Synopsis of the genus Chlorophytum (Asparagaceae) in Central Africa (Democratic Republic of the Congo, Rwanda, Burundi)

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1 This file is a preliminary version, and will be no longer available when the final version is published online Plant Ecology and Evolution FastTrack: 1 37, 2012 REGULAR PAPER Synopsis of the genus Chlorophytum (Asparagaceae) in Central Africa (Democratic Republic of the Congo, Rwanda, Burundi) Pierre Meerts 1,* & Charlotte Sletten Bjorå 2 1 Herbarium et bibliothèque de botanique africaine, Université Libre de Bruxelles, CP 169, Avenue F.D. Roosevelt 50, BE-1050 Brussels, Belgium 2 Natural History Museum, University of Oslo, P.O. Box 1172 Blindern, NO-0318 Oslo, Norway Author for correspondence: pmeerts@ulb.ac.be Background and aims The genus Chlorophytum is revised for the Democratic Republic of the Congo, Rwanda and Burundi, in order to prepare the treatment of the genus for the Flore d Afrique Centrale. Methods Herbarium taxonomy and SE micrographs of seeds. All the material kept in BR, BRLU, BRVU, FT, K, B and KIP has been examined, supplemented with recent field observations in S Katanga. Nuclear and chloroplast DNA markers have been used to clarify the taxonomic position of some species. Key results A total of 65 taxa (: 62, Burundi: 19, Rwanda: 8) are reported for the study area, i.e. fifty species, two subspecies, twelve varieties and one form. A total of 27 taxa are the, five to Burundi and two to Rwanda. Three taxa are endemic to the. Three new varieties are described (Chlorophytum africanum var. nordalianum, C. colubrinum var. upembense, C. cameronii var. ruziziense). Twelve new combinations, 16 new synonyms and one new name (Chlorophytum sphagnicolum) are proposed. An identification key is included. SEM pictures of seeds are included for fifteen taxa and colour photographs of living plants for fourteen taxa. A brief phytogeographic outline is provided. Upper Katanga, with 47 taxa, appears as a prominent center of diversity of the genus. Key words taxonomy, Chlorophytum, identification key, Central Africa, phytogeography, revision, Katanga, seed coat, SEM. INTRODUCTION [ ] Les Chlorophytum paraissent des végétaux extrêmement variables et [ ] les caractères sur lesquels on croirait pouvoir se baser pour les grouper d une façon sûre, peuvent être, eux-mêmes, peut-être, sous la dépendance des conditions extérieures. [ ] Il ne nous est possible que de décrire, comme nouveautés, les plantes qui ne peuvent être rapportées, sans le moindre doute à un type ancien. Ce sera aux monographes de l avenir à assigner à toutes ces espèces provisoires leur véritable place dans la classification." (De Wildeman 1921b: 11) The circumscription of the genus Chlorophytum Ker Gawl. (Asparagaceae in APG III 2009) was revised by Ober meyer (1962), Marais & Reilly (1978), Nordal et al. (1990) and Kativu & Nordal (1993) to include all the species formerly included in Acrospira Baker, Debesia Kuntze, Dasystachys Baker, Verdickia De Wild. and most of the African species formerly included in Anthericum L. The genus Chlorophytum has recently been revised for the Flora of Tropical East Africa (FTEA) (Nordal et al. 1997), the Flora Zambesiaca (FZ) (Kativu et al. 2008) and the Flore du Gabon (Bjorå & Nordal 2010). In order to prepare the treatment of the genus for the Flore d Afrique Centrale (FAC), we have critically revised all the material of Chlorophytum from the Democratic Republic of the Congo (), Rwanda and Burundi. For a number of taxa, phylogenetic relationships within the genus have been investigated using molecular markers, the results of which are presented elsewhere. We here present a key to the species, a check-list with a revised synonymy, and a brief phytogeographic sketch of the genus within the study area. When our taxonomic treatment departs from FZ and FTEA, a concise justification is provided. New combinations are coined when necessary and new synonyms are reported. Detailed accounts of the taxa, including citation of all specimens studied, will be published in the Flore d Afrique Centrale. All rights reserved National Botanic Garden of Belgium and Royal Botanical Society of Belgium ISSN

2 Pl. Ecol. Evol. FastTrack, 2012 Material and methods All the relevant material from, Rwanda and Burundi in BR, BRLU, BRVU, B, FT, K, KIP and LSHI has been examined. This material was supplemented with our own collections and field observations made in Upper Katanga in 2009 and No material pertaining to Anthericum L. was found in the study area. No less than 69 names have been found that are typified by specimens collected in and Burundi. Conventional methods of herbarium taxonomy have been applied. Seed shape and details of seed testa have been studied with SEM. To reveal sister relations available species were analyzed phylogenetically using nuclear ITS and chloroplast rps16 and trnl-f sequences (for methods see Bjorå et al. 2008). This information was also used to check consistency of our taxonomic views with other recent taxonomic publications. Our work has been considerably facilitated by the publication of the revision of the genus Chlorophytum in FTEA (Nordal et al. 1997) and FZ (Kativu et al. 2008), completed by Poulsen & Nordal (2005) for Guineo-Congolian taxa. Important references for the study area include De Wildeman ( , 1913c), Troupin (1955), the Flore des Spermatophytes du Parc National Albert (Robyns & Tournay 1955), the Flore des Spermatophytes du Parc National de la Garamba (Troupin 1956) and the Flore du Rwanda (Troupin 1988). Results and discussion In total, 65 taxa are recognized for the study area, i.e. fifty species, two subspecies, twelve varieties and one form. One species science was published separately (Meerts 2011). Three new varieties, one new name, twelve new combinations and sixteen new synonyms are proposed. No less than 27 taxa are, five to Burundi and two to Rwanda (table 1). A key to identify all taxa is included. Photographs of living plants are included for fourteen taxa (figs 1 3). SE micrographs of seeds and seed coat structure are provided for fifteen taxa (figs 4 6). Seed shape and testa anatomy are variable within the genus Chlorophytum and provide important taxonomic characters (Marais & Reilly 1978, Nordal et al. 1990). The shape of the seed varies from disc-shaped to deeply cup-shaped, more or less folded. The periclinal walls of the testa cells vary from flat to convex, in a few cases with a distinct papilla up to 10 µm long, separated by more or less deep furrows. In some cases, seed traits are useful to discriminate species with very similar floral and vegetative traits in particular in the complex of Guineo-Congolian rain forest species (Poulsen & Nordal 2005). In many cases, variation in the studied material was more extensive and complex than in adjacent regions (especially so for C. blepharophyllum, C. cf. brachystachyum, the C. cameronii complex, C. macrophyllum, C. sparsiflorum, C. rubribracteatum), and occasionally difficult to reconcile with other recent revisions. Phytogeographic outline Table 1 gives the distribution of taxa in Central Africa as presence/absence in Rwanda, Burundi and the nine phytogeographic regions of as defined by Robyns (1948). Species richness in, Rwanda and Burundi (50 species, 65 taxa) is very close to that in FTEA (52 species, 58 taxa) and FZ (56 species, 65 taxa). Upper Katanga stands out as an important centre of diversity of the genus for C Africa, with 47 taxa. Zambezian species are restricted mostly to that region of. Only few species occur both in the Guineo-Congolian region and in Katanga (e.g. C. blepharophyllum, C. sparsiflorum, C. stolzii, C. subpetiolatum). The occurrence in Katanga of disjunct populations of Southern African species is remarkable (C. recurvifolium, C. calyptrocarpum, possibly also C. macrosporum). A few W African species are represented in Bas-Congo (C. debile, C. staudtii, C. sphagnicolum, C. warneckei). Endemic taxa are surprisingly few (two species: C. cordifolium, C. arcuatoramosum and two varieties: C. ca meronii var. ruziziense and C. colubrinum var. upembense) considering the large area covered by the flora. However, fifteen taxa have a restricted area, just extending to adjacent countries (table 1). Upper Katanga alone counts 47 taxa i.e. by far the largest richness of all phytogeographic districts of. The region of Lubumbashi, in particular, appears as an Eldorado with 29 taxa within a 1000 km² area. Although sampling bias cannot be ruled out, the high species density in that region fits in well with the high species richness in the adjoining regions of N Zambia (FZ), pointing to the horn of Congo as a most important centre of diversity for the genus Chlorophytum in tropical Africa. That region is currently under heavy anthropic pressure due to the rapidly increasing human population and dramatic deforestation, and therefore qualifies as a hotspot (sensu Myers et al. 2000) for the genus, with several taxa probably at risk. Finally, it is significant that 25 taxa are known from less than six collections and six taxa by only one collection (table 1) highlighting the urgent need for new plant collecting efforts in. Concerning Chlorophytum, future sampling effort should be directed to the Katangan highlands (Kundelungu, Mitwaba, Biano, Manika, Marungu), and the regions adjoining the Congo-Angola borderline. Key to Chlorophytum species in, Rwanda and Burundi The identification of species requires fully developed material comprising underground parts, leaves, flowers, and mature fruits. Underground parts must be collected carefully as tuber shape, size and arrangement have high taxonomic value. Flower colour, and stamen arrangement (actinomorphic or zygomorphic) should be observed on fresh materials. In some cases, examination of seed surface at high magnification (20 40 ) is required. Plant size and habit often change dramatically throughout the growing season. In particular, leaves are often incompletely developed or lacking altogether ( hysteranthous plants) at the time of flowering and leaf measurements at that stage may be grossly underestimated. 2

3 Meerts & Bjorå, Chlorophytum in Central Africa Key to Chlorophytum species in, Rwanda and Burundi 1. A single flower at each inflorescence node...group A 1. Flowers 2 5( 9) together, at least in part of the inflorescence nodes (look carefully for flower buds or naked pedicels at the base of flowers at lowermost inflorescence nodes) Peduncle very short or lacking (< 3( 5) cm, to be measured from below the lowermost inflorescence branch or flower)...group B 2. Peduncle > 5 cm Peduncle with bracts or leaves...group C 3. Peduncle naked, without bracts (except 1 2 immediately below inflorescence) (leaves can have a long sheathing base forming a pseudostem) Plants having the following combination of characters: leaves distichous, pedicel articulated under the middle, mature fruit transversally ridged, seed coat irregularly folded...group D 4. Plants not having that combination of characters Inflorescence: a branched panicle...group E 5. Inflorescence: a raceme...group F Group A: 1 flower at all inflorescence nodes 1. Peduncle with (1 )2 12 bract-like leaves Peduncle naked Perianth narrowly campanulate, constricted above the ovary; inflorescence generally a dense raceme; pedicels 1 3 mm; tepals 1-nerved...C. colubrinum 2. Perianth widely open, not constricted; inflorescence lax, with rachis visible between flowers; pedicels 4 14 mm; tepals 3 5-nerved Leaves developed at flowering; bracts auriculate at base and glossy; pedicels c. 5 mm long in fruit......c. recurvifolium 3. Leaves developing after flowering; bracts rounded at base and not glossy; pedicels 9 14 mm long in fruit...c. nubicum 4. Inflorescence branched Inflorescence a raceme Plant < 20 cm high, with a rosette of 2 4 ovate leaves > 15 mm wide...c. perfoliatum 5. Plant (20 ) cm high, with leaves distichous, graminiform, < 12 mm wide (Fig. 2D)......C. vestitum 6. Leaves with ovate-elliptic lamina, contracted into a canaliculate petiole as long as the lamina and forming an angle with it; < 7 flowers to a plant; rain forest plant...c. occultum 6. Leaves with filiform to ovate-elliptic lamina, not petiolate; > 7 flowers to a plant; grassland and woodland plants Leaves filiform to grass-like, 1 4( 6) mm wide Leaves linear to ovate-elliptic, 5 40 mm wide Fruit mm; leaves and peduncles spreading in all directions; rachis smooth...c. warneckei 8. Fruit 6 10 mm; leaves distichous and peduncles not spreading in all directions; rachis papillate......c. affine var. curviscapum 9. Leaves distichous; raceme lax, with rachis visible; perianth 7 9 mm long; roots wiry with big distal tubers...c. affine var. affine 9. Leaves rosulate; raceme dense, with rachis most often not visible; perianth 4 7 mm long; roots more or less fleshy, without distal tubers (but sometimes with subsessile tubers) Lower (abaxial) leaf surface glabrous; fruit glabrous; peduncle glabrous or rarely pubescent; stolon lacking; rhizome very short without clavate tubers; plant cm high at flowering......c. africanum 10. Lower (abaxial) leaf surface pubescent; fruit papillate-hairy; peduncle pubescent; rhizome horizontal, elongate, occasionally stolon-like, bearing normal roots and clavate tubers; plant 3 10 cm at flowering...c. leptoneurum var. katangense Group B: Flowers 2 5 to a node; peduncle < 3( 5) cm 1. Inflorescence developing before the leaves; leaves distichous without a petiole; cataphylls with brown margin (fig. 2E)...C. hysteranthum 1. Inflorescence developing with or after the leaves; leaves rosulate, or, when distichous, with a distinct petiole; cataphylls uniform in color without brown margin Peduncle densely covered with long papillae; leaf margin strongly crispate...c. minor 2. Peduncle smooth or scabrid; leaf margin smooth or undulate...3 3

4 Pl. Ecol. Evol. FastTrack, Raceme very lax, with internodes longer than flowers and rachis clearly visible between them Raceme dense with internodes shorter than flowers and rachis not visible Leaves filiform or linear, without a petiole Leaves with ovate lamina contracted into a canaliculate petiole Leaves attenuate in lower third, 5 12( 20) mm wide, laxly veined (space between nerves much wider than nerves themselves); bracts 5 10 mm long, longer than pedicels...c. debile 5. Leaves parallel-sided, not attenuate at base, 1 4( 6) mm wide, closely veined (space between nerves narrower than nerves); bracts 1 5 mm long, shorter than pedicels Fruit 6 10 mm long; inflorescence a raceme, much shorter than leaves; peduncle and rachis papillate.....c. affine var. curviscapum 6. Fruit mm long; inflorescence most often branched, equalling or longer than leaves; rachis smooth Roots wiry with big distal tubers; flower 5 8 mm long with anther mm; leaves filiform, 1 2 mm wide; leaves erect...c. angustissimum 7. Roots without tubers; flower mm long with anthers mm; leaves linear, 1 mm wide; leaves and peduncules spreading in all directions...c. warneckei 8. Inflorescence < 6 cm long (including peduncle); pedicel with joint in upper half; flowers fewer than 7 to a plant; lamina < 8 cm long; tepals greenish; seed testa without apicules...c. occultum 8. Inflorescence 5 30 cm long (including peduncle); pedicel with joint in lower half; flowers more than 7 to a plant; lamina > 8 cm long (at least in some leaves); tepals whitish; seed testa apiculate...c. alismatifolium 9. Pedicel articulated at tip; raceme much longer than broad; bracts persisting, prominent, partly concealing young fruits, smooth and glabrous...c. stenopetalum 9. Pedicel without a joint; raceme usually no longer than broad; bracts neither persisting nor concealing young fruits, often finely papillate on margin or abaxial surface and occasionally ciliate Peduncle and rachis virtually zero; inflorescence lax, with flowers more or less umbellate and pedicels much longer than flowers; leaf margin smooth...c. sp. B near tetraphyllum 10. Peduncle and rachis distinct, though very short; inflorescence more congested, not umbellate, with pedicels shorter than flowers (but sometimes elongating in fruit); leaf margin often ciliate at places Leaves (in herbarium) thin, membranous and translucent, without a pseudopetiole; rhizome with small subsessile clavate tubers; fruiting pedicels < 10 mm long, straight; bracts thin, membranaceous, with nerves not prominent; seed testa with periclinal walls verrucose (figs 3A & 6B)...C. pusillum 11. Leaves (in herbarium) thick, more or less coriaceous, opaque, distinctly contracted into a pseudopetiole as long as inflorescence; roots with distal tubers; fruiting pedicels > 10 mm long, often recurved; bracts opaque, with prominent nerves; seed testa with periclinal walls smooth (fig. 6A)......C. geophilum Group C: Flowers 2 5 at a node; peduncle > 5 cm long with bracts or leaves 1. Flowers narrowly campanulate, not widely open, white; outer surface of tepals papillate Flowers more or less open (star shaped), green or white, not papillate Inflorescence a raceme or rarely with 1 2 branches; rachis densely papillate; leaves < 2.5 cm broad; plant < 1.2 m high...c. longifolium 2. Inflorescence with 3 8 branches curved upwards; rachis smooth (rarely with sparse papillae); leaves > 2.5 cm broad; plant m high (fig. 1A & B)...C. arcuatoramosum 3. Leaves < 5 mm wide; plant < 50 cm high Leaves > 5 mm wide; plant > 50 cm high Inflorescence developing after the leaves; tepals tightly pressed against the ovary after anthesis; roots spongy but not dilated into fusiform tubers; seeds c. 1 mm...c. calyptrocarpum 4. Inflorescence developing before the leaves; tepals free from the ovary after anthesis; roots dilated into spindle-shaped tubers; seeds c. 2 mm...c. nubicum 5. Anthers > 8 mm long; inflorescence simple or basally with 1 5 short erect branches Anthers < 8 mm long; inflorescence a panicle with long spreading branches Leaves glabrous; shoot base abruptly dilated into a corm more than 4 cm broad, with concentric ridges bearing regularly arranged fibrous remains < 1 cm long...c. stolzii 6. Leaves with velvety pubescence; shoot base little dilated, less than 3 cm thick, without concentric ridges but with irregularly arranged, fibrous remains > 1 cm long (fig. 3E)...C. velutinum 7. All leaves spaced along the stem; fruit < 4.5 mm long...c. ruahense 7. All leaves in a large rosette; fruit 4 15 mm long...8 4

5 Meerts & Bjorå, Chlorophytum in Central Africa 8. Leaves, peduncle and pedicels pubescent...c. hirsutum 8. Leaves, peduncle and pedicels glabrous Perianth greenish, campanulate with lower part of tepals enclosing ovary; tepals 3-nerved, > 15 mm long...c. andongense 9. Perianth white, stellate; tepals 5 7-nerved, c mm long...c. sp. A near comosum Group D: Flowers 2 5 to a node; peduncle > 5 cm long, naked, often flat; leaves distichous; pedicel articulated below middle; mature fruit with transversal ridges; seed coat irregularly folded 1. Cataphylls and outer leaf sheaths with purplish spots or stripes...c. cameronii var. cameronii 1. Cataphylls and outer leaf sheaths without spots or stripes, but sometimes uniformly reddish Tip of outer tepals blackish (fig. 3B)...C. sphacelatum 2. Tip of outer tepals white, or sometimes pink or greenish Inflorescence with 1 5 branches Inflorescence a simple raceme Inflorescence branches 1 5, spreading, divaricate; tepals 8 15 mm long; lowest bract not foliaceous, < 15 mm long; leaves 4 15 mm wide; plant not forming dense tufts...c. galpinii 4. Inflorescence branches (0 )1( 2), erect, not spreading and divaricate; tepals < 9 mm long; lowest bract foliaceous, 1 4 cm long; leaves 2 4 mm wide; plant forming dense tufts...c. staudtii 5. Leaf margin and lower surface of midrib with very long cilia...c. pilosicarinatum 5. Leaf margin and lower surface of midrib glabrous or with short hairs Cataphylls (and, often, bracts) with reddish tinge; leaves graminiform, < 5( 10) mm broad (fig. 2F)......C. rubribracteatum 6. Cataphylls green; leaf width variable Leaves filiform to linear, mm wide, conduplicate Leaves linear 3 30 mm wide, flat to folded Stamens not in two groups, much shorter than perianth, with anthers 2.5 mm; tepals 5 10 mm long; leaves filiform mm wide, erect; bracts ± scariose with prominent nerves; peduncle flattened to terete, with prominent ribs (in herbarium); plant in dense tufts; in moist soil, W Congo......C. sphagnicolum 8. Stamens in two groups, slightly shorter than perianth and with anthers > 2.5 mm; tepals 7 15 mm; leaves 2 3 mm wide, spreading; bracts not scariose with nerves not prominent; peduncle flattened; plant not cespitose; in dry soil, E Congo and Burundi...C. cameronii var. ruziziense 9. Cataphylls with brown band at margin (fig. 2E); inflorescence developing before the leaves, not exceeding 15 cm long including the peduncle...c. hysteranthum 9. Cataphylls without brown band at margin; inflorescence generally > 13 cm including peduncle, developing before or after the leaves Roots thick, carrot-like, without distal tubers...c. subpetiolatum 10. Roots wiry, with distal tubers Leaves (3 )4 30 mm wide, with margin and lower surface of mid-nerve smooth to papillose, not denticulate; inflorescence unbranched, without a foliaceous bract; tepals 7 15 mm; anther 4 6 mm; fruit 5 7 mm long...c. cameronii var. pterocaulon 11. Leaves 2 4 mm wide, with margin scabrid-denticulate to ciliate and lower surface of midrib denticulate-scabrid; inflorescence often with one slender basal branch subtended by a foliaceous bract 1 4 cm long; tepals 7 10 mm; anther 3 4 mm; fruit < 5.5 mm long...c. staudtii Group E: Flowers 2 5 to a node; peduncle > 3 cm long, without bracts; fruit smooth, not transversally ridged; inflorescence branched (exceptionally robust specimens of group F, with branched inflorescence, may key out here) 1. Inflorescence with sterile plantlets Inflorescence without sterile plantlets Tepals 8 10 mm long, 5-nerved; anthers 4 5 mm long; inflorescence (with peduncle) cm high; leaves mm wide, linear, parallel-sided, without a pseudopetiole...c. sp. A near comosum 2. Tepals 5 8 mm long, 3-nerved; anthers 2 3 mm long; inflorescence (with peduncle) cm high; leaves mm wide, linear or elliptic with a pseudopetiole...c. sparsiflorum 3. Leaves hairy or ciliate at places Leaves glabrous or papillate Leaves < 3 mm wide, filiform to linear Leaves > 3 mm wide, of variable shape Roots wiry with terminal tubers; fruit about as long as broad, emarginate...c. angustissimum 5. Roots fleshy without tubers; fruit longer than broad, not emarginate...c. calyptrocarpum 5

6 Pl. Ecol. Evol. FastTrack, Leaf surface glabrous; margin ciliate (cilia sometimes caducous)...c. blepharophyllum 6. Leaf surface hairy Leaves graminiform, < 12 mm wide, distichous...c. vestitum 7. Leaves elliptic, mm wide, rosulate...c. hirsutum 8. Flower > 15 mm long, greenish, urn-shaped; tepals with short ligule above perianth constriction...c. andongense 8. Flower 3 10 mm long, greenish or white, not urn-shaped; tepals without a ligule Leaves graminiform, < 15 mm wide, neither attenuate in lower half nor petiolate, distichous Leaves narrowly to broadly ovate or elliptic, petiolate or attenuate, or strap-shaped but, then, rosulate (not distichous) Pedicels and flowers papillate-pubescent...c. pubiflorum 10. Pedicels and flowers glabrous...c. polystachys 11. Seeds 3 5 mm; leaves distichous; lamina angular-ovate, forming an angle with the petiole and more or less horizontal...c. alismatifolium 11. Seeds < 3.5 mm; leaves rosulate; lamina elliptic, ovate, or linear, not forming an angle with the petiole Plant with leaves developing after the inflorescence, linear, < 10 mm wide......c. gallabatense var. micranthum 12. Plant with leaves developing before the inflorescence, linear or elliptic, > 10 mm wide Pedicels and rachis with hairs or cylindric papillae c. 0.4 mm long...c. gallabatense var. floribundum 13. Pedicels glabrous or rough with papillae < 0.2 mm Tepals 7 10 mm long, 5-nerved, anthers 4 5 mm...c. sp.a near comosum 14. Tepals 4 7 mm long, 3-nerved; anthers mm Lamina abruptly contracted into a distinct petiole, with base truncate or rounded Lamina progressively attenuated towards base, or strap-shaped, without a distinct petiole Petioles sheathing peduncule and forming a pseudostem (fig. 3F)...C. zingiberastrum 16. Petioles not sheathing peduncle; pseudostem lacking Robust plants, with at least one leaf > 45 6 cm, drying black...c. orchidastrum 17. Medium robust plants, with all leaves < 38 6 cm, drying blackish or greenish Lamina widest in the lower one-third, cm; petiole longer than lamina; plants most often drying blackish green to black; rachis smooth; anther mm...c. lancifolium 18. Lamina broadest near the middle, cm (measurements only for forms with petiolate leaves); pseudopetiole shorter than lamina; plants drying greenish; rachis most often scabrid; anther 2 3 mm...c. sparsiflorum 19. Tubers on lateral rootlets; plant not blackening in herbarium; inflorescence generally with 2 branches or more...c. gallabatense var. gallabatense 19. Tubers on main roots or lacking; plants blackening or not; inflorescence generally with 0 1 branch (when more than 3 branches then leaves petiolate) Robust plants, drying black, with at least one leaf > cm; capsules 7 12 mm long......c. filipendulum 20. Less robust plants, with all leaves < cm; leaves blackening or not; capsules mm long Lamina widest in the lower one-third, equalling pseudopetiole or shorter than it; plants drying dark green to blackish; rachis smooth; anthers mm...c. lancifolium 21. Lamina widest near the middle, longer than the pseudopetiole; plant blackening or not (when blackening: leaves strap-shaped); rachis most often scabrid; anthers 2 3 mm...c. sparsiflorum Group F: Flowers 2 5 to a node; peduncle > 3 cm long, without bracts; fruit not transversally ridged; inflorescence a raceme (depauperate specimens of group E with simple raceme may occasionally key out here) 1. Inflorescence with sterile plantlets Inflorescence without sterile plantlets Tepals 8 10 mm long, 5-nerved; anthers 4 5 mm long; inflorescence (with peduncle) cm high; leaves linear, mm wide, not attenuate in lower half...c. sp. A near comosum 2. Tepals 5 8 mm long, 3-nerved; anthers 2 3 mm long; inflorescence (with peduncle) cm high; leaves linear or elliptic with a pseudopetiole...c. sparsiflorum 3. Leaf margin ciliate; leaves not distichous Leaf margin not ciliate or rarely ciliate but then leaves distichous...5 6

7 Meerts & Bjorå, Chlorophytum in Central Africa 4. Flowers pure white; fruits generally verrucose or tuberculate, slightly trigonous; leaves thin and membranaceous, spreading, rosulate, not sheathing the peduncle at fruiting stage (though sheathing when flowering); leaf margin generally undulate or crispate, neither thickened nor recurved (fig. 1D & E)...C. cf. brachystachyum 4. Flower colour variable, never pure white: yellowish, greenish, brownish, purplish; fruit not verrucose, sharply triquetrous; leaves thick and somewhat fleshy to fibrose, tightly sheathing the base of the peduncle at fruiting stage, thus appearing superposate; leaf margin not undulate, often recurved or thickened...c. blepharophyllum 5. One leaf, broadly ovate, abruptly contracted into a canaliculate petiole which tightly clasps the peduncle...c. cordifolium 5. Almost always 2 leaves or more; when a single leaf, petiole not clasping the peduncle Rachis and upper part of peduncle shortly pubescent; leaves generally distichous Rachis and upper part of peduncle glabrous or papillate; leaves rosulate or, more rarely, distichous but then rachis not pubescent Roots carrot-like, i.e. thick at base and progressively attenuate, without tubers; leaves narrowly ovate-elliptic, more or less contracted into a pseudopetiole (sometimes very short < 5 mm); seed coat irregularly folded...c. subpetiolatum 7. Roots wiry, with distal tubers; leaves linear, never attenuate near base; seeds disk- or saucershaped Cataphylls with brown margin; plant < 15 cm high at flowering, with leaves appearing after the flowers (fig. 2E)...C. hysteranthum 8. Cataphylls without brown band; plant cm high at flowering, with leaves appearing before the flowers...c. affine var. affine 9. Leaves markedly distichous; lamina angular-ovate, broadest under middle, abruptly contracted into a canaliculated petiole and forming an angle with it; seeds 3 5 mm...c. alismatifolium 9. Leaves rosulate; lamina linear to narrowly elliptic, progressively attenuate at base, without a canaliculated petiole and usually not forming an angle with it; seeds mm Leaves linear to narrowly elliptic, < 6 mm wide; bracts purplish-brown; roots thick at base, distally attenuate; leaves appearing before the flowers (fig. 1F)...C. burundiense 10. Leaves linear to elliptic 5 80 mm wide (when 5 mm wide, then leaves appearing after the inflorescence); bract green to whitish; roots not thick at base, generally with distal tubers Tepals 5 6-nerved, 7 13 mm long; bracts 10 30( 50) mm long, often whitish and very conspicuous before anthesis, soon withering and turning brown; anthers 3 5 mm long Tepals 3( 5)-nerved, mm long; bracts 2 10 mm long, not whitish before anthesis; anthers mm Inflorescence (with peduncle) cm high, often pseudoviviparous, very lax with rachis visible; leaves linear, mm wide, not attenuate in lower half; bracts 5 10 mm long, never whitish and conspicuous...c. sp. A near comosum 12. Inflorescence (with peduncle) < 60 cm high, never pseudoviviparous, relatively dense with rachis often concealed by fruits; leaves mm broad, attenuate in lower half; bracts 10 30( 50) mm long, whitish and very conspicuous before anthesis Fruit obtusely trigonous to rounded in cross-section, not emarginate; seeds variable in shape, from saucer-shaped to strongly plicate (fig. 5A); seed testa with raised periclinal walls (fig. 5B); raceme 5 20 cm long, dense, with fruits concealing rachis and lowermost nodes not distant; tepals 7 9 mm; anthers 3 4 mm, equalling filament; tubers lateral; pedicel articulated at tip (fig. 2A C)...C. clarae 13. Fruit trigonous to triquetrous, emarginate; seeds flat to slightly saucer-shaped (fig. 5E); seed testa with periclinal walls shallowly convex or flattened (fig. 5F); raceme cm long, lax to moderately dense, with rachis visible and lowermost nodes often distant; tepals 8 13 mm; anthers 4 5 mm, longer than filament; tubers on main roots; pedicels articulated in upper half...c. macrophyllum 14. Plant with inflorescence appearing before the leaves; leaves linear, < 1.0 cm wide; rootstock with fibres...c. gallabatense var. micranthum 14. Leaves completely developed at flowering, narrowly to broadly elliptic mm wide; rootstock not fibrous Pedicel articulated at the tip; raceme densely flowered, with rachis not visible; rachis smooth......c. stenopetalum 15. Pedicel articulated below the tip; raceme less densely flowered, generally with rachis visible; rachis smooth or papillate

8 Pl. Ecol. Evol. FastTrack, At least one leaf longer than 38 cm or wider than 5 cm; plant blackish in herbarium; peduncle robust and stiff, generally > 2.5 mm thick below lowermost flower (in herbarium), always rough......c. filipendulum 16. All leaves < 38 cm long and with lamina < 5 cm wide; plant blackish or green in herbarium; peduncle more slender, generally < 2.5 mm thick below the lowermost flower (in herbarium), rough or smooth Lamina ovate, broadest below the middle, abruptly contracted into a petiole as long as or longer than lamina; plant blackish-green in herbarium...c. lancifolium 17. Lamina elliptic to sublinear, broadest at the middle, progressively attenuate at base; plant blackish or green in herbarium (when blackish, leaves linear) Seed testa with periclinal walls flat to convex; pedicel articulated near or above the middle, straight; fruit (3 )4 7( 8) (4 )5.5 7( 9) mm; leaves mm wide, with more than (17 )21 nerves......c. sparsiflorum 18. Seed testa with periclinal walls apiculate; pedicel articulated below the middle, often recurved when fruiting; fruit c mm; leaves 5 15( 25) mm wide, with nerves...c. debile Chlorophytum species are often polymorphic. The key allows to some extent for variation of polymorphic species, which key out at different places. However, it does not allow for exceptional character states. The key is based mostly on original observations on material from, Rwanda and Burundi, supplemented by literature data when the material collected in the study area was too scarce. For the description of leaf shape, we follow the recommendations of Anonymous (1962) and, in particular, we avoid the word lanceolate. For conduplicate leaves, leaf width is expressed as twice the apparent width. Check-list of species with synonyms In the following check-list, citation of synonyms is limited to (i) new synonyms, (ii) all the names typified by type specimens collected in the study area (, Burundi, Rwanda), (iii) all the names that were used in one of the following references: Durand & Durand (1909), De Wildeman ( , 1921a), Lebrun et al. (1948), Robyns & Tournay (1955), Troupin (1956, 1988), Schmitz (1971), Lewalle (1972). In this account, only one representative specimen is cited for each phytogeographic district of as defined by Robyns (1948). Occasionally, more specimens are cited for taxa of particular interest. Full species descriptions and extensive lists of specimens studied will be published in the Flore d Afrique Centrale. 1. Chlorophytum affine Baker (1875: 160 & tab. 104); Baker (1876: 327); Baker (1898: 507); Hanid (1974: 586); Kativu (1994: 49); Nordal et al. (1997: 23); Kativu et al. (2008: 60). Type: Tanzania, Tabora Ditrict, Unyamwezi, Rubugwa, 1860, Speke & Grant s.n. (holo-: K, barcode K ). Two varieties can be recognized: 1. Peduncle erect, straight, > 10 cm long; inflorescence about as long as the leaves; leaves > 10 mm wide......c. affine var. affine 1. Peduncle curved at base, more or less flexuous, < 6 cm long; inflorescence shorter than the leaves; leaves < 8 mm wide...c. affine var. curviscapum 1a. Chlorophytum affine Baker var. affine Other representative specimen examined : XI: Lubumbashi, 1939, Salésiens 1147 (BR). Bodenghien 331 (BR) is an extremely robust specimen with a raceme of up to 22 cm long. 1b. Chlorophytum affine Baker var. curviscapum (Poelln.) Hanid (Hanid 1974: 588); Nordal et al. (1997: 23); Kativu et al. (2008: 61). Chlorophytum curviscapum Poelln. (Poellnit z 1942: 122). Type: Tanzania, Dodoma dist., Uyansi, near Chaya, 3 Jan. 1926, Peter (holo-: B). Other representative specimens examined Burundi: Territ. Bubanza, Plaine de la Rusizi, 800 m, 30 Oct. 1973, Reekmans 2872 (BR). Rwanda: Parc Nation. Kagera, Gabiro, savane herbeuse à Themeda-Hyparrhenia, alt m, 3 Dec. 1944, Germain 2858 (BR, K). Cultivated: Yangambi, originaire de la Kagera, Germain 7499 (BR, K). A dwarf plant, 5 13 cm high, with grass-like leaves 2 5 mm wide, very short papillate peduncle, and more or less prostrate inflorescence. The cited collections are somewhat atypical in having smooth leaf margins and recurved fruiting pedicels. They are superficially similar to C. inconspicuum (Baker) Nordal, but with ovoid distal tubers, not progressively dilated roots. C. affine var. curviscapum, though widespread in Zambia and Tanzania, has apparently not been collected yet in. Duvigneaud 4380 (BRLU) approaches var. curviscapum, but it has erect peduncle, filiform leaves, and tubers with a reticulate surface. More material is needed. 2. Chlorophytum africanum (Baker) Engl. (Baker & Engler 1892: 470); Kativu (1994: 51); Nordal et al. (1997: 21); Kativu et al. (2008: 66). Type: Tanzania, Tabora Dist., Rubuga (Rubugwa) 1860, Grant s.n. (lecto-: K, barcode K ). 8

9 Meerts & Bjorå, Chlorophytum in Central Africa Table 1 Check-list of Chlorophytum species in the Democratic Republic of the Congo, Rwanda and Burundi, with their geographic distribution. Numbering of phytogeographic districts of after Robyns (1948): I: Côtier; II: Mayombe; III: Bas-Congo; IV: Kasaï; V: Bas-Katanga; VI: Forestier central; VII: Ubangi-Uele; VIII: Lac Albert; IX: Lacs Edouard & Kivu; XI: Haut-Katanga. I II III IV V VI VII VIII IX XI Burundi Rwanda Observations affine var. affine x affine var. curviscapum africanum var. africanum africanum var. nordalianum africanum var. silvaticum x x x x Burundi and Rwanda number of collections if 5 Distribution 4 Sudano-Zambezian 2 Sudano-Zambezian Zambia, Tanzania, S Katanga S Katanga, N Zambia x x x Burundi Zambezian-E African alismatifolium x x x Guineo-Congolian andongense x x Sudano-Zambezian angustissimum x x Sudanian E trop. African arcuatoramosum x x Endemic blepharophyllum Katanga, N Zambia, W Tanzania var. x x x amplexicaule blepharophyllum var. x x x x x x x x x x Afro-tropical blepharophyllum cf. x brachystachyum Sudano-Zambezian burundiense x Burundi, W Tanzania calyptrocarpum x cameronii var. cameronii cameronii var. grantii cameronii var. pterocaulon cameronii var. ruziziense Zambezian-S African x x x x x x Afro-tropical x 5 E tropical Africa x x x x Afro-tropical x x and Burundi E, Burundi 9

10 Pl. Ecol. Evol. FastTrack, 2012 Table 1 (continued) Check-list of Chlorophytum species in the Democratic Republic of the Congo, Rwanda and Burundi, with their geographic distribution. I II III IV V VI VII VIII IX XI Burundi Rwanda Observations clarae x x colubrinum var. colubrinum colubrinum var. upembense number of collections if 5 Distribution Katanga, N Zambia, SW Tanzania x x x x x x Broad Zambezian x cordifolium x 5 debile x x 3 3 W African Endemic (Upper Katanga) Endemic (Upper Katanga) filipendulum x x x x x x Guineo-Congolian gallabatense var. floribundum gallabatense var. gallabatense gallabatense var. micranthum galpinii var. matabelense x 2 Zambezian x x x x x Sudano-Zambezian geophilum x x x x x x x E African-Sudanian x 3 Zambezian - S African Sudanian, E African hirsutum x x Albertine rift hysteranthum x 4 S Katanga, N Zambia lancifolium subsp. cordatum x x x C African lancifolium subsp. lancifolium lancifolium subsp. togoense leptoneurum var. katangense longifolium var. aureum x x 3 C African x W Guineo-Congolian x 3 Katanga, Zambia, Malawi x 3 Sudanian macrophyllum x x? x Sudano-Zambezian minor x 1 SW Tanzania, N Zambia, S Katanga 10

11 Meerts & Bjorå, Chlorophytum in Central Africa Table 1 (continued) Check-list of Chlorophytum species in the Democratic Republic of the Congo, Rwanda and Burundi, with their geographic distribution. I II III IV V VI VII VIII IX XI Burundi Rwanda Observations nubicum x number of collections if 5 Distribution 2 Sudano-Zambezian occultum x 1 N, Uganda orchidastrum x x x x x x Guineo-Congolian perfoliatum x pilosicarinatum x polystachys x pubiflorum x 1 Zambia, Malawi, Katanga 2 Angola, Katanga 1 Sudano-Zambezian 1 Zambezian pusillum x Sudano-Zambezian recurvifolium x ruahense x 3 S African 4 Katanga, W Tanzania, N Zambia rubribracteatum x x Zambezian sparsiflorum x x x x x x x x x x x x Afro-tropical sphacelatum x x x x Zambezian sphagnicolum x x staudtii x x stenopetalum var. latifolium stenopetalum var. stenopetalum x Angola-Congo 5 W C African S Katanga, N Zambia x x x x Sudano-Zambezian stolzii x x x x x x x broad Zambezian subpetiolatum var. pilosifolium subpetiolatum var. subpetiolatum x S Katanga, N Zambia, SW Tanzania x x x x x x x x x Sudano-Zambezian 11

12 Pl. Ecol. Evol. FastTrack, 2012 Table 1 (continued) Check-list of Chlorophytum species in the Democratic Republic of the Congo, Rwanda and Burundi, with their geographic distribution. Distribution number of collections if 5 I II III IV V VI VII VIII IX XI Burundi Rwanda Observations, Congo, Sudan, Cameroon x x 4 subpetiolatum var. subpetiolatum f. variegatum velutinum x Zambia, Malawi, S Katanga vestitum subsp. pilosissimum x Zambezian 1 W African warneckei x Zambezian and Burundi zingiberastrum x x? Burundi and Rwanda x x 2? x Sp. A near comosum Sp. B near tetraphyllum The pattern of pubescence is variable and of taxonomic value. Three varieties can be recognized, of which the first two were treated as distinct species by FTEA and FZ. 1. Leaf margin with dense, crispate, reddish hairs......c. africanum var. africanum 1. Leaf margin glabrous or, more rarely, with short pallid cilia Peduncle and abaxial surface of cataphylls pubescent throughout...c. africanum var. nordalianum 2. Cataphylls glabrous; peduncle glabrous or sparsely pubescent in upper part...c. africanum var. silvaticum 2a. Chlorophytum africanum (Baker) Engl. var. africanum Caesia africana Baker (Baker 1875: 160 & tab. 103a). Dasystachys africana (Baker) T.Durand & H.Durand (Durand & Durand 1909: 569). Dasystachys grantii Benth. (Bentham & Hooker 1883: 789); Baker (1898: 513); De Wildeman (1921a: 29), nom. illeg. Acrospira giorgii De Wild. & Ledoux (De Wildeman 1930: 92). Type: Env. Lubumbashi, 1923, De Giorgi s.n. (holo-: BR, barcode BR ), synon. nov. Other representative specimens examined : XI: Paturage de la Karavia, ferme CSK, Dec. 1933, Quarré 3645 (BR). A rare taxon in the study area apparently restricted to the region of Lubumbashi. 2b. Chlorophytum africanum (Baker) Engl. var. nordalianum Meerts, var. nov. A typo differt pedunculo et rachide omnino pubescente, cataphyllis extus pubescentibus et foliis marginibus non ciliatis. Type:, Haut-Katanga, 5 km NW of Lubumbashi, dembo de la Kiamalale, 14 Nov. 1961, Schmitz 7498 (holo-: BR). Fig. 4A & B. Other representative specimens examined : XI: Lubumbashi, Katuba, dalle latéritique, Dec. 1952, Schmitz 4261 (BR, KIP); Keyberg, forêt claire sur sol relativement frais, Dec. 1947, Schmitz 1250 (BR); Lubumbashi, Kasapa, 23 Oct. 1961, Poelman 74 (BR); Lubumbashi, Keyberg, Vallée Katapa, 21 Nov. 1956, Detilleux 135 (BR, KIP); Kasumbalesa, savane dérivée en contrebas de la colline boisée, 3 Dec. 1959, Duvigneaud 4377L (BRLU). Other collections examined Zambia: Ndola, 1957, Duvigneaud s.n. (BRLU); Ndola, 27 Nov. 1953, Fanshawe F518 (BR, K); Kitwe, gravelly roadsides, 25 Dec. 1966, Mutimushi 1696 (K); Kitwe, miombo on laterite, 23 Nov. 1968, Mutimushi 2836 (K). This new variety is well characterized by a pubescent peduncle, rachis and abaxial side of the cataphylls. The cataphylls have a network of dark green veins. It also differs from var. silvaticum in its seed testa structure, having strongly buldging, hemisphaeric periclinal walls (Fig. 4A & B) (vs. flat to conical periclinal walls in var. silvaticum). C. africanum var. nordalianum, having a pubescent peduncle, can be confounded with C. leptoneurum. The latter species is usually less robust, has a pubescent abaxial leaf surface and a creeping rhizome. Plants with the distinctive traits of var. nordalianum had already been reported from N Zambia by Kativu et al. (2008: 66). Var. nordalianum appears to be endemic to the extreme south-east of and adjoining regions of Zambia. 12

13 Meerts & Bjorå, Chlorophytum in Central Africa 2c. Chlorophytum africanum (Baker) Engl. var. silvaticum (Dammer) Meerts, comb. & stat. nov. Chlorophytum silvaticum Dammer, Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflan zengeographie 48: (Dammer 1912); Kativu (1994: 75); Nordal et al. (1997: 20); Kativu et al. (2008: 65). Type: Tanzania, Kilwa Dist., Donde, near Kwa Mpanda, 22 Dec. 1900, Busse 1310 (holo-: B, iso-: EA). Other representative specimens examined : IX: Plaine de la Rusizi, Feb. 1950, Germain 6084 (BR); XI: Territ. Mitwaba, Simama, sur riv. Dikuluwe, plaine alluviale Dikuluwe-Lufira, 19 Jan. 1956, Brynaert 436 (BR). Burundi: Plaine de la Rusizi, Germain 5546 (BR); Territ. Bujumbura, Gihungwe, savane, alt. 850 m, 26 Nov. 1972, Reekmans 2182 (BR). Intermediates between var. africanum and var. silvaticum (short, pallid marginal hairs) are occasionally found (e.g. Germain 5524 (BR, K)). 3. Chlorophytum alismatifolium Baker (Baker 1876: 324); Baker (1898: 496); Engler (1908: 308, fig. 208 A, B); Troupin (1956: 188); Hepper (1968b: 102); Poulsen & Nordal (2005: 13, figs 1H, 4, 5, 28); Bjorå & Nordal (2010: 4). Type: Equatorial Guinea, banks of the Gaboon river, Jul. 1861, Mann 1030 (holo-: K). Other representative specimens examined : V: Upemba, Rivière Kanonga, galerie forestière, De Witte 5588 (BR); VI: Penghe, forêt ombragée au bord de l Ituri, 5 Feb. 1914, Bequaert 2312 (BR); VII: Parc Nat. Garamba, Galerie à Irvingia, 15 May 1950, Noirfalise 387 (BR). The material from is more variable than reported by Poulsen & Nordal (2005). Leaf length often exceeds 25 cm, pedicels are up to 10 mm (vs. 5 8 mm), bracts are larger (10 3 mm vs. 5 1 mm), and tepals are often longer (up to 7 mm vs. 3 4 mm). Some collections from Sudanian Congo (Garamba) have lateral tubers [De Saeger 1207, 1342 (BR)], a trait typical of the East African, closely related, C. holstii Engl. More material is needed. Troupin 1802 (BR, K) has obovoid fruits which are much longer than wide (9 5 mm) and pedicels articulated above the middle, but other traits correspond to those of C. alismatifolium. Keay 58 (K), from a plant cultivated at Kew, is atypical in having leaves up to 42 cm long, and an 80 cm long inflorescence with most flowers replaced by plantlets. Louis (BR, BRLU, K) with very long (up to 20 cm) foliaceous lowermost bracts has sometimes been misidentified as C. bracteatum Hua, but other traits are typical of C. alismatifolium. 4. Chlorophytum andongense Baker (Baker 1878a: 260); Baker (1898: 506); Troupin (1956: 188); Obermeyer (1962: 698); Hepper (1968b: 102); Kativu (1994: 52); Nordal et al. (1997: 10); Kativu et al. (2008: 70, fig ). Type: Angola, Pungo Andongo, Dec. 1856, Welwitsch 3770 (holo-: BM, iso-: K). Fig. 4C. Chlorophytum longipes Baker (Baker 1878b: 325); De Wildeman (1913c: 15; 1921a: 28). Types: Sudan, Jur Ghattas, 30 May 1869, Schweinfurth 1801 (syn-: K) & 10 Jul. 1869, Schweinfurth 2045 (syn-: K). Other representative specimens examined : VII: Parc Nat. Garamba, Bagbele, Noirfalise 521 (BR); XI: Lubumbashi, termitière en forêt claire, 13 Jan. 1954, Schmitz 4545 (BR). C. andongense is closely related to the southern African species C. macrosporum Baker. The latter is said to have longer fruits (11 15 mm vs. 5 9 mm in C. andongense) and narrower leaves ( cm wide vs cm in C. andongense). A few collections from Katanga have long fruits and relatively narrow leaves (e.g. Billiet & Jadin 4150 (BR)). Katangan collections are variable in leaf width from narrowly strap-shaped with more or less undulate margin (traits of C. macrosporum) to elliptic (trait of C. andongense) (e.g. Quarré 2733 (BR, P). Duvigneaud 5416A&C (BRLU) is another critical specimen with macrosporum-like fruits and andongense-like leaves. We feel that C. andongense and C. macrosporum might actually not be distinct at species rank. The group definitely needs revision and, after much hesitation, we provisionally keep all Katangan collections under C. andongense. 5. Chlorophytum angustissimum (Poelln.) Nordal (Nordal et al. 1997: 42). Anthericum angustissimum Poelln. (Poellnitz 1942: 26). Type: Tanzania, Tabora District, Ngulu, from Malongwe to Nyahua, Jan. 1926, Peter (holo-: B, not seen). Fig. 4E & F. Anthericum nigericum Hepper (Hepper 1968a: 459, 455 fig. 5/4-6); Hepper (1968b: 97). Type: Nigeria, Zaria Prov., Anaria Forest reserve, 1 Jul. 1957, Keay (holo-: K). Other representative specimens examined : IX: Vallée de la Rusizi, groupement de Luberidi, Centre zootechnique, Paddock III, 14 Jan. 1954, J.F. Laurent 1015bis (BR); Plaine de la Ruzizi, savane à Themeda, alt. 800 m, Jan. 1950, Germain 5541 (K). Burundi: Prov. Bubanza, Gihungwe, steppe à Bulbine abyssinica, 8 Mar. 1981, Reekmans 9802 (BR, BRVU, K). This species was erroneously referred to as Anthericum calyptrocarpum Baker by Lewalle (1972). Collections from the study area often have peduncle > 5 cm, i.e. longer than reported in FTEA. Lukuesa 941 (K), from Katanga, is an exceptionally large specimen with inflorescence up to 56 cm including cm peduncle. It may represent a distinct taxon, but more material is needed. 6. Chlorophytum arcuatoramosum R.B.Drummond (Drummond 1953: 120). Type:, Mutuy, Feb. 1931, Quarré 2465 (holo-: BR). Fig. 1A & B, fig. 4G & H. Other representative specimens examined : V: Kisamba, ferme Selemo, Jan. 1931, Quarré 2350 (BR, K); XI: Tenke, colline Shimbidi, savane steppique sur sol faiblement cuprifère, Feb. 2010, Meerts 2010/54 (BRLU). One of the largest Chlorophytum species in tropical Africa. It is obviously related to C. longifolium (perianth papillate) and to C. colubrinum (perianth constricted above ovary). Seed testa (fig. 4G & H) is similar to that of C. longifolium, with strongly convex, rugose, periclinal walls and perforate furrows (though perforations are narrower: com- 13

14 Pl. Ecol. Evol. FastTrack, 2012 pare with Nordal et al. 1990: 548, fig. 6E). Extraordinary dimensions (1.7 3 m high), inflorescence architecture (3 7 branched) and smooth rachis justify species rank. C. arcuatoramosum forms a well-supported clade with species that were formely assigned to the genus Dasystachys Baker (cf. Bjorå 2008). 7. Chlorophytum blepharophyllum Baker (Baker 1876: 327); Baker (1898: 501); Kativu (1994: 53); Troupin (1955: 234); Troupin (1956: 189); Hepper (1968b: 100); Troupin (1988: 42; 45, fig. 15/1); Nordal et al. (1990: 547); Nordal et al. (1997: 51); Kativu et al. (2008: 74). Types: Sudan/Ethiopian border, Gallabat, around Matamma, 14 Jun. 1865, Schweinfurth 9 (lecto-: K, here designated, isolecto-: P); Zimbabwe, Lower Gwelo R., 22 May 1870, Baines s.n. (syn-: BM, n.v.). The C. blepharophyllum complex appears to be much more variable in Katanga than in most other regions of tropical Africa. Kativu et al. (2008) recognized subsp. rubropygmaeum (dwarf plants with reddish cataphyll-like leaves) and subsp. pendulum (pendulous inflorescence) and separated C. amplexicaule (broad leaves clasping peduncle) as a distinct species. In Katanga, plant size, intensity of purplish colours, leaf width and inflorescence habit are combined in various ways sometimes within a single population. None of the aforementioned traits appears to be of high taxonomic value in the abundant materials we have seen. Pending a phenetic analysis with a populational sampling, we feel that the best option at the moment is a very synthetic treatment, recognizing amplexicaule only at varietal rank. 1. Cataphylls prominent with spreading lamina broadly ovate to cordate, clasping stem; leaves 4 10( 12) cm broad; plant cm high......c. blepharophyllum var. amplexicaule 1. Cataphylls not prominent, with small lamina not clasping stem; leaves cm broad; plant 5 40 cm high...c. blepharophyllum var. blepharophyllum 7a. Chlorophytum blepharophyllum Baker var. blepharophyllum Chlorophytum fibrosum Engl. & K.Krause (Engler & Krause 1910: 132); De Wildeman (1912a: 344). Types: Cameroon, between Duka and Dangadji, May 1909, Ledermann 3647 (syn-: B); Korrowalplateau, May 1909, Ledermann 3890 (syn-: B); near Balda, May 1909, Ledermann 4031 (syn-: B). Chlorophytum longebracteatum De Wild. (De Wildeman 1911a: 275); De Wildeman (1912a: 345). Type:, Vankerckhovenville, Apr. 1906, Seret 539bis (holo-: BR) Chlorophytum nigrescens De Wild. (De Wildeman 1911a: 275); De Wildeman (1912a: 346). Type:, Gunbari, 26 Feb. 1906, Seret s.n. (holo-: BR, barcode BR ). Chlorophytum kerstingii Dammer (Dammer 1912: 363); De Wildeman (1913c: 14; 1921a: 28). Type: Togo, Sokodé- Basari, near Aledyo, Feb. 1901, Kersting 314 (holo-: B, not seen).? Anthericum ruwense De Wild. (De Wildeman 1913a: 508); De Wildeman (1913c: 9; 1921a: 27). Type:, Katanga, Ruwe, Oct. 1911, Hock s.n. (holo-: BR, barcode BR ). Other representative specimens examined : III: Région de Kisantu, 1907, J. Gillet s.n. (BR); IV. Mabete, savane guinéenne, 2 Apr. 1955, Devred 1781 (BR); V: 8 S 23 E, groupement plus ou moins forestier en bordure d une galerie, 9 Oct. 1957, Liben 3834 (BR); VII: Parc National de la Garamba, mont Bawesi, savane arbustive, alt. 700 m, 11 Mar. 1941, Germain 627 (BR); VIII: Kerekere (Aru), alt m, savane, 4 Apr. 1960, D. Froment 692 (BR); IX: Plaine de la Rusizi, rives Sange, galerie, Jan. 1950, Germain 5819 (BR); XI: 20 km NE Lubumbashi, forêt claire sur sol argileux rouge, 23 Nov. 1955, Schmitz 5092 (BR). Rwanda: Préfect. Kibungo, route Lulama-Gabiro, km 6, savane arborée, 13 Jan. 1972, Bamps 2957 (BR). Burundi: Territ. Bubanza, Randa, alt. 900 m, savane boisée, 29 Oct. 1968, Lewalle 3061 (BR). A very variable taxon. Size is more variable than previously reported, with many collections < 10 cm high and with leaves < 1 cm wide. Hysteranthous forms are sometimes observed [Reekmans 7091 (BR)]. Plants from Rwanda tend to have more densely ciliate leaf margin (up to 200 cilia/cm vs. c. 50 cilia/cm in general). Bouxin & Radoux 1054 (BR), from Rwanda, with small (5 mm long), hanging flowers probably deserves taxonomic recognition, but more material is needed. 7b. Chlorophytum blepharophyllum Baker var. amplexicaule (Baker) Meerts, comb. & stat. nov. Chlorophytum amplexicaule Baker, Journal of the Linnean Society 15: (Baker 1876); Baker (1898: 501); P oellnitz (1946: 277). Type: Tanzania, near Lake Tanganyika, Feb. 1875, Cameron s.n. (holo-: K, barcode K ). Fig. 1C. Chlorophytum homblei De Wild. (De Wildeman 1913a: 514); De Wildeman (1913c: 13, Plate III); De Wildeman (1921a: 28; 1930: 95). Type:, Lubumbashi, Feb. 1912, Homblé 153bis (holo-: BR), synon. nov. Other representative specimens examined : VII: Près de la rivière Ndondo, savane à Lophira, 9 Jul. 1955, Boutique 201 (BR). IX: Plaine de la Rusizi, Feb. 1950, Germain 266 (BR); XI: Kiswishi, monastère des sept sources, forêt claire à Marquesia, Jan. 2010, Meerts 2010/14 (BRLU). Var. amplexicaule occurs mostly in miombo woodland and derived savannas in Katanga, Tanzania and Zambia. We have maintained var. amplexicaule with some scepticism, as intermediates exist with var. blepharophyllum in Upper Katanga. It also appears to be extremely variable in marginal cilia density (cilia sometimes almost lacking) and extent of reddish tinge; the most vigorous forms occasionally have a pendulous inflorescence [Meerts 2010/36 (BRLU); Duvigneaud 4432L (BRLU)]. It may eventually turn out to be an aggregate of convergent forms of C. blepharophyllum having little in common beyond extreme robustness. Hysteranthous forms with deep purple leaves are occasionally observed [Meerts & Muding 58 (BRLU); Schmitz 4523 (BR)] (fig. 1C); such forms approach subsp. rubropygmaeum Bjorå & Nordal, but intermediates exist with normal forms. 14

15 Meerts & Bjorå, Chlorophytum in Central Africa In preliminary phylogenetic analyses this complex has strong support as a clade, but little internal structure. Both the Congolese [Meerts 2010/36 (BRLU)] and the Zambian [Hoell & Nordal 134 (O)] specimens of var. amplexicaule cluster within the clade, as do purple-leaved ones [Meerts & Muding 58 (BRLU); Nordal & Bjorå 4578 (O)]. 8. Chlorophytum cf. brachystachyum Baker (Baker 1893: 710); Baker (1898: 502); Kativu (1994: 54); Nordal et al. (1997: 52); Kativu et al. (2008: 77, fig ). Type: Malawi, Shire Highlands, fl. at Kew 1893, Buchanan s.n. (holo-: K, barcode K ). Fig. 1D & E; fig. 4I & J. Other representative specimens examined : XI: Sur la piste de la ferme de la Kando, sur termitières, Oct. 1988, Schaijes 4139 (BR); Kiubo, savane arborée sur le rebord des grès de Kiubo dominant les chutes, 15 Jan. 1960, Duvigneaud 5047 (BRLU); Lubumbashi, Campus de la Kasapa, termitière dans l arboretum, Nov. 2009, Meerts et al. 77 (BRLU). Specimens from are undoubtedly members of the C. brachystachyum-c. pauper complex (fruit verrucose to tuberculate, leaf margin ciliate), but they are variable and their status is critical. They are often much more robust than both of these species, and the rachis is occasionally papillose (e.g. Duvigneaud 5047Ch), a trait not mentioned in previous descriptions of those species. Fruit surface is also variable in roughness, sometimes within the same population. Bracts are generally not ciliate. Leaf margin is undulate and has long cilia (traits of C. brachystachyum) but tubers are lateral (trait of C. pauper). In preliminary phylogenetic analyses the Congolese C. cf. brachystachyum [Meerts & Muding 77 (BRLU)] clusters with C. pauper, however, weekly supported. It does not form a monophyletic group with the Zambian specimen of C. brachystachyum [Bjorå 615 (O)] and further studies are therefore needed to clarify its taxonomic status. 9. Chlorophytum burundiense Meerts (Meerts 2011: 233). Type: Burundi, Territ. Muramvya, sommet du Mont Teza, prairie d altitude, 8 Dec. 1972, Baudet 304 (holo-: BR; iso-: K). Fig. 1F. Other representative specimens examined Burundi: Territ. Bururi, Tora, steppe, alt m, 7 Feb. 1971, Lewalle 5131 (BR). 10. Chlorophytum calyptrocarpum (Baker) Kativu (Kativu & Nordal 1993: 62); Kativu (1994: 55); Kativu et al. (2008: 53, Fig ). Anthericum calyptrocarpum Baker (Baker 1878: 258); Baker (1898: 480); Poellnitz (1943a: 62). Type: Angola, Huilla, between Mumpulla and Lopollo, Dec. 1859, Welwitsch 3786 (holo-: BM, iso-: K). Other representative specimens examined : XI: Lubumbashi, Mine de l Etoile, Cu-Co mineralized substrate, 2 Apr. 1990, Tropmetex 50 (BR, MO, K, MPN, WAG, C, IBE, P, PRE). Almost all collections come from the Lubumbashi area, most often on Cu-mineralised substrate. All records of C. calyptrocarpum from Burundi (e.g. Lewalle 1972) are erroneous and belong in C. angustissimum. Congolese collections lack the glands on the peduncle reported in the FZ area. 11. Chlorophytum cameronii (Baker) Kativu (Kativu & Nordal 1993: 62); Nordal & Thulin (1993: 262); Kativu (1994: 55); Nordal (1997: 102); Nordal et al. (1997: 30); Kativu et al. (2008: 44). Anthericum cameronii Baker (Baker 1876: 314); Troupin (1988: 31, fig. 9/2; 32). Type: Tanzania, Kigoma Dist., S of Kawele, Feb. 1875, Cameron s.n. (holo-: K, barcode K ). A very variable taxon. Four varieties are here recognized. 1. Base of leaves and cataphylls with purple spots or stripes...c. cameronii var. cameronii 1. Base of leaves and cataphylls unicolourous Lower leaf surface, peduncle and rachis pubescent......c. cameronii var. grantii 2. Leaves, peduncle and rachis glabrous (but leaf margin occasionally shortly ciliate) Leaves tightly conduplicate, filiform to narrowly graminiform, 2 3 mm wide, with margin generally ciliate at base; inflorescence 2 5 cm long with 1 5 nodes...c. cameronii var. ruziziense 3. Leaves flat or folded, not tightly conduplicate, linear, 4 25 mm wide, generally not ciliate; inflorescence (4 ) 7 12 cm long with 4 20 nodes......c. cameronii var. pterocaulon 11a. Chlorophytum cameronii (Baker) Kativu var. cameronii (Nordal et al. 1997: 31); Kativu et al. (2008: 44). Anthericum uyuiense Rendle (Rendle 1895: 415); Baker (1898: 485); Troupin (1956: 185 & 186, fig. 30); Hepper (1968b: 97). Type: Tanzania, between Uyui and the coast, 1886, W.E. Taylor s.n. (holo-: BM, barcode BM ). Anthericum congolense De Wild. & T.Durand (De Wildeman & T. Durand 1899: 60); De Wildeman & T. Durand (1901: 242); Durand & Durand (1909: 568); De Wildeman (1921a: 26; 1930: 94). Type:, environs de Lùbunda (Loubounda), Sep. 1896, Dewèvre 1029 (holo-: BR). Anthericum congolense De Wild. & T.Durand var. elongatum De Wild. (De Wildeman 1910b: 265); Troupin (1955: 233). Type:, Kasaï, Katola, 8 Apr. 1906, Sapin s.n. (holo-: BR, barcode BR ). Other representative specimens examined : IV: Kapanga, savane boisée, Nov. 1933, Overlaet 1089 (BR); V: La Pastorale, section 2, Oct. 1931, Quarré 2739 (BR); VII: Parc nation. Garamba, Crête Congo-Nil, forêt claire à Isoberlinia doka, 16 Jun. 1952, Troupin 1246 (BR); XI: Upemba, rive gauche de la Lufira vers Mabwe, alt. 920 m, 27 Oct. 1947, de Witte (BR). Rwanda: Territ. Kibungu, colline Muhororo, alt m, savane à bosquets, 26 Nov. 1958, Troupin 8860 (BR). Burundi: Territ. Cibiteke, savane boisée, alt m, 26 May 1967, Lewalle 1964 (BR). Collections from Rwanda generally have leaves narrower than in other regions (2 6 mm). 11b. Chlorophytum cameronii (Baker) Kativu var. grantii (Baker) Nordal (Nordal et al. 1997: 32). Anthericum grantii Baker (Baker 1875: 160 & tab. 103b); Baker (1898: 488). Type: Mozambique, Mgunda Mkali, Bass Rock, Dec. 1860, Speke & Grant s.n. (holo-: K, barcode K ). 15

16 Pl. Ecol. Evol. FastTrack, 2012 A B C D E F Figure 1 Photographs of living plants. A, B, Chlorophytum arcuatoramosum (, Tenke, Shimbidi, Feb. 2010, Meerts 2010/54); C, C. blepharophyllum var. amplexicaule, hysteranthous purple-leaved phenotype approaching C. blepharophyllum subsp. rubropygmaeum (, Kiswishi, Nov. 2009); D, C. cf. brachystachyum (, Lubumbashi, Kasapa, termite mound, Nov. 2009); E, C. cf. brachystachyum in fruit (, Kipopo, Jan. 2010); F, C. burundiense, cultivated, from Mt. Teza (Burundi). 16

17 Meerts & Bjorå, Chlorophytum in Central Africa Anthericum durum Poelln. (Poellnitz 1941: 232). Type:, Katanga, Kapiri, Oct. 1911, Hock s.n. (lecto-: BR, barcode BR , here designated) &, Katanga, Lukoni, 6 Oct. 1911, Hock s.n. (syn-: BR, barcode BR ), synon. nov. Anthericum rigidum De Wild. (De Wildeman 1913a: 507; 1913c: 8); De Wildeman (1921a: 27; 1930: 94); nom. illeg. non Anthericum rigidum Krause. A. rigidum De Wild. var. breviscapum De Wild. (De Wildeman 1915: 5) nom. illeg. Type:, Katanga, Plateau de la Manika, environs de Katentania, Nov. 1912, Homblé 754 (holo-: BR). Other representative specimens examined : XI: Lubumbashi, galerie forestière, Nov. 1927, Quarré 903 (BR); s.l., s.d., Quarré 1512 (KIP); s.l., s.d., Quarré 3684 (KIP); Territ. Mitwaba, Vibijo, alt m, savane herbeuse, 27 Mar. 1970, Coget 154 (BR); Tilwizembe, steppe sur pente rocheuse, légèrement métallifère en Cu et Co, 8 May 1957, Duvigneaud 3077L (BRLU); env. Fungurume, Kavifwafwaulu, sous forêt claire, 24 Nov. 2008, Malaisse et al. 428 (BR). Anthericum durum Poelln., described from Congolese materials, was erroneously reduced to C. galpinii var. galpinii by Kativu et al. (2008). The latter has a branched inflorescence (sometimes a raceme, but with a long aristate lower bract), and 3-veined tepals. In Anthericum durum, the inflorescence is always simple, with a short lower bract and the tepals have 5 widely spaced veins. Collections from Katanga are variable in size, pubescence and bract shape. Pubescent forms of C. cameronii might represent a heterogeneous assemblage. Duvigneaud 3082L (BRLU) from Tilwizembe copper hill with a forked raceme and scabrid leaves may belong here. 11c. Chlorophytum cameronii (Baker) Kativu var. pterocaulon (Baker) Nordal (Nordal et al. 1997: 32); Kativu et al. (2008: 45). Type: Angola, Pungo Andongo, s.d., Welwitsch 3795 (holo-: BM, iso-: K). Anthericum korrowalense Engl. & K.Krause (Engler 1910: 125); Troupin (1956: 184). Type: Cameroon, Korrowal plateau, May 1909, Ledermann 3832 (B, holo-, not seen). Anthericum speciosum Rendle (Rendle 1895: 413); Baker (1898: 486); Lebrun et al. (1948: 35). Type: Tanzania, between Zanzibar and Uyui, Feb. 1887, W.E. Taylor s.n. (holo-: BM, barcode BM ). Other representative specimens examined : V: Kuakidi, alt. 730 m, savane arbustive, 7 Dec. 1957, Lukuesa 464 (BR); VII: Parc Nat. de la Garamba, savane herbeuse sur alluvions de vallée, 1 Jun. 1951, De Saeger (BR); X: Makamba, 42 km S de Bururi, savane herbeuse sur argile rouge, 6 Jan. 1938, Robert 20 (BR); XI: Kilata, zone Sakania, 11 Nov. 1986, D Hose 34b (BR). Burundi: Territ. Ruyigi, Kigamba, savane arborée, alt m, 2 Jan. 1975, Reekmans 4108 (BR). In its typical form (peduncle flat, rachis undulate, inflorescence distichous, pedicels shorter than flowers), C. cameronii var. pterocaulon is easily recognized. However, many collections depart from that description in one or several traits. Var. pterocaulon is more variable than var. cameronii and is possibly a paraphyletic aggregate. Some collections have an inflorescence with 1 3 branches [De Witte 3088 (BR) and 5557 (BR) from l Upemba]. A few collections from Upper Katanga (region of Fungurume) [FC 12 (BR)] have red-coloured cataphylls and leaf margins. They may well deserve taxonomic recognition, but more material is needed. 11d. Chlorophytum cameronii (Baker) Kativu var. ruziziense Meerts, var. nov. A typo differt foliis tenuissimis, conduplicatis, 2 3 mm latis, nervatis, marginibus saepe ciliatis, cum vaginibus non purpureo maculatis. Type: Burundi, Bubanza, Gihungwe, alt. 800 m, 23 Nov. 1974, Reekmans 3955 (holo-: BR, iso-: K). Other representative specimens examined : IX: Plaine de la Ruzizi, piste Kabunambo-rivière Ruzizi, savane herbeuse, Jan. 1950, Germain 5517 (BR, FT); Plaine de la Ruzizi, env. Kihombo, savane à Themeda et Bulbine, Feb. 1950, Germain 6008 (BR, K); Plaine de la Rusizi, Jan. 1929, Humbert 7301 (BR, K, B). Burundi: Bubanza, Plaine de la Ruzizi, km 30, savane à Bulbine, alt. 800 m, 25 Nov. 1973, Reekmans 2929 (BR). This new variety has extremely narrow, conduplicate leaves 2 3 mm wide, with nerves, most often ciliate in lower third (though not in the membranaceous sheath), not purple-spotted. It is variable in size, from 20 cm in collections from high altitude in Burundi to 90 cm in collections from the Ruzizi plain. This variety is close to C. zanguebaricum (Baker) Nordal, another narrow-leaved variant of the C. cameronii complex, but this does not have tightly conduplicate leaves and leaf margin is not ciliate. More research is needed on the narrow-leaved forms within the C. cameronii complex. Germain (1952) referred to this plant as C. gracillimum V.D., a nomen nudum (see under C. warneckei). In the preliminary phylogenetic analyses the Congolese C. cameronii var. ruziziense [Reekmans 3955 (K, BR)] resolves as sister to the Ethiopian C. cameronii in a well supported clade of species that previously were included in the genus Anthericum. 12. Chlorophytum clarae Bjorå & Nordal (Bjorå et al. 2008: 232, fig. 5; 233 figs 7 9; 236 fig. 21). Type: Zambia, Northern region, Mansa District, 4 Dec. 2002, Nordal & Bjorå 4542 (holo-: O, not seen). Fig. 2A C; fig. 5A & B. Representative specimens examined : V: Kiala, Feb. 1954, Thiébaud 383 (BR); XI: Kiswishi, monastère ND des sept sources, base de termitière en forêt claire, Jan. 2010, Meerts 2010/23 (BRLU); rivière Mashinji, S E, forêt galerie, 19 Feb. 1987, Billiet & Jadin 4191 [BR, with photographs of plant cultivated in greenhouse (S 1975; )]; Tenke, fossé ombragé dans le dembo d empoisonnement de la colline cuprifère de Kabwelunono, Feb. 2010, Meerts 2010/37 (BRLU); Galerie de la Lofoï, Jul. 1939, Quarré 5724 (BR). This taxon has long been confounded with C. macrophyllum (see discussion under that species). It is clearly distinct by its seeds, variable in shape, from more or less saucershaped to very irregularly angulose-plicate (flat to slightly saucer-shaped in C. macrophyllum), and by its seed coat with raised periclinal walls and wide furrows (shallowly convex 17

18 Pl. Ecol. Evol. FastTrack, 2012 periclinal walls and narrow furrows in C. macrophyllum, fig. 5C F). Other differences include lateral tubers (terminal in C. macrophyllum), more or less rounded fruits (strongly trigonous in C. macrophyllum), and a denser raceme. Therefore, collections without mature fruits or well-collected belowground organs cannot be identified. Other characters are variable, including anther length (3 5 mm), tepal length ( mm), leaf width (3 8 cm), raceme length (4 25 cm) and fruit surface (smooth to slightly rugose-verrucose). Collections from Katanga have a more variable seed shape than reported for Zambia (Bjorå et al. 2008). It is still unclear whether genuine C. macrophyllum occurs in Katanga. 13. Chlorophytum colubrinum (Baker) Engl. (Engler 1892: 162); Poellnitz (1946: 353); Kativu (1994: 56); Nordal et al. (1997: 19); Kativu et al. (2008: 66, fig ). Dasystachys colubrina Baker (Baker 1878a: 256 & tab. 35, figs 5 10); De Wildeman (1913c: 12; 1921a: 28; 1930: 96). Type: Angola, Huilla, Empalanca, Jan. 1860, Welwitsch 3784 (lecto-: BM; isolecto-: K, B, P). Two varieties can be recognized. 1. Raceme cylindrical, 4 30 cm long, 4 20 times as long as wide; leaves 3 40 mm wide; bracts not or only shortly ciliate; cataphylls glabrous......c. colubrinum var. colubrinum 1. Raceme subglobose to ovoid, cm long, less than 2 times as long as wide; leaves 2 4 mm wide; bracts long ciliate; cataphylls pubescent......c. colubrinum var. upembense 13a. Chlorophytum colubrinum (Baker) Engl. var. colubrinum Dasystachys verdickii De Wild. (De Wildeman 1902: 10); Durand & Durand (1909: 569); De Wildeman (1921a: 29; 1930: 96) Chlorophytum verdickii (De Wild.) Poelln. (P oellnitz 1945: 232) Type:, Katanga, Lofoi, 1899, Verdick s.n. (holo-: BR, barcode BR ). Dasystachys hockii De Wild. (De Wildeman 1911a: 264); De Wildeman (1913c: 11; 1921a: 29). Type:, Katanga, Vallée de la Luembe, Feb. 1910, Hock s.n. (holo-: BR, barcode BR ). Dasystachys bequaertii De Wild. (De Wildeman 1913b: 294); De Wildeman (1921a: 28). Type:, Katanga, Elisabethville [Lubumbashi], 15 Apr. 1912, Bequaert 333 [published as 533] (holo-: BR). Chlorophytum subpapillosum Poelln. (Poellnitz 1945: 232). Type:, Lubembe, 27 Jan. 1908, Kassner 2394 (holo-: B, iso-: BR). Chlorophytum poggei Engler. ex Poelln. (Poellnitz 1945: 232). Type:, Mukenge, 27 Apr. 1882, Pogge 1491 (holo-: B). Dasystachys pulchella P.A.Duvign. & Dewit (Duvigneaud & Denaeyer-De Smet 1963: 149, fig. 11). Chlorophytum linearifolium Marais & Reilly (Marais & Reilly 1978: 660). Type:, Kolwezi, Dikuluwe, 21 Jan. 1960, Duvigneaud 5141D (holo-: BRLU). Chlorophytum decoratum (Baker) Marais & Reilly (Marais & Reilly 1978: 659); Troupin (1988: 44). Type: Zambia, Fwambo, Sep. 1893, Carson 26 (holo-: K). Other representative specimens examined : III: Route de Kenge, Vallée de la Black River, 30 Jul. 1974, Pauwels 5142 (BR); IV: Mayi a Kantshia, Kazumba; savane, Oct. 1978, Mabika 0168 (BR); V: Gandazjika (Kasaï), savane, 24 Feb. 1953, Risopoulos 37 (BR); XI: Katanga, Plateau de la Manika, riv. Potopoto, savane steppique, 24 Dec. 1988, Schaijes 4232; Mine de Luiswishi, 14 Feb. 1982, Malaisse & Robbrecht 2118 (BR). Rwanda: Rusumo, route vers Nyarubuye, savane arbustive sur sol rocailleux, 25 Jan. 1998, Bridson 306 (BR). Burundi: Prov. Bururi, Gitwe (Rumonge), forêt claire à Brachystegia, dans un petit ravin, 24 Feb. 1980, Reekmans 8622 (BR). The species is extremely variable in the size of all its parts. Part of the variation is correlated to habitat and seems to be genetic. The most vigorous forms with a very dense raceme, long tepals (up to 11 mm), big fruits (up to 10 mm) and a compact infrutescence mm wide [Dasystachys verdickii De Wild.; Chlorophytum poggei Engl. ex Poelln.; e.g. Quarré 8045 (BR)] are striking. However, they are linked to normal forms by a range of intermediates. At the opposite extreme of the variation range, slender few-flowered forms from copper-rich soil in Upper Katanga were described as a distinct species (Dasystachys pulchella P.A.Duvign. & Dewit; Chlorophytum linearifolium Marais & Reilly) [Cu-soil: e.g. Malaisse & Robbrecht 2398 (BR, WAG, MO, K, P), Duvigneaud 4670 (BRLU); non mineralized soil: Duvigneaud 5292D (BRLU)]. However, intermediates exist between those and more robust forms even on copper-rich soil and it does not seem possible to maintain that taxon even at varietal rank (Faucon et al. 2010). Some collections from Katanga have ovate-elliptic leaves with length/width ratio unusually low [ mm: de Witte 65 (BR); Malaisse 4907 (BR)]. Small forms with hysteranthous flowers sometimes occur [Quarré 1990 (BR)]. A variant with a ramified inflorescence is rarely encountered [e.g. env. Kibunda, forêt sèche, 28 Apr. 1953, Callens 4074 (K)] corresponding to C. pleiostachyum (Baker) T.Durand & Schinz, but this does not seem to warrant taxonomic recognition. Many collections have purple-spotted leaf sheaths but the taxonomic value of this trait is unclear. 13b. Chlorophytum colubrinum (Baker) Engl. var. upembense Meerts, var. nov. A typo differt racemo sugloboso aut ovoideo, mm longo, cataphyllis pubescentibus et bracteis longe ciliatis. Type:, Haut-Katanga, Parc national de l Upemba, Buyé-Bala, affl. Muyé, savane marécageuse, alt m, 30 Mar. 1948, de Witte (holo-: BR). Other representative specimen examined : XI: Parc nation. Upemba, savane herbeuse d altitude autour de la tête de source de la riv. Kalumengongo, alt m, 31 Mar. 1949, de Witte (BR). This variety is well-characterized by a subcapitate to ovoid inflorescence, long ciliate bracts and hairy cataphylls. It has very narrow leaves (2 4 mm), but this trait is occasionally found in var. colubrinum, especialy in populations on copper-rich soil. 18

19 Meerts & Bjorå, Chlorophytum in Central Africa 14. Chlorophytum cordifolium De Wild. (De Wildeman 1914: 111); De Wildeman (1921a: 27). Type:, Katanga, env. Katentania (Plateau de la Manika, Biano), Nov. 1912, Homblé 766 (holo-: BR, barcode BR ; iso-: BR, barcode BR ). Chlorophytum unifolium Malaisse & Bamps (Malaisse & Bamps 2009: , figs 1 & 2). Type:, Katanga, Kavifwafwaulu, Nov. 2006, Malaisse, Kisimba & Saad 598 (holo-: BR), synon. nov. Other representative specimens examined : XI. Biano Hôtel, Forêt claire à Brachystegia et Monotes sur sol profond et léger, 8 Dec. 1959, Duvigneaud 4459L (BRLU). A remarkable, very distinct species collected only on few occasions since its description in Rediscovered in 2006 it was then erroneously described as a new species. Apparently a narrow endemic of the Tenke-Biano area. 15. Chlorophytum debile Baker (Baker 1878a: 260). Type: Angola, Ambriz, ad rupes, Welwitsch 3769 (holo-: BM; iso-: K, P). Fig. 5G & H. Chlorophytum gilletii Compère (Compère 1962: 211). Chlorophytum gracile De Wild. (De Wildeman 1911a: 274); De Wildeman (1912a: 344) nom. illeg. Type:, Bas-Congo, environs de Kisantu, 1906, Gillet 3963 (holo-: BR), synon. nov. Chlorophytum laxum auct. W Afr. (e.g. Baker 1898: 503; Hepper 1968b: 100; Vanden Berghen 1988: 425, fig. 349; 426) non Chlorophytum laxum R.Br. Other representative specimen examined : III. Territ. Songolo, Kiandu, galerie forestière de la Lunionzo, 16 Dec. 1959, Compère 1049 (BR, K). This species can be confounded with slender forms of C. sparsiflorum. The seed testa however is very different, with smooth periclinal walls in C. sparsiflorum and apiculate periclinal walls in C. debile (fig. 5G & H). The species is very variable in size (6 40 cm high, with inflorescence more or less prostrate in the smallest forms). The type specimen originates from NW Angola, i.e. not far from Congolese collections. Specimens from and the type of C. debile perfectly match collections from W Africa to which the name C. laxum has long been wrongly applied based upon a type specimen from Australia (Poulsen & Nordal 2005). In our opinion, C. debile may be the correct name for most if not all collections of C. laxum auct. from W Tropical Africa. Xylem vessel anatomy also indicated that Asian and African collections of C. laxum auct. are not conspecific (Naik & Nirgude 1981). Donis 177 (BR), cited in Compère (1962), has blackening leaves, an unusual trait for C. debile. As it lacks flowers and fruits, identification is uncertain. 16. Chlorophytum filipendulum Baker (Baker 1878a: 260); Baker (1898: 499); Rendle (1899: 54); Nordal et al. (1997: 59); Poulsen & Nordal (2005: 16, figs. 12 & 13); Bjorå & Nordal (2010: 4). Type: Angola, Golungo Alto, s.d., Welwitsch 3776 (holo-: BM, iso-: K, P). Chlorophytum macrophyllum sensu Robyns & Tournay (1955: 354) non C. macrophyllum (A.Rich.) Aschers. Other representative specimens examined : IV: Kamembele, galerie forestière, 12 Jan. 1957, Liben 2257 (BR); V: Kamalenge, galerie forestière, 10 Feb. 1948, Mullenders 2039 (BR); VI: Yangambi, km 13 route N gazi, sous-bois forêt primitive ombrophile, 12 Jan. 1938, Louis 7466 (BR, K); VII: Mapere- Suronga, forêt, 20 Dec. 1905, Seret 399 (BR); VIII: Kima Kima (Mahagi), alt m, galerie forestière, 27 May 1959, D. Froment 459 (BR); IX: Mutsora, galerie forestière de la Ngokoye, endroit fort ombragé, 2 Aug. 1952, de Witte 7816 (BR). C. filipendulum has been much underrecorded until now due to confusion with C. macrophyllum (see discussion under the latter species). C. filipendulum is closely related to and sometimes difficult to discriminate from C. sparsiflorum. In particular, small forms with incompletely blackening leaves are puzzling. Introgression might explain the existence of more or less intermediate forms. 17. Chlorophytum gallabatense Schweinf. ex Baker (Baker 1876: 325); Baker (1898: 504); Troupin (1955: 235); Troupin (1956: 190); Hepper (1968b: 99); Troupin (1988: 43, fig. 4/2; 44); Kativu (1994: 60); Nordal et al. (1997: 45); Kativu et al. (2008: 81). Type: Sudan/Ethiopian border, Gallabat, Matamma, Jul. 1865, Schweinfurth 10 (lecto-: K; isolecto-: B, P). Three varieties, treated as distinct species in FZ and FTEA. 1. Rachis and pedicels densely covered with hairs or cylindric papillae mm long; lower leaf surface often papillate on veins; fruits c. 7 mm long, generally longer than broad...c. gallabatense var. floribundum 1. Rachis and pedicels glabrous or scabrid with conical papillae < 0.3 mm long; lower leaf surface glabrous; fruits 4 6 mm long, no longer than broad Leaves developing after inflorescence, linear, < 1 cm wide; inflorescence: a raceme or a poorly ramified panicle with 1 2 branches; rootstock with copious fibrous remains of leaves; tubers mm long, mostly terminal...c. gallabatense var. micranthum 2. Leaves well developed at flowering, narrowly elliptic to almost linear, attenuate towards base, mm wide; inflorescence with 3 15 branches; rootstock most often without fibers; tubers < 12 mm, mostly on lateral rootlets...c. gallabatense var. gallabatense 17a. Chlorophytum gallabatense Schweinf. ex Baker var. gallabatense. Fig. 5I & J. Chlorophytum psammophilum Engl. & Gilg (Engler & Gilg 1903: 188). Type: Angola, between Kiubango and Kuito, alt m, 9 Dec. 1899, Baum 517 (holo-: B; iso-: Z, K), synon. nov. Chlorophytum ramulosum De Wild. (De Wildeman 1911a: 275); De Wildeman (1912a: 346; 1921b: 19). Type:, route Bambili-Amadi, 4 Mar. 1905, Seret 244 (holo-: BR). Chlorophytum katangense De Wild. (De Wildeman 1911a: 277); De Wildeman (1913c: 14; 1921a: 28; 1930: 95). Type:, Katanga, Vallée de la Luembe, Feb. 19

20 Pl. Ecol. Evol. FastTrack, 2012 A B C D E F Figure 2 Photographs of living plants (photos P. Meerts except otherwise stated). A, C. clarae, in flower (, Kiswishi, termite mound in miombo, Nov. 2009); B, C. clarae, cultivated in greenhouse (, from Billiet & Jadin 4191; photo F. Billiet); C, C. clarae, in fruit (, Fungurume, Feb. 2010); D, C. vestitum subsp. pilosissimum (, Kipopo, Jan. 2010); E, C. hysteranthum (, Kipopo, Nov. 2009; Meerts & Muding 73); F, C. rubribracteatum (, Kiswishi, Nov. 2009). 20

21 Meerts & Bjorå, Chlorophytum in Central Africa 1910, Hock s.n. (holo-: BR, barcode BR ), synon. nov. Chlorophytum breviflorum De Wild. (De Wildeman 1913a: 513; 1913c: 12); De Wildeman (1921a: 27; 1930: 95). Type:, Lubumbashi, Feb. 1912, Homblé 153 (holo-: BR). Chlorophytum hockii De Wild. (De Wildeman 1913a: 514); De Wildeman (1913c: 13; 1921a: 27). Type:, Katanga, Lubumbashi, Oct. 1911, Hock s.n. (holo-: BR, barcode BR ). Other representative specimens examined : IV: Topo-Guilu-Kwango, savane guinéenne, 28 Mar. 1955, Devred 1713 (BR); V: Kiala, savane ensoleillée, Dec. 1954, Thiébaud 397 (BR); VII: Faradja, Kibale-Ituri, savane arbustive, Jul. 1931, Lebrun 3409 (BR); VIII: Kasengi, Nov. 1935, Brédo 1893 (BR); XI: Plateau de la Manika, Biano, env. Katentania, termitière, Nov. 1912, Homblé 760 (BR); Kimilolo, 10 Feb. 1927, Quarré 128 (BR); Piste Kolwezi-Musokantanda, 0.35 km avant la piste du détournement, S E, 1459 m, sur haute termitière en zone déboisée près d un village, 7 Dec. 1986, Schaijes 3228 (BR, MO). Var. gallabatense is variable. It is often said to have glabrous rachis and pedicels (e.g. Kativu et al. 2008: 39, 82), thus contrasting with the papillate var. floribundum. This is not true, as many specimens of var. gallabatense have a papillate rachis too (e.g. C. katangense De Wild.), but the size and shape of papillae is different (see key) and the distal part of pedicels is usually smooth in var. gallabatense. In FZ, var. gallabatense is said to have long anthers (4 mm), longer than filaments, but materials from have variable anther size (1 3.5 mm), often shorter than filaments. Articulation of pedicel is also variable, from lower half to upper half. FZ recognizes Chlorophytum psammophilum Engl. & Gilg, said to have narrower leaves, shorter pedicels and tepals, stamens longer than perianth (vs. shorter in C. gallabatense) and white petals (vs. greenish in C. gallabatense). Some collections from Katanga resemble the type specimen of that taxon [e.g. Duvigneaud 4455Ch (BRLU)]. However, we have not been able to find reliable diagnostic characters and C. psammophilum is here reduced to C. gallabatense var. gallabatense. 17b. Chlorophytum gallabatense Schweinf. ex Baker var. floribundum (Baker) Meerts, comb. & stat. nov. Chlorophytum floribundum Baker, Bulletin of Miscellaneous Information, Kew 1897: (Baker 1897b); Baker (1898: 505); Kativu (1994: 60); Nordal et al. (1997: 47); Kativu et al. (2008: 82). Type: Malawi, Zomba, Dec. 1896, Whyte s.n. (holo-: K, barcode K , iso-: B, barcode B ). Other representative specimen examined : XI: Vallée de la Lofoi, Lukafu, sol forestier peu ombragé d éboulis de pente, 17 Oct. 1948, Schmitz 2082 (BR). Symoens 12825j (BRVU), from Lubumbashi, may well belong here but the plant is very young. Var. floribundum has never been collected in the fruiting state in and the character of fruit length mentioned in the key is based on material from Zambia. 17c. Chlorophytum gallabatense Schweinf. ex Baker var. micranthum (Baker) Meerts, comb. & stat. nov. Chlorophytum micranthum Baker, Journal of Botany, British and foreign 16: (Baker 1878b); Baker (1898: 507); Troupin (1956: 191); Nordal & Thulin (1993: 269). Type: Sudan, Jur, s.d., Schweinfurth 1745 (holo-: K, iso-: B). Chlorophytum bequaertii De Wild. (De Wildeman 1921b: 14); Robyns & Tournay (1955: 356, Plate XLIX); Hepper (1968b: 102). Type:, Kabare, 22 Aug. 1914, Bequaert 5397 (lecto-: BR, isolecto-: K, here designated) & Bequaert 5410 [syn-: BR (3 sheets)], synon. nov. Other representative specimens examined : VII: Kurukwata (Aba), savane, endroit pierreux, 20 Feb. 1957, Gérard 3100 (BR); VIII: Parc national de la Garamba, crête Congo-Nil, km 10, forêt à Isoberlinia, 4 Mar. 1952, Troupin 286 (BR); IX: env. d Uvira, Plaine de la Rusizi, alt. 800 m, Jan. 1929, Humbert 7307 (BR, P). Burundi: Territ. Bubanza, Gihungwe, steppe à Bulbine, alt. 800 m, 30 Nov. 1974, Reekmans 3954 (BR, K). Rwanda: Parc Nation. Kagera, Runyonza, savane à Acacia et Aloe brûlée, 20 Aug. 1948, Robyns 3434 (BR). All collections from Rwanda previously referred to as C. gallabatense (e.g. in Troupin 1988) belong here. This taxon was treated at species rank by FTEA, but it was indicated to be a small and hysteranthous (fire adapted?) form, not deserving more than subspecific rank, and this view is supported here. Tepal length, often used to discriminate C. micranthum and C. gallabatense, shows broad overlap between the two taxa. Var. micranthum often has yellowish flowers, but this is not a constant character. Collections from the Rusizi plain show fully developed leaves, c. 9 mm wide and cm long (fully developed leaves not observed in FTEA). C. bequaertii was reduced to C. micranthum by Marais & Reilly (1978) and to C. gallabatense by FZ. Narrow leaves, sparingly branched inflorescence, copious leaf remains and terminal tubers suggest it belongs here. The taxonomic significance of tuber position needs further investigation, as tubers are rarely collected. 18. Chlorophytum galpinii (Baker) Kativu var. matabelense (Baker) Kativu (Kativu & Nordal 1993: 63); Kativu (1994: 6); Kativu et al. (2008: 47, fig ). Anthericum matabelense Baker (Baker 1898: 484). Type: Zimbabwe, Bulilamangwe Dist., on banks of Matengwe R., May 1883, Holub s.n. (holo-: K, barcode K ). Anthericum kapiriense De Wild. (De Wildeman 1915: 4); De Wildeman (1921a: 26). Type:, Katanga, Vallée Kapiri, Feb. 1913, Homblé 1092 (holo-: BR). Other representative specimens examined : XI: Plateau de la Manika, savane steppique au nord de la tête de source de la rivière Luilu, 1500 m, 8 Feb. 1987, Schaijes 3322 (BR); Piste Nzilo-Kyamasumba à 10.5 km par la piste du centre urbain de Nzilo, forêt claire, 01 Aug. 1987, Schaijes 3575 (BR). C. galpinii var. galpinii was reported by error from by FZ (Kativu et al. 2008) (see discussion under C. cameronii var. grantii). 19. Chlorophytum geophilum Peter ex Poelln. (Poellnitz 1943b: 127; 1946: 292); Hepper (1968b: 100); Kativu (1994: 21

22 Pl. Ecol. Evol. FastTrack, 2012 A B C E D F Figure 3 Photographs of living plants (photos P. Meerts except otherwise stated). A, C. pusillum (cultivated in greenhouse, from Billiet & Jadin 4151; photo F. Billiet); B, C. sphacelatum (, Kiswishi; photo J. Piqueray); C, C. stenopetalum var. latifolium (, Kiswishi; photo J. Piqueray); D, C. subpetiolatum var. pilosifolium (, Lubumbashi, Kasapa, termite mound, Nov. 2009); E, C. velutinum (, Fungurume, copper hill, Feb. 2010); F, C. zingiberastrum (cultivated in greenhouse, from Billiet & Jadin 4195; photo F. Billiet). 22

23 Meerts & Bjorå, Chlorophytum in Central Africa 63); Nordal et al. (1997: 54, fig. 11); Kativu et al. (2008: 87, fig ). Type: Tanzania, Tabora Dist., Unyamwesi, Kombe, 24 Jan. 1926, Peter (holo-: B). Fig. 6A. Chlorophytum pusillum sensu Troupin (Troupin 1956: 191). Other representative specimens examined : VII: Parc Nation. Garamba, savane arbustive, 5 Jun. 1950, Noirfalise 441 (BR). XI: Bunkeya, plaine, Dec. 1948, Hoffmann 889 (BR). C. geophilum and C. pusillum are two closely related species and intermediate specimens exist. The collections from the Parc National de la Garamba are critical. 20. Chlorophytum hirsutum Poulsen & Nordal (Poulsen & Nordal 1999: 941). Type: Uganda, Bundibugyo Distr., Bwamba County, near Sempaya, 22 Jul. 1938, Thomas 2311 (holo-: K; iso-: BR). Other representative specimens examined : IX: Territ. Uvira, Lubarika, savane à Loudetia, 29 May 1959, A. Léonard 4463 (BR). Burundi: Territ. Bubanza, alt m, bord de forêt, Dec. 1967, Lewalle 2549 (BR). 21. Chlorophytum hysteranthum Kativu (Kativu & Nordal 1993: 63); Kativu (1994: 64); Kativu et al. (2008: 49). Anthericum breviscapum De Wild. (De Wildeman 1913a: 507); De Wildeman (1913c: 7; 1921a: 26), non Chlorophytum breviscapum Dalzell or Chlorophytum breviscapum Dammer. Type:, Katanga, Lubumbashi, Sep. 1911, Hock s.n. (holo-: BR, barcode BR ). Fig. 1K. Other representative specimens examined : XI: Kipopo, 25 km NO Lubumbashi, dembo d argile grise alternativement marécageuse et très sèche, 28 Sep. 1952, Schmitz 4093 (BR); ibid., Nov. 2009, Meerts & Muding 73 (BRLU). A rare species with a restricted distribution range in S Katanga and N Zambia. The few collections from all come from a restricted area in the vicinity of Lubumbashi. Rogers (K) has unusually large leaves up to 20 cm long suggesting that C. hysteranthum is closely related to C. affine. 22. Chlorophytum lancifolium Baker (Baker 1878a: 260); Baker (1898: 498); Kativu (1994: 65); Nordal et al. (1997: 60); Poulsen & Nordal (2005: 16-17); Kativu et al. (2008: 83). Type: Angola, Pungo Andongo, Nov. 1856, Welwitsch 3772 (lecto-: BM, isolecto-: K); Welwitsch 3773 (syn-: BM, K). 1. Seed testa apiculate...c. lancifolium subsp. cordatum 1. Seed testa not apiculate Leaf base most often attenuate, flowers greenish to yellowish...c. lancifolium subsp. togoense 2. Leaf base most often truncate to cordate, flowers whitish...c. lancifolium subsp. lancifolium 22a. Chlorophytum lancifolium Baker subsp. lancifolium [Poulsen & Nordal (2005: 17, figs 1J; 18, 19, 35)]. Other representative specimens examined : XI: Sakania, forêt claire mélangée sur terre rouge, 31 Jan. 1960, Duvigneaud 5357 (BRLU). Burundi: Ruyigi, colline Nyabitangu, sous-bois de forêt à Julbernardia, 8 Feb. 1979, Reekmans 7661 (BR, K). Subsp. lancifolium has been much less collected than the other two subspecies. 22b. Chlorophytum lancifolium Baker subsp. cordatum (Engl.) Poulsen & Nordal (Poulsen & Nordal 2005: 16, fig. 1K; 16, 17, 34). Chlorophytum cordatum Engl. (Baker & Engler 1892: 468). Type: Soudan, Jebel Bangenze, Niamniamland, am Baginse, 25 May 1870, Schweinfurth 173 (holo-: B). Other representative specimens examined : VII: Montagne Kundungu, 560 m, village Masabe, à 40 km au N de Bondo, forêt dense primaire sur rocher granitique, 3 Jul. 1978, Lejoly 4118 (BRLU, BR, K, WAG); Parc nation. Garamba, Dedegwa, galerie à la tête de source de la Dedegwa, 17 May 1952, De Saeger 2606 (BR, K); VIII: Bunia, Mont Hoyo, ravin, sur rochers humides, 5 Aug. 1976, Pauwels 5751 (BR); XI: Route Kolwezi-Kyamasumba, rivière Mushingi, 19 Feb. 1987, Billiet & Jadin 4192 (BR; cultivated in greenhouse: S 1945). Subsp. cordatum mostly occurs in NE Congo (Poulsen & Nordal 2005); however, the record from Upper-Katanga [Billiet & Jadin 4192 (BR)] (seed testa checked with SEM) is widely disjunct from the main area of the subspecies indicating that subsp. lancifolium and subsp. cordatum are not completely allopatric. 22c. Chlorophytum lancifolium Baker subsp. togoense (Engl.) Poulsen & Nordal (Poulsen & Nordal 2005: 17, fig. 1L; 20, 21, 36). Chlorophytum togoense Engl. (Engler 1902: 92); Hepper (1968b: 99). Type: Togo, Jaggebach, bei Misa Höhe, 5 Sep. 1890, Büttner 172 (lecto-: B, here designated); Togo, Fasugu, 30 May 1891, Büttner 665 (syn-: B). Chlorophytum ealaense De Wild. (De Wildeman 1911a: 274); De Wildeman (1912a: 343; 1916: 164; 1921b: 15). Type:, Eala, Oct. 1907, Pynaert 1040 (lecto-: BR; isolecto-: B);, Eala, s.d., Pynaert 557 (syn-: BR);, Eala, s.d., Pynaert 1777 (syn-: BR). Other representative specimens examined : VI: Prov. Equateur, près de la rivière Ikilemba, 26 Aug. 1914, Nannan 63 (BR, K). Subsp. togoense has a western Guineo-Congolian distribution, having its eastern limit in W in the region of Eala. Some collections [(e.g. Gombe (Equateur), Aug. 1903, M. Laurent s.n. (BR, barcode BR )] are more or less intermediate between C. lancifolium and C. sparsiflorum, and might be of hybrid origin. C. lancifolium is often said to have a scabrid leaf margin (Nordal et al. 1997, Nordal & Poulsen 2005), but in most collections examined leaf margin is smooth. 23. Chlorophytum leptoneurum C.H.Wright var. katangense (De Wild.) Meerts, comb. & stat. nov. Verdickia katangensis De Wild. Annales du Musée du Congo, Botanique, série 4,1: (De Wildeman 1902); Engler (1908: 309); Durand & Durand (1909: 569); De Wildeman (1921a: 27). Type:, Katanga, Lukafu, terrains 23

24 Pl. Ecol. Evol. FastTrack, 2012 Figure 4 Scanning electron microscope images of seeds and seed testa. A & B, Chlorophytum africanum var. nordalianum, seed and seed testa (Schmitz 4261); C, C. andongense (short-fruited form), seed testa (Schmitz 4545); D, C. andongense (long-fruited form), seed testa (Billiet & Jadin 4150); E & F, C. angustissimum, seed and seed testa (Lewalle 6452); G & H, C. arcuatoramosum, seed and seed testa (Mullenders 142); I & J, C. cf. brachystachyum, seed and seed testa (Quarré 1950). A C B D E F G H I J 24

25 Meerts & Bjorå, Chlorophytum in Central Africa A B Figure 5 Scanning electron microscope images of seeds and seed testa. A & B, C. clarae, seed and seed testa (Meerts 2010/37). C & D, C. macrophyllum, seed and seed testa (Schimper 1837); E & F, C. macrophyllum, seed (Allard 370) and seed testa (Vanderyst 38806); G & H, C. debile, seed and seed testa (Compère 1049); I & J, C. gallabatense var. galabatense, seed and seed testa (Homblé 1150). C D E F G H I J 25

26 Pl. Ecol. Evol. FastTrack, 2012 argileux, Dec. 1899, Verdick 329 (holo-: BR); non Chlorophytum katangense De Wild. Fig. 6C, D & I. Other representative specimens examined : XI: Keyberg, Katuba, prairie sur sol superficiel à dalle latéritique, 28 Nov. 1947, Schmitz 1052 (BR); pâturage artificiel du dembo de la Katuba, Jan. 1934, Quarré 3741 (BR). Other specimens examined Malawi: N. Prov., Mzimba dist., 6 miles NE Mzambazi, edge dambo, very sandy, 22 Jan. 1978, Pawek (BR, K). Zambia: dist. Luanshya, 23 Jan. 1956, Fanshawe 2708 (BR, K, NDO) [cited by Kativu et al. (2008) as C. macrorhizum]. The three collections from are indisputably members of the C. leptoneurum-c. macrorhizum complex. A striking finding is the observation of hairs on the fruits [a mixture of obtuse flattened papillae c mm high on faces and elongate acute hairs up to 0.7 mm long on angles (fig. 6I)], apparently a unique condition in African species of Chlorophytum. This observation prompted us to check fruits of collections stored under C. leptoneurum and C. macrorhizum in BR. Fanshawe 2708 from Zambia, and Pawek from Malawi proved to have the same fruits and very similar habit compared to the Congolese collections. These five collections share many traits and obviously represent a distinct variety with an original combination of traits in the complex. These collections also highlight the narrow affinities between C. leptoneurum and C. macrorhizum. They have a striking horizontal rhizome, but of variable thickness from very thin and leptoneurum-like to thick and more or less macrorhizum-like. Leaves and bracts often have a reddish tinge, a trait said to be typical of C. macrorhizum. Seed testa has convex periclinal walls, without spines (fig.6 C & D). See also note under C. africanum var. nordalianum. 24. Chlorophytum longifolium Baker (Baker 1876: 327); Baker (1898: 507); Kativu (1994: 66); Nordal et al. (1997: 17); Kativu et al. (2008: 63). Type: Sudan/Ethiopian border, Gallabat, Matamma, Aug. 1865, Schweinfurth 8 (lecto-: P, designated here). Chlorophytum papillosum Rendle (Rendle 1895: 422); De Wildeman (1921a: 28); Nordal et al. (1990: 547 fig. 5e, 548 fig. 6E). Type: Tanzania, Between Zanzibar and Uyui, 1886, Taylor s.n. (holo-: BM, barcode BM ). This species is represented in by a variety with golden-yellow pubescence on young flower buds. 24a. Chlorophytum longifolium Baker var. aureum (Engl.) Meerts, comb. & stat. nov. Chlorophytum aureum Engl., Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflan zengeographie 15: (Baker & Engler 1892); Troupin (1955: 234; 1956: 189); Hepper (1968b: 99). Type: Sudan, Niamniamland, river Huuh, 21 May 1870, Schweinfurth 3752 [lecto-: B (2 sheets: B & B ), here designated; isolecto-: K]; Sudan, Dschurland, bei der grossen Seriba Kutschuk Ali, 29 Apr. 1896, Schweinfurth 1504 (syn-: K); Wando, steppen in der Wälden, s.d., Schweinfurth 3570 (syn-: K). Other representative specimens examined : VII: Parc Nation. Garamba, savane arbustive, 26 May 1950, Noirfalise 429 (BR); Uele, Guvane, savane arbustive, May 1931, Lebrun 2881 (BR, K). This variety has a Sudanian distribution, while the typical variety is mostly Zambezian. 25. Chlorophytum macrophyllum (A.Rich.) Aschers. (Schwein furth 1867: 294); Baker (1876: 323); Baker (1898: 498); Engler (1908: 308, fig. 208 C & D); Kativu (1994: 67); Nordal et al. (1997: 49, fig. 9); Bjorå et al. (2008: 233, fig. 6, 10, 11); Kativu et al. (2008: 85). Anthericum macrophyllum A.Rich. (Richard 1851: 334). Type: Ethiopia, Djeladjekanne, Jun. 1840, Quartin-Dillon s.n. (lecto-: P, barcode P ); Ethiopia, Djeladjeranne, 24 Oct. 1840, Schimper 1837 (syn-: P, isosyn-: BR, B, MPU). Fig.5 E & F. Chlorophytum fuchsianum De Wild. (De Wildeman 1904: 102 as fuchsianus); De Wildeman (1905: 38 & Plate XVI; 1909: 54 & Plate 1, figs 1 & 2); Durand & Durand (1909: 568). Type:, Kisantu, s.d., Gillet 902 (lecto-: BR). Other representative specimens examined : III: Bas-Congo, Mbanza Ngungu, entrée de la petite grotte, 3 Mar. 1987, Billiet & Jadin 4264 (BR); Mvuasi, îlot forestier, 17 Dec. 1947, Devred 4 (BR, K); VI: Eala, s.d., Goossens s.n. (BR). Burundi: Ubimbi, prairie, 8 Dec. 1965, Lewalle 63 (BR). C. macrophyllum has long been misinterpreted in. Our revision shows that actually only few collections belong to that species, nearly all originating from SW. Most other records of that species actually belong in C. filipendulum. At the flowering stage, C. macrophyllum and C. filipendulum cannot be confounded. At the fruiting stage, C. filipendulum is recognized by its blackish colour in herbarium (pale brownish to brownish-green in C. macrophyllum), papillate rachis (smooth in C. macrophyllum), and strongly saucer-shaped seeds (disk-shaped to slightly saucer-shaped in C. macrophyllum). The name C. macrophyllum has been applied to plants with large flowers (> 8 mm long), with 5-nerved tepals and long anthers (> 3 mm) and large whitish bracts. We have found that two closely related taxa with that combination of traits have been confounded in until now. i.e. C. macrophyllum and C. clarae. See discussion under the latter species. Fig. 5C F shows that seed shape and seed coat details are very similar for a syntype of C. macrophyllum from Ethiopia and for a number of collections from W. 26. Chlorophytum minor Kativu (Kativu 1993: 502); Kativu (1994: 68); Kativu et al. (2008: 81). Type: Zambia, Mbala Dist., Itembwe Gap and Gorge, 7 Jan. 1968, Richards (holo-: K). Other representative specimens examined : XI: Route Mitwaba-Manono, km 45, riv. Kalumengongo, 3 Feb. 1986, Bamps & Malaisse 8619 (BR). A rare species, restricted to SW Tanzania, N Zambia and Katanga. 27. Chlorophytum nubicum (Baker) Kativu (Kativu & Nordal 1993: 63); Kativu (1994: 69); Nordal et al. (1997: 15); Kativu et al. (2008: 55). Anthericum nubicum Baker (1876: 26

27 Meerts & Bjorå, Chlorophytum in Central Africa 301); Baker (1898: 484). Type: Sudan, Jur, White Nile, Nyangara, s.d., Petherick s.n. (holo-: BM, n.v.). Other representative specimens examined : XI: ancienne piste Kolwezi-Likasi, S E, savane arborescente en forêt claire, alt m, 21 Aug. 1986, Schaijes 3051 (BR). Two collections from l Upemba [de Witte & (BR)] key out here but are atypical, having a short peduncle with only 1( 2) bracts and inflorescence nodes with only one flower. They probably represent a distinct taxon, but more material, especially with fruits, is needed. C. nubicum is very close to C. nidulans. The former can be distinguished by spindle-shaped tubers and anthers > 2 mm. 28. Chlorophytum occultum Poulsen & Nordal (Poulsen & Nordal 1999: 944, figs. 2C F, 3, 4); Poulsen & Nordal (2005: 17, fig. 1G; 22, 23, 37). Type: Ouganda, Masindi Dist., Budongo Forest Reserve, 2 Oct. 1995, Poulsen, Nkuutu & Dumba 975 (holo-: C, iso-: MHU, EBB; not seen). Other representative specimens examined Not seen. Key after description in Poulsen & Nordal (2005). 29. Chlorophytum orchidastrum Lindl. (Lindley 1824: 79); Baker (1898: 500); Hepper (1968b: 101, 102 fig. 351); Panigrahi (1975: 563); Poulsen & Nordal (2005: 18, fig. 1M, 24); Bjorå & Nordal (2010: 5). Type: Cultivated at the garden of the Horticultural Society, from a plant sent from Sierra Leone, in 1822, by their collector Mr. George Don H.HS., 1824 (holo-: CGE). Chlorophytum seretii De Wild. (De Wildeman 1909: 56; Plate 2 & 3); De Wildeman (1921b: 21). Type:, env. de Gombari, 26 Apr. 1906, Seret 577 (holo-: BR). Chlorophytum seretii De Wild. var. likimensis De Wild. (De Wildeman 1911b: 289); De Wildeman (1912b: 526). Type:, env. de Likimi, 21 Apr. 1910, Malchair 262 (holo-: BR). Other representative specimens examined : II: Territ. Seke-Banza, Gimbi, vieille forêt secondaire à Terminalia superba, 15 Sep. 1959, Compère 384 (BR, K); III: Gimbi, jachères secondaires humides, 15 Jun. 1948, J. Laurent 691 (BR); V: Région de Pene-Yumbi, sous-bois de forêt primitive, Jun. 1952, Germain 7810 (BR); VI: Panga, forêt humide sur terre riche en humus, 16 Dec. 1913, Bequaert 1500 (BR); VIII: Irumu, galerie forestière dans la savane, 13 Mar. 1914, Bequaert 2966 (BR). 30. Chlorophytum perfoliatum Kativu (Kativu 1993: 503); Kativu (1994: 71); Kativu et al. (2008: 69, Fig ). Type: Zambia, Kasama Dist., 95 km E of Kasama, among rocks, s.d., Robinson 4344 (holo-: K). Other representative specimen examined : XI: Tilwizembe, végétation d une termitière dans la plaine au nord du gisement, 15 Dec. 1959, Duvigneaud 4601T (BRLU). A rare species restricted to Zambia, Malawi and Katanga, with only one collection in. 31. Chlorophytum pilosicarinatum (Poelln.) Meerts, comb. nov. Anthericum pilosicarinatum Poelln., Boletim da Sociedade Broteriana. XVII, 2A serie: (Poellnitz 1943a). Type: Angola, Bié, Ufer des Cubango, Vila da Ponte, 1906, Gossweiler 4018 (lecto-: K, here designated). Other representative specimen examined : XI: Tilwizembe, alignement métallifère rocheux, steppe sur schiste cobaltifère, 15 Dec. 1959, Duvigneaud 4590L1 (BRLU). The other syntypes, Gossweiler 2039 & 2663, have not been found. The cited collection is apparently the first record of that taxon since its description. C. pilosicarinatum belongs in the C. cameronii complex and might best be treated at varietal rank within this latter species, but more material is needed. It is easily recognized by its unique pattern of leaf pubescence, with glabrous leaf surface and long ciliate leaf margins and lower mid nerve; the hairs are retrorsely oriented at their base; bracts have the same pattern of pubescence in the examined collection, but less clearly so in the type specimen. Chlorophytum galpinii var. norlindhii has a more or less similar pattern of pubescence, but in the latter taxon the inflorescence is lax and branched (vs. unbranched, < 5 cm long in C. pilosicarinatum), bracts are attenuate in a long fine point (vs. bracts ovate without a long point), tepals are 3-veined (vs. 5-nerved) and pedicels are terete (vs. narrowly winged in the upper part). Mortelmans 142 (BR) may belong here: it is a dwarf plant with three very narrow cataphyll-like leaves with the characterictic pattern of pubescence of the species. 32. Chlorophytum polystachys Baker (Baker 1878b: 326; 1898: 509); Hepper (1968b: 496); Kativu (1994: 71); Nordal et al. (1997: 43); Kativu et al. (2008: 61). Type: Sudan, Jur Ghattas, 2 Jul. 1869, Schweinfurth 1838 (holo-: K, 2 sheets, iso-: B, 3 sheets). Other representative specimens examined : XI: Lubumbashi, dembo de la Kiboko, 20 Nov. 1957, Schmitz 6037 (BR). Apparently a rare species with a single collection in the study area. This specimen is remarkable by its big fusiform distal tubers, a character rarely reported for that species. 33. Chlorophytum pubiflorum Baker (Baker 1876: 329); Baker (1898: 509); Kativu (1994: 73); Nordal et al. (1997: 44). Type: Mozambique, Delta du Zambèze, Dec. 1862, Kirk s.n. (holo-: K, barcode K ). Other representative specimens examined : XI: Musosa, 28 Feb. 1940, Brédo 4019 (BR). The single collection examined originates from the Congo-Zambia borderline, possibly on Zambian territory. The species should be sought for in the floodplain of the Luapula River. 34. Chlorophytum pusillum Baker (Baker 1878b: 325); Baker (1898: 502); Hepper (1968b: 100); Kativu (1994: 73); Nordal et al. (1997: 55); Kativu et al. (2008: 87, 88, fig ); non C. pusillum sensu Troupin (1956: 191). Type: Sudan, Jur, 10 Jul. 1869, Schweinfurth 2043 (holo-: K, iso-: B (3 sheets), P, PRE). Figs 3A & 6B. 27

28 Pl. Ecol. Evol. FastTrack, 2012 Other representative specimens examined : XI: Lubumbashi, arboretum de l Etoile, sur termitière, 17 Jan. 1948, Schmitz 1241 (BR); Mont Mamuntamba, 1987, Billiet & Jadin 4151 (BR, MO, P; cultivated in greenhouse: S 2072, Photo ). Specimens from have papillate bracts, which had not been noticed previously. Seed testa has raised periclinal walls which are verrucose (fig. 6B) and distinct from the convex periclinal walls of C. geophilum (fig. 6A). See also under that species. 35. Chlorophytum recurvifolium (Baker) C.Archer & Kativu (Archer & Kativu 2001: 31); Kativu et al. (2008: 17). Anthericum recurvifolium Baker (Baker 1906: 28). Type: Zimbabwe, 6-miles spruit, near Harare, s.d., Cecil 143 (holo-: K). Anthericum longistylum Baker (Baker 1876: 305); Obermeyer (1962: 686, 687: Fig. 2). Type: South Africa, Gold fields, 1870, Baines s.n. (holo-: K, barcode K ). Other representative specimens examined : XI: Kisenge, colline de Mn, bosquet d Uapaca robynsii, 1956, Duvigneaud 2342Li (BRLU); s.l. Mn, s.d., Joris D30 (BRLU); Bunkeya, Dec. 1942, s. rec. 457 (BRLU). A phytogeographically very interesting record, as this species has a southern African distribution, reaching its northern limit in Zimbabwe. Katangan populations are widely disjunct. 36. Chlorophytum ruahense Engl. (Engler 1900: 361); Poellnitz (1946: 332); Kativu (1994: 75); Nordal et al. (1997: 13); Kativu et al. (2008: 74, fig ). Type: Tanzania, Iringa Dist., Uhehe, hanging cliff on Ruaha R., 10 Jan. 1899, Goetze 461 (holo-: B). Other representative specimens examined : XI: Baya-Pierkat, végétation d une colline rocheuse de calcaires de la Mofya, 7 Jan. 1960, Duvigneaud 4953L1 (BRLU); Route Nguba- Mokabe Kasadi, à 2 km au S de Kate, forêt claire, 7 Nov. 1988, Pauwels 7211 (BR). Katangan collections extend the distribution area of this rare species, otherwise confined to SW Tanzania and N Zambia, somewhat to W. The seeds are echinulate (seed not seen in FZ). The leaves of the Congolese collections are variable in width. 37. Chlorophytum rubribracteatum (De Wild.) Kativu (Kativu & Nordal 1993: 64); Kativu (1994: 75); Nordal et al. (1997: 36); Kativu et al. (2008: 50). Anthericum rubribracteatum De Wild. (De Wildeman 1913a: 508; 1913c: 9); De Wildeman (1921a: 27). Type:, Lubumbashi, Nov. 1911, Hock s.n. (holo-: BR, barcode BR ). Fig. 2F. Anthericum roseum Poelln. (Poellnitz 1942: 114); Lewalle (1972: 173). Type: Tanzania, Songea dist., E of Ungoni, Mampyuy, s.d., Busse 727 (holo-: B, not seen; iso-: EA). Other representative specimens examined : XI: Luiswishi, colline cuprifère, 18 Nov. 1980, Malaisse (BR, WAG, MO); Kipopo, dembo sur plateau latéritique, 30 Jan. 1962, Schmitz 7570 (BR). Burundi: Territ. Bubanza, ferme de la Randa, savane boisée, s.d., Lewalle 1254 (BR). The species is well characterized by the red-coloured cataphylls, narrowly graminiform leaves, and lax inflorescence often with remote lower nodes. Bracts are generally carinate, erect, and often clasping the peduncle. However, C. rubribracteatum is much more variable than mentioned in FZ and FTEA, especially in intensity of red coloration (often lacking on bracts), pubescence (from glabrous to shortly pubescent on lower leaf surface and peduncle), height (5 80 cm) and leaf width (1 10 mm). Size tends to be negatively correlated with pubescence. In Katanga, the largest specimens occur in tall vegetation on moist soil (e.g. Duvigneaud 5055 (BRLU)). All collections from Burundi also belong to the robust form. More research is needed to decide if robust forms deserve taxonomic recognition. Some collections with broad leaves, pale coloration and dense pubescence on lower leaf surface and peduncle approach C. cameronii var. grantii [e.g. Van Meel 805 (BR); Schmitz 3633 (BR); Duvigneaud 4769C & 4772A (BRLU)]. Robyns 1646 (BR) is an extreme dwarf form with prostrate peduncle, which may belong here. More material is needed. 38. Chlorophytum sparsiflorum Baker (Baker 1876: 325); Baker (1898: 498); Hepper (1968b: 100); Troupin (1988: 43, fig. 4/1; 44); Bjorå & Nordal (2010: 6). Type: Equatorial Guinea, Fernando Po, Apr. 1860, Mann 388 (holo-: K). Chlorophytum semlikiense De Wild. (De Wildeman 1921b: 20); Robyns & Tournay (1955: 356). Type:, Kasonsero, Vallée de Semliki, s.d., Bequaert 5041 (holo-: BR). Chlorophytum beniense De Wild. (De Wildeman 1921b: 13); Lebrun et al. (1948: 36); Robyns & Tournay (1955: 354); Troupin (1988: 42; 45, fig. 15/2). Type:, Mayolo (Beni), s.d., Bequaert 3981 (holo-: BR). Chlorophytum elongato-fusiforme De Wild. (De Wildeman 1921b: 15). Type:, en aval de Panga, 15 Dec. 1913, Bequaert 1485 (holo-: BR). Chlorophytum ituriense De Wild. (De Wildeman 1921b: 17); Troupin (1956: 191). Type:, Penghe, 17 Feb. 1914, Bequaert 2513 (syn-: BR), Irumu, 12 Mar. 1914, Bequaert 2398 (syn-: BR). Chlorophytum butaguense De Wild. (De Wildeman 1921b: 19); Robyns & Tournay (1955: 354). Type:, Ruwensori (Butagu), 15 Apr. 1914, Bequaert 3686 (lecto-: BR, barcode BR , here designated); Ruwensori, 16 May 1914, Bequaert 4292 (syn-: BR). Chlorophytum kirkii sensu Robyns & Tournay (1955: 356) non Chlorophytum kirkii Baker. Chlorophytum comosum sensu Baker (1876: 329); Baker (1897a: 400); Troupin (1988: 42; 45, fig. 15/3); Nordal et al. (1997: 55); Poulsen & Nordal (2005: 15), Kativu et al. (2008) non Chlorophytum comosum (Thunb.) Jacques. Other representative specimens examined : II: Luki, est du village de Kimbuya, association végétale sur le replat du somet du rocher de granit, lisière de la forêt à Gilletiodendron, 28

29 Meerts & Bjorå, Chlorophytum in Central Africa A B C D E F G H I Figure 6 Scanning electron microscope images of seeds and seed testa. A, C. geophilum, seed testa (Noirfalise 441); B, C. pusillum, seed testa (Billiet & Jadin 4151); C & D, C. leptoneurum var. katangense, seed and seed testa (Quarré 3741); E & F, C. stenopetalum var. stenopetalum, seed and seed testa (Seret 196); G & H, C. stenopetalum var. latifolium, seed and seed testa (Detilleux 570); I, C. leptoneurum var. katangense, fruit (Quarré 3741). 2 Aug. 1957, Wagemans 1594 (BR); IV. Kwango, Kisantu, Panzi, 1925, Vanderyst (BR); VI: Prov. Orientale, Territ. Mambasa, rivière Lemda, région Ndesa, 700 m, clairière humide dans forêt à Gilbertiodendron dewevrei, 8 Jun. 1956, Christiaensen 1763 (BR); Yangambi, au pied du plateau de l Isalowe, alt. 470 m, forêt primitive ripicole le long du fleuve Congo, 18 May 1938, Louis 9437 (BR); Rivière Dhi, pont de l ancienne route Djugu-Nioka, sous-bois ombragé humide de la forêt galerie, 19 Apr. 1991, Dhtechuvi 1128 (BR); VIII: Mont Aboro, pied du versant W, alt m, forêt de montagne à Chrysophyllum fulvum, 15 Apr. 1958, Bamps 154 (BR); Ituri, Djugu (Kibali), alt m, forêt, Sep. 1931, Lebrun 3942 (BR); IX: Territ. Kabare, Kahusi, le long d un caniveau, alt m, 11 Feb. 1959, A. Léonard 2983 (BR); XI: Parc Nation. Upemba, Katongo, alt m, galerie forestière, 16 Apr. 1948, de Witte 3713 (BR). Rwanda: Gisakura (Cyangugu), alt m, plantation de thé, 23 Jul. 1954, Auquier 3330 (BR); Territ. Shangugu, Rugege, alt m, forêt de montagne, 21 Mar. 1956, Christiaensen 1488 (BR). Burundi: Territ. Bururi, Rumonge, alt m, talus ombreux, 16 Mar. 1969, Lewalle 3338 (BR). This taxon has been known as C. comosum auct. A recent molecular phylogenetic study (Bjorå et al. 2008) has shown that this taxon is not monophyletic. In particular, accessions from C Africa and S Africa belong in different clades. Since the type specimen originates from S Africa, the epithet comosum can no longer be applied to C African material. C sparsiflorum is an extremely polymorphic taxon. A multivariate statistical analysis (Poulsen & Nordal 2005) concluded that three infraspecifc taxa can be recognized, i.e. one with strap-shaped, blackening leaves ( Chlorophytum comosum var. comosum ), a second with elliptic, non blackening, leaves more or less attenuate into a pseudopetiole ( var. sparsiflorum ) and a third with elliptic, non blackening leaves, abruptly contracted into a petiole ( var. bipindense ). We have not been able to settle the objective limits of those taxa in the abundant material examined. Leaf width and shape and colour in herbarium (blackening or not) show an almost continuous pattern of variation in material from Central Africa. Therefore, the placing of many collections is problematic. Molecular markers may hopefully provide new insight into this taxonomic conundrum. 29