A synopsis of the genus Ficus (Moraceae) in southern Africa

A synopsis of the genus Ficus (Moraceae) in southern Africa

S.AfrJ.Bot., 1990,56(6): 599-630 599 A synopsis of the genus Ficus (Moraceae) in southern Africa J.V. van Greuning Department of Botany, University ...

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S.AfrJ.Bot., 1990,56(6): 599-630

599

A synopsis of the genus Ficus (Moraceae) in southern Africa J.V. van Greuning Department of Botany, University of Pretoria, Pretoria, 0002 Republic of South Africa Accepted 15 August 1990

The genus Ficus L. in southern Africa is delimited into infrageneric taxa. Keys to subgenera, sections, subsections and species, as well as a brief description of each species are given. Distribution maps, line drawings, notes on habitat and a concise synonymy are supplied for each species. Subgenus Ficus L. is represented by the section Sycidium Miq. with species F. capreifo/ia Del. and F. pygmaea Welw. ex Hiern. Ficus sycomorus L. and F. sur Forssk. comprise subgenus Sycomorus (Gasp.) Mildbr. & Burr. The remaining species belong to subgenus Urostigma (Gasp.) Miq. and include the following: F. cordata Thunb. subsp. cordata, F. cordata subsp. sa/icifo/ia (Vahl) C.C. Berg, F. ingens (Miq.) Miq. and F. verrucu/osaWarb. in sect. Urostigma; F. /utea Vahl in sect. Ga/og/ychia (Gasp.) Endl. subsect. Ga/og/ychia; F. abuti/ifo/ia (Miq.) Miq., F. g/umosa Del., F. stuh/mannii Warb., F. tettensis Hutch. and F. trichopoda Bak. in sect. Ga/og/ychia subsect. P/atyphy//ae (Mildbr. & Burr.) C.C. Berg; F. burtt-dat-yiHutch., F. craterostoma Warb. ex Mildbr. & Burr., F. fischeri Warb. ex Mildbr. & Burr., F. i/icina (Sond.) Miq., F. nata/ensis Hochst. subsp. nata/ensis and F. thonningiiBlume in sect. Ga/og/ychiasubsect. Ch/amydodorae(Mildbr. & Burr.) C.C. Berg; F. bizanaeHutch. & Burtt-Davy, F. bubuWarb., F. sanslbarica Warb. subsp. sansibarica, F. po/ita Vahl subsp. po/ita and F. tremu/a Warb. subsp. tremu/ain sect. Ga/og/ychiasubsect Cau/ocalpae(Mildbr. & Burr.) C.C. Berg. Die genus Ficus L. in Suider-Afrika is geklassifiseer in infrageneriese taksons. Sleuteis tot die subgenusse, seksies, subseksies en spesies sowel as 'n kort beskrywing van elke spesie word gegee. Verspreidingskaarte, Iyntekeninge, habitatgegewens en 'n verkorte sinonimie word vir elke spesie verskaf. Subgenus Ficus L. word deur die seksie Sycidium Miq. met die spesies F. capreifo/ia Del. en F. pygmaea Welw. ex Hiern verteenwoordig . Ficus sycomorus L. en F. sur Forssk. verteenwoordig die subgenus Sycomorus (Gasp.) Mildbr. & Burr. Die oorblywende spesies behoort tot die subgenus Urostigma (Gasp.) Miq. en sluit die volgende in: F. cordata Thunb. subsp. cordata, F. cordata subsp. sa/icifo/ia (Vahl) C.C. Berg, F. ingens (Miq.) Miq. en F. verrucu/osaWarb. (seksie Urostigma); F. /utea Vahl [seksie Ga/og/ychia (Gasp.) Endl. subseksie Ga/og/ychia] ; F. abuti/ifo/ia (Miq.) Miq., F. g/umosa Del., F. stuh/manmiWarb., F. tettensis Hutch. en F. trichopoda Bak. [seksie Ga/og/ychia subseksie P/atyphy//ae (Mildbr. & Burr.) C.C. Berg] ;F. burtt-davyiHutch., F. craterostoma Warb., F. fischeriWarb. ex Mildbr. & Burr., F. i/icina (Sond.) Miq., F. nata/ensis Hochst. subsp. nata/ensis en F. thonningli"Blume [seksie Ga/og/ychia subseksie Ch/amydodorae (Mildbr. & Burr.) C.C. Berg] ;F. bizanae Hutch. & Burtt-Davy, F. bubuWarb., F. sansibaricaWarb. subsp. sansibarica, F. po/ita Vahl subsp. po/I~a en F. tremu/a Warb. subsp. tremu/a[seksie Ga/og/ychiasubseksie Cau/ocalpae(Mildbr. & Burr.) C.C. Berg] . Keywords: Ficus, Moraceae, southern Africa, taxonomy

Introduction Taxonomically the genus Ficus L. is a large and complex one. In southern Africa it is well represented and notorious for taxonomic problems, particularly at the species level. This brief taxonomic review of the genus Ficus in southern Africa emanates from a doctoral dissertation on the group (van Greuning 1982). The principal aim of the present paper is to give a concise account of the current classification of the native taxa, particularly their delimitation and diagnostic characters. Little attention is given to nomenclatural aspects such as synonyms and misapplied names. Notes on the taxonomic value of the leaf anatomy in the genus Ficus has previously been published (van Greuning et al. 1984). The classification proposed in the present paper benefited considerably from the work of Mildbraed & Burrett (1911), Hutchinson (1915), Comer (1965), and particularly Berg et al. (1984, 1985) and Berg (1986a, b). Organographic descriptions are based on observations in the field and from spirit material, as well as the examination of herbarium specimens from BM, BOL, BR, GRA, K, L, MPU, PRE, PRU, SRGH, U, UPS and WIND. Southern

Africa has been taken as the region south of the Okavango and Zambezi Rivers, excluding Zimbabwe and Mozambique. Distribution maps were compiled from over 2 500 herbarium specimens and are based on quarter-degree grid references for collecting localities (Leistner & Morris 1976). Some of the type specimens not seen by the present au thor were kindly studied by C.C. Berg (University of Bergen, Norway) and their identity verified. Only the place of original publication of accepted correct names are supplied. Synonyms are limited to the most recent ones used in southern African literature, or those based on types of southern African origin. Representative herbarium specimens are cited for each taxon. A list of all specimens examined by the author is available on request. Taxonomic treatment

Delimitation of the genus Ficus in southern Africa Genus Ficus L. Subgenus Ficus Section Sycidium Miq. F. capreifolia Del. F. pygmaea Welw. ex Hiern

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S.-Afr.Tydskr.Plantk., 1990,56(6)

Subgenus Sycomorus (Gasp.) Mildbr. & Burr. F. sycomorus L. F. sur Forssk. Subgenus Urostigma (Gasp.) Miq. Section Urostigma F. cordata Thunb. subsp. cordata F. cordata Thunb. subsp. salicifolia (YaW) C.C.Berg F. ingens (Miq.) Miq. F. verruculosa Warb. Section Galoglychia (Gasp.) Endl. Subsection Galoglychia F. lutea Vahl Subsection Platyphyllae (Mildbr. & Burr.) C.C.Berg F. abutilifolia (Miq.) Miq. F. glumosa Del. F. stuhlmannii Warb. F. tettensis Hutch. F. trichopoda Bak. Subsection Chlamydodorae (Mildbr. & Burr.) C.C. Berg F. burtt-davyi Hutch. F. craterostoma Warb. ex Mildbr. & Burr. F.fischeri Warb. ex Mildbr. & Burr. F. ilicina (Sond.) Miq. F. natalensis Hochst. subsp. natalensis F. thonningii Blume Subsection Caulocarpae (Mildbr. & Burr.) C.C.Berg F. bizanae Hutch. & Burtt-Davy F. bubu Warb. F. sansibarica Warb. subsp. sansibarica F. polita Vahl subsp. polita F. tremula Warb. subsp. tremula

Key to the subgenera, sections and subsections la Leaf margin crenate, toothed or lobed; syconia relatively large, mostly more than 15 mm in diameter with outer ostiolar bracts imbricate; stamens mostly 2 per male flower ................. .............. 2 1b Leaf margin entire; syconia relatively small, mostly less than 10 mm in diameter with outer ostiolar bracts imbricate or all descending into ostiolar cavity, when larger then ostiolar bracts all descending into syconial cavity; stamen usually 1 per male flower ......................................................Subgenus Urostlgma 3 2a Shrubs, growing in marshy areas; leaves and syconia scabrous; syconia borne axillary, single or in pairs; basal bracts scattered over peduncle and syconium; stipules not covering the stem apex ........................................ Subgenus Ficus, section Sycidium 2b Trees; leaves and syconia not scabrous; syconia mostly borne in panicles on trunk and main branches; basal bracts in a whorl at the base of the syconium; stipules covering the stem apex ..................................................................... Subgenus Sycomoros 3a Syconia with outer ostiolar bracts imbricate; male flowers ostiolar ........................ ...... ...............................Section Urostlgma 3b Syconia with bracts all descending into ostiolar cavity; male flowers dispersed among female flowers ........................................ ................................................................... Section Galoglychia 4

4a Syconia borne on spurs on trunk and main branches, pedunculate, relatively large (15-40 mm in diameter) ................................................................Subsection Caulocarpae 4b Syconia axillary, pedunculate or sessile, small, mostly less than 20 mm in diameter .............. '" .......................................................5 5a Syconia sessile, 18-26 mm in diameter; leaves large (more than 125 mm long) ..........................................Subsection Galoglychia 5b Syconia sessile or pedunculate, mostly less than 20 mm in diameter; Leaves small (less than 125 mm long) when large then syconia pedunculate ..................................................................... 6 6a Branchlets, leaves and syconia densely tomentose, when somewhat tomentose then leaves relatively large and syconia pedunculate; bark mostly exfoliating ..... Subsection Platyphyllae 6b Branchlets and leaves glabrous, syconia sometimes tomentose; leaves relatively small; bark not exfoliating .................................. . .......................................................... Subsection Chlamydodorae

Key to the species la

1b 2a 2b

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Syconia with outer ostiolar bracts arranged transversely over ostiole, inner descending into ostiole; male flowers occurring in the vicinity of the ostiole ..................................................... 2 Syconia with ostiolar bracts all descending into ostiole; male flowers dispersed among female flowers ................................9 Syconia axillary; stipules not forming a sheath covering stem apex and young leaves ................ ............................................3 Syconia axillary or borne in panicles on trunk and main branches; if axillary, stipules form a sheath around stem apex and young leaves .....................................................................4 Leaf margin dentate or lobed; lamina with adaxial surface scabrous, abaxial surface pubescent .................. 2. F. pygmaea Leaf margin entire or somewhat crenate; lamina with both surfaces scabrous ............................................ 1. F. capreifolia Syconia borne mainly in panicles on trunk and main branches, 20 mm or more in diameter; leaf margin dentate or eroded .....5 Syconia axillary, less than 15 mm in diameter; leaf margin entire .......................................................................................6 Leaves and syconia puberulous, sometimes scabrous; leaf margin eroded ...........................................•.... .3. F. sycomorus Leaves and syconia glabrous or sometimes somewhat villous but not scabrous; leaf margin coarsely dentate ........... .4. F. sur Syconia sessile ............................. 5. F. cordal3 subsp. cordal3 Syconia pedunculate .................................................................7 Syconia 9-12 mm in diameter; young leaves reddish ................. ........................................ ........................................7. F. ingens Syconia less than 9 mm in diameter; young leaves green ......... 8 Trees occurring in dry, rocky habitats; tepals fused; lamina 3-6 times the length of the petiole ............................................ . ...................................................6. F. cordatasubsp. salcifolia Shrubs, usually less than 3 m high, occurring in wet, marshy habitats; tepals free; lamina more than 7 times the length of the petiole .....................................................8. F. verruculosa Syconia borne on trunk and main branches, usually in clusters on short, arrested outgrowths ............................... .................10 Syconia axillary ......................................................................14 Syconialess than 15 mm in diameter ........................................ . ..................................................25. F. tremulasubsp. tremula Syconia more than 20 mm in diameter ................................... 11 Petiole extremely long, lamina rarely more than 1.5 times the length of the petiole; leaves more or less cordate ...................... .........................................................24. F. polil3subsp.polil3

S.Afr.I.Bot., 1990,56(6) 11 b Petiole relatively short, lamina rarely less than twice the length of the petiole ............................................................... 12 12a Lamina elliptic, base cuneate .............................. 21 . F. bizanae 12b Lamina ovate, base rounded to cordate ..................................13 13a Lamina relatively narrow, rarely less than twice as long as wide, apex obtusely acuminate; plants occurring in northern and eastern Transvaal ............................................................... . ........ .............................•23. F. sansibaricasubsp. sansibarica 13b Lamina relatively wide, rarely up to 1.8 times as long as wide; plants occurring in coastal region of Zululand ............... . ................................................................................ 22.F.bubu l4a Syconia and/or branch lets conspicuously tomentose ............. 15 l4b Syconia and branchlets glabrous or somewhat pubescent ......18 l5a Lamina cordate, as long as wide or shorter than wide ............... . ...........................................................................13. F. re/rensis l5b Lamina elliptic or obovate, 1.2-2.4 times as long as wide ..... 16 l6a Leaves conspicuously tomentose .................................. .........17 16b Leaves glabrous or abaxially somewhat tomentose on the veins ...............................................................20. F. Ihonningii l7a Leaves relatively narrow, rarely less than twice as long as wide; venation abaxially conspicuously reticulate .......... ......... . ..................................................................... 12. F. s/Uh/rpannii l7b Leaves .relatively wide, rarely more than 1.6 times as long as wide; venation abaxially not conspicllous ........ 11. F.g/umosa l8a Lamina 1-2 times the length of the petiole ............................ 19 18b Lamina more than 3 times the length of the petiole ............... 20 19a Principal lateral veins 5-7 pairs; lamina as long as wide .......... . .. ... ..................................................................10. F. abulilifolia 19b Principal lateral veins 9-11 pairs; lamina 1.3-1.5 times as long as wide ........................................................17. F. fischeri 20a Syconiasessile ....................................................................... 21 20b Syconia pedunculate .............................................................. 22 2la Syconialarge (18-26 mm in diameter) ...................... 9. F./utea 21 b Syconia small (5-10 mm in diameter) ....... 16. F. crarerostoma 22a Syconia large (11-20 mm in diameter); lamina rarely less than 100 mm long ......................................... 14. F. trichopoda 22b Syconia small (5-13 m,m in diameter); lamina rarely longer than 70 mm ............................................................. ............... 23 23a Petiole stout, 3 mm or more in diameter; lamina chartaceous; plants occurring in the arid western regions of southern Africa .............................. ...................................... 18. F. i/icina 23b Petiole slender, 2.5 mm or less in diameter; lamina coriaceous; plants occurring in the eastern coastal regions of southern Africa ...................................................................... 24 24a Large trees; syconia 7-13 mm in diameter ............................... .. ........................................... 19. F. nala/ensissubsp. nalalensis 24b Shrubs or lianes; syconia5-7 mm in diameter .......................... . ...................................................................... 15. F. burt-davyii

Description of species 1. F. capreifolia Del. in Annales des ' Sciences Naturelles 2(20): 94 (1843). Type: Ethiopia, Calinier s.n. (MPU, holo.!). F. antithetophylla Steud.: 236 (1848).

601

each side of the midvein, diverging at angles of 40-70°, conspicuously curved towards apex, basal pair prominent; lamina 20-122 x 11--43 mm, 2.3-4.6 times as long as wide, 7.3-23.3 times as long as petiole; petiole 3-18 mm long. Syconia axillary, single or in pairs 11-20 mm in diameter; basal bracts scattered over syconium and peduncle; peduncle 9-15 mm long; outer ostiolar bracts imbricate; red when ripe . Male flowers ostiolar, stamens 1-2 per flower; pedicel and filament villous, rudimentary pistil sometimes present; tepals 4-5, slender, separate or only basally fused. Female flowers sessile to pedicellate; tepals 4-5(6), basally fused (Figure 1). Slender shrubs or lianes of wet areas, mainly along river banks in subtropical and tropical regions. Leaves and fruit often heavily parasitized by gall-forming insects. Elsewhere in Africa its distribu'tion ranges as far as Guinea in the west and lower Sudan in the east. Representative specimens -1712 (Posto Velho): Otjinungwa (-AB), Giess & Wiss 3323 (WIND).

-2431 (Acornhoek): Skukuza (-DC), De Winter & CoM 5058 (PRE). -2430 (Pilgrim's Rest): Blyde River Bridge (-BD), Van Greuning 215 (PRU). -2732 (Louwsburg): Iozini (-AC), Van Greuning 169 (PRU).

2. P. pygmaea Welw. ex Hiern in Catalogue of the African plants collected by Dr. Friedrich Welwitsch 1(4): 1009 (1900). Type: Angola, Welwitsch 6370 (GRA, iso.!). F. brevicola Hiern: 1009 (1900) .

Shrubs up to 1 m high; bark greenish-grey; branchlets villous; stipules not covering the stem apex. Leaves oval-elliptic; base rounded, apex acute; margin dentate or lobed; principal lateral veins 4-7 on each side of the midrib, diverging at angles of 50-70°, basal pair prominent; lamina adaxially scabrous, abaxially pubescent, 33-79 x 18-44 mm, 1.8-2 times as long as wide, 9-12 times as long as petiole; petiole 2-7 mm long. Syconia borne singly in leafaxils; 7-14 mm in diameter; peduncle 8-13 mm long; basal bracts scattered over syconium and peduncle; outer ostiolar bracts imbricate; red when ripe. Male flowers ostiolar; tepals 4, distinct; 1 stamen per flower, sometimes with rudimentary pistil. Female flowers with 4 distinct tepals (Figure 2). F. pygmaea resembles F. capreifolia in the scabrous adaxial leaf surface, the occurrence of a pistil rudiment, and syconia and leaves often infested with galls. Both species prefer a moist habitat. Ficus pygmaea differs from F. capreifolia in leaf shape, the hispid abaxial leaf surface and the single stameri per male flower. The species ranges from central Namibia and northern Botswana to as far north as the plateau area of Huilla in Angola.

F. palustris Sim: 99 (1909) .

Representative specimens

Shrubs up to 5 m high; bark greenish-grey; branchlets villous; stipules not covering the stem apex. Leaves oblong-ovate; base rounded, apex acute or trifid; margin entire or somewhat crenate; both surfaces scabrous; principal lateral veins 5-9 on

-1719 (Rundu) : Rundu (-DC), De Winter & Marais 5039 (WIND).

-1821 (Andara): Andara (-AB), Van Greuning 264 (PRU). -1923 (Maun) : Moremi Game Reserve (-BC), Lent 29 (PRE).

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-2416 (Maltahohe): Naukluft (-AA), Merxmuller & Giess 28175

F. gnaphalocarpa (Miq.) A. Rich.: 270 (1851).

(WIND).

F. sycomorus L. subsp. gnapha/ocarpa (Miq.) C.C.Berg: 272 (1980).

3. F. sycomorus L. in Species Plantarum 2: 1059 (1753). Type: not yet traced.

Trees up to 20 m high with spreading branches; bark cream

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coloured; branchlets hispid. Leaves ovate-cordate; base cordate, apex rounded to obtuse; margin entire to crenate; principal lateral veins 5-8 on each side of the midrib, diverging at angles of 40-50°, basal pair reaching to more than half of the length of the lamina; lamina puberulous on both sides, scabrous adaxially; 46-100 x 24-90 mm, 3.3-9.3 times as long as petiole; petiole 3-36 mm long. Syconia borne on leafless branchlets on stem and main branches or in leaf

axils or both; 20-22 mm in diameter, villose; peduncle 6-20 mm long; outer ostiolar bracts imbricate; bright orange when ripe. Male flowers ostiolar; tepals 4, fused or free; stamens 2(1) per flower. Female flowers with 3 tepals, fused or free (Figure 3), On alluvial soil along river banks and streams, Distribution in the rest of Africa ranges from Central and East Africa to as far north as Egypt.

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Representative specimens

4. F. sur Forssk. in Flora Aegyptiaco-Arabica: 180 (1775). Type: Yemen, Forsskdl s.n. (C, halo., verified by C.C. Berg).

-1913 (Sesfontein): Sesfontein (-BA), Theron 3362 (PRU). -1922 (Nokaneng): Thoakhe River (-AB), Erens 247 (PRE). -2017 (Waterberg): Waterberg Plateau (-AC), De Winter 2815a (PRE). -2330 (fzaneen): Albasini Dam (-AA), Van Greuning 208 (PRU).

F. capensis Thunb.: 13 (1786).

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villous, Leaves elliptic to ovate; base rounded or cordate, apex acuminate, rarely rounded; margin somewhat to prominently crenate or dentate; principal lateral veins 3-6 on each side of the midrib, diverging at angles of 30-S0°, basal pair prominent; lamina glabrous except for basal part of midrib, 40-1S0 X 2S-120 mm, 1.2-2 times as long as wide, 2.5-6.7 times as long as petiole; petiole 8-60 mm long. Syconia borne mostly in racemose panicles on trunk: and main branches,

rarely also axillary; glabrous or villpus; 20-40 mm in diameter; peduncles S-13 mm long; outer ostiolar bracts imbricate; red when ripe. Male flowers arranged near ostiole; tepals 4, free or basally fused; stamens 2; rudimentary pistil sometimes present. Female flowers sessile or pedunculate; tepals 3, fused (Figure 4). In moist places along streams and in ravines. Generally the syconia are borne on leafless branchlets on the trunk and

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E, branchlet and syconia; F. distribution.

606

S.-Afr.Tydskr.Plantk., 1990,56(6)

main branches, or even underground on the roots. In addition, several plants with axillary syconia are found along the east coast of the Cape Province and in Natal.

-2340 (Pilgrim's Rest): Blyde River (-BD), Van Greuning 215 (PRU).

Widely spread in central Africa, Angola, Zimbabwe, Mozambique and as far north as Yemen.

-2632 (Bela Vista): Kosi Bay (-DD) , Vahrmeijer 481 (PRE) . -2832 (Mtubatuba): Fani's Island (-CB), Theron 321 (PRE).

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S.Afr.J.Bot., 1990,56(6)

607

-3227 (Stutterheim): Stutterheirn (-CB), Van Greuning 135 (PRU).

attain heights of up to 20 m. It also occurs as far north as Ethiopia and Sudan.

5. F. cordata Thunb. subsp. cordata in Dissertationes Academicae Upsaliae Habitae: Figure 8 (1786). Type: Thunberg 24343 (upS, microfiche!).

Representative specimens

F. cordata var. tristis Warb.: 137 (1906).

Trees up to 7 m high; bark ash grey; branchlets somewhat tomentose. Leaves ovate; base cordate or rounded, apex acuminate; margin entire; principal lateral veins 6-9 on each side of the midrib, diverging at angles of 50--60° from midrib, basal pair prominent; both surfaces glabrous; lamina 28-116 x 10-53 mm, 1.6-2.4 times as long as wide, 2.5-5.6 times as long as petiole; petiole 7-40 mm long. Syconia 5-7 mm in diameter; sessile, axillary; glabrous or somewhat villous; outer ostiolar bracts imbricate; yellowish-green when ripe. Male flowers arranged near ostiole; tepals 3, basally fused; 1 stamen per flower. Female flowers with 3 basally fused tepals (Figure 5). Common in the dry, mountainous parts of Namibia, and the northern and north-western Cape Province. It occurs inland along the banks of the Orange River and as far south as Porterville. This subspecies closely resembles F. cordata subsp. salicifolia, but can be clearly distinguished by its sessile syconia and different geographical distribution. The northernmost distribution seems to be the Kunene River. Representative specimens -1712 (Posto Velho): Otjihipa Mountains (-BC) , Vahrmeijer & Du Preez 2554 (PRE) . -2416 (Maltahohe) : Bergzebra Park (-AA), Merxmuller & Giess 28185 (WIND). -2917 (Springbok): Springbok (-DB), Rosch & Le Roux (PRE). -2922 (Prieska) : Kransfontein (-DA), Acocks 3513 (BOL).

6. F. cordata Thunb. subsp. salicifolia (Vahl) C.G. Berg in Kew Bulletin 43(1): 82 (1988). Type: Yemen, ForsskaJ s.n. (C, holo., verified by C.C. Berg). F. salicifoliaVahl: 82, tab. 23 (1790). F. pretoriae Burtt-Davy: 365 (1912).

Trees up to 20 m high, low-growing shrubs in cold areas; bark dark grey, rough; branchlets glabrous. Leaves ovate to oblong; base cordate or rounded, apex acuminate to obtuse; margin entire; principal lateral veins 7-15 on each side of the midrib, diverging at angles of 50-80°, basal pair prominent; lamina glabrous; 23-130 x 14-57 mm, 2-2.7 times as long as wide, 3-5.8 times as long as petiole; petiole 6-50 mm long. Syconia axillary; 5-8 mm in diameter; glabrous, dotted; peduncle 1-2 mm long; outer ostiolar bracts imbricate, inner desc~nding;.red when ripe. Male flowers ostiolar; tepals 3, fused; 1 stamen per flower. Female flowers with 3 tepals, fused to form a cup (Figure 6). Plants usually occur against slopes and rocky hills where they maintain a shrubby growth form. When growing on deep loamy soil, such as the 'wonderboom' at Pretoria, they may

-2229 (Waterpoort) : Greefswald (-AB), Van Greuning 133 (PRU) . -2527 (Rustenburg): Rustenburg (-CA), Leistner 669 (PRE). -2530 (Lydenburg) : Lowveld Botanical Gardens (-BD) , Buitendag 893 (PRE). -2830 (Dundee) : Ncamboshe (-DA), Edwards 2267 (PRE).

7. F. ingens (Miq.) Miq. in Annales Musei Botanici Lugduno-Batavi 3: 288 (1867). Type: Ethiopia, Schimper 1771 (L, holo.!). F. cordata Sim non Thunb.: 99, tab. 94, Fig. A (1909). F. lutea Mildbr. & Burr. non Vahl: 209 (1911) .

Trees up to 15 m or higher, often low-growing shrubs; branch lets glabrous or shortly villous; bark greyish. Leaves ovate to oblong-ovate; base cordate or rounded, apex shortly acuminate to rounded; margin entire; principal lateral veins 7-10 on each side of the midrib, basal pair inconspicuous; lamina glabrous, 38-140 x 17-81 mm, 1.7-2.3 times as long as wide, 3.7-4.4 times as long as petiole; petiole 5-37 mm long. Syconia pedunculate, axillary; 9-12 mm in diameter; glabrous or slightly pubescent; outer ostiolar bracts imbricate, inner descending into ostiolar cavity; pink when ripe. Male flowers ostiolar with 3-4 fused tepals; 1 stamen per flower. Female flowers with 3-4 fused tepals (Figure 7). F. ingens closely resembles F. cordata subsp. salicifolia, especially in the dried state. It differs, however, in the size of the syconia and the shape and colour of the leaves. Young leaves of F. ingens are reddish, those of F. cordata subsp. salicifolia are green. Generally low shrubs on rock faces. Exceptionally large plants occur in the vicinity of Rustenburg in the Transvaal. Widespread in the rest of Africa up to Eritrea in Ethiopia. Representative specimens -2329 (Pietersburg) : Louis Trichardt (-BB), HuJchinson 2006 (PRE). -2525 (Mafeking) : Gopane (-BD), Snyman 162 (PRE). -2829 (Harrismith): Bergville (-CB), Killick 2309 (PRE). -3228 (Butterworth) : Komga (-CB), Flanagan 850 (BOL).

8. F. verruculosa Warb. in Botanische lahrbUcher 20: 166 (1894). Type: Angola, Welwitsch 6375 (BM, holo.!). F. praeruptorum Hiem: 1004 (1900).

Shrubs, 1-3 m high, rarely small trees up to 8 m high; bark greyish-green; branchlets glabrous. Leaves elliptic to ellipticoblong; base and apex rounded; margin entire; principal lateral veins 5-18 on either side of midvein, diverging at angles of 50-70°, basal pair inconspicuous; lamina glabrous, 85-117 x 22-61 mm, 2-3.8 times as long as wide, 6.9-11 times as long as petiole; petiole 8-15 mm long. Syconia axillary, glabrous, 6-9 mm in diameter; peduncle 4 mm long; outer ostiolar bracts imbricate, inner descending; red when ripe. Male flowers arranged near ostiole; tepals 2-3, free; 1 stamen per flower. Female flowers with 3 unfused tepals per

608

S.-Afr.Tydskr. Plantk., 1990,56(6)

flower (Figure 8). Plants grow in marshy soil along rivers and near lakes. Although no specimens higher than 3 m were found in

southern Africa during this survey, Hutchinson & Reundle (1917) reported heights of up to 8 m. Ranges to West Africa as far north as Nigeria and in East Africa up to Uganda.

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S.Afr.J.Bot., 1990,56(6)

609

Representative specimens

F. nekhudu Warb.: 6, tab. 4 (1904).

-1820 (farikora): Kapupahedi (-AB), Van Greuning 267 (PRU). -2022 (Lake Ngami): Toakhe River (-AD), Pole-Evans 4048 (PRE). -2227 (Palapye): Sofala (-DC), Miller E250 (PRE). -2832 (Mtubatuba): Cape Vidal (-BA), Van Greuning407 (PRU).

F. vogelii (Miq.) Miq.: 288 (1867).

Trees up to 15 m high; bark rough, grey; branchlets glabrous or somewhat villous. Leaves ovate to elliptic; base rounded to somewhat cuneate, apex rounded to obtusely acuminate; margin entire; principal lateral veins 6-8 on either side of the midvein, diverging at angles of 40-70°, basal pair prominent; lamina glabrous, 125-179 x 70-135 mm, 1.6-1.8 times as long as wide, 3-3.9 times as long as petiole; petiole 23-75

9. F. lutea Vahl in Enumeratio Plantarum 2: 185 (1805). Type: Ghana, Hall 47207 (U, neo., verified by C.C. Berg). F. quibeba Hiern: 1000 (1900).

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610

S.-Afr.Tydskr.Plantk., 1990,56(6)

mm long. Syconia sessile; axillary; 18-26 mm in diameter; sparsely tomentose or glabrous; ostiolar bracts descending into syconial cavity; red when ripe. Male flowers dispersed among the female flowers; tepals 3, free or fused at base; 1 stamen per flower; rudimentary pistil rarely present. Female flowers with three tepals, fused at base (Figure 9).

Representative specimens -2632 (Bela Vista): Ingwavuma (-CC), Vahrmeijer & Taiken 958 (PRE).

-2732 (Ubombo): Kwangwanase (-BA), De Winter & Vahrmeijer 8628 (PRE). -2832 (Mtubatuba): Dukuduku (-AB), Wells 2099 (PRE). -2931 (Stanger): Stanger (-AD), Stewart 75 (PRE).

Occurs in southern Africa mainly along the Natal coast, with a few records from northern Namibia. Also reported from Zimbabwe, Zaire, Tanzania and Senegal.

10. F. abutilifolia (Miq.) Miq. in Annales Musei Botanici Lugduno-Batavi 3: 288 (1867). Type: Ethiopia, Kotschy 462 (K, lecto.!).

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S.Afr.J. Bot., 1990,56(6)

611

Trees or shrubs up to 8 m high; bark cream coloured, exfoliating; branchlets glabrous or somewhat pubescent. Leaves cordate; base cordate, apex rounded or obtusely acuminate; margin entire; principal lateral veins 5-7 on either side of the midvein, diverging at angles of 40-70°, basal

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612

S.-Afr.Tydskr.Plantk .• 1990.56(6)

ostiolar bracts descending into sycooial cavity; red when ripe. Male flowers dispersed among female flowers; repals (2)3(4), fused; 1(2) stamens per flower. Female flowers with 3(4) fused tepals (Figure 10).

Representative specimens -2229 (Waterpoort): Greefswald (-AB), Eider 212 (PRU). -2330 (Tzaneen): Gravelotte (-OC), Galpin 13865 (PRE) . -2531 (Komatipoort): Noordkaap (-CA), Acocks 12870 (PRE). -2830 (Dundee): Weenen (-CC), West 1813 (PRE).

Mainly small trees or shrubs growing on rocky hills. On deep, loamy soil, they may attain heights of up to 8 m. Occurs in Zimbabwe and Mozambique as well.

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S.AfrJ.Bot., 1990,56(6)

613

Type: Ethiopia, Caillaud s.n. (MPU, holo.!).

21-96 mm, 1.2-1.7 times as long as wide, 3.1-7.9 times as long as petiole; petiole 5-34 mm long. Syconia 8-15 mm in diameter; warted; sessile, axillary; ostiolar bracts descending into syconial cavity; red when ripe. Male flowers dispersed among female flowers; tepa Is 2-3, fused at the base; 1 stamen per flower. Female flowers with (2)3(4) tepals, fused at the base (Figure 11). On rock faces in dry and temperate areas. Plants from the arid, western parts of Namibia have smaller, less tomentose

F. sonderi Miq.: 295 (1867).

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leaves and smaller syconia than those of the more mesic areas of Natal and Transvaal. It also occurs in Senegal, Nigeria, Mozambique, Zaire and Zimbabwe.

S.-Afr.Tydskr. Plantk., 1990,56(6)

with 3 tepals, mostly fused (Figure 13). Plants occur on rocks where the roots penetrate crevices and aid in soil formation. The only known locality outside southern Africa is near Tete in Mozambique.

Representative specimens -1712 (Posto Velho): Baines Mountains (-BB), Giess 8982 (WIND). -1814 (Otjitundua): Otjivasandu (-CB), Van Greuning 429 (PRU). -2229 (Waterpoort): Wyllie's Poort (-DD), Schweickerdl & Verdoorn 669 (PRE). -2930 (Pietermaritzburg): Pinetown (-DD), Johnson 1468 (PRE).

12. F. stuhlmannii Warb. in Botanische Jahrbiicher 20: 161 (1894). Type: Tanzania, Stuhlmann 4141 (B, holo., verified by C.C. Berg). F. howardiana Sim: 100, Figure 92A (1909).

Representative specimens -2227 (Palapye): Topsi (-AB), Miller B804 (PRE). -2229 (Waterpoort): Mapungubwe (-AB) , Van Greuning 134 (PRU). -2330 (Tzaneen): Klein Letaba (-AB) , Van der Schijff 3317 (PRU). -2430 (Pilgrim's Rest): Erasmus Pass (-BC), Van der Schijff6744 (PRU).

14. F. trichopoda Bak. in The Journal of the Linnean Society of London 20: 261 (1883). Type: Madagascar, Baron 1663 (K, lecto.!). F. hippopotami Gerstn.: 151 (1943).

Trees up to 9 m high; bark ash-grey, rough; branchlets lomentose. Leaves obovate, ovate or elliptic; base rounded or somewhat cordate, apex rounded or obtusely acuminate; margin entire; principal lateral veins 4-6 on either side of midvein, diverging at angles of 50-70°, basal pair prominent, venation prominently reticulate below; lamina tomentose on both surfaces, 28-83 X 13-56 mm, 1.6-2.4 times as long as wide, 3.2-4.6 times as long as petiole; petiold 8-25 mm long. Syconia sessile, axillary; tomentose; 8-15 mm in diameter; ostiolar bracts descending into syconial cavity; red when ripe. Male flowers dispersed among female flowers; tepals 3(2), fused at the base; 1 stamen per flower. Female flowers with 3(2) tepa Is, fused at the base (Figure 12). Plants favour deep, sandy soil in woodland and dry forest. Also found in Zaire, Kenia, Tanzania and Mozambique. Herbarium specimens of this species are often confused with those of F. glumosa. The prominent reticulate venation on the lower leaf surface of this species helps to separate it from the latter. Representative specimens -2230 (Messina): Punda Milia (-DB), Van Rooyen 1340 (PRU). -2431 (Acornhoek): Satara (-BD), Codd 4295 (PRE). -2732 (Ubombo): Ingwavuma (-AA), Van Greuning 280 (PRU). -2831 (Nkandla): Ngwelezani Hills (-DD), VenJer 3812 (PRE).

13. F. tettensis Hutch. in Kew Bulletin 7: 341 (1915). Type: Mozambique, Kirk s.n. (K, holo.!). F. smuJsi Verdoom: 205 (1935).

Trees up to 7 m high; bark cream coloured to almost white, exfoliating; branchlets tomentose. Leaves cordate; base cordale, apex rounded to obtusely acuminate; margin entire or unevenly crenate; principal lateral veins 4-6 on either side of the midrib, diverging at angles of 40-70°, basal pair prominent; lamina tomentose on both surfaces, scabrous adaxially, 37-90 X 33-115 mm, 0.8-1 times as long as wide, 4-5.6 times as long as petiole; petiole 9-30 mm long. Syconia axillary, sessile to pedunculate; 7-10 mm in diameter; tomentose; ostiolar bracts descending into syconial cavity; red when ripe. Male flowers dispersed among female flowers; tepals 3, mostly fused; 1 stamen per flower. Female flowers

Shrubs or trees up to 9 m high with many prop roots; bark brownish-grey; branchlets somewhat pubescent or glabrous . Leaves cordate; base cordate, apex rounded to obtusely acuminate; margin entire; principal lateral veins 7-11 on either side of the midrib, diverging at angles of 60-70°, basal pair inconspicuous; lamina glabrous, 68-170 X 56-128 mm, 1.3-1.4 times as long as wide, 3.7-8.8 times as long as petiole; petiole 19-45 mm long. Syconia axillary; somewhat pubescent or glabrous, freckled; 11-20 mm in diameter; peduncle 7-12 mm long; ostiolar bracts descending into syconial cavity; red when ripe. Maleflowers dispersed among female flowers; tepals 3, fused; 1 stamen per flower. Female flowers with 3(4) tepals, fused (Figure 14). Plants grow in wet, marshy areas and range from central West Africa, through central Africa, Madagascar to as far south as Ballito Bay in Natal. Representative specimens -2632 (Bela Vista): Kosi Bay (-DD), Van Greuning 282 (PRU). -2732 (Ubombo): Mseleni River (-BC), De WinJer & Vahrmeijer 8579 (PRE) . -2831 (Nkandla): Mtunzini (-DD), VenJer 1865 (PRE). -2931 (Stanger): Stanger (-AD), Moll 2187 (PRE).

15. F. burtt-davyi Hutch. in Kew Bulletin 9: 232 (1916). Type: Kowie West, Burtt-Davy 7954 (PRE, holo.!). Shrubs or lianes, 0.5-5 m high; bark greyish, rough; branchlets somewhat pubescent. Leaves elliptic; base rounded, apex rounded to obtusely acuminate; margin entire; glabrous on both surfaces; principal lateral veins 4-9 on either side of the midrib, diverging at angles of 40-70°, basal pair more or less prominent; lamina 13-76 X 7-36 mm, 1.5-2 times as long as wide, 3-10 times as long as petiole; petiole 2-20 mm long. Syconia axillary; 5-7 mm in diameter; villous; ostiolar bracts descending into syconial cavity; peduncle 3-6 mm long; basal bracts 2-3, fused. Maleflowers few, dispersed among female flowers; tepals 3(4), mostly unfused, sometimes basally fused; 1 stamen per flower. Female flowers with 3(4) tepals, mostly unfused, sometimes fused at base (Figure 15).

S.An-.J.Bot., 1990,56(6)

615

The habit of F. burtt-davyi varies according to habitat. On mountains and rocky outcrops such as at Robberg, Oudtshoom and Graaff Reinet, plants are shrubs with relatively small leaves, while in forest and coastal dune forest, they occur as lianes with slender stems and relatively large leaves. This species does not appear to occur north from Kosi Bay in Zululand.

Representative specimens -2732 (Ubombo): Sibaya (-BC), Vahrmeijer 699 (PRE). -2930 (Pietermaritzburg): Shongweni Dam (-DC), Morris 602 (PRE). -3227 (Stutterheim): Fort Cox (-CC), Robbertse 840 (PRU). -3423 (Knysna): Plettenberg Bay (-AA) , Van Greuning 345 (PRU).

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Figure 12 F. stuhlmannii. A. Male flower; B-D. female flowers; E. branchlet with syconia; F. distribution.

616

S.-Afr.Tydskr.Plantk., 1990,56(6)

16. F. craterostoma Warb. ex Mildbr. & Burr. in Botanische lahrbiicher 6: 247 (1911). Type: Tanzania, Stuhlmann 8995 (B, lecto., verified by C.C. Berg).

descending into ostiole. Male flowers dispersed among female flowers; tepals 3, fused; 1 stamen per flower. Female flowers with 3 fused tepals (Figure 16). F. craterostoma is closely related to, and resembles F. natalensis subsp. natalensis. It differs from the latter by its sessile syconia and semipersistent stipules. Plants occur mainly in forests, from Komga in the eastern Cape Province to Mariepskop in the eastern Transvaal and as far north as Uganda.

Trees or shrubs up to 15 m high; bark grey, rough; branchlets glabrous. Leaves obovate to obtriangular; base cuneate or rounded, apex rounded or truncate; margin entire; veins adaxially inconspicuous, principal lateral veins 5-11 on either side of midvein, diverging at angles of 40-70° from midvein, basal pair inconspicuous; lamina glabrous, 27-90 x 12-37 mm, 1.6-2.8 times as long as wide, 3.8-6.9 times as long as petiole; petiole 5-24 mm long. Syconia 5-10 mm in diameter; axillary, sessile; glabrous; ostiolar bracts

Representative specimens -2430 (Pilgrim' s Rest): Mariepskop (-DB), Van der Schijff 6214 (PRU).

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S.AfrJ.Bot.. 1990.56(6)

617

-2731 (Louwsburg): Steilrand (-CC). Coda

1~77

(PRE).

yet traced).

-3030 (Port Shepstone): Oribi Gorge (-CB). Van Greuning 413 (PRU) .

F. kiloneura Hornby: 1007 (1948).

-3228 (Butterworth): Kentani (-CB). Pegler 1126 (PRE).

Trees up to 20 m high. starting life as epiphytes; bark grey. rough; branchlets glabrous. Leaves ovate to cordate; base cordate to round. apex obtusely acuminate; margin entire; glabrous; principal lateral veins 9-11 on either side of

17. F. fischeri Warb. ex Mildbr. & Burr. in Botanische lahrbiicher 46: 227 (1911). Type: Tanzania. Fischer 545 (not

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Figure 14 F. trichopoda. A & B. Male flowers with perianth partly removed; C. sessile and D. pedicellate female flowers; E. branchlet with syconia; F. distribution.

S.-Afr.Tydskr.Plantk .• 1990.56(6)

618

midvein, diverging at angles of 50-65°, basal pair conspicuous; lamina 85-132 x 60-104 mm, 1.3-1.5 times as long as wide, 1.5-2.3 times as long as petiole; petiole 53-77 mm long. Syconia 13-15 mm in diameter; axillary; glabrous; ostiole umbonate, ostiolar bracts descending into ostiole; peduncle 10-15 mm long. Male flowers dispersed among female flowers; tepals 2(3), fused; 1 stamen per flower. Female flowers with 2(3) fused tepa Is (Figure 17). Occurs mainly north of the Zambesi River in Angola,

Zambia, Zimbabwe and Mozambique. The two localities in Namibia represent the southernmost distribution of the species. Representative specimens -1722 (Chirundu): Bwabwata (-DC). Van Greuning 307 (PRU). Tinley 1482 (WIND). -1724 (Katirna Mulilo): Katima Mulilo (-CB). Vahrmeijer s.n. (PRE). Geldenhuys 207 (PRE).

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Figure 15 F. burlt-davyi. A & B. Male flowers. B. with perianth partly removed; C. sessile and D. pedicellate female flowers; E. branchlet with syconia; F. distribution.

S.AfrJ.Bot.. 1990.56(6)

619

18. F. i1icina (Sond.) Miq. in Annales Musei Botanici Lugduno-Batavi 3: 289 (1867). Type: Kamiesberg. Zeyher 3869 (PRE, holo.!). F. gurichiana Eng!.: 130 (1894).

Shrubs or trees up to 5 m high; bark white; branch lets glabrous. Leaves elliptic; base and apex rounded; margin entire; principal lateral veins 7-10 on either side of the midvein, diverging at angles of 40-70°, basal pair

inconspicuous; glabrous; coriaceous; 30-90 x 25-45 mm, 1.9-2.7 times as long as wide, 5-7 times as long as petiole; petiole 7-18 mm long. Syconia axillary; 5-8 mm in diameter; glabrous or somewhat villous, peduncle 2-5 mm long; ostiolar bracts descending into ostiole. Male flowers dispersed among female flowers; tepa Is 3(4), basally fused; 1 stamen per flower. Female flowers with 3(2) basally fused tepals (Figure 18). On rocky outcrops of dolomite or granite in the arid areas

A

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620

S.-Afr.Tydskr.Plantk., 1990, 56(6)

of western Namibia and the Cape Province. Not recorded outside southern Africa.

19. F. natalensis Hochst. subsp. natalensis in Flora 28(1): 88 (1845). Type: Durban, Krauss 276 (BM, tecto.!).

Representative specimens

Trees up to 20 m high; stranglers; bark dark grey; branchlets glabrous. Leaves elliptic or obtriangular; base cuneate, rarely rounded, apex rounded or truncate; margin entire; principal lateral veins 6-9 on either side of midvein, diverging at angles of 50-70°, adaxially inconspicuous; glabrous; lamina 28-71 X 13-37 mm, 1.4-3 times as long as wide, 3.6-5.5

-2016 (Otjiwarongo): Otjiwarongo (-BC) , Van Greuning 273 (PRU) . -2217 (Windhoek) : Ositoto (-AA), Gibson 24 (WIND). -2616 (Aus): Aus (-BA), Acocks 15644 (PRE). -2917 (Springbok): Springbok (-DB), Van Greuning 387 (PRU).

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Figure 17 F. fischeri . A. Male flower with one tepal removed; B. sessile and C. pedicellate female flowers; D. branchlet with syconia; E. distribution.

S.AfrJ. Bot., 1990,56(6)

621

times as long as petiole; petiole 4-20 mm long. Syconia

fused; 1 stamen per flower. Female flowers with 3 fused tepals (Figure 19). In southern Africa mainly confined to the coastal forests of Natal and Zululand. Also in East Africa as far north as Uganda.

axillary; 7-13 mm in diameter; glabrous; peduncle 3-7 mm long; ostiolar bracts descending into syconium; red when ripe. Male flowers dispersed among female flowers; tepa Is 3,

2 mm Imm

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622

S.-Afr.Tydskr.Plantk., 1990,56(6)

Representative specimens

Trees up to 12 m high; bark grey; branchlets villous, sparsely tomentose or glabrous. Leaves mostly elliptic to obovate; base rounded to cuneate; apex rounded to acuminate; margin entire; venation prominent on both surfaces, principal lateral veins 6-11 on either side of midvein, diverging at angles of 50-70°, basal pair inconspicuous; lamina glabrous or somewhat tomentose on veins, 24-109 x 10-42 mm, 1.7-3.4 times as long as wide, 2-3 times as long as petiole, petiole 7-45 mm long. Syconia 7- 11 mm in diameter; tomentose, rarely glabrous; sessile or shortly to long pedunculate; axillary; ostiolar bracts descending into syconial cavity; ostiole often umbonate; red when ripe. Male flowers dispersed among female flowers; tepals 3, fused at the base; 1

-2632 (Bela Vista); Kwangwanase (-DD) , Van Greuning 339 (PRU).

-2832 (Mtubatuba); Dukudulcu (-AD), Van Greuning 406 (PRU). -2931 (Stanger); Maidstone (-"{:A) , Van Greuning 326 (PRU). -3030 (Port Shepstone) ; Scottburgh (- BD), Van Greuning 150 (PRU) .

20. F. thonningii Blwne. in Rumphia 2: 17 (1836). Type: Ghana, Thonning 325 (not yet traced). F. burkei (Miq.) Miq.; 289 (1867). F. pelersii Warb.; 164 (1894) .

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flowers; E. branchlet with syconia; F. distribution.

S.Afr.J.Bot., 1990,56(6)

623

stamen per flower. Femaleflowers with 3 tepals, fused at the base (Figure 20). Ficus thonningii occurs in tropical and temperate regions of southern Africa, mainly in savannah. woodland where the plants start life as epiphytes and develop into stranglers. Plants are also found on rocks, in ravines and in forests. In southern Africa its distribution ranges from Alexandria in the eastern Cape Province to Natal, Swaziland, Transvaal and

northern Namibia. It has also been recorded from Angola, Botswana, Zimbabwe and Mozambique. Ficus thonningii is probably the most variable of all fig species. The variation is mainly manifested in the length of the peduncles, the length of the petioles and the shape of the leaves. There is, however, no discontinuity in variation to justify recognition of different species or even intraspecific taxa.

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624

S.-Afr.Tydskr.Plantk., 1990, 56(6)

Plants may often be found with large syconia, up to 20 mm in periods when the plant yields very few syconia and may result from heavy infestation by gall-forming wasps.

-2329 (Pietersburg): Louis Trichardt (-BB), VerMorn 2226 (PRE). -3326 (Grahamstown): Alexandria (-CB), Van Greuning 138 (PRU).

Representative specimens

21. F. bizanae Hutch. & Burtt-Davy in Kew Bulletin 2: 52 (1921). Type: Umtamvuna River, Burtt-Davy 15291 (K, holo.!).

-1721 (Mbambi): Shamangoro Island (-CC), De Winter 4256 (PRE). -1917 (Tsumeb) : Elefanteberg (-CD), Kinges 2946 (PRE) .

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S.AfrJ.Bot., 1990,56(6)

625

somewhat villous; ostiolar bracts descending into syconial cavity; basal bracts 2-3; yellowish-green when ripe. Male flowers dispersed among female flowers; tepals 3, free or fused at the base; 1 stamen per flower, with conspicuous apical outgrowth. Female flowers with 3 tepals, free or fused at the base (Figure 21).

Trees up to 18 m high; bark ash-grey; branch lets glabrous. Leaves elliptic; base cuneate, apex acuminate; margin entire; glabrous; principal lateral veins 5-9 on either side of midvein, diverging at angles of 40-50°, basal pair prominent; 33-88 X 15-44 mm, 1.7-3 times as long as wide, 1.6-3 times as long as petiole; petiole 14-41 mm long. Syconia borne in fascicles on short, arrested branchlets arising from the trunk and main branches; 25-46 mm in diameter; peduncle 7-20 mm long; surface warty, glabrous or

Restricted to the eastern coastal area of southern Africa where it has been recorded from Pondolancl, southern Natal and the Ngoye Forest in Zululand. No records from outside

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Figure 22 F. bubu. A. Pedicellate female flower and B. sessile female flower with perianth removed; C. male flower; D. branchlet and syconium; E. distribution.

626

S.-Afr.Tydskr. Plantk., 1990,56(6)

-3130 (Port Edward): Umtarnvuna Nature Reserve (-AA), Van Greuning 335 (PRU).

these mentioned areas were reported. It closely resembles F. po/ita subsp. po/ita, but differs from it in respect of the cuneate lamina base and the relatively long lamina which is up to three times as long as the petiole and two to three times as long as wide.

22 .. F. bubu Warb. in Annales du Musee Congo Beige, Botanique ser. 6: 3, Tab. 8 (1904). Type: Zaire, Gillet 1167 (BR, halo.!).

Representative specimens Trees up to 11 m high; bark grey; branch lets glabrous. Leaves elliptic to ovate; base rounded or slightly cordate, apex rounded; margin entire; principal lateral veins 6-7 on either

-2831 (Nkandla): Ngoye Forest (-DC), Muller 6530 (PRE). -3129 (Port St. Johns): Old Bunting (-CA), Van Greuning 146 (PRU); Port St. Johns (-DA), Wells 3496 (PRE).

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Figure 23 F. sansibarica subsp. sansibarica. A. & B. Male flowers, B. with perianth partly removed; C. pedicellate and D. sessile female flowers; E. branchlet and syconia; F. distribution.

S.Afr.J.Bot., 1990,56(6)

627

side of mid vein, diverging at angles of 40-60°, basal pair prominent; glabrous; 85-170 X 67-104 mm, 1.4-1.8 times as long as wide, 2-3.9 times as long as petiole; petiole 30-56 mm long. Syconia in fascicles on short, arrested branchlets on main branches; 20-21 mm in diameter; glabrous; ostiolar bracts descending into ostiole; peduncle 11-13 mm long; yellowish-green when ripe. Male flowers dispersed among female flowers; tepals 3, free or fused at the base; 1 stamen per flower. Female flowers with 3 tepals, free or fused at the base (Figure 22). In southern Africa this species is rather rare with the only known localities at Masengwenya and Lake Sibaya in northern Zululand. It has also been reported from Angola,

Malawi, Mozambique, Swaziland and as far north as Uganda and the Ivory Coast. In the original description of F. bubu the syconia were not described. Both Mildbraed & Burrett (1911) and Hutchinson & Reudle (1917) mentioned that the syconia had not been seen in position, but assumed that they are borne axillary. Representative specimens -2631 (Mbabane): Manzini (-BA), Compton 27928 (PRE). -2632 (Bela Vista): Umbuluzi Poort (-AA) , Culverwell 152 (PRE). -2732 (Ubombo): Lake Sibaya (-BC), Van Greuning 288 (PRU); Mazengwenya (-BC), De Winter & Vahrmeijer 8541 (PRE).

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23. F. sansibarica Warb. subsp. sansibarica in Botanische lahrbticher 20: 171 (1894). Type: Tanzania, Stuhlmann 793 (B, holo., verified by C.C. Berg). F. delagoensis Sim: 99, Figure 92 (1909).

Trees up to 16 m high; bark grey; branchlets glabrous. Leaves ovate; margin entire; base rounded, apex obtusely acuminate;

principal lateral veins 7-9 on either side of midvein, diverging at angles of 40-70°, basal pair conspicuous; lamina glabrous, 43-170 x 20-104 mm, 2-3.2 times as long as wide, 2.4-7.9 times as long as petiole; petiole 11-56 mm long. Syconia borne in clusters on stout, wartlike branch lets on the trunk and major branches; 20-44 mm in diameter; glabrous; peduncle 11-25 mm long; ostiolar bracts descen-

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S.Afr.J.Bot. , 1990,56(6)

ding into ostiole; yellowish-green when ripe. Male flowers dispersed among female flowers; tepals 3, fused; 1 stamen per flower, with apical protuberance. Female flowers with 3 basally fused tepals (Figure 23). Grows in sandy loam soil in the savannah vegetation of northern and eastern Transvaal. Trees start life as semiepiphytes. Widely distributed throughout tropical Africa. Representative specimens -2230 (Messina): Sibasa (-CD), Van Greuning 211 (PRU). -2231 (Pafuri): Punda Milia (-CA), Codd & Dyer 4583 (PRE). -2330 (fzaneen): Westfalia Estate (-CA), Scheepers 1254 (PRE). -2430 (Pilgrim's Rest): F.H. Odendaal Resort (-BD), Van Greuning 292 (PRU).

24. F. pol ita Vahl subsp. polita in Enumeratio Plantarum 2: 182 (1805). Type: Ghana, Isert s.n. (C, holo., verified by C.C. Berg.

629

petiole; petiole 10-50 mm long. Syconia borne in fascicles on short, curved branch lets on trunk and main branches; spotted; glabrous; 13-18 mm in diameter; ostiolar bracts descending into ostiole; peduncle 6-8 mm long; yellowish-green when ripe. Maleflowers dispersed among female flowers; tepals 3, free; 1 stamen per flower. Female flowers with 3-4 tepals, free or fused at the base (Figure 25). Occurs in the northern parts of Zululand, from False Bay Park to as far north as Kenya. In southern Africa it usually reaches a height of about 4 m and are predominantly semiepiphytic. Hutchinson (1917) recorded plants of up to 10 m along the Mozambique coast. Representative specimens -2632 (Bela Vista): Muzi (-DC), Ross & Moll 5098 (PRE), Van Greuning 302 (PRU). -2732 (Ubombo): Makani's Pont (-AB), Moll 4513 (PRE); Kwangwanase (-BB), Van Greuning 401 (PRU).

F. umbrosa Sim: 100, Figure 88 (1909) .

Acknowledgements Trees up to 15 m high; branchlets glabrous; bark grey; main branches warty where syconia abscised. Leaves cordate to elliptic-ovate; base cordate, rarely rounded, apex acuminate; margin entire; principal lateral veins 5-8 on either side of midvein, diverging at angles of 40-60°, basal pair inconspicuous; lamina glabrous, 34-92 x 25-55 mm, 1.6-1.8 times as long as wide, 1.2-1.9 times as long as petiole; petiole 22-78 mm long. Syconia borne in clusters on short, arrested branchlets on trunk and main branches; 18-35 mm in diameter; peduncle 10-15 mm long; glabrous, sometimes villous; ostiolar bracts descending into ostiole; yellowishgreen when ripe. Male flowers dispersed among female flowers; tepals 3, fused at the base; 1 stamen per flower, with conspicuous apical protuberance. Female flowers with 3 tepa Is, fused at the base, sometimes free (Figure 24). Although widely distributed in tropical Africa, this species has a limited distribution in southern Africa, ranging from Swaziland to as far south as Port Shepstone. Similar to F. bizanae and F. sansibarica subsp. sansibarica, but differs from the latter by its fewer lateral veins and the shorter and broader lamina. Representative specimens -2631 (Mbabane): Mlawula (-AC), Van laarsveld 930 (PRE). -2732 (Ubombo) : Jozini Dam (-AC), Van Greuning 281 (PRU) . -2832 (Mtubatuba): Mapelane Forest (-AD), Venter 5580 (PRE). -2931 (Stanger): Stanger (-AD), Van Greuning 322 (PRU).

25. F. tremula Warb. subsp. tremula in Botanische lahrblicher 20: 171 (1894). Type: Tanzania, Stuhlmann 274 (B, holo., verified by C.C. Berg). F. pulvinata Warb.: 169 (1894).

Trees up to 10 m high; bark grey; stem with short, curved branchlets upon which syconia are borne; branches glabrous. Leaves elliptic to oblong; base rounded to somewhat cordate, apex acuminate; margin entire; principal lateral veins 7-10 on either side of mid vein, diverging at angles of 50-70°, basal pair inconspicuous; lamina glabrous, 30-95 x 12-33 mm, 2-3 times as long as wide, 1.9-3.7 times as long as

I am grateful to the C.S.I.R. and the University of Pretoria for financial support, to Martmari van Greuning for the drawings and to Prof. A.E. van Wyk for his invaluable comments on this paper. Prof. C.C. Berg's valuable discussions and comments on some of the type specimens are highly appreciated.

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