Coccinia rehmannii Cogn. in Schinz, Bull. Herb. Boiss. 3: 418. 1895.

Holstein, Norbert, 2015, Monograph of Coccinia (Cucurbitaceae), PhytoKeys 54, pp. 1-166 : 104-107

19. Coccinia rehmannii Cogn. in Schinz, Bull. Herb. Boiss. 3: 418. 1895.

Coccinia rehmannii Type: South Africa. Transvaal: boshveld ad Klippan [according to Meuse (1962) in Limpopo: Greater Sekhukhune District Municipality, Doornpoort; 24°37'S, 29°26'E], 1875-1880. A. Rehmann 5156 (Syntype: Z! [Z-000004445, digital image: Z]; syntype: BR! [BR0000005111602, digital image: BR, JPS]).

Coccinia rehmannii Type: South Africa. Ibid., A. Rehmann 5157 (Syntype: Z! [Z-000060807, digital image: Z]).

Coccinia rehmannii Type: South Africa. Ibid., A. Rehmann 5168 p.p. (Lectotype, designated by Meeuse (1962: 102): Z! [Z-000060808, digital image: Z]; isolectotype: BR! [BR0000005111930, digital image: BR, JPS], K! [K000313196, digital image: JPS, K]).

Coccinia rehmannii Type: South Africa. At Eland river, A. Rehmann 4944 [sic, must be A. Rehmann 4954, see Taxonomic remarks] (Syntype: Z! [Z-000060806, digital image: Z]).

Coccinia rehmannii Cogn. var. littoralis A.Meeuse, Bothalia 8: 104. 1962. pro parte ex R. de Carvalho s.n. (Paratypes: COI (2)!).

Coccinia rehmannii Type: South Africa. [Eastern Cape]: [Amatole District Municipality], Komgha, Kei Mouth, H.G. Flanagan 457 (Holotype: PRE [PRE0190559-0, digital image: JPS], isotypes: BOL?, NBG?).

Coccinia rehmannii Type: South Africa. [Eastern Cape]: Cape Morgan, H.G. Flanagan 457 (Paratype: GRA [GRA0002852-0, digital image: JPS], BOL?, NBG?).

Coccinia rehmannii Type: South Africa. [Eastern Cape]: East London, Nahoon, M.W. Nanni 151 (Paratype: PRE!).

Coccinia rehmannii Type: South Africa. [Eastern Cape]: Coffee Bay, W. Tyson 24 (Paratypes: B!, COI!, GRA, MO!, NY!, PRE!, S! [S08-12380]).

Coccinia rehmannii Type: South Africa. [KwaZulu-Natal]: 10 mls NW of Mtubatuba, L.E.W. Codd 9620 (Paratypes: COI!, M! [M-0198513], PRE).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: Umhlanga Rocks, A. Dohse & B. de Winter 223 (Paratypes: NH, PRE!).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: Manaba Store, J. Gerstner 3407 (Paratype: NH).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: Dhlebe, J. Gerstner 4261 (Paratypes: NH, PRE).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: near Durban, T.J. Jenkins TRV7092 (Paratype: PRE).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: Mtunzini, S.M. Johnson 612 (Paratype: NBG).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: Stella Bush, W.E. Marriott herb. no. 24341 (Paratype:?).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: ibid., W.E. Marriott herb. no. 27143 (Paratype: NH).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: Shelly Beach, A.O.D. Mogg 11941 (Paratype:?).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: ibid., A.O.D. Mogg 12070 (Paratypes: M! [M-0198511], M! [M-0198512], PRE!).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: without detailed location, T.B. Oatley C 15 (Paratype: PRE).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: Berea, Small herb. no. 34714 (Paratype: NH).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: Ubombo coastal veld, P.A. Tosh 28 (Paratype: NU).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: Ndumu Game Reserve, Ward 3169 (Paratype:?).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: ibid., Ward 3170 (Paratype:?).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: Umvoti, Thorns near Greytown, J.M. Wood 5318 (Paratype: NH).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: Durban, Berea, J.M. Wood 6350 (Paratypes: BOL, L!, NBG, NH, PRE).

Coccinia rehmannii Type: South Africa. [Kwa-Zulu Natal]: Doonside, J. Wylie herb. no. 23299 (Paratype: NH).

Coccinia rehmannii Type: Mozambique. Maputo: Lourenço Marques [Maputo], J.M. Borle 253 (Paratypes: M! [M-0198510], MO!, P! [P05620807, digital image: P]).

Coccinia rehmannii Type: Mozambique. Maputo: ibid., J.M. Borle 427 (Paratype:?).

Coccinia rehmannii Type: Mozambique. Maputo: ibid., J.M. Borle 442 (Paratype: PRE!).

Coccinia rehmannii Type: Mozambique. Maputo: Inhaca Island, H.G. Breyer TRV20506 (Paratype: PRE).

Coccinia rehmannii Type: Mozambique. Maputo: Inhachingo, A.W. Exell et al. 630 (Paratype: SRGH).

Coccinia rehmannii Type: Mozambique. Maputo: Massinga, A.W. Exell et al. 645 (Paratype: SRGH).

Coccinia rehmannii Type: Mozambique. Maputo: Lourenço Marques [Maputo], A.J.W. Hornby 4599 (Paratype: PRE!).

Coccinia rehmannii Type: Mozambique. Maputo: Delagoa Bay [Maputo Bay], H.A. Junod 20 (Paratypes: BR!, G (2)!, Z! [Z-000073406, digital image: Z]).

Coccinia rehmannii Type: Mozambique. Maputo: Inhaca Island, 6 Jul 1958, A.O.D. Mogg s.n. (Paratype: PRE!).

Coccinia rehmannii Type: Mozambique. Maputo: ibid., 14 Dec 1955, A.R.A. Noel s.n. (Paratype: PRE!).

Coccinia rehmannii Type: Mozambique. Maputo: Lourenço Marques [Maputo], R. Schlechter 11555 (Paratypes: BOL, COI!, G (3)!, GRA, HBG! [HBG518897], PR! [PR 801378], WAG! [WAG0234182], Z! [Z-000073405, digital image: Z]).

Coccinia rehmannii Type: Mozambique. Maputo: Katembe [Catembe], R. Schlechter 11614 (Paratypes: G (2)!, GRA, PRE!, Z! [Z-000073407, digital image: Z]).

Coccinia ovifera Dinter & Gilg in Dinter, Veg. Veldkost Südw.-Afrik.: 16. 1912.

Coccinia rehmannii Type: [Namibia]. Karas: Sandverhaar, M.K. Dinter 1214 (Syntype:?).

Coccinia rehmannii Type: [Namibia]. Otjozondjupa: Otjiwarongo, female, fl, fr, Jan 1912, M.K. Dinter s.n. (Syntype: SAM [SAM0072115-0, digital image: JPS]).

Coccinia rehmannii Type: [Namibia]. Waldau, female, fr, 3 Feb 1917, M.K. Dinter 432 (Lectotype, designated here: SAM [SAM0066515-0, digital image: JPS]).

Coccinia rehmannii Further possible syntypes (cited in Dinter 1919/20) if collected before end of 1912: Oshikoto: Gaub, M.K. Dinter 2412 (?); Tsumeb, M.K. Dinter s.n. (?). Unknown: Hereroland, Palmenwald, M.K. Dinter s.n. (?); Hereroland, Wilhelmsberg, M.K. Dinter s.n. (?).

Description.

Perennial climber or creeper. Stems up to 4 m, glabrous or with broad-based trichomes, when old often densely white pustulate (esp. in drier areas). Petiole 0.2-4.2 cm, glabrous or with erect, broad-based or often up to 1.5 mm long, articulate trichomes or only wart-like, when old sometimes dense white pustulate (esp. in drier areas). Leaves 0.9-9.7 × 1.4-16.6 cm, shallowly to deeply 3- or 5-lobate, auriculate, sometimes lobulate, rarely cordate. Lobes and lobules usually extending, rarely pointing towards tip, narrowly to broadly triangulate to lanceolate. Leaf margin rather remotely denticulate. Apex acute to obtuse, apiculate. Upper leaf surface more or less densely white pustulate, pustules sometimes with a thick, small trichome, on nerves often with thick, small trichomes. Lower leaf surface glabrous, sometimes with small, blackish glands between nerves, nerves usually with erect trichomes, sometimes wart-like. Probract usually absent, if present then up to 3.5 mm. Tendrils simple. Male flowers 1-3 solitary, if fasciculate or in few-flowered racemes then accompanied by 1-2 flowers. Common peduncle 0.7-4.5(-8.5) cm, glabrous or with long, articulate trichomes. Pedicel of flowers in inflorescences 0.6-2.8 cm, bracts up to 2.5 mm or missing. Pedicel of solitary flowers (0.2-)0.5-5(-9) cm, glabrous or especially at apex with long, articulate trichomes. Perianth tube usually with long (> 0.5 mm) trichomes, rarely almost glabrous. Calyx lobes 0.2-7 mm, narrowly lanceolate or lineal, when young erect, later also spreading to reflexed. Corolla 0.8-2.5 cm long, buff to more or less pale yellow, sometimes with green venation. Lobes 0.3-1.1 cm. Filament column pale buff, anthers buff, pollen sacs yellow (Fig. 10a). Female flower solitary. Pedicel 0.4-1.5 cm, glabrous or with long, articulate trichomes. Hypanthium usually with long (> 0.5 mm) trichomes, rarely almost glabrous, calyx lobes and corolla like in male flowers. Ovary rarely glabrous, often more or less densely covered with articulate trichomes. Style columnar, green. Stigmas 2-lobed, yellow (Fig. 11b). Immature fruit, rarely also at maturity, with whitish, longitudinal mottling that develops a dark green corona during ripening. Ripe fruit globose to elliptical 1-6.2 × 1-2.8 cm, glabrous or with few articulate trichomes, bright orange to red. Seeds 4.5-7 × 2-3.5 × 1-1.2 mm (L/W/H), asymmetrically obovate to somewhat falcate, face flat.

Phenology.

Flowering time: January-April, June, October-December.

Distribution.

Fig. 37. South Africa (except Western Cape and SW Eastern Cape), Namibia (except hyperarid regions), Swaziland, southern Mozambique (Gaza, Inhambane, Maputo), Botswana, Zimbabwe (Manicaland, Masvingo, Matabeleland South), southern Angola (Namibe, Huila, Cunene, Cuandocubango). Elevation sea level to 1850 m. Limestone, dolomitic, quartzitic, granitic, and ultrabasic soils. Possibly some tolerance to Ni and Cu. On loam, clay, white and red sand, sandstone, and gravel, but prefers sandy (well drained) soils ( Meeuse 1962). Full sun to shade. Coastal dunes, riverbanks, Acacia sclerocarya - Acacia caffra woodland, Combretum apiculatum bushland, Grewia flava bushland, mopane, Kalahari thornveld, grassland, semi-desert, dunes. Grazing is tolerated. Light frost seems to be tolerated ( Meeuse 1962).

Use.

Tuber edible after baking ( Dinter 1912, M.E. Keith 50, B. de Winter & O.A. Leistner 5598). Leaves used as spinach by Shangane people in Gaza province of Mozambique (K.L. Tinley 3000). Fruits edible (R.H.W. Seydel 938, R. Story 5029, B. de Winter & O.A. Leistner 5598).

Vernacular names.

Muchope [Xichope?]: fuculumué (L.A. Grandvaux Barbosa & F. de Lemos 8502), Otjiherero: otjimaga (M.K. Dinter s.n. Jan 1912), Ronga dialect [of Xitsonga language]: inyamgwazi (A.O.D. Mogg 31308), shiracarana (L. Macuácua 73 and 75), Tsonga [Xitsonga]: inyamwazi (A.O.D. Mogg 31538), nyampape (C. Liengme 491), Zulu [isiZulu]: uselwa-iwenyoka ( Sewram et al. 2006).

Remarks.

The (sub-)glabrous " littoralis " form can be easily confused with the polymorphic Coccinia adoensis (Hochst. ex A.Rich.) Cogn., which differs in shorter calyx lobes and lenticular seeds, and with Coccinia senensis , which also has lenticular seeds and usually long-peduncled male racemes.

Meeuse’s variety littoralis is hard to define as the paratypes are variable, and characters for delimitation are unclear. For example, although the variety should lack white speckles on the stems, there are some individuals with white speckles along with long pedicels or conspicuous racemes in KwaZulu-Natal and southern Mozambique as in the variety littoralis . The holotype of var. littoralis is, in the present author’s opinion, rather intermediate between the holotype of var. rehmannii and the subglabrous forms, e.g., from Inhaca Island. However, the tendency that Meeuse describes is apparent. Other characters in the collections of his variety, viz. relatively long calyx lobes and petals, also occur in the high mountains of Namibia but also in the whole periphery of the Coccinia rehmannii distribution range. Strangely, Meeuse does not mention the most striking difference between Coccinia rehmannii collections from the inland/dry areas and coastal/peripheral collections being the globose fruit in inland/dry area individuals or long elliptical fruit in coastal/peripheral individuals (shown and mentioned in this treatment as " Coccinia rehmannii aff. var. littoralis "; Fig. 10a) respectively. Collections with this fruit morph also occur in Angola and Zimbabwe but do not follow Meeuse’s other characters of the var. littoralis . On the other hand, collections with long calyx lobes and long petiolate flowers can have globose fruits. Like the area of the southeastern coast of South Africa, areas in the north of southern Africa receive more and longer rainfall per year than the inland, so there is a possible correlation between precipitation and fruit morphology. Elliptical fruits also occur in the closely related Coccinia trilobata from mountainous and thus more humid habitats but not in Coccinia microphylla from the dry bushlands of NE Africa whose fruit is globose. However, the characterization by Meeuse that Coccinia rehmannii is an aggregate species seems to be true. It might be interesting to link morphological characters with haplotypes and to test the fitness of these morphs in the different habitats. In any case, the morphological differentiation seems to be stable in cultivated individuals, and artificial crosses between different forms (inland vs. subglabrous from the Southeast) result in the onset of a normal fruit (resulting seeds were not used for cultivation).

The placement of Coccinia rehmannii var. littoralis forms with other Coccinia rehmannii forms in plastid and nuclear phylogenies (Figs 17, 18) also rejects the hypothesis of a hybrid origin (e.g., with Coccinia adoensis or Coccinia senensis ), but rather suggests parallel evolution due to ecological factors.

Taxonomic remarks.

The protologue contains a literal mistake for the syntype from Eland River and must be corrected from 4944 to 4954. On the one hand, A. Rehmann 4944 (GRA, K) is a Malpighiaceae . On the other hand, there is A. Rehmann 4954 (a Coccinia rehmannii ) in Z from Eland river and with a remark by Cogniaux 'sp. nov.' Hence, A. Rehmann 4954 is the syntype of Coccinia rehmannii , not A. Rehmann 4944.

The GRA specimen of H.G. Flanagan 457, which is supposed to be the isotype (cited by Meeuse) of Coccinia rehmannii var. littoralis , is in fact merely a paratype. Meeuse stated clearly the location as "Komgha: Kei Mouth" and chose the PRE specimen from there as the holotype, but the GRA specimen is from the nearby located Cape Morgan. Apparently, Flanagan used the same collection number for different gatherings. The GRA specimen thus cannot be regarded as a duplicate despite the same number. As the GRA specimen does not have a label by Meeuse, he just cited the specimen without seeing it.

The similarity of Coccinia rehmannii var. littoralis to Coccinia senensis led to a misplaced paratype. One of the two specimens by R. de Carvalho is a syntype of Coccinia jatrophiifolia var. australis Cogn. and the two R. de Carvalho specimens from COI are paratypes of Coccinia subglabra , which are both synonyms of Coccinia senensis .

Coccinia ovifera is a validly published name, although the description is a little cryptic, hence the species name is not a nomen nudum. Dinter writes that he has found, viz. collected, the species around Grootfontein, in Hereroland (not in the narrow sense of the 1968 homeland) and in Sandverhaar (Namaland). Therefore, the requirements for validity are met (37.3 Note 2). The latter site is cited by him explicitly in a later publication ( Dinter 1919/20). The collections designated as syntypes above match the description as given in the protologue. As the present author did not see the specimens designated as "possible syntypes", they are tentative and might have been destroyed in Berlin.

Specimens examined.

(Selection, in total: 321) Angola. Huíla: Cambos, near Chiange, A. Menezes 3629 (LISC [LISC 031347], P). Namibe: andados ca. 55 km de Moçamedes [Namibe] para Dois Irmãos [Caraculo], E.J. Mendes 3969 (COI, LISC). Botswana. Ghanzi: 200 mls [320 km] NW of Molepolole, R. Story 5029 (COI (2), EA (2), PRE (2), Z). Kgalagadi: c. 50 mls [80 km] NNW of Tsabong, O.A. Leistner 3120 (LISU, M, PRE). Ngamiland: 80 km W of Tsau on Cae Cae road, D.G. Long & D.A.H. Rae 416 (E). Mozambique. Gaza: Vila de João Belo [Xai-Xai], entre Chicumbane e a Barra do Limpopo: próximo da povoação commercial da Barra, F. de Lemos & A. Balsinhas 131 (BM, COI) and 133 (BM, COI (2)). Inhambane: Pomene, in hotel area, P.C.M. Jansen et al. 7533 (G n.v. [G00305107], MO, WAG [WAG0234198], WAG [WAG0234199]). Maputo: between Costa do sol and Marracuene, Mutanhane, A. Balsinhas 230 (BM, COI, PRE). Namibia. Erongo: Farm Anschluss, 150 km E of Swakopmund on Khomas road to Windhoek, B. de Winter & D. Hardy 8001 (M, PRE, WAG [WAG0234173]). Hardap: c. 20 mls [32 km] from Kalkrand on road to Rehoboth, B. de Winter 3538 (COI (2), L, M, PRE). Karas: Dassiefontein Farm, 2-3 km E of highway, in foothills of Groot Karasberge, c. 64 km NNE of Grünau, G. Davidse & A. Loxton 6240 (M, MO, S). Omaheke: [Farm] Breitenberg, border of Kalahari, R.H.W. Seydel 2513 (COI, M, WAG [WAG0234172]). Otjozondjupa: 160 mls [257.5 km] E of Grootfontein, Gautscha Pan, R. Story 6219 (PRE); ibid., R. Story 6238 (M (2), PRE (2), S [S08-12417]). South Africa. Eastern Cape: Port St. Johns distr., First Beach, M.J. Wells 3434 (MO). Gauteng: Pretoria, Brummeria: BRI, A. Balsinhas 3474 (MO, PRE, WAG [WAG0234189]). KwaZulu-Natal: Lower Tugela Valley, below Maqumbi, D. Edwards 3053 (B, M, HEID, PRE); 5 mls [8 km] on Nkonkoni-Pongola road, M.J. Wells 2162 (B, EA, M, Z [Z-000060820]). Limpopo: Immerpan, near post office on roadside, A.D.J. Meeuse 9452 (B, COI, L, PRE, Z [Z-000060813]). Mpumalanga: Impala, siding, E. Retief 1260 (PRE); ibid., E. Retief 1261 (MO, PRE); Steelport, Burgersfort, 2 km E of town, 24°40'S, 30°22'E, H.J. Venter & A. Venter 10260 (S [S08-12382], WAG [WAG0234178]). Northern Cape: 25 ml. [40.2 km] W of Kimberley, H.J.E. Schlieben & H.R. Tölken 11017 (G (2), HEID, M, PRE, S). Northwest: Rooisloot, 6 Apr 1935, A.O.D. Mogg s.n. (B, COI, L, PRE); Farm Welgevonden, 6 Apr 1935, A.O.D. Mogg s.n. (B, L, PRE [PRE43077], Z [Z-000060812]). Swaziland. Lubombo: Thsaneni [Tjaneni], I.F. La Croix 4909 (MO, WAG [WAG0234170]). Zimbabwe. Manicaland: Chipinga district, Sabi Valley Experimental Station, C. Soane 162 (COI (3)). Matabeleland South: Beit Bridge [Beitbridge], A.W. Exell et al. 425 (LISC); ibid., L.C. Leach 10700 (COI, MO).