The of the Rhoicissus tridentata () complex in southern Africa

Noel. R. Urton, Maria C. Olivier and B.L. Robertson Department of Botany, University of Port Elizabeth, Port Elizabeth

The taxonomy of the polymorphic Rhoicissus tridentata Introduction (L.f.) Wild & Drumm. complex in southern Africa is revised. Historical background It is concluded that two subspecies are involved, viz. subsp. tridentata and subsp. cuneifo/ia (Eckl. & Zeyh.) N.R. Urton, The earliest reference to the now known as Rhoicissus stat. et comb.nov. tridentata was in 1781 under the name of Rhus tridentatum S. Afr. J. Bot. 1986, 52: 389- 396 by the younger Linnaeus. This name was based on the trifolio­ late specimen collected by Thunberg (s.n.) at the Cape of Die taksonomie van die polimorfe Rhoicissus tridentata (L.f.) Good Hope. Two additional specimens of Thunberg were Wild & Drumm.-kompleks in suidelike Afrika is hersien. described in the same publication as R. cirrhijlorum and R. Daar is tot die slotsom gekom dat twee subspesies ter digitatum both of which are now recognized as Rhoicissus sprake is, nl. subsp. tridentata en subsp. cuneifolia (Eckl. & digitata (L.f.) & {fi/.) Zeyh.) N.R. Urton, stat. et comb.nov. Gilg Brandt. Linnaeus separated the specimen he called Rhus tridentatum from the other two on S.-Afr. Tydskr. Plantk. 1986, 52: 389-396 the grounds of it having serrated leaflets covered with short Keywords: Rhoicissus, southern Africa, taxonomy, Vitaceae whitish hairs whereas the other two had entire, glabrous leaflets. In 1803 Thunberg describ~d plants collected by himself and Masson as Rhus dimidiatum, the specific epithet referring to the asymmetrical nature of the lateral leaflets. Burchell (1824) described a similar to Rhus tri­ dentatum L.f. as Cissus paucijlora but it had almost entire leaflets. In 1835 Ecklon & Zeyher named their No. 433 Cissus tridentata, quoting the Thunberg specimen used by Linnaeus (fil.) for his description of Rhus tridentatum and thus recog­ nizing that it was not a Rhus species. At the same time they described their No. 432 (which is clearly a form of Rhoicissus tridentata) as Cissus cirrhijlora based on the Thunberg specimen used for the description of Rhus cirrhijlorum L.f. This was the start of a nomenclatural error that persisted down the years until exposed by Wild & Drummond (1963). Two other Ecklon & Zeyher specimens, differing in leaflet shape and margin, No. 434 and No. 435, were described by them as Cissus dimidiata (based on Thunberg's Rhus dimi­ diatum) and Cissus sericea respectively. A fifth related , No. 431, was named C. cuneifolia and is distinct in having relatively large, wide leaflets and long petioles. Harvey (1860) referred to Ecklon and Zeyher's No. 433 as Cissus paucijlora var. tridentata, maintaining that it was merely a variety of the plant which Burchell (1824) had named C. paucijlora. Harvey distinguished two varieties of C. paucijlora viz. var. tridentata with leaflets two- to three-toothed and var. cirrhijlora with leaflets 'very entire'. The latter variety was based on Ecklon and Zeyher's No. 432 and thus per­ Noel. R. Urton*, Maria C. Olivier and B.L. Robertson petrated the nomenclatural error already mentioned. Harvey Department of Botany, University of Port Elizabeth, P.O. Box recognized both C. dimidiata and C. sericea but noted that 1600, Port Elizabeth, 6000 Republic of South Africa the former might be merely a variety of the latter. *To whom correspondence should be addressed When Planchon (1887) established Rhoicissus as a separate from Cissus, he recognized Rhoicissus paucijlora but Accepted 5 May 1986 gave it a different interpretation to Burchell, not confining 390 S.-Afr. Tydskr. Plantk., 1986, 52(5)

the name to plants with almost entire leaflets, but including structure and marked similarity in flowers, fruits and seeds as synonyms, Cissus tridentata, C. paucijlora var. cirrhijlora, throughout the complex (Figure 1). Pollen morphology, insect C. paucijlora var. tridentata and C. cirrhijlora. He too pollinators and reproductive phenology are similar. Thus it accepted Ecklon & Zeyher's interpretation of the epithet is concluded that one species is involved. cirrhijlora and thus also perpetuated the nomenclatural error. In contrast an extremely high degree of polymorphism is In addition he recognized Rhoicissus sericea and agreed with shown in the leaves regarding petiole length, the size, shape Harvey (1860) regarding Cissus dimidiata being a mere form and margin of leaflets as well as the degree and colour of of it. Like Harvey he upheld C. cuneifolia and called it the indumentum. Rhoicissus cuneifolia. There are two trends in hypocotyl and primary root Planchon recognized yet another species incorporated in development (Figure 2) as well as in leaf development where the R. tridentata (L.f.) Wild & Drumm. complex namely R. juvenile foliage is markedly different from adult foliage in erythrodes (Fres.) Planch. This was based on Vitis erythro­ one group. These features form the basis of the recognition des, the name given by Fresenius (1837) to a plant collected of two subspecies. by Dillon & Petit in Ethiopia. Its distinctive features include a long petiole and leaflets which are tomentose abaxially and Phytogeography have well-defined teeth. Planchon drew attention to the These plants, commonly known as wild grape (Smith 1966) affinity between Rhoicissus erythrodes and R. cuneifolia of are widespread in southern Africa, occurring from Riversdale the Cape. in the west, extending eastwards and then northwards through­ Planchon also recognized R. erythrodes var. jerruginea out tropical Africa (Figures 3 & 7). A southern group is which Baker (1868), under the name of Vitis erythrodes var. represented by subsp. tridentata and a northern group by ferruginea, separated on account of its more brightly fer­ subsp. cuneifolia. The two groups are partially sympatric and ruginous silky pubescence in contrast to the grey to slightly the greatest diversity of form occurs where distributions ferruginous colour of the usual form. It was based on a plant overlap such as in the Grahamstown division. The inter- collected by Welwitsch in Lower Guinea (Angola). . mediate forms encountered here could indicate hybridization. Szyszylowicz (1888) did not recognize Rhoicissus and applied the names Vitis paucijlora, V. cuneifolia and V. Relationships erythrodes to plants of the Rhoicissus tridentata complex. According to Lawrence (1966) and others, the family Vitaceae Kuntze (1898) agreed with Szyszylowicz in recognizing Vitis is widely distributed in the tropics and subtropics with ranges cuneifolia which he divided into three varieties, namely var. extending into the north and south temperate zones. Since scandens from the Cape, var. intermedia from Natal and var. the R. tridentata complex extends northwards into tropical erythrodes from Abyssinia and Central Africa. Africa and beyond (Wild & Drummond 1966), the species In 1912 Gilg and Brandt gave preference to the name as a whole may have had a northern origin. In this case the Rhoicissus cirrhijlora and their concept of the whole complex ancestral form is likely to have been similar to the northern involved no less than four nomenclatural types on which the form of subsp. cuneifolia (Figure 8.1). With migration of the following names were based: Rhus cirrhijlorum L.f., R. tridentatum L.f., Cissus paucijlora Burch. and C. cuneifolia Eckl. & Zeyh. In addition they considered Rhoicissus sericea to be conspecific with the plant named Cissus dimidiata by Ecklon and Zeyher (1835) and correctly applied the name Rhoicissus dimidiata to them. Gilg & Brandt (1912) retained Rhoicissus erythrodes as a b separate species and included var. ferruginea, R. holstii Engl., R. usambarensis Gilg and R. verdickii De Wild., all referring to plants beyond the borders of southern Africa, as synonyms. Suessenguth (1953) agreed with Planchon rather than with Gilg and Brandt in recognizing R. paucijlora and R. cuneifolia as well as R. erythrodes. Like Gilg and Brandt (1912) he gave preference to the name k. dimidiata instead of R. sericea. R. usambarensis was retained as a separate species. In 1963 Wild & Drummond combined three previously recognized species as Rhoicissus tridentata (L.f.) Wild & Drumm., namely R. cuneifolia (Eckl. & Zeyh.) Planch., R. erythrodes (Fres.) Planch. and R. cirrhijlora (L.f.) Gilg & Brandt pro parte. Since their work was based on material found north of the Zambesi River, Wild and Drummond did not consider previously recognized species such as R. pau­ cijlora and R. dimidiata. The aim of the present investigation was to subject this long a list of taxa to a critical taxonomic examination to determine their individual or collective status.

Evaluation of characters Figure I Rhoicissus tridentata subsp. tridentata. (a) Branch with flower Examination of herbarium material, of live material from a buds on tendrils; (b) late flower bud; (c) flower; (d) longitudinal section wide range of localities and of plants grown from seed under of flower with petals and stamens removed; (e) ripe fruit; (f) seed. All similar conditions indicated little difference in anatomical drawn from Urton 603 (PEU). S. Afr. J. Bot., 1986, 52(5) 391

complex southwards has come diversification of form, related Reproductive phenology at least partly to different climatic conditions, and adaptations The main flowering period throughout the complex is in subsequently becoming genetically established. The forms of summer and autumn (December to April) and fruiting occurs subsp. tridentata occupying the most arid regions show what from late autumn to summer (April to December). can be considered the most extreme adaptations such as reduction of leaf size, degree of dentation and density of Taxonomy indumentum (Figure 4. 7). Rhoicissus tridentata (L.f) Wild & Drumm. in Kirkia 3: 19 (1963). Type: Cap. b. spei [Cape Province], Thunberg s.n. (UPS, bolo.; PEU photo!, PRE, phot9!) (Figure 5). Shrub with or without scandent branches, or a woody climber up to 10 m or more tall, with tuberous roots and a series of slender rhizomes giving rise to aerial shoots at intervals; young branches glabrous, or fulvous- or cinereous-pubescent, or more rarely ferruginous-tomentose; tendrils glabrescent and usually unbranched except when bearing an inflorescence.

h

pr

40 mm

• Figure 4 Rhoicissus tridentata subsp. tridentata: variation in leaves. (1) Taylor 9443 Suurberg; (2) Ecklon & Zeyher 435 Uitenhage; (3) Bayliss A 1422 Baviaanskloof; (4) Rogers 28600 Port Alfred; (5) Urton 654 Fraser's Watch Tower; (6) Muir 2464 Langeberg; (7) Urton 607 Springs Reserve Uitenhage. B

10mm J

Figure 2 Seedlings showing differences in hypocotyl (h) and primary root (pr) development at I month. A - Rhoicissus tridentata subsp. tridentata; B - Rhoicissus tridentata subsp. cuneifolia.

v--- 1'--- 22' v t\ __/ 24' \ ~~ !'----~ lv / 26° I "" '- ( v ; ( ..-/ ·' ' ' I / ' -- ~ / I / 28° /" -~· ~- j I v ) ' 1/ 30° - "'-·-..- 6. ;;': llc; J ~f'v'>.- ~ / . 32° 'J 1-••...... ; v .. ·;;;. ;;;: v .~ f- ~ 34° ~ ~-v Ur5 ...... 225 18' 20' 22' 24' 26' 28' 30' 32' 34' Figure 5 Photograph of the holotype of Rhoicissus tridentata subsp. Figure 3 Distribution of Rhoicissus tridentata subsp. tridentata. tridentata (Photograph from UPS). 392 S.-Afr. Tydskr. Plantk., 1986, 52(5)

Leaves spirally arranged, sessile or petiolate, glabrous or Cissus cirrhijlora (L.f.) Eckl. & Zeyh. : 56 (1835) pro parte. fulvous- or cinereous-pubescent, or more rarely ferruginous­ Rhoicissus cirrhijlora (L.f.) Gilg & Brandt : 438 (1912) pro parte. tomentose, 3-foliolate; petioles almost cylindrical, (0-) Cissus paucijlora Burch. var. cirrhijlora (L.f.) Harv. : 251 (1860) 2- 20(- 70) mm long; terminal leaflets petiolulate, petiolule pro parte. (1-)2- 4( - 12) mm long; lamina narrowly elliptic to broadly C. sericea Eckl. & Zeyh. : 57 (1835); Harv. : 251 (1860). obovate, ( 12-)20- 45(- 120) mm long, (3 - )5 - 20(- 100) Rhoicissus sericea (Eckl. & Zeyh.) Planch. : 466 (1887); Gilg : mm broad, apex truncate, obtuse or subacute, base cuneate to 450 (1895). Type: Winterhoecksberge [Winterhoeksberg, Uitenhage, acute; lateral leaflets petiolulate or sessile, petiolule if present Cape Province] Ecklon & Zeyher 435 (SAM! lecto., here designated; shorter than that of terminal leaflet, (0- )1 ~ 2(- 8) mm long, GRA!). lamina narrowly elliptic to ovate, (9-) 15 - 30(- 85) mm long, (3 - )5 - 20(- 55) mm broad, apex obtuse to acute, base usually Description asymmetrical with outer side acute to obtuse and inner side Shrub up to I m or more; branches scandent or erect, cuneate to acute; margin of all leaflets entire to variously toothed occasionally having a cinereous appearance. Leaves small, but teeth lack a distinct apiculus; acarodomatia sometimes glabrous, more rarely cinereous pubescent; petiole (0-)2- 3 present abaxially in axils of lower lateral veins; stipules ligulate (- 6) mm long; lamina of terminal leaflet (12-) 15 - 25(- 35) to conical, caducous, ca. 2- 3 mm long. Inflorescence a cyme, mm long, (3- )5 -15( -18) mm broad; lamina of lateral (1 - )5-9(- 60)-flowered, leaf-opposed or on tendrils, young leaflet (9-) 12-25(- 30) mm long, (- 3)6 -10(- 15) mm parts fulvous- or cinereous-pubescent or ferruginous-tomentose, broad; margin entire or with up to four (rarely six) dentations peduncle (1-)4- 5( -14) mm long; bracts deltoid, caducous, or crenations in all leaflets; juvenile foliage differing from ca. I mm long. Pedicel (1 - )2- 3 (- 4,5) mm long. Calyx cup­ adult foliage in having a longer petiole and larger leaflets with shaped with ca. 7- 10 ill-defined obtuse ferruginous teeth and more and deeper dentations. Petals often with indumentum pubescent-tomentose. Petals (4)5(6), ovate, 2- 3 mm long, on outer side. Hypocotyl in seedling swelling before primary I ,5- 2,5 mm broad, cream to green, sometimes tinged with root, apparent above ground (Figure 2). pink or red, glabrous on inside, sometimes pubescent-sericeous on outside. Stamens (4)5(6), ca. 2 mm long, anthers dorsifixed Discussion and versatile, filaments widening towards base. Disc (4)5(6)­ Distribution and habitat lobed, annular, 2-3 mm wide. Ovary immersed in disc, style­ This subspecies occurs along the southern coast from the stigma ca. 1 - 2 mm long, stigma simple, obtuse but becoming Riversdale district eastwards to Port St Johns where it grows slightly bilobed. Fruit a globose to ovoid berry, (5-)6- 8( -13) chiefly in sandy soils on permanent dunes in coastal bush. mm long, (5-)8- 10 (- 15) mm broad, turning red and It extends inland in the south-eastern Cape to the Karoo where finally purplish-black, sometimes spotted with lenticels. Seeds it is a constituent of Valley Bushveld and other karroid I - 4, globose with one or two sides flattened when more than vegetation types (Figure 3). one seed is present, 4- 6 mm long, 3- 6 mm broad, testa smooth with a longitudinal furrow. Variants Considerable variation occurs within this subspecies, especially Key to subspecies regarding length of petiole, leaflet margin (the number and Ia Leaflet margin entire or nearly so ...... I. subsp. tridentata form of dentations) and indumentum (density and colour). I b Leaflet margin with dentations or crenations: A number of forms or variants are recognizable. The Cape 2a Terminal leaflet with up to four dentations or crenations; leaves glabrescent or cinereous-pubescent ...... coastal variant corresponds most closely to Thunberg's ...... I. subsp. tridentata specimen which Linnaeus.fi/. named Rhus tridentatum. This 2b Terminal leaflet with more than four dentations or ere­ form has several teeth per leaflet (Figure 4.6). nations; leaves glabrescent or fulvous- or ferruginous-pub- A form with glabrous, almost entire leaflets (Figure 4.7) escent ...... 2. subs. cuneifolia is widespread in the drier parts of the eastern Cape and Karoo 1. subspecies tridentata and corresponds most closely to the specimen named Cissus paucijlora by Burchell. A minor variation is evident among Rhus tridentatum L.f. : 184 (1781); Thunb. : 52 (1794); 266 (1823). Type: Cap. b. spei [Cape Province] Thunberg s.n. (UPS, specimens collected in the Suurberg area (Taylor 9443) where halo.; PEU, photo!, PRE, photo!) (Figure 5) . the margin of the lateral leaflets sometimes shows a thumb­ like lobe along the outer side (Figure 4.1). Cissus tridentata (L.f.) Eckl. & Zeyh. : 56 (1835). Along the coast at Port Alfred a form with the smallest C. paucijlora (Burch.) var. tridentata Harv. : 251 (1860). and finest leaves occurs and in these the petioles are relatively Rhoicissus tridentata (L.f.) Wild & Drumm. : 19 (1963); 446 long (Figure 4.4). (1966) pro parte. Among the forms with entire leaflets are some which have Rhus dimidiatum Thunb. : 28 (1803); 266 (1823). been in the past, mistakenly identified as Rhoicissus digitata, Cissus dimidiata (Thunb.) Eckl. & Zeyh. :57 (1835); Harv. : 251 especially on account of their narrowly elliptical leaflets (Figure (1860). 4.3). This applies to plants collected in the Humansdorp and Rhoicissus dimidiata (Thunb.) Gilg & Brandt: 437 (1912); Schonl. Steytlerville areas. The identity of this form was uncertain until : 75 (1919); Suesseng. : 330 (1953); Martin: 73 (1960). Types: Cap. the juvenile stage was found and showed clearly that it [Cape] Masson s.n. (UPS, lecto., here designated; PEU, photo­ belonged to the R . tridentata complex. copy!); Cap. [Cape] Thunberg s.n. (UPS, PEU, photocopy!). Although in most variants the leaves are glabrescent, two Cissus paucijlora Burch.: 630 (1824). closely related forms occur in which the indumentum is Rhoicissus paucijlora (Burch.) Planch. : 468 (1887) pro parte; retained at least abaxially on the leaves and is cinereous. Gilg : 450 (1895); Suesseng. : 330 (1953) pro parte. Because of the density of this indumentum especially on all Vilis paucijlora (Burch.) Szyszyl. : 44 (1888). Type: Cape young parts of the plant including flower buds, the plants as Province, Somerset East, Bruintjies Hoogte, Burchell 3009 (K, holo.; a whole present a characteristic silvery-grey appearance. The PEU, photo!). more easterly form is represented by Thunberg's type specimen S. Afr. J . Bot., 1986, 52(5) 393

of Rhus dimidiatum and by Ecklon and Zeyher 434. This Rhoicissus erythrodes (Fres.) Planch. var. jerruginea Bak. : 466 form has smaller leaves with a few distinct dentations on each (1887); Gilg & Brandt : 440 (1912); C.A.Sm. : 473 (1932). leaflet and the lateral leaflets are markedly asymmetrical Vitis cuneijolia (Eckl. & Zeyh.) Szyszyl. var. intermedia Kuntze (Figure 4.5). It occurs in a limited zone in the Grahamstown : 40 (1898). Type: Natal, Charlestown, Kuntze s.n. (NY, lecto·. , here division, where it grows on exposed rocky hillsides sometimes designated; PEU, photocopy!). associated with 0/denburgia arbuscu/a. This latter plant is Rhoicissus cirrhiflora (L.f.) Gilg & Brandt : 438 (1912) pro parte. associated with outcrops of Witteberg quartzite (Gledhill R. tridentata (L.f.) Wild & Drumm. : 19 (1963) pro parte; 1981). The second silvery-grey form has a more westerly 444 - 446, t. 94 (1966) pro parte; Gibson : t. 62 (1975). distribution and is represented by Ecklon and Zeyher 435 which was described as Cissus sericea. It has larger leaves, Description less obvious dentations and the lateral leaflets are not so Shrub with scandent or erect branches or a woody climber markedly asymmetrical (Figure 4.2). up to 10 m or more. Leaves usually large, with or without Transitional forms between these variants within the a fulvous-ferruginous indumentum, petiolate more rarely subspecies occur where distributions overlap, indicating that subsessile; petiole ( 1 -)10- 30(-70) mm long; lamina of hybridization could have taken place. terminal leaflet (15- )25- 50( -120) mm long, (8- )15- 40 ( -100) mm broad; lamina of lateral leaflet (12- )20- 40 Representative specimens (- 85) mm long, (4-)15- 25(- 55) mm broad; margin with A copy of the complete list of specimens examined is available more than four dentations or crenations in terminal leaflet on request. but fewer in lateral leaflets; juvenile and adult foliage similar. Petals glabrous. Hypocotyl not swollen in seedling (Figure 2). - 3128 (Umtata): Nduli ( - DB), Johnson 633 (PEU). - 3129 (Port St Johns): Egossa district ( - BC), Sim 1822, 2396 Discussion (NU). Typification - 3224 (Graaff-Reinet): 48 km south of Graaff-Reinet ( - BC), Bolus 291 (BOL, GRA). In their description of Cissus cuneifolia, Ecklon & Zeyher - 3225 (Somerset East): Somerset East district ( - DA), van der (1835) name three different localities as follows: Walt 482 (PRE). 'Sylvis in "Adow" (Uitenhage), tum inter frutices hiatuum - 3226 (Fort Beaufort): Bedford ( - CA), Scott Elliott 129 (PRE). montium prope "Grahamstown" altit. II (Albany) saltibusque - 3324 (Steytlerville): Baviaanskloof ( - CA), Bayliss 1422 (PRE). montis "Chumiberg" altit. III (Kafferland).' - 3325 (Port Elizabeth): Addo Elephant Park ( - BC), Hall-Martin 5940 (PRE); North foothills of Suurberg, 500 m from Ann's Villa ( - BD), Taylor 9443 (PRE, STE); Uitenhage, Coega River near Winterhoeksberg ( - CB), Eck!on & Zeyher 435 (GRA, SAM); Springs Nature Reserve ( - CD), Urton 607; Bluewater Bay ( - DC), Urton 603 (PEU). - 3326 (Grahamstown): Highlands, 'Mill Hills' ( - AD), Comins 955 (GRA, PRE); Bloukranz, Albany ( - BC), Bayliss 1007 (PRE); Fraser's Watch Tower ( - BD), Urton 654 (PEU); Port Alfred (-DB), Rogers 28600 (J, PRE, SAM). - 3421 (Riversdale): Lange berg, Riversdale district ( - AB), Muir 2464 (PRE). - 3422 (Mossel Bay): Sedgefield ( - BB), Taylor 663 (NBG). - 3423 (Knysna): Knysna ( - AA), Duthie 570 (BOL, STE). - 3424 (Humansdorp): Eersterivier ( - AA), Urton 630 (PEU). - 3425 (Schoenmakerskop): Cape Receife ( - BA), Olivier 2798 (PEU). 2. subspecies cuneifolia (Eckl. & Zeyh.) N.R. Urton, stat. et comb. nov. Cissus cuneijolia Eckl. & Zeyh. : 56 (1835); Harv. : 251 (1860). Types: Cape Province, precise locality uncertain, Ecklon & Zeyher 431 (SAM!, lecto., here designated -sheet 2356) (Figure 6). Rhoicissus cuneijolia (Eckl. & Zeyh.) Planch. : 466 (1887); Gilg : 450 (1895); C.A.Sm. : 473 (1932); Codd : 114 (1951); Suesseng. : 330 (1953); Martin : 74 (1960). Vitis cuneijo!ia (Eckl. & Zeyh.) Szyszyl. : 44 (1888); Sim: t. 36, fig. 3 (1907) . Vilis erythrodes Fres. : 284 (1837); Bak. : 401 (1868); Szyszyl. : 44 (1888). Type: Ethiopia, Adowa, Dillon & Petit s.n. (P, holo.; PEU, photocopy!). Rhoicissus erythrodes (Fres.) Planch. : 465 (1887); Gilg : 450, t. 219 J- L (1895); Gilg & Brandt : 442, t. 3 (1912); C.A.Sm. : 473, t. 75 A (1932); Suesseng. : 332, t. 97 (1953); Willems : 565 (1960).

Vilis cuneijolia (Eckl. & Zeyh.) Szyszyl. var. erythrodes (Fres.) \ .. a,/'; I ~~~ Kuntze : 41 (1898) . t..-., ~~.tf,.% V. erythrodes (Fres.) var. jerruginea Bak. : 402 (1868); Szyszyl. l ,. ·~J:. ~ l-. (. : 44 (1888). Type: Angola, Huilla, Welwitsch 1455 (K, lecto., here designated, PEU, photo!). Figure 6 Lectotype of Rhoicissus tridentata subsp. cuneijolia (SAM). 394 S.-Afr. Tydskr. Plantk., 1986, 52(5)

km 100 100 200 300 400 500 600 700 BOO km at the present time in all three localities. There are two sheets in SAM of Eck!on & Zeyher 431, the type of subsp. cuneifolia which they named Cissus cuneifolia. The sheet numbered 2356 (at the top right) is here designated lectotype because the several plant portions are uniform in features and could clearly be parts of one plant. The sheet numbered 2355 shows four plant portions, two of which match those of the lectotype but the other two are clearly subsp. tridentata. One of these latter specimens is labelled No. 2033 and is from a different locality from those of No. 431. The four specimens are unfortunately not arranged on the sheet according to their two identities. The specimen at S under the name Cissus cuneijolia and indicated as Typus was not chosen as lectotype as the locality given on a label attached to the sheet indicates that the specimen was collected in the Somerset East district which is not one of the localities given in the original description.

Description and habitat This subspecies has a wide distribution, extending from the eastern Cape to the northern limits of the Republic of South Africa (Figure 7). It occurs in forest along forest margins or in high bush where it scrambles freely over other plants. Along the eastern coast of southern Africa it may grow as an erect shrub in open grassveld.

Variants 20° 220 24° 26° 28° 30° 32° 34° As in the case of subsp. tridentata considerable variation Figure 7 Distribution of Rhoicissus tridentata subsp. cuneijolia. occurs in the size and shape of the leaflets, the number and form of dentations and in the colour and density of the indumentum. The shape of the terminal leaflet, especially at the base and apex, is particularly variable. The base may be cuneate to obtuse and the apex truncate to acute. Dentations may sometimes be apiculate (Figure 8.1) but if so, the apiculus is less than 1 mm long. The colour of the indumentum is usually fulvous but may be distinctly golden or even fer­ ruginous. It is however never cinereous. From north to south in general throughout the distribution range, there is a decrease in leaf size and in density of indumentum. Southern forms are usually glabrescent. The length of petiolules is also very variable but, with the exception of a variant along the east coast, the petiole is very well defined being usually not less than half the length of the lamina of the terminal leaflet. Plants from the eastern Cape Province (Figure 8.3 & 5) correspond most closely to the type specimen named Cissus 40mm cuneijolia by Ecklon & Zeyher. Most plants from the Transvaal, Orange Free State and Figure 8 Rhoicissus tridentata subsp. cuneijolia: variation in leaves. Natal interior correspond more with plants previously known (!) Luckhojj s.n. Letaba; (2) Urton 671 Bulwer; (3 ) Urton 642 Hogs­ back; (4) Ross 1398 Mkuzi road; (5) Urton 609 Grahamstown Nature as Rhoicissus erythrodes, having numerous well defined Reserve; (6) A cocks 20937 Lyden burg; (7) Ward 7550 Pietermaritzburg. dentations and often having a fulvous indumentum at least abaxially on the leaves (Figure 8.1 & 2). These two forms grade into one another and intermediates There are two possible interpretations of this information. are common. One is that the specimens were collected in the first locality, An unusual variant restricted to parts of the eastern namely forests at Addo, but that, in addition, similar plants Transvaal is notable for its relatively large leathery leaves with were found near Grahamstown and in the Tyumie mountains a ferruginous indumentum (Figure 8.6). . (Amatolas). In this case, Addo is the type locality and the Along the coast of Natal and Transkei there is yet another different specimens of No. 431 are isotypes. The other distinctive form. In contrast to other forms it grows in open interpretation is that No. 431 represents plants collected in grassveld as a nearly erect shrub and the leaves are almost the three different localities in which case the specimens are sessile (Figure 8. 7). Intermediates between this and the inland syntypes. The present authors favour the first interpretation Natal form are common. because it seems unlikely that plants collected in three widely An unusual variant found in northern Zululand has small separated localities, and therefore collected at different times, markedly ferruginous and pubescent leaves (Figure 8.4). Apart would bear the same collectors' number. Similar plants occur from the colour of the indumentum it shows affinities with S. Afr. J. Bot., 1986, 52(5) 395

the silvery-grey form of subsp. tridentata from the Grahams­ - 2832 (Mtubatuba): Hluhluwe Game Reserve (-AA), Ward 2089 town division. Because the two populations are so widely (NH, NU, PRE). separated and because intermediates between this Zululand - 2926 (Bloemfontein): Thaba Nchu (- BB), Moster! 1190 (PRE). form and other forms of subsp. cuneifolia occur, it is included - 2927 (Maseru): Leliehoek near Lady brandt (-AB), Olivier 2001 (PEU); Roma (- BD), Schmitz 609 (PRE). under this subspecies. - 2929 (Underberg): Bulwer, behind Mountain Park Hotel ( - DO), Transitional forms between all the variants within this Urton 671 (PEU). subspecies occur where distributions overlap indicating that, - 2930 (Pietermaritzburg): Lidgetton, Lions River District (-AC), as in the case of subsp. tridentata, hybridization could have Mogg 6690 (PRE); Sarnia Beacon Hill (- DD), Ward 7550 (NU, taken place. PRE). - 2931 (Stanger): Umhlanga (-CA), Wood 1187 (NH). Representative specimens - 3030 (Port Shepstone): Oribi Gorge (- CA), Pienaar 113 (PRE). - 3126 (Queenstown): Madeira Hill ( - DD), Urton 683 (PEU). A copy of the complete list of specimens examined is available - 3127 (Lady Frere): Near Cala (- DA), Royffe 226 (GRA). on request. -3128 (Umtata): Pot River, Maclear district ( -AB), Galpin 6598 - 2229 (Waterpoort): Soutpansberg, 10 km west of main road (GRA, PRE). (-DC), Rodin 4063 (PRE). - 3129 (Port St Johns): Port St Johns, 2nd Beach (-DA), Pienaar - 2230 (Messina): Tate Vondo Forest Reserve ( - CD), Hemm 71 149 (PRE). (PRE). - 3130 (Port Edward): Bizana (-AA), A cocks 13398 (PRE). - 2231 (Pafuri): Kruger National Park, Hippo pool, south bank - 3225 (Somerset East): Farm Bergvliet, Bruintjies Hoogte (- CB), of Sabie River ( - AC), Mogg 16750 (PRE). Urton 668 (PEU). - 2329 (Pietersburg): Louis Trichardt, Soutpansberg (- BB), - 3226 (Fort Beaufort): Hogsback, I ,5 km from Inn on road to Galpin 14216 (NH, PRE). Alice ( - DB), Urton 642 (PEU). - 2330 (Tzaneen): Westfalia Estate (-CA), Scheepers 269 (PRE); - 3227 (Stutterheim): Kubusie Forest (- CB), Urton 641 (PEU). Letaba (-CD), Luckhoff s.n. (STE). - 3325 (Port Elizabeth): Suurberg Pass, near Hotel (-BD), Bayliss - 2425 (Gaborone): Mannyelanong Hill, 32 km south of Gaborone 487 (PRE). ( - DB), Matt 651, 655 (PRE). - 3326 (Grahamstown): Grahamstown Nature Reserve at Dassie-· - 2427 (Thabazimbi): Witpoortjie (- CA), Gilmore T.M. 26363 krans (- BC), Urton 609 (PEU). (PRE). - 3327 (Peddie): East London ( - BB), Thode 10430 (STE). - 2428 (Nylstroom): Mosdene, Naboomspruit ( - DA), Galpin M60 (GRA, PRE). Excluded names - 2429 (Zebediela): 24 km north-north-west of Schoonoord, Cissus inaequilatera E. Mey. A nomen nudum quoted by Harvey Lydenburg district (-DB), A cocks 20937 (PRE). in Harvey & Sonder Fl. Cap. I : 251 (1860) under Cissus cuneifo!ia. - 2430 (Pilgrims Rest): Mount Sheba Nature Reserve (-DC), Kerfoot 8165 (PRE). Rhoicissus grosseserrata Gilg. A nomen nudum quoted as a - 2431 (Acornhoek): Sabiesand Game Reserve (-CD), Botha 1920 synonym for R. erythrodes by Smith in Burtt Davy, F.P.F.T.2 : (PRE). 473 (1932). - 2525 (Mafeking): Gopane, 40 km west of Zeerust (- BD), R. stenophyl/us C.A. Smith & Shaw. A nomen nudum applied Snyman 10 (PRE). to specimens representing the juvenile stage of some forms of subsp. - 2526 (Zeerust): Zeerust (-CA), Marloth 10192 (PRE). tridentata. - 2527 (Rustenburg): Buffelspan, Pilansberg (-AA), Pole Evans 628 (PRE). Sp.non.sat.cog. - 2528 (Pretoria): Pretoria (- CA), Moss 4503 (J); Fountains Cissus tridentata (L.f.) Eckl. & Zeyh. var. robusta Eckl. & Zeyh. Valley (-CC), Verdoorn 622 (PRE). in Enum : 57 (1835). Typification uncertain: Ecklon & Zeyher's - 2529 (Witbank): Loskop Dam Nature Reserve (-AD), Mogg type (SAM) has the same number (433) as their Cissus tridentata, 30415, 31041 (J, PRE). is mounted on the same sheet and there is no indication as to which - 2530 (Lydenburg): Farm Zwagershoek ( - AB), Obermeyer 182 of the several plant portions is the variety. (PRE); Lowveld Botanic Garden, Nelspruit (-BD), Buitendag 105 Vitis cuneijolia (Eckl. & Zeyh.) Szyszyl. var. scandens Kuntze, (NBG, PRE). Rev.Gen. 3,2: 40 (1898). Typification uncertain: Kuntze's specimen - 2531 (Komatipoort): 16 km north-east of Barberton on Bar­ s.n. (NY) from Kingwilliamstown consists of stems only. berton-Belfast road (-CA), Jordaan 6 (PRE). - 2627 (Potchefstroom): Koedoesfontein, Potchefstroom district ( - CD), Louw 1674 (PRE). Acknowledgements - 2628 (Johannesburg): Melville Koppies ( - AA), Macnae 1142 The authors wish to thank the Council for Scientific and (J); Klipriviersberg, 14 km south of Johannesburg (-AC), Mogg Industrial Research, P·retoria for their financial support. 17671 (J); Suikerbosrand, Kareekloof (-CA), Bredenkamp 146 (PEU), 1021 (PRE). References - 2630 (Carolina): Near The Brook (-BA), Strey 8012 (NH, PRE). BAKER, J.G. 1868. Vitis. In: Oliv., Fl. Trop.Afr. I : 401-402. - 2631 (Mbabane): Usutu Forests ( - CA), Compton 27768 (NBG, BURCHELL, W.J. 1824. Catalogue of South African plants. In: PRE). DC., Prod. I : 630. - 2726 (Odendaalsrus): De Kroon (-BD), Putteri/116169 (PRE). CODD, L.E. 1951. Trees and shrubs of the Kruger National Park. - 2729 (Volksrust): Charlestown ( - BD), Kuntze s.n. (NY). Mem. bot. Surv. S. Afr. 26 : 114. - 2730 (Vryheid): Vryheid ( - DD), Galpin 9752 (PRE). ECKLON, C.F. & ZEYHER, K.L. 1835. Enumeratio plantarum - 2731 (Louwsburg): 14 km from Jozini Dam on Mkuzi road africae australis extropicae. I. Hamburg. (-DB), Ross 1398 (NBG, NH, NU). FRESENIUS, G. 1837. Vitis Linn. In: Mus. Senckenb. 2 : 284. - 2732 (Ubombo): False Bay (-CD), Ward 1137 (NV). GIBSON, J.M. 1975. Wild flowers of Natal (coastal region). - 2827 (Senekal): Willem Pretorius Reserve ( - A C), Leistner 3006 Durban: Trustees of the Natal Publishing Trust Fund. (PRE). GILG, E.F. 1895. Vitaceae. In: Engl. & Pratt, Pf!anzenjam. 3, 5 : - 2828 (Bethlehem): Witsieshoek (-DB), Junod 17423 (PRE). 540 (t 219, J - L). - 2829 (Harrismith): Farm Nolens Volens east of van Reenen GILG, E.F. & BRANDT, J.F. 1912. Rhoicissus. In: Engl., Bot. ( - AC), Jacobsz 1584 (PRE). Jahrb. 46 : 437, 438, 440 & t 3. - 2830 (Dundee): Krantzkop (-AA), Thode 4805 (STE). GLEDHILL, E. 1981. Eastern Cape veld flowers. 2nd edn. Cape - 2831 (Nkandla): Umfolozi Game Reserve ( - BD), Ward 1469 Town: The Department of Nature & Environmental (NU). Conservation of the Cape Provincial Administration. 396 S.-Afr. Tydskr. Plantk., 1986, 52(5)

HARVEY, W.H. 1860. Ampelideae. In: Flora Capensis, eds SMITH, C.A. 1966. Common names of South African plants. Harvey, W.H. & Sander, O.W. Vol. I p. 251. Lovell Reeve, Mem. bot. Surv. S. Afr. 35 : 585 . London. SUESSENGUTH, K. 1953. Rhoicissus. In: Engl. & Prantl, KUNTZE, C.E.O. 1898. Ampelidaceae (Vitaceae). Rev. Gen. 3,2 : Pflanzen/am. 2nd edn, 20d : 330- 332. 40-41. SZYSZYLOWICZ, I. 1888. Vilis. In: Polypet. Disc. Rehm. 18 : 44. LAWRENCE, G.H.M. 1966. Taxonomy of vascular plants. New THUNBERG, C.P. 1794. Prodromus plantarum capensium. York, Macmillan Co. Uppsala. LINNAEUS, C. Fil. 1781. Supplementum plantarum. THUNBERG, C.P. 1803. Rhus dimidiatum. In: Hoffm. Phyt. Braunschweig. Bliitt. : 28. Gottingen. MARTIN, A.R.H . & NOEL, A.R.A. 1960. The flora of Albany THUNBERG, C.P. 1823. Flora capensis, ed. Schultes J.A. p. 266. and Bathhurst. Grahamstown, Rhodes University. Stuttgardt. PLANCHON, J.E. 1887. Arnpelideae. In: A. DC., Monogr. URTON, N.R. 1983. A revisionary study of the polymorphic Phan. 5, 2 : 465-468. Rhoicissus tridentata (L.f.) Wild & Drumm. in southern Africa. SCHONLAND, S. 1919. Phanerogamic flora of Uitenhage and Ph.D. thesis, Univ. of Port Elizabeth. Port Elizabeth. Mem. bot. Surv. S. Ajr. I : 75. WILD, H. & DRUMMOND, R.B. 1963. Rhoicissus. Kirkia 3 : SIM, T.R. 1907. The forests and forest flora of the colony of the 19. Cape of Good Hope. Aberdeen, Taylor & Henderson. WILD, H. & DRUMMOND, R.B. 1966. Vitaceae. Fl. Zamb. 2, 2 SMITH, C.A. 1932. Arnpelidaceae. In: A manual of the flowering : 439, 444 - 446. plants and ferns of the Transvaal with Swaziland. ed. Burtt WILLEMS, L. 1960. Rhoicissus. In: De Wit & Willems in Flore Davy, J. Vol. 2 p. 473. London, Longrnans Green. du Congo Beige et du Ruanda - Urund: 9 : 565.