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The Taxonomy of the Rhoicissus Tridentata (Vitaceae) Complex in Southern Africa

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The Taxonomy of the Rhoicissus Tridentata (Vitaceae) Complex in Southern Africa The taxonomy of the Rhoicissus tridentata (Vitaceae) 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 plants 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 species 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 plant, 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 genus 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
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