool, either white or dirty brown. Guinea-Nigeria D.pallidum 3 a Stipulesdeeplypinnatisect, atleast dentate,veryrarelyentire,usually long persistent. Liberia-Nigeria D.angolense b Stipules entire,veryrarelywith afewteeth,or stipulesearlycaducous 4 4 a Indumentum of ovary like cotton-wool, i.e. with waved or strongly curled hairs, eitherwhite or dirtybrown 5 b Indumentum of ovary consisting of erect or nearly erect, usually straightorslightly curved hairs, either shortandusuallystiff orlonger and moreflexible,orovary glabrous 19 5 a Leaves coriaceous, glabrous or nearly so, (6.5)12-16 x (3)7-8cm; flowers glomerate fasciculate, stamens longer than petals. S.E. Nigeria D.obanense b Above characters notcombined 6 6 a All or nearlyallflowersof aspecimen with2-lobedstyles and2-locular ovaries 7 b All or nearly all flowers of a specimen with 3(4)-lobed styles and 3(4)-locular ovaries 10 7 a Sepalsreflexed, atleast mostsepalsof asingleflower 8 b Sepals erectornearlyso 9 8 a Inflorescence sessile in the leaf axil or nearly so (peduncle at most 0.5cm long).liberia-nigeria D.choristilum b Inflorescence distinctlypedunculate.s.e.nigeria D.zenkeri 9 a Branches and branchlets hollow;peduncle of inflorescence adnate to petiole.s.e.nigeria D.gabonense b Branches and branchlets solid; peduncle free from petiole. Sierra Leone-Nigeria D.oblongum 10(6) a Specimenfromwestof Togo 11 b Specimenfrom Togoorfrom eastof it 13 11a Styledeeplysplit,lobes 1-3 mmlong.liberia-ivory Coast D.choristilum b Styleveryshortlylobed a Sepalsreflexed; fruits glabrous. IvoryCoast... D.dictyospermum b Sepals erect;fruits hairy.guineabissau-ghana.... D.toxicarium 13 a Sepalsreflexed, glabrousoutsideornearly so, the margin puberulous. S.E.Nigeria D.mundense b Sepalsreflexed or not,completelyhairyoutside a Branches andbranchletshollow.s.e.nigeria D.gabonense b Branches andbranchletssolid a Flowers arranged in an indistinctly branched (rarely once distinctly so)head or subumbel,thestalkoftheinflorescence shortorlong,free oradnate to petiole 16 b Flowers arranged in distinctly branched inflorescences, itspeduncle, if present,freefrom petiole ornot Agric. Univ. Wageningen Papers 86-3 (1986)

24 16 a Peduncle short, (4)7-11mm long, at least partly adnate to petiole; branchlets with a brown, short-hairy, somewhat papillate-like indumentum.s.e.nigeria D.rudatisii b Peduncle slender, (0.5) (4.5)cm long, free from petiole; indumentumof branchletsdifferent. S.E.Nigeria D.tomentosum 17 a Inflorescence sessile in the leaf axil or nearly so (peduncle at most 0.5cm long);styledeeplysplit, lobes1-3mm long.s.e. Nigeria.... D.choristilum b Inflorescence usuallydistinctlypedunculate;styleveryshortlylobed a Sepalsreflexed. S.E.Nigeria D.fructuosum b Sepals erect ornearly so.s.e.nigeria D.unguiculatum 19(4) a Pistil glabrous, atmostwith averyfew,sparse hairs (ovarysoonvelutinousafter fertilisation).nigeria D.umbellatum b Pistil, atleastthe ovary,ditinctlyhairy a Petallobesdenselyhairyoutside. Nigeria D.reticulatum b Petallobesglabrous outside, at most with afew hairs orpetalsentire ornearlyso 21 21a Upper part of pedicel more of less as long as the reflexed sepals, at leastas longas thelower part.ivorycoast-nigeria... D.parvifolium b Upper part ofpedicel shorter than thelower part,whenequalatmost half aslong as thesepals, or0, orflowerssessileornearlyso a Erectshrubortree 23 b Lianescent shruborliana a Sepalsreflexed, atleast mostsepalsof asingleflower. S.E.Nigeria.. D.obanense b Sepals erectornearlyso a Sepals 4-6(7)mm long;petals retuse to 1 mm split, lobes rather flat. IvoryCoast-Nigeria D.barteri b Sepals (1) (3)mm long;petals distinctlybilobed,lobes concave. GuineaBissau-Nigeria D.madagascariense 25 a Petals entire to emacginate,ifshallowlylobedlobes notconcave, with stamens unitedinto a mm long tube.liberia-nigeria D.crassifolium b Petals lobed, if united with filaments into a distinct tube, then lobes concave a Fruits glabrous; thin liana or lianescent shrub usually not more than 1 cm diam.; midrib of leaves prominent above. Ivory Coast-western Ghana D. filicaule b Fruitshairy; other charactersusually notassociated a Petals thin,fragile, 1.7-3mm long;stylealmosthairy to the top.liberia-western IvoryCoast D.lofense b Petalsdifferent, usuallylonger;styleif hairyonlysoinlowerhalf. 28 Agric. Univ. Wageningen Papers 86-3 (1986) 21

25 28 a Leaves coriaceous, glabrous ornearly so;flowers glomerate-fasciculate; sepalsreflexed atleast most sepalsof asingleflower; lower part ofstyle hairy. S.E.Nigeria D.obanense b Above charactersnotassociated a Petals and stamens slightly spreading at base, upper parts usually curvedinwards; petallobesflat or nearlyso.ivorycoast, Nigeria... D.dewevrei b Petals andstamens erect,withoutinflexed upper parts; petal lobes concave a Stylehairyinlowerhalf;inflorescenceusually distinctlystalked and/or branched.guinea Bissau-Nigeria D.madagascariense b Style glabrous inlower half, rarely notso;inflorescence a glomerule or fascicle, usually sessile andindistinctly branched (some inflorescenceof aspecimenmaybevery shortlystalked and/or branched) a Leaf acumen with a distinct mucro, if not theleaf blade beneath at least with distinct, large ( mm diam.) glands along midrib, especially near base and/or on theacumen or top (sometimes hidden by indumentum). GuineaBissau-Nigeria D.heudelotii b Leafacumen usually broad, obtuse,without adistinct mucro;glands usuallysmallerthanabove.ivorycoast,nigeria D.staudtii KEY II: CENTRALAFRICANSPECIES 1 a Pistil glabrous, at most with averyfewsparse hairs (ovarymaybecome denselyhairysoonafter fertilisation) 2 b Pistil, at leasttheovary, hairy 3 2 a Leaves and branchlets glabrousornearly so;fruits glabrous. Gabon, Congo D.bellum b Leaves andbranchlets hairy;fruitsvelutinous.western CentralAfrica D.umbellatum 3 a Indumentumof the ovarylike cotton-wool,i.e. withwaved orstrongly curled hairs, eitherwhite ordirty brown 4 b Indumentum ofthe ovary consisting oferect ornearly erect, usually straight orslightlycurved hairs,either short and usually stiff or long andusuallyflexible., 39 4 a Leaves, atleastwhen young,with long arachnoid (cobweb) hairsbeneath,eitherforming alooseweb or aclosefelt 5 b Leaves withoutarachnoidhairsbeneath 9 5 a Stipules,atleast some,palmately divided into 2-4(5) parts,the parts leavingseparateor quitenearlyseparatescars 6 b Stipules entire 8 6 a Sepalsreflexed. Gabon D.nyangense b Sepals erectofnearlyso 7 22 Agric. Univ. Wageningen Papers 86-3 (1986)

26 7 a Youngleaves above andoften alsothe branchletswith a cobweb indumentum.whole area D.lujae b Young leavesabove and branchlets lacking thecobweb hairs.whole area D.bangii 8 a Arachnoid hairs on leaves beneath forming a persistent close felt. Western CentralAfrica D.pallidum b Arachnoid hairsonleaves beneathdeciduous. WesternCentral Africa D.arachnoideum 9 a Stipules pinnatisect,atleast dentate,usuallylong persistent b Stipules entire, orstipules earlycaducous a Styles predominantly 3(4)-lobed,ovary3(4)-locular.Whole area... D.angolaise b Styles predominantly2-lobed, ovary2-locular.sao Tomé, Principe.. D.bocageanum 11 a Fullydevelopedleaves with strongly and rather abruptlyrevolute margins at theextreme base, more orlesscoveringoneor a pair of rather largeglandsoneachsideof the midrib.whole area.. D.congoense b Baseofleaf-blade notasabove a Sepals rathersharplyreflexed infully developed flowers 13 b Sepals not sharply reflexed in fully developed flowers, at least never most sepals of a single flower but sepals usually erect or slightly or looselyspreading a All ornearlyallflowersof aspecimenwith2-lobedstyles and2-locular ovaries 14 b All ornearlyallflowersof aspecimenwith3-lobedstyles and3-locular ovaries a Inflorescence sessile in the leaf axil or nearly so; petals dark-brown toblackindryspecimens. Whole area D.choristilum b Inflorescence usually distinctly pedunculate; petals yellowish to pale brownin dryspecimens. Whole area D.zenkeri 15 a Stylesdeeply split,lobes1-3mmlong 16 b Stylesvery shortlylobed a Leaves elliptic to obovate, acuminate at top, rarely not so, usually tapering to a cuneate ornarrowlyroundedbase. Whole area D.choristilum b Leaves ovate to elliptic, top rounded, rarely acuminate and then not distinctlyso, broadly rounded to obtuse or cordate at base. Cameroun, Gabon D.potamophilum 17 a Flowerswithan up to 15mmlong slender pedicelwith 3-6 mmlong upper part, and aggregated on a knob-like or very shortly branched base; peduncle, at most as long as the petiole, usually adnate to it. EasternZaire D.ugandense b Flowers notarranged asabove and/orpedicelsdifferent a Sepals glabrous outside or nearly so, the margin puberulous. Whole area D.mundense Agric. Univ. Wageningen Papers 86-3 (1986) 23

27 b Sepals completelyhairyoutside a Inflorescence a slender-stalked usually indistinctly branched (sometimes distinctlybranched apically, i.e. justbelow the head)subglobose head;leaveswithrather long, moreorlessappressed (butusuallyperpendicular to the nerves) hairs on midrib and main lateral nerves beneath.s.e. Gabon,Congo, westernzaire D.gillerii b Inflorescence notas above;leaves glabrous ornearlyso,if hairy hairs appressed on mainnerves beneath a Petiole (5)7-10(12)mm long; branchesusuallyhollow. Gabon... D.beilschmiedioides b Petiole (1)2-5(10)mmlong; branchessolid 21 21a Leaf-top distinctlyglandular and tomentose beneath.whole area... D.librevillense b Leaf-top notas above a Fruitsbeaked toapiculate,indéhiscent. Whole area.. D.fructuosum b Fruits rounded toobtuseat top, exocarpdehiscent a Inflorescence a long-stalked,shortly butdistinctly branched, rusty peberulous cyme; flowerswith long slender pedicels, aggregated on a fewknob-likebases. Congo, Zaire D.pedicellatum b Inflorescence long-stalked or not, usually long-branched, pale-hairy; flowers notaggregated a Leaf-blade (2)3-4(5) times as long as wide, (4)6-10(14) x (1)2-4(5)cm; exocarp notreticulatelyfissured. Gabon,Congo, Zaire D.acuminatum b Leaf-blade timesaslong as wide, (6)8-12(16) x (2)3-4(6)cm; exocarpreticulatelyfissured. Cameroun, Gabon... D.altescandens 25(12) a Inflorescence a slender-stalked subglobose head, indistinctlybranched or once distinctly so just below the head, peduncle free from petiole 26 b Inflorescence either very distinctly and more than once branched or notslender stalked or both, orpeduncle adnate topetiole or inflorescencesessile a Stamens distinctly longer than petals; fruits with dehiscent exocarp andstinging caducous hairs.s.e. Gabon, Congo,western Zaire... D.gillerii b Stamens as long as the petals or nearly so;fruitsindéhiscent, velutinous to tomentose. Western CentralAfrica D.tomentosum 27 a Peduncleof inflorescence adnate to petiole (the bladeof the supporting leafmaybestronglyreduced) 28 b Peduncle of inflorescence free from petiole or inflorescence sessile or nearlyso a Branches andbranchletshollow, atleastbetween thenodes b Branches andbranchletssolid a Fresh leaves papery, usually with domatia in some lateral nerve axils beneath; glands on lower surface less than 0.2mm diam.; petals 24 Agric. Univ. Wageningen Papers 86-3 (1986)

28 3-4.5mm long, usually slightly shorter than the stamens; pistil 2-3-merous, 3-6 mmlong.westerncentralafrica.. D.gabonense b Fresh leaves coriaceous, without domatia; glands on lower surface mm diam.; petals mm long, usually slightly longer than stamens; pistil 3-merous, mm long. Gabon, Congo, western Zaire, northern Angola (Cabinda) D.thollonii 30 a Fruitsglabrous 31 b Fruitshairy (hairsmaybeveryshort!) 32 31a Fruitslenticellate, beaked.whole area D.mombuttense b Fruits smooth, at most with afew lenticells, obtuse to apiculate apically. Cameroun,Equatorial Guinea, Gabon D.affine 32 a Branchletsglabrous,if puberuloussoonglabrescent. Gabon D.pierrei b Branchlets with a brown,powdery-like indumentum,oftenmixed with some normally developed hairs, not soon glabrescent. Cameroun, northern Gabon D.rudatisii 33(27) a Style deeplysplit,lobes 1-3 mm long.whole area... D.choristilum b Styleveryshortlylobed a Petalsand stamens 2-3 mm long. Cameroun, Gabon D.tetrastachyum b Petals andstamens (3.5)4-6mm long a Style2-lobed,ovary2-locular (afew3-merous pistilsmaybe present). Cameroun, Gabon D.oblongum b Style3(4)-lobed,ovary3(4)-locular (afew2-merouspistils maybepresent) a Petalstomentoseoutsideandonkeel inside. Gabon.... D.pierrei b Petals glabrousor with averyfewhairs justbelowsplitoutside a Sepals mm long. Cameroun, Gabon D.witianum b Sepals 2.5-4mm long a Petals andstamens basally unitedinto a 2-3mm long tube. Angola.. D.sumbense b Petalsfreefrom stamensornearlyso. Whole area. D.unguiculatum 39(3) a Leaf-blade atbase with 2very distinctpearshapedpouches. Gabon.. D.gassitae b Leaf-blade atbase notas above a Stipulesdeeplylobed, atleast dentate,usuallylong persistent b Stipules entireorstipulesearly caducous a Flowersarranged in a stalked subglobosehead; petallobes hairy outside. Gabon D.trichocephalum b Flowersglomerate; petallobes glabrousoutside a Stipules ovate to elliptic in outline, deeply pinnatisect with almost threadlikelobes; pistil 3-4.5mm long. Central Zaïre... D.germainii b Stipules triangular in outline, dentate; pistil mm long. Cameroun, Gabon,Congo D.pulchrum 43 a Flowers predominantlywith2-lobedstyles 44 Agric. Univ. Wageningen Papers 86-3 (1986) 25

29 b Flowers predominantly with3(4)-lobed styles a Petals geniculate below the spreading lobes; stamens at most half as long as the petals. CentralZaire, Gabon (?) D.staminellatum b Petalsgeniculateor not,stamensrelativelylonger a Petalsand stamens atbaseunited into adistinct 0.5-1mmlong tube, the stamens usually distinctly shorter than petals, the anthers sometimes almostsessileon tube 46 b Petalsand stamensfree from eachother ornearlyso a Branchlets sparsely hairy when young, soon glabrescent. Cameroun, Gabon,easternZaïre D.montanum b Branchlets densely villous-tomentose, the indumentum long persistent.westerncentralafrica D.insigne 47 a Branchlets glabrous tosparsely puberulous,verysoonglabrescent;inflorescence an up to 7-flowered cymule; fruits glabrous. Cameroun, Gabon D.minutiflorum b Branchlets more or lessdensely hairy, glabrescent or not, if glabrous ornearlyso theinflorescence different; fruits hairy a Petiole 2-11(13)mmlong;sepalserect;petals suberect; stylehairyin lowerhalf. Whole area D.madagascariense b Petiole 0-3(5)mm long; sepalserect to reflexed; upper part of petals often curved;styleusuallyglabrous a Inflorescence up to ca 25-flowered; pedicel usually slender, (1.5)3-5(10)mm long; leaves usually with a cordate to subcordate base.westerncentralafrica D.dewevrei b Inflorescence upto4-flowered;pedicelstiff, up to 1.5mmlong; leaves rounded to cuneateatbase.s.e. Cameroun D.oliganthum 50(43) a Hispid liana or lianescent shrub with hollow branchlets; stipules, bracts and bracteoles slender, usually curved; flowers large (up to 10mmlong) arrangedin up to20-floweredinflorescences;petals hairy inside,usually black when dry. Western Central Africa south of Cameroun D.chalotii b Above characters notassociated 51 51a Upper part ofpedicelcaaslongas thereflexed sepals, at leastaslong as thelower part.whole area D.parvifolium b Upper part ofpedicelshorter than thelower part,whenequal atmost half aslongasthesepals,or 0,orflowers sessile ornearly so,or joint inpedicelabsent a Thinhispid liana orlianescent shrubwith solid branchlets and sessile inflorescence of 1-3 flowers only; sepals spreading; petals glabrous inside, lobes flat, spreading; petals and stamens distinctly united at base. N.W. Gabon D.geminostellatum b Above characters notassociated a Petalsdistinctlylobed, thelobes completelyhairyoutside 54 b Petalsdistinctlylobedor not,thelobes glabrousoutside or apical part glabrousoutside Agric. Univ. Wageningen Papers 86-3 (1986)

30 54 a Inflorescence a subumbel, flowers distinctly stalked; petiole (4)6-9(15)mm long, theblade timesas long aswide. Cameroun D.reticulatum b Inflorescence a subglobose head, flowers sessile or nearly so; petiole (2)4-7mm long, the blade times aslongas wide. Gabon... D.ruficeps 55 a Bracts and bracteoles 3-5(6)mm long, ca as long as or longer than the stoutpedicels; youngvegetative parts barbateby afurry orhirsute indumentum; leaves usually cordate at base, above with prominent midribandmainlateralnerves. Cameroun, Gabon... D.barbatum b Above characters notassociated a Pedicels without a joint;flowerspersistent 57 b Pedicels jointed;flowers,atleast partly,caducous 58 57a Stipules l-5(6)mm long; leaves (4)10-16(24) x (1.5)3.5-6(12)cm with 8-12(16) pairs of main lateral nerves; sepals mm long; petals (3)4-6mm long. Whole area D.glomeratum b Stipules (4)8-17(22)mm long; leaves (15)20-35(42) x (6)7-11(14)cm with (10)11-14(16) pairs of main lateral nerves; sepals 3-5mm long;petals 4.5-8mm long. Western CentralAfrica D.pulchrum 58 a Fruitsglabrous,ifsparsely puberulousthen3(4)-lobedand 1-seeded. 59 b Fruitshairy,ifsparselyso notas above a Leaves glabrousornearly so,leaf marginrevoluteattheextremebase and more or less covering large glands beneath; fruits 3(4)-lobed, 1-seeded. Cameroun, Gabon, northernangola (Cabinda) D.integripetalum b Leavesdifferent; fruitswhen lobedmore than 1-seeded a Leaves drying greenish to pale-brown, (8)11-15(18) x 3.5-6(9)cm, midrib glabrous above ornearlyso;petals (2.5) mm long;pistil (3.5)4-5.5mm long. Gabon, Congo,western Zaïre D.bodyi b Leaves drying dark-brown to black, (3)8-12(15) x (1)3-4(5)cm,midrib always distinctly hairy above; petals 2.5-3(3.5)mm long; pistil 3-4(4.5)mm long.s.e.cameroun D.cymulosum 61a Brancheswhenfreshlycut exuding a reddishslime turning dark-brown to black indrying;leavesusually coriaceous, glabrous or rather soon glabrescent;petalsentire toemarginate, at basedistinctly united with stamens into a mmlong tube.whole area.. D.crassifolium b Above characters notassociated a Flowersca 3mmlong;stamensdistinctly shorter thanpetalsandunited with them, the anthers almostsessile on tube. Cameroun, Gabon. D.melanocladum b Petals and stamens free or nearly free from each other, if distinctly united theflowersmuchlonger a Flowersinglomerules or fascicles (someinflorescences of aspecimen Agric. Univ. Wageningen Papers 86-3 (1986) 27

31 maybeveryshortlystalked and/or branched) 64 b Flowers in distinctly branched and/ordistinctly stalked inflorescences a Petals and stamens slightly spreading at base, upper parts usually curvedinwards;petallobes flat ornearlyso. WesternCentralAfrica. D.dewevrei b Petals and stamens erect without incurved upper parts; petal lobes concave a Style hairyinlowerhalf.whole area D.madagascariense b Style glabrousinlower halfornearlyso a Leaf acumen with a distinct mucro, if not the leaf blade beneath at least with distinct, large ( mm diam.) glands along midrib, especially near base and/or on the acumen or top (sometimes hidden by indumentum). Whole area D.heudelotii b Leaf acumen usually rounded or obtuse, without a distinct mucro; glands usuallysmaller.whole area D.staudtii 67 a Style hairyinlowerhalf; sepals (1) (3)mmlong. Whole area.. D.madagascariense b Style glabrous in lower half, or with a very few hairs only; sepals (2.5)3-6(7)mm long a Petal lobes distinct, concave, not incurved to inflexed. Cameroun, easternzaïre D.dewildei b Petal lobes flat or nearly so, often incurved or folded one over the other,orpetals emarginate a Lianescent shrub or liana; fruit obovoid, velutinous, firmly walled, endocarp filmy. Angola,Zaire D.ruhlandii b Erect shrub or tree; fruits subellipsoid to subglobose, velutinous to tomentellous,usually echinate to tuberculate or rugose-wrinkled, rarely smooth,wallitself ratherthin, endocarppergamentaceous, fibrous a Petals mm long; stamens (4.5)5-6.5(7)mm long; pistil 6-7.5mm long. Western CentralAfrica D.barteri b Petals andstamens 3-4 mmlong; pistil mm long. EasternZaire D.stuhlmannii KEYIII: EASTAFRICANANDSOUTH (SUB)TROPICALAFRICANSPECIES 1 a Pistil glabrous;fruits glabrous; plants glabrous or nearlyso.s.e. Tanzania D.braunii b Pistil, at least the ovary, hairy;fruitshairy at least partly; plants usually hairy, atleastyoung parts 2 2 a Stipules, at least someof them,pinnately orpalmately lobed, at least dentate, or completely palmately or pinnately divided, usually long persistent 3 b Stipulessimple,entireornearlyso, orearly caducous 6 28 Agric. Univ. Wageningen Papers 86-3 (1986)

32 3 a Indumentum of ovary short-erect-hairy (velutinous), if hairs longer andmoreflexible thennotwaved.s.e. Tanzania, N.E. Mozambique. D.edule b Indumentum of ovary like cotton-wool, i.e. with waved or strongly curledhairs, eitherwhite ordirtybrown 4 4 a Flowerswitherectorslightlyspreadingsepals. Zambia.. D.bangii b Flowers withreflexed sepals 5 5 a Petiole (3)4-10(15)mm long, (6)12-20(30)mm longwhen unitedwith stalkofinflorescence; leaves (7)12-25(53) x (2.5)5-11(21)cm. Western Uganda D.angolense b Petiole 1-3(10)mm long,alsowhenunitedwithstalkofinflorescence; leaves 5-15(18) x (2)3-7(10)cm.Coastal area ofs.kenya, Tanzania, and N. Mozambique D.mossambicense 6 a Indumentum of ovary short-erect-hairy (velutinous), if hairs longer andmoreflexible then notwaved 7 b Indumentum of ovary like cotton-wool, i.e. with waved or strongly curvedhairs, eitherwhite ordirtybrown 13 7 a Petals entire,with stamensunited into a mmlongtube;stems and thick branches exuding a pale reddish slime, dark-red to black when dry. WesternTanzania D.crassifolium b Petals lobed or emarginate, not or very shortly united with stamens; stemsandthickbranches withoutexudate 8 8 a Petiole 2-4 mm long;leaves usually cordate at base,sometimes obtuse, both sides with rather long, subappressed hairs;inflorescence sessile ornearly so.s.e.tanzania, N.E.Mozambique D.edule b Petiole usually longer, if not leaves cuneate at base; other characters notassociated 9 9 a A rhizomatous suffrutex, annually sprouting from a woody, largely subterranean base. N.E. Zambia, Angola, S.W. Africa, Zimbabwe, Botswana,SouthAfrica D.cymosum b Trees, shrubsorlianas 10 10a Sepals (1) (3)mm long; petals (0.2)l-2(3)mm split, the lobes concave 11 b Sepals 3-5(6)mm long; petals emarginate or up to 1 mm split, lobes flat,often inflexed 12 11a Petals (1.7)2.5-4(5.5)mm long, (0.5)l-2(3)mm split; stamens and pistil atleastas longas the petals,usuallylonger, (2)2.5-6(7)mm long and (1.7)2.5-6(9)mm long respectively. Uganda, Kenya, Tanzania, Zimbabwe, Mozambique. D.madagascariense var.madagascariense b Petals mm long, mm split; the stamens almost as long asthe petals, mmlong,thepistildistinctly shorter, mm long. Kenya, Tanzania D.madagascariense var.brevisrylum 12 a Lianescent shrub or liana (rarely tree-like?); leaves, at least when young,sparsely hairy, rathersoon glabrous or nearly so;inflorescences often grouped on leafless axillary shoots or leafless parts of such Agric. Univ. Wageningen Papers 86-3 (1986) 29

33 shoots, distinctly 3-4 times branched, usually many flowered;fruits obovoid, velutinous, firmlywalled, endocarp filmy not fibrous. Kenya, Tanzania D. ruhlandii b Shrub or tree; leaves, at least when young, rather densely hairy, especially so beneath, longpersistent on midrib above; inflorescences usually singlein theleaf axil,more compact, indistinctly branched or basally once distinctly so, up to ca40-flowered; fruits subellipsoid to subglobose, velutinous to tomentellous, usually echinate to rugosewrinkled,rarelysmooth,wallitself rather thin, endocarp pergamentaceous, fibrous, finely striateinside. Tanzania,Mozambique D.stuhlmannii 13(6) a Flowers arranged in asessile to subsessile subumbellate inflorescence with slender, up to 15mm long, pedicels, with 3-6mm long upper parts. Uganda,W. Tanzania D.ugandense b Flowers notarranged as above; pedicelsmuchshorter a Inflorescence a glomerule or fascicle, up to 5-flowered; flowers with mmlong petals and 7-8 mm long stamens;stipules rather long persistent, (3)7-14(17) x 1-3mm.S.E.Tanzania. D.macrocarpum b Inflorescences different, at least more flowered; flowerssmaller; stipulesusuallysmaller a All or nearly all flowers of a specimen with 2-locular ovaries and 2-lobedstyles.S.W.Kenya, N.E.Tanzania D.zenkeri b All or nearly all flowers of a specimen with 3-locular ovaries and 3-lobedstyles a Petals and stamens mm long, the pistil 1-2mm long. S.E. Kenya, N.E.Tanzania D.eickii b Petals, stamens, and/orpistillonger a Petalsinsidehairybelowthelobes 18 b Petals inside glabrousbelow the lobes a Leavesusuallypapery,hairy above.s. Angola, Zambia, S.W. Africa, Zimbabwe D.rhodesicum b Leaves coriaceous, if hairy above then only so on the main nerves. Kenya, Tanzania, Mozambique a Leavesnevercompletely glabrous beneath but usually shortlypubescent allover thelower surface, margins usually strongly revolute; peduncle (4)6-11(13)mmlong; petals mmlong,less than 1 mm split.s.e.kenya, Tanzania D.arenarium b Leaves glabrous beneathorsparsely, shortly appressed-haired on midriband main lateral nerves,margins not or only slightly revolute; peduncle 2-5 mm long;petals (4) (6)mm long, mm split. S.E. Tanzania, Mozambique D.barbosae 20 a Inflorescence very shortly stalked (1-3mm), branched into a few to several, many flowered,spreading scorpioid arms; sepals mm long.s.e.kenya D.fadenii 30 Agric. Univ. Wageningen Papers 86-3 (1986)

34 b Inflorescence a normal compound dichasium, usually with much longer peduncle; sepals mm long 21 21a Plants densely hairy all over, leaves not glabrescent. S. Angola, Zambia, S.W. Africa, Zimbabwe D.rhodesicum b Plants glabrous or glabrescent, if densely hairy only so on young parts. Kenya, Mozambique a Branchlets grey-hairy; leaves (2)4-8 x 1-3(4) cm; peduncle of inflorescence (2)4-5(9) mm long. Mozambique D.deflexum b Branchlets brown-hairy; leaves (5)9-14(20) x 3-6(8) cm; peduncle of inflorescence (0)5-20(50) mm long. S.E. Kenya... D.frucruosum KEY IV: SPECIES OF MADAGASCAR AND THE COMORES 1 a Petals distinctly bilobed 2 b Petals entire, at most emarginate 3 2 a Pedicel above articulation mm long; sepals reflexed; ovary shortly velutinous D.chlorinum b Pedicel above articulation 0-0.5(1) mm long; sepals erect of nearly so; ovary soft-hairy D.madagascariense 3 a Leaves very unequal-sided at base D.virchowii b Leaves not or only slightly unequal sided at base 4 4 a Indumentum of ovary like cotton wool 5 b Indumentum of ovary absent or ovary very shortly velutinous a Sepals usually reflexed in mature flowers; petals (2) mm long; pistil (2.5) 3-4 mm long D.bojeri b Sepals erect in mature flowers; petals mm long; pistil mm long D. vondrozanum 6 a Flowers mm long D.rufum b Flowers 5-7 mm long D.leucosia ADDITIONS, CORRECTIONS AND EMENDATIONS TOTHE PREVIOUS PARTS III-VIII PUBLISHED IN , IN ALPHABETICAL ORDER. D. albidum Chev.ex Pellegr. This species has in 1974 been collected near Mampong-Akwapim in Ghana by A. A. ENTI under number F. E D. barbosae Torre As could be expected, this species so far only known from N.E. Mozambique has recently been collected in the Lindi District in S.E. Tanzania (coll.: Magogo & Rose Innés RRI397 (K)). Agric. Univ. Wageningen Papers 86-3 (1986) 31

35 D.brauniiEngl. &Krause When treating thisspecies (BRETELER,1973:103) maturefruitswere unknown. Recentmaterial from thelindi DistrictinS.E.Tanzania (coll.: Magogo & Rose Innés RRI383 (K)) has maturefruits whicharedescribedbelow. Fruits orange, 1-seeded, glabrous, obovoid-ellipsoid, beaked, 4.5cm long (beakof 1.5cminclusive), cm diam.; mesocarp juicy (?); endocarppergamentaceous, glabrous and finely striateinside.seedovoid,2.5 x cm, glabrous, testa thin. D. brazzaepellegr. = D.librevillensePellegr.See p.38. D.choristilumEngl. Recentcollectionsfrom Gabon included smallflowered D. choristilum specimens. As has been done before in other species (D. crassifolium, D. dewevrei, D. heudelotii, D. lujae, D. madagascariense) these specimens are classified in a separatevariety.thetwo varieties canbekeyed outasfollows: Petals (1.5)2.5-4mm long, mm split; stamens and pistil (2)2.5-4(5)mm and 2-4.5mmlongrespectively... D.choristilum var.choristilum Petals mm long,0.5mm split; stamens and pistil mmand 1 mm longrespectively D.choristilum var.louisii D.choristilum Engl. var.louisiibret. var.nov. Map 2 Haec varietas a D. choristilum var. choristilumdiffert floribus parvisstaminibus petalisque maximaliteraequilongiset pistillodistincte breviore. Type: Gabon,ChailluMassif, 15kmMimongo-Mbigou Rd, Naguila Mt,alt. 760 m, J. J. de Wilde, Arenas, Bouman, Karper and Louis 519,fl. (holotype: WAG); 20km E. of Mimongo, Songou Mt., alt m, Louis, Breteler and de Bruijn 1002, fr. (paratype: WAG). Congo, near Kouyi, Sita 4120 (paratype: WAG). Besides the small flowers which are in fact the only sound character to distinguish thisnewvariety, thetypematerial hasshorter petiolesand larger fruits than previously seeninthisspecies.consequently thespeciesdescription (BRE TELER,1978:12)has tobeemendedasfollows: Change the length of the petiole into: 1-11(18)mm, the length of the petal incision into mm, the stamen and pistil length into 1-4(5)mm and mm respectively, the fruit length into cm; add after fruit diam.: Beaked or not, sometimes stipitate;replacein thedescription of thefruit indumentum 'villous' by 'velutinous'; change the seed measurements into 30 x 15mm. 32 Agric. Univ. Wageningen Papers 86-3 (1986)

36 lh >. c ^^a"*.. ~\,' ^ ^ II 'T; 20 ^ ' ^ 11 1 r^ri / \ (^ Ai (i ' ' \ 20 MAP 2. O. choristilum var. louisii. D.fadeniiBret, sp.nov. Fig. 3Map 3 Liana. Rami cortice brunneo vel nigro, lenticellati. Ramuli tomentelli, glabrescentia. Folia elliptica, subtus persistens puberulo-tomentelli, costis et nervis lateralibus principalibus utrinque 6-8 impressis.inflorescentiae basipauci - vel pluribrachiatae, brachiis patentibus, multifloris, scorpioideis. Sepala suberecta vel patentia, mmlonga.petala mmlonga, 1 mm fissa, glabra. Staminapetalisaequilonga.Pistillum 3-merum, ovariovilloso. Endocarpium intus villosum. Type: Kenya, Kilifi District, Chasimba, R. B. & A. J. Faden, J. B. Gillett & N. Gachathi 77/416 (holotype: WAG; isotype: K, US; other duplicates (not seen)inb,br,c,ea, F, MO, P, PRE, UPS). Diagnostic characters. Woody climberwithdark-brown to black,lenticellatebranchesand tomentellous,glabrescent branchlets.leaves elliptic, persistently short-hairy beneath, midrib and the 6-8 pairs of main lateral nerves impressed above.inflorescence branched near base into afew toseveral, many flowered, spreading, scorpioid arms. Sepals suberect to spreading, mm long. Petals 3-3.5mm long, 1 mmsplit, glabrous.stamensas long as the petals. Pistil3-merous,ovary villous. Endocarplong hairy inside. Description. Woody climber. Branches dark-brown to black, lenticellate. Branchlets tomentellous, glabrescent. Stipules rather long persistent, entire, narrowly triangular to subulate, (1)2-5(6)mm long,hairy as branchlets. Leaves: petiole subterete, (2)3-4(6)mm long, tomentose; blade papery, elliptic, timesas longaswide, (4)6-11 x (2)3-5 cm, rounded toobtuse andoften somewhat unequalsided at base, top acutish to rounded, mucronate or not, or obscurely shortly acuminate, midribandthe 6-8 pairsof main lateralnerves impressed Agric. Univ. Wageningen Papers 86-3 (1986) 33

37 FIG. 3. D. fadenii: 1. branchlet with inflorescences, x ; 2. inflorescence with some flowerbuds and young fruits, 2 x ; 3. flower partly, 15 x ; 4. fruit in cross section (seed removed) showing hairy endocarp, 10 x ;5. part of fruit (seed removed) with protruding hairs, 7 x (1-5. R. B. & A. J. Faden, Gillett & Gachathi 77/416). 34 Agric. Univ. Wageningen Papers 86-3 (1986)

38 above, prominentbeneath;densely tomentoseabovewhen young, soon glabrescent, beneath persistently short-hairy allover,moredensely soonmainnerves; glands absent. Inflorescence branchednearbase into afew toseveral many flowered, spreading, scorpioid arms, tomentellous; peduncle 1-3mm long; bracts andbracteolesnarrowlytriangular, mm long. Pedicel2-4 mm long, tomentellous, the upper part at most 0.5mm long. Sepals suberect to spreading orevensomemore orlessreflexed, ovate-oblong, x ca 1 mm,tomentellousoutside and onapical partinside. Petalssuberect, oblong tonarrowly obovate in outline, mm long, 1 mm split, glabrous both sides, at base very shortly adnate to filaments. Stamens suberect, as long as the petals, glabrous; anthers small,ca0.3mmlong. Staminodes subquadrate totransversely oblong, 0.5 x mm, slightly bilobed, glabrous or hairy at base inside. Pistil 3-merous, mm long;ovary villous, style glabrous or with a few hairs in basal part, apically distinctly lobed. Fruits (only immature fruits seen) subglobose to obovoid, slightly laterally compressed, tomentose; endocarp densely long-hairyinside. Seeds glabrous. Distribution: Onlyoncecollectedin Kenya (seetype). Ecology: Limestone outcrops MAP 3. D.fadenii. D.gassitaeBret.,sp.nov. Fig. 4Map 4 Liana tenuis, hispida. Stipulae subulatae, 5-9mm longae. Folia elliptica, x 7-10cm, basibimarsupiata, acuminata,omnibus hispida.inflorescencia florifera, subsessilia, compacta. Flores 7-8 mmlongi. Sepala erecta. Agric. Univ. Wageningen Papers 86-3 (1986) 35

39 36 Agric. Univ. Wageningen Papers 86-3 (1986)

40 Petalabreviterbiloba.Ovarium 3-loculare, velutinum. Fructus ignotus. Type: Gabon, 50km S.E. of Achouka (0.17S, E), Louis, Breteler and De Bruijn 761 (holotype:wag). Diagnostic characters. Slender hispid liana. Stipules subulate, 5-9mm long. Leaves elliptic, x 7-10cm, bimarsupiate at base, acuminate at top, hispid bothsides. Flowers 7-8 mm long arrangedin rather compact,subsessile,12-34-flowered inflorescences. Sepals erect. Petals shortly bilobed,sparsely pubescent onupperhalf bothsides. Ovary3-locular,velutinous. Discription. Slender liana. Stemwith intruding phloem, sparsely lenticellate. Branches and branchlets long hispid-hairy mixed with short curled hairs. Stipulessubulate, rather early caducous, 5-9mm long, hairy as branchlets. Leaves: petiole subterete, ca 4mm long, hairy as branchlet; blade elliptic, ca twice as long as wide, x 7-10cm, narrowed at base into 2 distinct pouches (bimarsupiate)more orlesshidingthepetiolefrom above,acutelyacuminate at top, the acumen 0.5-1cm long,with 7-9 main laterals each sideof midrib, hispid bothsides,usuallymixedwith short,often curled hairson midrib, mainlateralsand margin,glands rather small, dispersed, beneathonly. Inflorescence subsessile, rather indistinctly or oncedistinctly branched, ormore orless 2-4 armed, 12-34flowered, velutinous,in the basal partoften mixedwithsome longhispidhairs; peduncle atmost 4mm long;bractsand bracteoles triangular, 1-6 mm long, velutinous outside,puberulous inside. Pedicel up to 5mmlong, velutinous, the upper part0. Flowers 7-8 mm long. Sepals erect,concave,ovateelliptic, x 4mm, topacutish,moreorlessribbed and velutinousoutside, tomentellous inside. Petals erect, at base mm unitedwith stamens, oblong, mm long,0.5mmsplit,sparselypubescent on upperhalfbothsides,lobes concave. Stamens erect, as long as the petals or slightly shorter, glabrous; anthers 1 mm long connective distinct.stammofifes subquadrate,ca0.5 x 0.5 mm, rather thick, glabrous. Pistil 3-merous, 7mm long; ovary velutinous, style glabrous shortly3-lobedapically;stigmas ratherlarge,reniform. Fruits unknown. Distribution: Onlyonce collectedin Gabon (seetype). Ecology: Rainforest. Notes. This species has been named after DR. JEAN NOEL GASSITA, former headofthegabonese National ResearchOrganisation (CENAREST), to honourhisefforts to promote the botanical explorationofhis country andin establishing a national herbarium atlibreville. FIG. 4. D. gassitae: 1. flowering branchlet, x ;2-4. detail of leaf base: 2.from above, 1 x ; 3.cut lengthwise, 1 x; 4. from beneath, 2x; 5. flower, 4x; 6. flower partly, 4x (1-6. Louis, Breteler & de Bruijn 761). Agric. Univ. Wageningen Papers 86-3 (1986) 37

41 MAP 4. D. gassitae. Itisdifficult to indicatethe positionof D. gassitaeamong theafrican species. By its indumentumof the vegetative parts, its stipules andinflorescencesitseems to be close to D. chalotii. Many characters of the flower, like sepals, petals, stamens, and stigma indicate affinity with D. thollonii, but ovary indumentum does not. Maturefruitsmayclarify its position. D.heudelotii (Planch,ex Oliv.) Baill. var.heudelotii & var.ndongense Asdescribed the fruits of D. heudelotiimay beprominently veined. A recent collectionfrom Gabon,southofBoouéin the 'Régiondes Abeilles' {Louis, Breteler & De Bruijn WAG)showsfruits with adensely, irregularly, narrowly ridged to tuberculate skin.the same taxon hasbeencollected in thesame area in flower and fruit (Louis, Breteler & De Bruijn WAG). It is somewhat doubtful whetherthesespecimenshave tobeplacedin var. heudelotii or in var. ndongense. Theflowers aresmall,havingpetals,stamens,and pistilsof2.5mm lengthwhich means that they should beclassified asvar. ndongense According tothefruits, however,thespecimensare betterplacedinvar. heudelotii, aseven strongly nerved fruits have never been observed in var. ndongense. Moreover, the soft-hairy indumentum of the vegetative parts, although of minor importance, fits better invar. heudelotii thaninvar. ndongense. For thesereasons the twospecimenshavebeenidentified as D. heudelotii var. heudelotii. D.librevillensePellegr. Whentreatingthisspecies and D. brazzae, it hasbeenstated that bothspecies areveryclosely related, but that in absence of sufficient material, especially of D. librevillense, thesetwospecies,simultaneouslypublished bypellegrin,were maintained separate. More material with flowers and nearly mature fruits has been collected recently in Gabon in the 'Région des Abeilles' (Louis, Breteler 38 Agric. Univ. Wageningen Papers 86-3 (1986)

42 & De Bruijn 664, WAG). This material has been compared accurately with the already available material of both species. As a result the two taxa cannot be maintained as separate species and are here united under the name D. librevillensepellegrin. D.madagascariensePoir.var.madagascariense Inthe 'Région des Abeilles', the part of Gabon south of Booué, the forests proved to be extremely rich in Dichapetalum species (see also D. gassitae sp. nov.).theretwospecimensof D. madagascariensevar. madagascariense (Louis, Breteler & De Bruijn 707 & WAG) were collected with narrowly ridged to tuberculate fruits, more or lesslike those of D. heudelotiimentioned above. Similarly strongly nerved fruits had already been observed in D. madagascariense var. madagascariense aswell, butneverstrongly ridged to tuberculate ones. D.pedicellatum Krause Thepistilofthisspecies hasbeenmentioned as 2 (-4)-merous inthedescription, but as 3 (-4)-merous under the diagnostic characters. The last statement iscorrect,2-merouspistils have notbeenrecordedsofar. D.potamophilumBret, sp.nov. Fig. 5Map 5 Liana cylindrico ligneo profunde lobato provisum. Ramuli breviter pubescentes. Petiolus (2)3-5(6)mm longus. Folia ovato-elliptica 4-11(17) x (2)3-5(6)cm,basi late rotundata usque cordata, apice rotundata vel interdum obscure acuminata, juvenilia sparse pubescentis, nervis principalibus utrinque prominentibus. Inflorescentia sessilis vel fere sessilis, ramosa, multiflora, breviterpubescens.sepalareflexa. Petalasuberectausque patentia, 2-3 mmlonga, mm fissa, staminibus pistilloque aequilonga. Ovarium villosum, stylis liberisvelfereliberis. Fructus ignotus. Type: Gabon, Ivindo R. near Mayibout I, periodically inundated forest, Breteler & J. J. F. E. de Wilde 604 (holotype: WAG). Diagnostic characters. Lianawith a deeplylobed woodcylinder. Branchlets rusty topale-brown short-hairy. Petiole (2)3-5(6)mm long. Leavesovateelliptic, 4-11(17) x (2)3-5(6)cm, broadlyrounded to cordate atbase, rounded rarely obscurely acuminate at top, sparsely hairy when young, main nerves prominent both sides.inflorescence sessileor nearly so,branched, many flowered, rusty to pale-brownshort-hairy. Sepalsreflexed. Petalssuberect tospreading, 2-3 mm long, mm split, aslong asstamens and pistil.ovary villous, stylesfree ornearly so. Fruitsunknown. Agric. Univ. Wageningen Papers 86-3 (1986) 39

43 FIG. 5 D. potamophilum: 1. branchlet with immature fruits, x; 2. flower, 8x; 3. pistil, 8x (1. Louis, Breteler & de Bruijn 588; 2-3. Breteler & J. J. de Wilde 604). Description. Liana. Sternswith a deeply lobed woodcylinder; lenticels usuallydistinct, moreorlessin 5rows. Branchesgreyish to pale-brown,sparsely lenticellate, glabrous orglabrescent. Branchlets rusty to pale-brown short-hairy. Stipulesearly caducous or not, narrowly triangular, 1-3mm long, hairy as 40 Agric. Univ. Wageningen Papers 86-3 (1986)

44 branchlet. Leaves: petiole subterete to semiterete, somewhat grooved above, (2)3-5(6)mmlong,hairy asbranchlet; bladepapery tocoriaceous,ovate-elliptic, 1.5-2(3) times as long as wide, 4-11(17) x (2)3-5(6)cm, sparsely rusty hairy on main nerves and margin when young, soon glabrescent, broadly rounded to obtuse tocordate at base,usually rounded (sometimes emarginate) and often mucronate at top,rarelyacuminateand then notdistinctlyso,midrib andthe 5-9 pairsof main laterals prominent bothsides,marginslightlyrevolute or not; glands both sides, more numerous beneath, rather distinct, dispersed. Inflorescencessessileornearlyso (peduncle up to 3mm long), 3-5 times distinctlybranched, many flowered, rusty to pale-brown short-hairy; bracts and bracteoles minute, ovate-triangular, up to ca 0.5mm long. Pedicel up to ca 3mm long, rusty short-hairy, the upper part at most 0.5mm long. Sepalsreflexed, shortly united at base, ovate-oblong to narrowly triangular, x mm,rustytomentoseoutside, glabrous orwithsome shorthairson apical part inside. Petals suberect to spreading, at basevery shortly adnate to filaments,narrowly obovate toellipticin outline, 2-3 mmlong, split,glabrous or with averyfewhairs just below split outside, lobesconcave, rounded to acutish apically. Stamenssuberect, 2-3mm long, glabrous; anthers up to 0.5mm long,connective distinctor not. Staminodes subquadrate totransversely oblong, up to0.2 x 0.2 mm, glabrous. Pistil 3-4-merous, 2-3 mmlong;ovary villous; styles 3-4, free, or partly united,slightlycurved, glabrous;stigmasmall. Fruits (only youngfruitsseen) 1 (- 3?)-seeded, velutinous, distinctly beaked,beak curvedor not. Distribution: Cameroun, Gabon. Ecology: Riverine forest. MAP 5. D. potamophilum. Agric. Univ. Wageningen Papers 86-3 (1986) 41

45 Specimens examined. CAMEROUN.70 kms.s.w,ofbafia,borderofsanaga R., Letouzey 9806 (P, WAG). GABON. 7 km S.W. of Makokou,borderof IvindoR., Breteler 7623 (WAG);7643 (WAG);Invindo R.near Mayibout I,periodically inundated forest, Breteler & J. J. F. E. de Wilde 604 (WAG,type); alongoffoué R.nearAchouka, Louis, Breteler & De Bruijn 588 (WAG). Note. This species has, so far, only been collected on riverbanks, hence its epithet potamophilum. Itisrelated to D. choristilum byitskind of inflorescence and its flowers with free or nearly free styles. It differs from D. choristilum by theleaves andbythedifferent habitat.theymaybe distinguished asfollows: Leaves elliptic-obovate, usually tapering to a cuneate or narrowly rounded base, acuminate at toprarely not so D. choristilum Leaves ovate-elliptic, broadly rounded to obtuse or cordate at base, top rounded,rarelyacuminateandthen notdistinctlyso D.potamophilum D. rudatisii Engl. This species, hitherto only known from S.E. Nigeria and S.W. Cameroun, hasnowbeencollected in N.W. Gabon,ca15kmS. of Cocobeachalong Ntoum roadin primaryforest (coll. J. J. F. E. de Wilde c.s WAG). D. witianum Bret. When thisspecieswas treated its fruitswere unknown. However, recent collectionsfrom Gabon included material withmaturefruits whicharedescribed below. Fruits 1-2-seeded, obtuse both ends, tomentellous, orange at maturity; 1-seeded fruits: subglobose to slightly ellipsoid, slightly laterally compressed, x 1.5-2cm; exocarp firm, mm thick; mesocarp juicy, 3-6mm thick; endocarp woody, up to 0.5mm thick, rugose outside, smooth and glabrousinside.seedssubellipsoid, laterally compressed, up to 1.5 x 1 x 0.8cm, seedcoat thin, brown, glabrous. Fruiting specimens: 20 km S.E. of Sindara, Louis, Breteler & De Bruijn 1212 (WAG);32 kms.e.of Sindara, Louis, Breteler & De Bruijn 1377 (WAG). UNIDENTIFIED MATERIAL, POSSIBLY NEW TAXA Although almost all sterile or incomplete material of Dichapetalum could be identified beyond doubt, afewspecimensremain unnamed. Someofthemmay represent new taxa or may be new records for large areas and therefore they arediscussedbelow. When treating D. corrugatum, asynonymof D. unguiculatum (BRETELER,1978: 42 Agric. Univ. Wageningen Papers 86-3 (1986)

46 23) the specimens Breteler 6434, 6469, and 6904 (WAG), all with woodsamples, were kept separate. Breteler 6306 (WAG), also from Gabon but without woodsample, belongs to the same taxon. Since then material with flowers and/or fruits which may elucidate the identity of these specimens has not been collected. Chapman 1290 (CÖI, SRGH) was longtime the first and only specimen of Dichapetalum from Malawi that had been examined. It was collected in the Nchisi Forest, Kota Kota, on the west side of Lake Malawi. The small flower buds of it have been analyzed. This proved that it represents the genus Dichapetalum but the flower characters together with those of the leafy branchlets do not fit any known species. Better material is required to see whether a new species is involved. Recently a second specimen (F. Dowsett-Lemaire 600), this time completely sterile, has been examined at K. It has been collected in the same forest and certainly belongs to the same species. Vollesen in MRC 4919 (K) from S.E. Tanzania is sterile. It fits best in D. edule from that area, but some differences, especially as regards indumentum and leaf nervation, withhold me from placing it there without further evidence. Crosse-Upcott 137 (K), collected in the Kilwa Ditrict, is another unidentified specimen from S.E. Tanzania. Although the specimen is in flower, I have not been able to identify it. It shares some characters of D. edule of that area and of D. deflexum from Mozambique. REVISION OF TAPURA HISTORY OF THE GENUS Tapura was described by AUBLET in 1775 who based it on T. guianensis, the most common species in America. On this continent 21 species are now recognized by PRANCE, the revisor of the American Dichapetalaceae. OLIVER described the first African species in 1868, namely T. africana from Fernando Po. Initially ENGLER used the same name for an East African species in 1895, but in an appendix to the same paper he changed the name into T. fischeri. This same species was described once more in 1915 by DE WILDEMAN naming it T. lujae. PELLEGRIN is in 1922 the author of a third African species named T. letestui from Gabon. From this country and adjacent Congo HALLE & HEINE in 1967 described two new species T. bouquetiana and T. neglecta. This was followed in 1970 by T. ivorensis published by the present author and at present by T. carinata, once more from Gabon and adjacent Congo. Agric. Univ. Wageningen Papers 86-3 (1986) 43

47 DESCRIPTION OF THE GENUS Tapura Aublet, 1775: 126, pi. 48; De Jussieu, 1789: 419; De Candolle, 1825: 58; Endlicher, 1840: 1105; Bentham & Hooker. 1862: 341; Oliver, 1868: 344; Bâillon, 1874:141;Engler, 1896: 351;Pellegrin, 1913:579; Hutchinson & Dalziel, 1928: 321; Engler & Krause, 1931: 10; Hauman, 1958-a: 347; Keay, 1958: 433; Torre, 1963: 328; Hutchinson, 1964: 219; Halle & Heine, 1967:43; Prance, 1972: 49; Punt, 1975:43. Type species: T. guianensis Aubl. Rohria Schreber, 1789: 30. Type species: R. schreberi Gmelin ( = T. guianensis Aubl.). Dischizolaena (Bâillon) Van Tieghem, 1903: 230. Basionym: Tapura section Dischizolaena Bâillon, 1873: 112. Type species: T. capitulifera Baill. Gonypetalum Ule, 1906: 174. Type species: G. juruanum Ule (= T. juruana (Ule) Rizzini). Trees or shrubs. Stipules entire, caducous. Leaves petiolate, usually acuminate. Inflorescences usually compact, pedunculate or not, the peduncle free or adnate to petiole. Flowers zygomorphic, (4-)5-merous, bisexual, pedicel articulate. Sepals unequal, free or nearly so. Petals partly united with the stamens into a distinct tube, unequal, 1-2 distinctly longer and bicuculate apically, the others usually entire or nearly so. Stamens 2-3 fertile, 3-2 without anther. Basal staminodes (or disc lobes) united or not, wanting with the large petals. Pistil 2-3-merous; ovary hairy or not; style slender, often curved or nodded, hairy in upper part, 2-3-lobed apically. Fruits (only partly known) 1-3-seeded; exocarp dehiscent or not; mesocarp juicy or mealy; endocarp woody to pergamentaceous. Seeds with or without endosperm. Distribution: Tropical America and tropical Africa. Notes. There are 7 species recognized in Africa and 21 in America. The most primitive species with 5 equal petals and 5 fertile stamens, occur in America, the most advanced ones with 2 fertile stamens only, are African. Of the African species T. fischeri has a large distribution, and T. ivorensisis only known from Eastern Ivory Coast and Western Ghana. The remaining 5 species are confined to Western Central Africa, all occurring in Gabon. Some species (T. carinata, T. letestui, T. neglectd) are only known from a single or only two collections while their fruits have still to be collected. It is expected that T. fischeri may occur in Gabon and that further exploration of this country may yield further new species (see also p. 61: Tapura sp.). 44 Agric. Univ. Wageningen Papers 86-3 (1986)

48 SECTIONAL ARRANGEMENT In the genus 4 sections have been distinguished, two are based on American species and two others accommodate each a single African species. The two American sections, the type section Tapura and the section Dischizolaena Bâillon, are commented upon by PRANCE (1972: 50). PRANCE does not adopt any sections in American Tapura 'since there are no clear cut divisions'. The distinction between these American sections, namely 5 fertile stamens in section Dischizolaena and 3 in section Tapura is, according to PRANCE, bridged by T. colombiana, a species with 4 fertile stamens. Both African sections are based on species with one large petal and two, rarely three, fertile stamens, which characters are not found in American Tapura. It concerns ENGLER'S (1895: 235) Trispermium with type species T.fischeri and HALLE & HEINE'S (1967: 44) Laratapura based on T. bouquetiana. According to ENGLER his section is characterized by flowers which are mostly 4-merous with one large petal and 2 fertile stamens, rarely 5-merous with 2 large petals and 3 fertile stamens. However, all the flowers of T. fischeri which have been analyzed for this revision are 5-merous, have one large petal and 2, rarely 3, fertile stamens (see also note p. 61). HALLE & HEINE (I.e.) produced a key to all sections and to the African species of the genus. The section Dischizolaena is not represented in Africa. T. africana, T. letestui, and T. neglecta, are placed in the section Tapura. T. ivorensis also clearly belongs to it. In their key the authors separate the section Laratapura and Trispermium as follows: Corolla with 3 bilobed petals (one large) and 2 entire petals; peduncle of inflorescence adnate to petiole (at least half the petiole length) and always shorter than it section Trispermium Corolla with 1 bilobed petal and 4 smaller, entire ones; peduncle of inflorescence free from petiole and always longer than it section Laratapura Laratapura, with its onlyspecies T. bouquetiana, is quite distinct from all other species which is supported by a distinct pollen type allotted to it by PUNT (1975: 45). According to the same author, however, T. fischeri is not different from African species of section Tapura, while T. ivorensisis placed in another separate pollen type of its own. The new species T. carinata, will fit into section Trispermium as demonstrated in the key given above. This is very acceptable, as T.fischeri is in the author's opinion the closest relative. In this revision the sections reviewed above are not adopted, because it is felt, that prior to this, a critical comparison of the African species with the American ones is necessary. For such an investigation knowledge of the evolution of the staminal tube in Dichapetalaceae seems to be basic. Agric. Univ. Wageningen Papers 86-3 (1986) 45

49 KEYTOTHEAFRICANSPECIES 1 a Peduncle ofinflorescence free from petiole orinflorescence sessilein theleafaxil 2 b Peduncleofinflorescenceadnate topetiole 3 2 a Inflorescence slender stalked;flowers 5-6mmlong;fertile stamens 2. Gabon,Congo T.bouquetiana b Inflorescence sessileinleafaxil;flowers 4-5mmlong;fertile stamens 3. Gabon T.neglecta 3 a Flowers mm long. West, Central,Eastand SouthernAfrica. T.fischeri b Flowers at least4.5mmlong 4 4 a Sepals distinctlykeeled. Gabon.Congo T.carinata b Sepals notkeeled 5 5 a Youngleaves without a sparsecobwebbyindumentumbeneath.ivory Coast, Ghana T.ivorensis b Young leaveswith asparse cobwebby indumentum beneath. Eastern Nigeria,Cameroun, Gabon,Congo 6 6 a Leaves with 3-5(8) pairs of main lateral nerves; flowers 4.5-7mm long with up to 3mm long pedicels; petals up to 5mm long, notgeniculate.eastern Nigeria,Cameroun, Gabon T.africana b Leaves with 6-7 pairs ofmain lateral nerves;flowers 7-8mm long, sessileornearly so; petals mmlong,geniculate. Gabon, Congo T.letestui T.africanaOliv. Fig. 6,7Map 6 T. africana Oliver, 1868:344; Engler, 1896:351;Pellegrin, 1913:648; Engler, 1915:849;Hutchinson &Dalziel, 1928:325; Keay, 1958: 439;Keay as.,1960: 327; N.Halle & Heine, 1967:44;Breteler, 1970:14; Punt, 1975:44;Breteler, 1982: 1. Type: Equatorial Guinea, Fernando Po, Mann 1161 (holotype:k;isotypes: LE,P). Diagnostic characters. Tree to 20mormore, up to 100cm diam.branchlets appressed-short-hairy, glabrescent. Stipulesusually earlycaducous.leaves papery to coriaceous, obovate-elliptic, (6)9-20(22) x 3-8(10) cm, cuneate, rarely roundedat base, acuminate, with 3-5(8) pairsof main lateral nerves,sparsely hairy, glabrescent. Flowers subumbellate, inflorescence indistinctly branched, peduncle adnate to petiole.pedicel up toca 3mm long. Flowers 4.5-7mm long. Petals 2large andbicuculate, 3 smaller.fertilestamens3.pistil3-merous,ovary shortlyvelutinous. Fruitsellipsoid, x 1-1.5cm, minutely velutinous. 46 Agric. Univ. Wageningen Papers 86-3 (1986)

50 FIG. 6. T. africana: 1. flowering branchlet, \ x ; 2. large leaf beneath, \ x ; 3. flower abaxially, with bracteoles, 5 x ; 4. flower adaxially, 5 x ;5. staminal tube with basal glands, 5 x ; 6. as 5 but without indumentum, 5 x ; 7. pistil, 8 x ; 8. leaf-base with 1-seeded fruit, J x (1, 3-7. van Meer 1553; 2. van Meer 1460; 8. Leeuwenberg 9910). Agric. Univ. Wageningen Papers 86-3 (1986) 47

51 Mf FIG. 7. T. africana: 1. seedling, x ; 2. detail of first node (en = endocarp, co = cotyledon) 5 x (1-2. Thomas 2470). Description. Tree up to at least 20m tall and 100cm diam., densely branched and often lowly so. Boleoften fluted, irregularly or not. Bark rather smooth. Slash white, turningyellow quickly. Wood whitish. Branches glabrous 48 Agric. Univ. Wageningen Papers 86-3 (1986)

52 or glabrescent. Branchlets appressed-short-hairy, glabrescent. Stipules usually early caducous, appressed, ovate-triangular, (1)2-5(7)mm long, densely appressed-silkyhairy outside, nearly glabrous inside. Leaves: petiole subterete to semi-terete, usually grooved above, (2)5-10(17)mm long, not grooved and (6)10-15(30)mm longwhen unitedwith peduncle, appressed-short-hairy;blade papery to coriaceous, obovate-elliptic, 2-3(3.5) times as long as wide, (6)9-20(22) x 3-8(10) cm,cuneaterarely roundedat base, gradually to abruptlyacuminate at apex, theacumen 0.5-2(2.5)cm long, roundedapically,sparsely subappressed-hairy onmidrib and the 3-5(8)pairs ofmain lateral nerves both sides as well as the basally revolute margin, glabrescent, very soon so above, beneath alsowithsome arachnoid indumentum;midrib andmainlateralsmore or less plane above, prominent beneath; glands usually few, both sides or beneath only, small, rather well dispersed, but beneath somewhat concentrated near base. Inflorescencesubumbellate-flowered, indistinctly branched orbasally once,more orless,distinctlyso, subappressed-short-hairy, up to ca50-flowered; peduncle usually completely or nearly completely adnate to and ca as long as petiole, rarelyfreefrom petiole;bracts and bracteolesovate-triangular, 1-2mm long. Pedicel up toca 3mmlong,theupper part up toca0.5mmlong. Flowers 4.5-7mm long. Sepalsfree, unequal, ovate-elliptic to obovate, ratherthick, concave, x 1-2mm, the 2 outer distinctly smaller, the 3inner somewhat asymmetric, glabrous to puberulousoutside, the marginciliate, glabrous inside. Petals up to 5mm long, with stamens united into a 2-4 mm long tube with freeapical parts, deepersplit atthesideof thefertilestamens, glabrous to sparsely puberulous onupper part oftubeand somewhatextendingonfree parts outside,moredenselyhairy onupper part inside;free apical part of 2 major petals mm split, bicuculate, that of the 3minor petals shorter, entire, hairy inside. Stamens 3fertile, 2 without anther, the fertile ones equalling the large petals or slightly longer, the two lateral ones at one side (proximal to minor petals) provided with a small, subacute lobe; anthers ca 0.5mm long, with a very distinct, often muriculate connective; thesterile filaments usually distinct, acute, hairyinside. Basalstaminode(s) thick,moreorless united, semi-annular, somewhat cupular, wanting with the large petals, mm high, glabrous. Pistil3-merous, 4-5.5mm long; ovary depressed subglobose, densely shortly velutinous; style glabrous in lower, hairy in upper part, 3-lobed apically, the lobes up to ca 0.5mm long; style often bent and by its hairs attached to tube opposite the fertile stamens. Fruits l(-3?)-seeded, narrowly ellipsoid, rounded both ends, x cm,minutely velutinous, like finesandpaper to the touch;exocarp thin;mesocarp firm, juicy 2.2-3mm thick; endocarp thin,pergamentaceous, more orless smooth, finely striate, andglossy inside. Seeds narrowly ellipsoid, acute both ends, ca 20 x 5mm, brown, glossy; testa rather firm. Seedling with epigeal germination. Hypocotylca 10cm long, appressed-pubescent. Cotyledons with distinct, 5-8mm long petioles, enclosed by endocarp, lifted above the first pairofoppositeleaves. Firstleaves elliptic, cuneate at base, acuminate at top,pubescent, densely soonpetiole,lesssoon midrib and main laterals beneath, only sparsely so on remaining parts, with sparse arachnoid Agric. Univ. Wageningen Papers 86-3 (1986) 49

53 20 20 MAP 6. T. africana hairsonlowersurface. Epicotylar stem part between attachment ofcotyledons andnodebearing the first pairofoppositeleaves indistinct. Distribution: EasternNigeria,Cameroun,Equatorial Guinea, Gabon. Ecology: Rain forest. Specimens examined. NIGERIA. Aboabam, Afi F.R., Latilo FHI30976 (K); (FHI, K); near Orem, Oban Group F. R., Latilo & Ogunlayo FHI (K, WAG); van Meer 1353 (WAG); 1460 (WAG); 1553 (WAG). CAMEROUN. Cross R. North F.R., ca km 250 Ikom-Mamfe, Latilo FHI31829 (K); km 11 Nkongsamba-Loum, Bakaka Forest, Leeuwenberg & Breteler 8763 (WAG); 8772 (WAG); 3 km E. of Yingui, Leeuwenberg 9133 (WAG); km 11 Nkongsamba-Loum, Leeuwenberg 9634 (WAG); 9910 (WAG); 30 km S.W. of Ndikiniméki, Letouzey (P); 3 km E. of Yingui, Letouzey ( = Leeuwenberg 9133) (P, WAG); 30 km N.W. of Eséka, Letouzey (P); 20 km N. of Bipindi, Letouzey (WAG); Victoria, Maitland611 (BR, K, WAG); 611 bis (K, WAG); 614 (BR, K); 38 km N. of Bafia, Ngoro Mt., Ngamenikamga 101 (P, WAG); sin.loc, Preuss 1234 (B, BREM, M); 5 km S. of Kumba, Thomas 2190 (MO); near Kumba, Thomas 2470 (WAG); Thomas 3449 (MO, WAG); Winkler 1281 (Z). EQUATORIAL GUINEA. Fernando Po, Mann 1161 (K, LE, P, type). GABON. Near Libreville, Mondah Forest, Breteler 7669 (WAG); Louis c.s. 837 (WAG); Malende, Le Testu 7821 (BM, BR, P, WAG); Divinda, Le Testu 8573 (BM, BR, P, WAG). Notes. The descriptionoftheseedlinggivenaboveisbased on Thomas 2470 consistingof 4seedlings. Thelabelofthis materialgives noreference tomothermaterial, so it is difficult to be sure whether these seedlings are indeed from T. africana. Careful investigation of the material, especially of the shape and textureof the endocarp aswellas theleaf indumentum (sparse arachnoid hairs) leads to the conclusion that these seedlings almost certainly belong to T. africana. Itis remarkable that this Tapuraspecies producesseedlingswith adistinct hypocotyl and without a distinct stem part between attachment of cotyledons and firstleaves. In Dichapetalum atleastas knownfrom ca20 species, the hypocotylis absent, i.e.germination is hypogeal,and whenthe firstleaves are oppo- 50 Agric. Univ. Wageningen Papers 86-3 (1986)

54 sitetheyarewellseparated from thenodebearingthecotyledons (see BRETELER 1973: 29). In these Tapura seedlings the apical bud is, as it seems, protected by the two subparallel petioles of the cotyledons, which petioles are just sufficientlystalked togiveroom to the first oppositepairofleaves. T.bouquetiana N.Halle &Heine Fig. 8Map 7 T. bouquetiana N.Halle &Heine, 1967:45; Punt, 1975:45. Type: Gabon,LaLara, Le Testu 9348 (holotype: P; isotypes:br,k, WAG). Diagnostic characters. Shrub or treelet. Branchlets glabrous. Stipules minute, with nectar glands. Leaves papery, obovate-elliptic, (6)9-14 x (2)3-4(6)cm, glabrous, cuneate to rounded at base, acuminate, with distinct (4)5-7 pairs of main lateral nerves. Inflorescence a slender stalked, glabrous, few-flowered, loose head, peduncle free from petiole. Flowersyellow,ca 6mm long.onelarge petal, 2fertile stamens. Fruits glabrous. Description.Shrub or treelet up to ca 10 mtall and 10cm diam. Branches greyish-brown, glabrous, lenticellate or not. Branchlets glabrous. Stipules minute, ovate-triangular to transversely oblong, up to 1.5mm diam., the margin withnectarsecreting glands, theapexoften with afewstiff hairs. Leaves:petiole subterete,grooved or canaliculate above, (1)2-6(10)mmlong,glabrous to appressed-short-hairy above,glabrescent;blade papery,obovate-elliptic, 2-3(3.5) times as long as wide, (3)5-14 x (1)2-5(6)cm, glabrous, cuneate to rounded at base, acuminate at apex, the acumen (0.5)1-2.5cm long, rounded to somewhat acutish apically;midrib and the (4)5-7 pairs ofmain lateral nervesplane to slightly prominent above, prominent beneath, the blade slightly bullatebetween the main laterals or not, margin often slightly revolute; glands present or not,few,small,beneathonly, atsome distancealongside the margin. Inflorescencesinglein theleafaxilorsometimesgrouped onvery short tonomalleafless axillary shoots, subcapitate, rather indistinctly branched, slender stalked, up to9-flowered, glabrous; peduncle (0.2)1-3.5(4.5)cmlong;bracts andbracteoles minute, ovate-triangular totransverselyoblong, up toca 1 x 1 mm,often much smaller, glabrous, themargin with nectar secreting glands. Pedicelup to 3mm long, slightly curved or not, the lower part up to 2mm long, the upper part up to 1 mm long. Flowers yellow, 5-6mm long. Sepals appressed, free, very unequal in size, slightly so in shape, the outer much smaller, subcircular to broadly ovate, 2-4 x 2-3 mm, concave, the inner ones strongly so, glabrous exceptfor thesparselyandfinely ciliate margin. Petalswithstamensunitedinto a 2.5-3mmlong, firm tube,withfree apical parts; tubeglabrousoutsideexcept apically,insidehairyinupper part;free partoflarge petal 0.7-1mmsplit,bicuculate, glabrous inside except at base, outside and especially on margin shorthairy; small petals 4, the free part ovate-triangular, sparsely hairy both sides, the two lateral ones (proximal to fertile stamens) sometimes very slightly bicu- Agric. Univ. Wageningen Papers 86-3 (1986) 51

55 FIG. 8. T. bouquetiana: 1. flowering branchlet; 2. stipule; 3. petiole; 4. detail of leaf nervation; 5. inflorescence; 6. flower; 7. flower diagram; 8. flower cut lengthwise; 9. outer sepal; 10. inner sepal; 11. staminal tube outside; 12. staminal tube inside; 13. details of anther; 14. basal staminode; 15. basal staminode from above; 16. overy cut lengthwise; 17. detail of style and stigma; 18. immature fruit (1-18. Le Testu 6392). For measurements see description. 52 Agric. Univ. Wageningen Papers 86-3 (1986)

56 10 10 MAP 7. T. bouquetiana. culate apically. Stamens 2 fertile; anthers almost sessile on tube with slightly unequal thecae, 1 mm long, the connective very distinct, muriculate, at bast strongly papillate;free partsof sterilefilaments 5, subtriangular,geniculate,densely hairy. Basal staminode one, across large petal, semiannular to horseshoe shaped, glabrous, ca as high as ovary, depressed apically. Pistil 2-3-merous, 4-5mmlong; ovarysemiglobose, glabrous;styleexserted,slightly curved,sparselyhairy inupper part, apicallyvery shortly 2-3-lobed, lobes papillate. Fruits (only immaturefruits seen) glabrous. Distribution: Gabon,Congo. Ecology: Rainforest, semi deciduous forest. Specimens examined. GABON. 50 km S.E. of Lambaréné, Breteler 5740 (BR, K, P, WAG); Ahiemé on upper Komo R., Chevalier (BR, P); Fougamou, Le Testu 6392 (BR, P, WAG); Koulamoutou, Le Testu 8459 (P, WAG); La Lara, Le Testu 9348 (BR, K, P, WAG, type); 7 km E. of M voum, Louis, Breteler & de Bruijn 244 (WAG), 261 (WAG); 5 km S.E. of Ekouk, Louis, Breteler & de Bruijn 312 (WAG); La Lara R. crossing Mitzic-Medouneu Rd., Louis, Breteler & de Bruijn 445 (WAG); Oveng, ca 40 km S.W. of Mitzic, J. & B. Reitsma, Breteler & Louis 914 (WAG). CONGO. M'Vouti, Bouquet 1896 (BR, K, P, WAG). TapuracarinataBret, sp.nov. Fig. 9 Map 8 Frutex ad 20 m altus. Ramuli breviter subappresse-pilosi. Folia chartacea, obovata, basi obtusa vel cordata, sparse pilosa, glabrescentia. Inflorescentia 3-25florifera, subappresse-pilosa; pedunculus petiolo adnatus.sepala carinata. Petalacum staminibusin tubum bilabiatum connata; tubusin tus annulo distinctostrigoso ornatus.staminafertilia 2, labioinferiore lateraliter adnata;stamina sterilia (0)1-2. Staminodium basale unicum. Pistillum 3-merum, ovarium glabrum.fructus ignotus. Agric. Univ. Wageningen Papers 86-3 (1986) 53

57 FIG. 9. T. carinata: 1. large leaf with inflorescence, 1 x ; 2. flowering branchlet with small leaves, 1 x ; 3. flower, 5 x ; 4. flower without calyx, S x ; 5. staminal tube with basal staminode, 5 x ; 6-7. sepals in cross section, 10 x ;8. pistil with basal staminode, 10 x (1. Sita 3630; 2-8. Pobeguin s.n.). 54 Agric. Univ. Wageningen Papers 86-3 (1986)

58 Type: Gabon, 'Bas Ogowé, arbuste de sous-bois', Pobeguin s.n. (holotype: P; isotype:wag). Diagnostic characters. Shrub to 20m tall. Branchlets subappressedshort-hairy.leaves papery, obovate, rounded to cordate atbase,sparselyhairy, glabrescent. Inflorescence 3-25-flowered, subappressed-hairy, peduncle completely adnate to petiole.sepalkeeled,stronglyunequalinsizeandshape. Petals andstamensunitedinto a bilabiatetubewithone large bicuculate petalinupper lip and 2-3 smaller petals in lower lip, the tube inside with a distinct ring of stiff, suberect hairs.fertilestamens2, laterally attached tolowerlip;sterilestamens (0)1-2. Basal staminode 1.Pistil 3-merous, ovary glabrous, upper part of style bent over lower lip and by its indumentum attached to it. Fruits unknown. Description. Shrub to20 m tall. Branchlets subappressed-short-hairy. Stipules caducous, narrowly triangular, 1-5mm long, appressed or not, subappressedhairy. Leaves: petiole subterete to semiterete, grooved above or not, 2-5mm long, 4-13mm long when united with stalk of inflorescence, subappressedhairy; blade papery, obovate, 2-3 times as long as wide, 5-17 x 2-7cm, rounded to cordate at base,with rounded to acute, up to 1.5cm long acumen apically, margin and nerves subappressed-hairy both sides or indumentum restricted to midrib both sides only, glabrescent, the midrib and the 5-6 pairs of main lateral nerves slightly prominent above, more distinctly so beneath; glands rather small, beneath only, a few near base and often a few in upper halfalongsidethe margin. Inflorescenceindistinctly branched oroncedistinctly soinolderinflorescences, 3-25-flowered, subappressed-hairy;peduncleas long as petioleandcompletelyadnate toit;bractsand bracteoles triangular-trullate, up to 1 mm long. Pedicel up to 2mm long, hairy as inflorescence, the upper part at most 0.5mm long. Sepals 5,free or nearly so, strongly unequal in size and shape,theoutermuch smaller than theinner,concaveinside,firmly keeled (carinate)outside,thekeel central toalmostmarginal,subovate-elliptic to obovateinoutline, 3-4 x 1-3 mm,short-hairy onmarginand sparselysoonkeel. Petals 3-4, withstamensunited into a bilabiate tube,laterally deepersplitthan adaxially or abaxially; tube 4mm long, at base bulging over the single basal staminode, slightly nodded in apical part, glabrous outside or only appressedshort-hairy towards the lips, inside with a distinct ring of stiff suberect hairs; upper lip (1large petal), bent over mouth of tube, bicuculate mm split, appressed-short-hairy outside, less hairy so inside especially the lobes; lower lip (2-3 petals) more or less hairy as upper lip, the two lateral petals usually bicuculate, ca 1 mm split with unequal lobes, the fourth, adaxial petal present or not,narrowly triangular apically. Stamens 2 fertile, 2-1(0) sterile;fertile stamenslaterally attached tolower lip,free part offilament ca mmlong, anthersbroadlyelliptic,ca 1 mmlong,connectivedistinct,muriculate;free part ofsterile filaments narrowly triangular, ca 1 mmlong,shortly pubescent. Basal staminode 1, adaxial,transversely oblong to broadly obcordateca0.5 x 1 mm, Agric. Univ. Wageningen Papers 86-3 (1986) 55

59 MAP 8. T. carinata. rather thick, glabrous. Pistil 3-merous; ovary and lower part of style glabrous, upper part bent over lower lip and byits hairs attached to it, lobesca 0.5mm long. Fruits unknown. Distribution: Gabon, N.W. Congo. Ecology: Rain forest. Specimens examined. GABON. Lower Ogooué R., Pobeguin s.n. (P, WAG, type). CONGO. N'gongo R., Sita 3630 (IEC, P, paratype). Note. Thisspeciesisverydistinct byitsbilabiate flowers with a remarkable ring of hairsin the tube.the epithet 'carinata'refers tothe stronglykeeled sepals. Tapura fischeri Engl. Fig.10Map 9 Tapura fischeri Engler, 1895:235,423; 1896-a: 351; 1912-b: 445; 1915:849 Engler & Krause, 1931: 11;Robijns, 1948: 434; Brenan & Greenway, 1949: 131 Andrews, 1952: 107;Keay, 1958:439; Hauman, 1958-a: 347;Keay c.s., ; White,1962: 185; Ross, 1972: 219; Punt, 1975:44. Type: Tanzania,Kagehi, Fischer 282 (holotype:bf;lectotype: K). Tapura fischeri Engler var. pubescens Verdcourt & Torre,in Torre, 1962: 69; Torre, 1963: 328; Gonçalves, 1980:104. Type: Moçambique,Mossurize, Simao 1173 (holotype:ea; isotype: LISC). Tapura lujae De Wildeman, 1915: 101 (as T. lujai); Hauman, 1958-a: 348. Type: Zaïre, Cataractes, Luja 149 (holotype: BR). Dichapetalum dummeri Moss, 1928:123.SeeBreteler, 1978: 62 forfulldetails. Dichapetalum corradii Chiovenda, 1952:232. See Breteler, 1978:23 for full details. 56 Agric. Univ. Wageningen Papers 86-3 (1986)

60 FIG. 10. T. fischen:1. flowering branchlet, \ x ;2. flower,10 x ; 3.staminal tube with basal staminode, 10 x ; 4.as3.without indumentum; 5. pistil,20 x ; 6.2-seeded fruit, 5 x ; 7.dehiscing3-seeded fruit, 5 x ; 8. pyrene, 5 x ;9. pyrene in cross section (en = endocarp, e = endosperm), 10 x ; 10.seed, lateral view, 5 x ; 11. embryo with exserted radicle, 5 x (1. Geerling & Bokdam 2538; 2-5. Semsei 1887;6,8-11. Wagemans 2157; 7. Can 906). Agric. Univ. Wageningen Papers 86-3 (1986) 57

61 Diagnostic characters. Shrub to much and often lowly branched small tree, up to ca 20 m tall and 45cm diam. Bark grey-brown. Sapwood whitish. Branchlets more orlesssubappressed-hairy. Stipules soon caducous.leaves papery,elliptic to obovate, (2)4-10(16) x (1)2-5(7)cm,with 4-7(8) pairsof main lateralnerves,glabrous tohairy,glabrescent or not. Inflorescence up to 35-flowered,peduncle partly tocompletely adnate topetiole,sometimesfree. Flowers mm long.flowertubewith 1 largebicuculatepetaland usuallv 2fertile stamens. Pistil 3-merous, ovary glabrous to densely hairy. Fruit 1-3-seeded, ovoid toellipsoid, up to 5 x 4 mm, glabrous todensely hairy,dehiscent. Description. Shrub to much and often lowly branched small tree, up to ca20 m tall and45cm diam.,branchesoften horizontal oreven drooping. Bark grey-brown, rather smooth. Sapwood whitish, soon turning yellow or darker onexposure. Branchesgrey-brown, glabrous or nearly so,often minutely lenticellate. Branchletsmoreorless subappressed-hairy, often soonglabrescent. Stipulestriangular, usuallynarrowly so, 1-5 mm long,subappressed-hairy, sometimeswithglandular margin, soon caducous. Leaves: petiolesemiterete tosubterete,groovedabove or not, (1)2-10(17)mm long, usuallyslightlylongerwhen united with stalk ofinflorescence than when not so,subappressed-hairy; blade papery, elliptic to obovate, (1.5)2-3(3.5) timesaslong aswide, (2)4-10(16) x (1)2-5(7)cm,cuneate to roundedandoftenunequalsided at base,usually shortlyacuminate at top, the acumen rounded to acute, up to 1(2)cm long; midrib and the 4-7(8) pairs of main lateral nervesplane or slightly prominent above, moredistinctlyso beneath; glabrous tohairyandoften soonglabrescentabove, oftenmoredensely hairy beneathespeciallyon midrib and main laterals,glabrescent or not,sometimeswithpersistent dotsofhairsintheaxilsofmainlaterals; glands when present beneath only, usually a few, rather large,well dispersed, sometimes near base only. Inflorescence a subumbel, sometimes once or twice distinctly branched, up to ca 35-flowered, subglabrous to more or lessdensely subappressed-hairy;peduncle partly tocompletely adnate to petiole,sometimes free, (4)5-8(13) mm long; bracts and bracteoles ovate-triangular, up to 1(1.5)mmlong. Pedicel up to 3(6)mmlong,theupper part 0-0.5(1)mmlong, usually glabrous. Flowers mmlong. Sepalserect, subappressed, free or nearly so, slightly unequal in shape, usually strongly so in size, suborbicular toovate-elliptic, up toca 1 x 1 mm,slightly concave, glabrous tosparselyhairy outside,usuallyglabrousinside, marginciliate. Petalswithstamensunited into a0.5-1(1.5)mmlong,thin, glabrous to hairy tubewithfree apical parts; large, abaxial petal 2-2.5mm long, basal parthyaline, upper partdeflexed, pubescent outside, mm split, lobes cuculate, slightly spreading; 2 lateral petals shorter, ca 2 mm long, mm split, the lobes usually both cuculate, at least the abaxial longer lobe, appressed-pubescent outside; 2 adaxial petals often slightly shorter than lateral ones, apical part entire, narrowly triangular, 0.5-1mmlong,appressed-pubescent outside. Fertile stamens 2,rarely 3, equal toslightly shorter,rarelylonger thanlarge petal, long unitedwithadjacent lateral petal, shortly so with large abaxial petal, free part of filaments up to ca 58 Agric. Univ. Wageningen Papers 86-3 (1986)

62 0.5(1) mm long, glabrous, anthers cordiform,ca 0.5 mm long,connective distinct or only slightly so,muriculate or not, sometimes slightlyproduced beyond the thecae apically; sterile stamens 1-3 (the adaxial one or the lateral ones may be wanting or rudimentary) the free part filiform, mm long, pubescent. Basal staminodes united into an often lobulate ring around the ovary, open at thesideof the large petal,rarelyfree, mm high, glabrous, basallyadnate to tube. Pistil 3-merous, rarely 2-merous, 2-3(3.5)mm long; ovary glabrous to densely hairy; style subappressed-pubescent in upper part, more sparsely so or glabrous in lower part, apically shortly (2)3-lobed, lobesusually unequal in length, papillate. Fruitsovoid to obliquely ellipsoid, sometimes subglobose, 1-3-seeded, more or less distinctly lobed when more than 1-seeded, glabrous todensely villous-tomentose, dehiscent, exposing the endocarp surrounded by somescarletmesocarp; exocarpandmesocarp thin; endocarp bony,slightly rugose outside, glabrousandfinely striateinside. Seedsubellipsoid 3-4 mmlong, mm diam., rounded to acutish both ends with a thin, brown seedcoat. Embryo enveloped by a 0.2mm thick layer of endosperm, cotyledons flat, ca 0.2mm thick,radicleexserted. Distribution: From Ivory Coast eastwards to Ethiopia and southwards to Natal. Not yet collected in Central African Republic, Gabon, Angola and Zambia. Ecology: Semideciduous to deciduous forest, gallery forest, dry evergreen forest, savannah woodland,savannahthickets, up toca 1300m altitude. Specimens examined. IVORY COAST. Daoukro region, Aké Assis.n. (WAG); Aka-Komoékro, Chevalier (P); Bouna Res., 9 km N. of Kakpin, Geerling & Bokdam 2538 (WAG). GHANA. Sin.loc, Irvine 966 (K); Tan F.R., Lloyd Williams255 (K, WAG). NIGERIA. Iwo-Oyo distr., Oba F.R., AdebusyiFHI40947 (K); Olokemeji, Forest SchoolFHI27291 (K); Olokemeji F.R.. Jones, Keay & Onochie FHI 4922 (B, BR); FHI 4926 (B, BR); Egbe distr., Latilo FHI (FHI); Okafor FHI54691 (K); Ondo prov., Owo F.R., Onochie FHI 33240; Olokemeji F.R., Ross 145 (K). CAMEROUN. Doumpello, Hedin 726 (P); 45 km E. of Yokadouma, Letouzey 5036 (BR, P, WAG). CONGO. 35 km N. of Ouesso, Sita 808 (P, WAG); Bolozo region, Sita 3424 (WAG). ZAÏRE. Mahagi Port, Bamps 286 (BR); Kifuku, Bequaert 2696 (BR); Irumu, Bequaert 2791 (BR, K, WAG); Malitawa (Lesse), Bequaert 3005 (BR); 3255 (BR); Beni, Bequaert 3405 (BR); Lesse, Bequaert 4141 (BR); Luki, Breyne 4683 (WAG); Baniari, J. de Wilde 147 (BR, WAG); Luki, Donis 57 (BR); 1543 (BR, PRE); 2066 (BR, K); 2282 (BR, EA, K); Kurukwata, Gérard 3134 (BR); Ahungbo, Germain 5284 (BR, LISU); Bankaie, Gilbert (BR); Ntolo-Ibaku, Jans 888 (BR); Kibati, Lebrun 4360 (BR, K); Cataractes, Luja 149 (BR^type of T. lujae); Luki, Maudoux 187 (BR, LISU, WAG); Kifuku, Mildbread 2885 (UPS); Luk\,~Nsimundele 175 (BR); Mahagi Port, van der Ben 1326 (BR, K, M, SRGH); Luki, Wagemans 730 (BR, LISC, WAG); 2157 (BR, LISC). SUDAN. Meridi distr., Andrews 1339 (K); Mongalla prov., Aylmer 27/48 (K); Imatong Mts, Talanga, Frits & Vollesen 508 (K); 6 km E. of Bor, Lock 81/66 (K); Lado, Yei R., Sillitoe 476 (K). ETHIOPIA. Lower Omo R., Carr & Brown 719 (BR, K); Can & Matolo 906 (K, WAG); Murle, Omo R., Corradi8401 (FI); 8402 (FI); 8403 (FI); 8404 (FI); Murle Lake, Corradi8400 (FI, WAG, lectotype ofdichapetalum corradii); 8405 (FI); 8406 (FI); 8407 (FI); 8408 (FI); 8409 (FI); km S. of Abobo, Friis, Woldu & Vollesen 2447 (K). UGANDA. Panyimur, Alonzi 16 (ENT); Mabira, Dummer 3974 (BM, Z); Luala, Dummer 3976 Agric. Univ. Wageningen Papers 86-3 (1986) 59

63 MAP 9. T.fischeri. fk); Mabira, Dummer 4469 (BM, BR, type of Dichapetalum dummeri); Budongo Forest, Eggeling 1158 (1283) (EA, K); Rumogi near Koich R., Eggeling 1828 (1652) (BR, K); Chua distr., Madi Mt., Eggeling 1735 (K); Okallo, Eggeling 1934 (K); Budongo, Eggeling 2289 (EA); Bunyoro, Eggeling 3042 (EA, K); Budongo, Eggeling 3076 (K); 3461 (EA); 3462 (EA); Bunyoro, Karani 33 (EA, ENT); Budongo, Myers (K); Maramagambo, Synott 212 (ENT); Busoga distr., Wood 57 (EA, ENT). KENYA. Taveta, Dale 3753 (BM, BR, EA, K, P); 14 km Kaldeni-Kilifi, Faden & Evans 71/730 (K); Kwale distr., Diani forest, Gillett & Kibuwa (K); Taveta, Greenway 4476 (EA, K); Polhill & Paulo 978 (BR, EA, K, P). TANZANIA. Ukerewe Isl., Conrads 5547 (BR, EA, K); 6248 (EA); Kagehi, Fischer 282 (K, type); Ruhembe, Goetze 404 (BM); Goetze s.n. (PRE); Rau forest, Greenway 4526 (EA, K); Lake Manyara Nat. Park, Greenway & Karani s.n., (K); EAH (EA); near Moshi, Lewis 7 (K); Rau forest, Lewis 40 (K); 225 (K); Handeni distr., Misufini, Semsei 569 (EA); Mtibwa F.R., Semsei 855 (BR, EA, K); Kilosa distr., Kidadi, Semsei 1016 (B, BR, K); Handeni distr., Romwe, Semsei & Gone FH2919 (BR, EA, K); Kilosa distr., Vigude, Semsei 1068 (BR, EA, K); Mtibwa F.R., Semsei 1426 (BR, K); 1887 (EA, K, WAG); Pemba, Vaughan 473 (EA); 897 (EA); Zanzibar, Vaughan 2036 (BM); Himo R., Volkens 2170 (BM, BR, E, G, K, LE); Magombera F.R., Vollesen 4151 (K); Morogoro R., Wigg 1024 (EA, K); Rau F.R., Wigg 1534 (EA, SRGH). 60 Agric. Univ. Wageningen Papers 86-3 (1986)

64 MALAWI. Blantyre, Buchanan 30 (E, K); Lengwe Game R., Hall-Martin 741 (K, SRGH); 744 (SRGH); 944 (K); 1211 (K, SRGH). ZIMBABWE. Chiredzi distr., Sabi R., Sherry 302/71 (BR, K, P, SRGH, WAG). MOZAMBIQUE. Inhafenga, de Carvalho 1178 (BR, LMU, WAG); between Tete and Lupata, Kirk s.n. (K, LISC); Inhamitanga, Müller & Pope 1901 (K); Simao 237 (LISC); Mossurize, Simao 1173 (EA, LISC, type of T.fischeri var. pubescens). SOUTH AFRICA. Ubombo distr., de Winter & Vahrmeyer 8351 (K); Hlabisa distr., Moll 2835 (K); Lake Sibayi, Moll 4926 (K); 5011 (SRGH); 10 km W. of Mazengwenya, Moll & Muller 5699 (K); Hlabisa distr., Strey 5475 (PRE); Lake Sibayi, Vahrmeyer 561 (PRE); 719 (EA, K); 740 (K). Notes. According to ENGLER (1896, 1915) and ENGLER & KRAUSE (1931) the two flowers depicted in figure 188, K-L, figure 401, K-L, and figure 4, K-L show a 5- and a 4-merous corolla respectively. Both flowers, however, have a 5-merous corolla. In part K, the flower has one large and 4 small bicucculate petals, while in part L one large and 2 small bicucculate petals with 2 entire petals are shown. The situation as depicted in part K has never been observed in the material used for this revision. Tapurafischeri var. pubescens is based by the authors on more hairy specimens with 2 fertile stamens. Two fertile stamens is usualin T.fischeri (see cited figures above) and only rarely flowers with 3 fertile stamens are observed. The very hairy leaves and branchlets, and the more hairy flowers, indeed more so than generally seen in this species, do not justify a separate taxonomie status for this material, especially not when the more hairy material of the former T. lujae is taken into account as well (see below). In the concept of T.fischeri as presented here, the ovary varies from completely glabrous to completely hairy, with all stages in between. The indumentum of the ovary usually vanishes completely in fruit, butit remains and does become even more dense in fruit in some specimens from Western Zaire as well as in two other specimens, Sita 808 from adjacent Congo and Letouzey 5036 from S.E. Cameroun. HAUMAN (1958: 348) classified the Zaire specimens with hairy fruits as T. lujae. In Hauman's key to distinguish between T. fischeri and T. lujae, glabrous ovaries and glabrous fruits lead to the former, hairy ovaries and hairy fruits to the latter. Although material bearing flowers and fruits at the same time is rather scarce, I noticed that hairy ovaries may produce glabrous fruits. The other character used by HAUMAN to distinguish between the two speciesis more or less of quantitative nature i.e. the hairiness of leaves and inflorescences. When all available material from Africa was examined it became clear that this character did not permit a satisfactory separation of T. lujae either. Considering this variation in characters, I am convinced that more material from Central Africa will reveal more intermediate specimens as regards ovary and fruit indumentum as well as regards the indumentum of leaves and inflorescences. Therefore T. lujae has been placed in synonymy of T.fischeri. T.fischeri is the first species of the African Dichapetalaceae in which the seeds have distinct endosperm (see fig. 10-9). Also the embryo is slightly different in having a protruding radicle. Agric. Univ. Wageningen Papers 86-3 (1986) 61

65 FIG. 11. T. ivorensis: 1. leaf beneath, J x ; 2. detail of leaf beneath with gland, 9 x ; 3. inflorescence, j x ; 4. flower, 4J x ; 5. part of staminal tube with basal staminodes, 6 x ; 6-7. staminal tube both sides, 4J x ;8-9. stipules,4 x ; 10. pistil, 6 x ; 11. length cut ovary cell, 22i x ; 12. ovule with obturator, 22± x ; 13. fruit, J x ; 14. pyrene, 2\ x (1-2,8-9, Breteler 5955; 3-7, Breteler 5964). 62 Agric. Univ. Wageningen Papers 86-3 (1986)

66 T.ivorensis Bret. Fig.11 Map10 T. ivorensisbreteler, 1970:12; Punt, 1975:46; Hall &Swaine, 1981: 346. Type: Ivory Coast, 5km E. of Aboisso, Breteler 5955 (holotype: WAG; isotypes:b,br,ea,k,lisc, M, P, UCI,W, Z). Diagnostic characters. Shrubtosmall tree.branchlets appressed-puberulous, glabrescent. Stipules early caducous or not, subtriangular, 2-4 x 1-2 mm.petiolecanaliculate above.leafbladepapery tocoriaceous,obovateelliptic, (8)12-17 x (3)5-8cm, with 5-7(8) pairs of main lateral nerves,glabrous above,nearlyso beneath.inflorescence glomerulous, peduncle completely adnate to petiole. Flowers sessile or nearly so, ca 8mm long. Fertile stamens 3.Pistil3-merous,ovary velutinous. Fruitminutely velutinous. Description. Shrub or small tree,up to ca 6m. Branches brown, glabrous or glabrescent. Branchletsappressed-puberulous,glabrescent. Stipules narrowly to broadly subtriangular, 2-4 x 1-2mm, appressed-puberulous, early caducous or not. Leaves:petiole (2)3-8(9)mm long, canaliculate above, subappressed-hairy, glabrescent; blade papery to coriaceous, obovate-elliptic, (8)12-17 x (3)5-8cm, cuneate to rounded at base, graduallyacuminate at top, the acumen (0.5)1-1.5(2)cm long, rounded; glabrous above, beneath with a fewappressed hairsonthe mainnerves andalong the margin,soonglabrescent; main lateral nerves 5-7(8) pairs, rather inconspicuous above, prominent beneath;glandsfew, scattered, beneathonly. Inflorescence glomerulous, indistinctly branched, cm diam., appressed-hairy; peduncle completely adnate to petiole;bractsandbracteolessmall, triangular,appressed-hairy outside. Pedicel 0-1 mm long,lower part wanting, upper part 0-1 mm long. Flowers ca 8 mm long, sessile or nearly so. Sepals unequal, the two outer smaller, ovate-elliptic to obovate-elliptic, x mm, top rounded to acutish, shortly united atbase,tomentose outside,glabrous inside. Petals 7mm long,withthestamens united into a 3-5 mm long tube with free apical parts, deeper split at the side of thefertilestamens, sparsely pubescent outside,villous inside,mainly on upper part of tube; the free part of the two major petals 1 mm split, bicuculate, glabrous or nearly so; the 3 minor petals rather flat, rounded at top. Stamens: 3 fertile with 2-4 sterile filaments, the fertile ones equalling the large petals, anthers ca 1 mm long with very distinct, muriculate connective; the free part of the sterile filaments rather short, sometimes inconspicuous, densely villous at top. Basal staminodes 2-3,across thelarge petals,free topartly united, flat, ca 1 mm high, glabrous. Pistil3-merous, 7.5-8mm long, slightly longer than stamens; style glabrous or almost so in lower part, pubescent in upper part, 3-lobedapically, thelobes up toca 1 mmlong;ovarydepressed-globose,sometimes slightly 3-lobed, velutinous. Fruit ovoid, laterally compressed 3 x 2 x 1.5cm,2(-3?)-seeded, yellow,minutely velutinous, finely scabrid to the touch; mesocarpslimy; endoearp bony,rugose outside, glabrousinside. Seed narrowly ellipsoid, 18 x 7mm;testabrown,glossy, glabrous. Agric. Univ. Wageningen Papers 86-3 (1986) 63

67 ZU- 1 f 1 Â t{- ;\1 q " < - * 1 ' j ( N ^ ay «4 >/ r 0 i i e ) 10 MAP 10. T. ivorensis. Distribution: EasternIvoryCoast,Western Ghana Ecology: Rain forest. Specimens examined. IVORY COAST. 5 km E. of Aboisso, Breteler 5955 (B, BR, EA, K, LISC, M, P, UCI, W, WAG, Z, type); 6 km E. of Aboisso, Breteler 5964 (C, COI, E, FI, HBG, LISU, NY, P, PRE, UCI, WAG, WU). GHANA. 4 miles N. Draw River F.R., Hall & Abbiw, GC (WAG). T.letestuiPellegr. Fig.12 Map11 T. letestui Pellegrin, 1922:91 (as T. Le Testui);1924:59; N. Halle & Heine, 1967:44,49; Punt, 1975:44. Type: Gabon, Inganga, Le Testu 1742 (holotype:p; isotypes:bm, K, WAG). Diagnostic characters. Tree.Branchletssubappressed-brown-hairy. Stipules early caducous. Leaves coriaceous, obovate-elliptic, (6)7-10(12) x (2)3-5 cm, cuneate at base, shortlyacuminateapically, glabrous above, beneath with hairy midrib and with some whitish, arachnoid hairs on blade, with 6-7 pairs of main laterals. Flowers subumbellate-glomerate at base of leaf blade, ca 7-8mm long. Sepals 2.5-3mm long, appressed-pubescent outside. Petals subequal in length, two bicuculate, the three others narrower, entire. Fertile stamens 3. Pistil3-merous. Description. Tree up to atleast40cm diam. Branchesglabrous orglabrescent. Branchlets shortly, subappressed-brown-pubescent. Stipulesearly caducous, ovate-triangular, 2-4 mm long, hairyas branchlet. Leaves: petiolesubterete tosemiterete, grooved above, (1)3-5(7)mmlong, notgrooved and 6-9 mm longwhenunitedwithpeduncle,hairy as branchlet;blade coriaceous, obovate- 64 Agric. Univ. Wageningen Papers 86-3 (1986)

68 FIG. 12. T. letestui: 1. flowering branchlet, ix; 2. flower, 5x; 3.staminal tube inside, 5x; 4.as 3,butwithoutindumentum; 5. largepetallateralview, 5 x ; 6.smallpetallateralview, 5 x ; 7. pistil, 10 x ;8. pistil partlywith basalstaminode(s),10 x (1-8. Le Testul742). Agric. Univ. Wageningen Papers 86-3 (1986) 65

69 elliptic, 2-3 times aslong aswide, (6)7-10(12) x (2)3-5cm, cuneate at base, shortly acuminate at apex, the acumen up to 1 cmlong, rounded apically,glabrousabove, beneathwithsparsely,subappressed-brown-pubescent midriband with aloose,whitish, arachnoid, caducousindumentum onblade; midribplane toslightlyimpressed above, prominent beneath,mainlaterals 6-7 pairs, rather thin, plane and rather indistinct above, slightly prominent beneath, margin slightly revolute;glands usually few, beneath only, mainly inlower part. Inflorescence subumbellate-glomerate-flowered, indistinctly branched, up to ca 10- flowered,subappressed-short-brown-hairy;peduncle as long as petiole and adnate toit;bractsand bracteolessubovate-triangular, concave,up to 1 x 1 mm,glabrous inside. Pedicelca 1 mm long, the upper part ca 0.3mm long. Flowers ca 7-8 mmlong. Sepals suberect,cafree, subequal, thetwo outer slightly shorter, ovate-oblong, x mm, appressed-pubescent outside, glabrous inside. Petals subequalinlength, mm long,slender,thin,rather fagile,with stamens united into a 1-2mm long tube with free, geniculate, apical parts, deeper split at the sideof thefertile stamens; thetwopetals between the fertile stamens up to 1 mm split, bicuculate; 3 narrower petals entire;petals of both types long-hairy inside except for 2.5-3mm length apically. Stamens 3fertile, 2 without anther; fertile stamens up to 8mm long, filaments flat, long-hairy inside,anthers rather flat, broadly ovate,ca0.5 x 0.7 mm,withdivergent thecae;sterilefilaments filiform, 5-6 mmlong, long-hairy, topoften curved. Basal staminode 1 only (or afew united ones), acrossbicuculate petals, subquadrate totransversely oblong,upto0.3 x 0.7mm, glabrous. Pistil3-merous, 5-7 mm long; ovarydepressedsubglobose,sparselyvelutinous to moreorless soft-erecthairy;style pubescent,apicallywith 3, up to 1.5mmlong, glabrouslobes. Fruits unknown. Distribution: Gabon, Congo. Ecology: Semi deciduous forest. 0 iq >! ) Q% ' t:\ r -p- y 7 ^ <v«: ''-*' 66 1 \ \ \ / \ KSI \ V 20 \ MAP 11. T.letestui. Agric. Univ. Wageningen Papers 86-3 (1986)

70 Specimens examined. GABON. Inganga, Le Testu 1742 (BM, K, P, WAG, type). CONGO. Haute Loukénené, Groulez-Morel, Service Forestier du Moyen Congo 34 (P) (see note). Note. The only specimen from Congo issterile and has therefore not been identified with complete certainty. The vegetative characters of this specimen, however,only fit thisspecies. T.neglecta N. Halle & Heine Fig. 13 Map 12 T. neglecta N. Halle & Heine, 1967:47; Punt, 1975:44. Type: Gabon, Mandji (Iboundji), Le Testu 6090 (holotype: P; isotypes: BR, WAG). Diagnostic characters. Shrub (or small tree?). Branchlets soon glabrescent. Stipules early caducous, ca 3mm long. Petiole canaliculate above (4)6-9mmlong;bladecoriaceous,obovate-elliptic, 7-12 x 2.5-5cm, cuneate at base, shortlyacuminate, glabrousornearly so, glands beneathonly. Inflorescence free from petiole, sessile or nearly so, usually compact. Flowers shortly stalked 4-5 mm long. Sepals thick, concave. Two large petals bicuculate apically,fertilestamens 3. Pistil3-merous,ovaryshortly velutinous. Description. Shrub (orsmalltree?). Branchletsbrownish, appressed-shorthairywhen young,soonglabrescent. Stipulesearly caducous,narrowly triangular to lanceolate, ca 3mmlong. Leaves: petiolecanaliculate above, (4)6-9mm long, glabrous tosparselyshort-hairy;bladecoriaceous,obovate-elliptic, 7-12 x 2.5-5cm, cuneate at base, shortly acuminate at top, the acumen rounded, 0.5-1cmlong,theleafmarginoften slightlyrevolute; midribandthe 4-7 pairs of main lateralnerves plane above, prominent beneath; glabrous above, sparsely subappressed-hairy onmidrib and mainlateralsbeneath,sometimeswithsome arachnoid hairs between the main nerves aswell;glands rather small, beneath only, mainly in lower half. Inflorescence axillary, free from petiole, sessile or nearly so,usually compact and rather indistinctly branched, consisting of few to several, subsessile to shortly stalked, subglobose, up to 10-flowered heads; axis of inflorescence very short to somewhat elongated, subappressed-pubescent; bracts and bracteoles narrowly triangular, up to ca 2mm long, subappressed-pubescent. Pedicel up to ca 1.5mm long, the lower part up to 0.5mm long, appressed-short-hairy, the upper part mm long, glabrous. Flowers 4-5mm long. Sepals unequal, appressed or nearly so, thick, concave, ovateelliptic, 2-3 x mm, margin ciliate, sometimes with a few hairs outside. Petals 5with the 5stamens united into a 2.5mm long tube, apical parts free, 1-1.5mm long; tube sparsely short-hairy in upper part outside, the free parts glabrous,inside tomentose mainly so on upper partof tube and partlyextending on free parts;largepetals 2, with a 1.5mm long,0.5mm split bicuculate apical part; the 3 minor petals with a narrowly triangular to lanceolate entire apical Agric. Univ. Wageningen Papers 86-3 (1986) 67

71 FIG. 13. T. neglecta: 1.floweringbranchlet;2.terminal partof branchletwithstipules;3.inflorescence;4. bract; 5.flower diagram; 6.flower cut lengthwise;7-8.outer and inner sepal, from outside, lateral view and cross section; staminal tube outside and inside; 11. anther both sides; 12. upper partofstyle (1-12.Le Testu 6090). Formeasurementsseedescription. 68 Agric. Univ. Wageningen Papers 86-3 (1986)

72 MAP 12. T. neglecta. part. Stamens 3 fertile, 2 without anther; thefertile onesequalling thelarge petals, the two lateral ones at one side (proximal to minor petals) provided with asmall,subacute lobe, anthers ca0.5mm long with a distinct,slightlymuriculate connective,free part of sterile filaments narrowly triangular to lanceolate,apically hairy. Staminode1, across the large petals, semi-annular, 0.7mm high, glabrous. Pistil3-merous,ca 5mmlong;ovarydepressed,densely shortlyvelutinous;style usually curved or nodded, glabrous inlower part, hairy in middle andupper part,theapical partattached tothe tubeacross thelarge petals,lobes 3, short,glabrous, exserted. Fruits unknown. Distribution: Gabon. Ecology: Rain forest. Specimens examined. Onlyknownfrom the type. Tapurasp. There remains one unidentified specimen collected near the waterfalls in the TchimbéléR., Crystal Mts, Gabon (Breteler & J. J. de Wilde 49 - WAG).This specimen, withleaves andvery young inflorescencesonly,almost certainly represents thegenus Tapuraand probablybelongs to anew species.theinflorescence issessile or nearlysoasin T. neglecta. From the latter the materialdiffers,however,byitsleaveswhichhave 8-12mainlateral nervesoneach sideof themidrib insteadof 4-7. More materialisneeded. Agric. Univ. Wageningen Papers 86-3 (1986) 69

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