Coccinia grandis (L.) Voigt, Hort. suburb. Calcutt.: 59. 1845.

Holstein, Norbert, 2015, Monograph of Coccinia (Cucurbitaceae), PhytoKeys 54, pp. 1-166 : 70-77

5. Coccinia grandis (L.) Voigt, Hort. suburb. Calcutt.: 59. 1845.

Bryonia grandis L., Mant. Pl.: 126. 1767. Coccinia indica Wight & Arn., Prodr. fl. Ind. orient.: 347. 1834. nom. illeg. [nom. superfl. as epithet has not been adopted]. Coccinia grandis M.Roem., Syn. Pepon.: 93. 1846. nom. illeg. [nom. superfl.] Cephalandra indica Naudin, Ann. Sci. Nat. Bot. ser. 5, 5: 16. 1866. nom. illeg. Cephalandra grandis (L.) Kurz, J. As. Soc. Beng. 46(2): 103. 1877. Type: Sri Lanka. Bryonia foliis subrotundis angulosis, momordicae facies Burm., Thes. zeylan.: 49, t. 19, fig. 2. 1737 (Type: drawing in l.c.). Type: Sri Lanka. Vitis alba indica Rumphius [G. E. Rumpf], Herb. Amboin. 5: 448, t. 166, fig. 1. 1747 (Type: drawing in l.c.).

Coccinia grandis Type: No detailed information, female, fl, Anon. in Herb. Linn. 1153.2 (Typolectotype, designated by Nazimuddin and Naqvi (1984): LINN! [digital image: LINN]).

Coccinia grandis Type: India. Gottori, male and female, fl, Anon. in Herb. Linn. 1153.3 (Type?: LINN! [digital image: LINN]).

Coccinia grandis Type: India. [Gujarat]: Suratt [Surat], female, fl, Anon. in Herb. Linn. 1153.12 (Type?: LINN! [digital image: LINN]).

Coccinia grandis Type: India. No detailed information, female, fl, Anon. in Herb. Linn. 1153.13 (Typotype: LINN! [digital image: LINN]).

Cucumis sativus var. arakis Forssk., Fl. aegypt.-arab.: 169. 1755.

Coccinia grandis Type: Yemen. [Al-Hudaydah Governorate]: Lohaja [Al-Luhayyah] [region?], Môr [Mawr], male, fl, P. Forsskål 660 (Holotype: C [C10002122, digital image: JPS, microfiche IDC: 35 II, 7-8]).

Turia moghadd Forssk. ex J.F.Gmel., Syst. nat. 2(1): 403. 1791. Turia moghadd Forssk., Fl. aegypt.-arab.: 166. 1755. nom. inval. Coccinia moghadd (Forssk. ex J.F.Gmel.) Asch. in Schweinf., Beitr. Fl. Aethiop.: 251. 1867. Cephalandra moghadd (Forssk. ex J.F.Gmel.) Broun et Massey, Fl. Sudan: 105. 1929.

Coccinia grandis Type: Yemen. [Al-Hudaydah Governorate]: Lohaja [Al-Luhayyah], female, fl, P. Forsskål 663 (Lectotype, designated here: C! [microfiche: IDC 110 I, 1-2]).

Coccinia grandis Type: ibid., male, fl, P. Forsskål 662 (Syntype: C! [microfiche: IDC 109 III, 7-8]).

Coccinia grandis Type: ibid., P. Forsskål 666 (Syntype: C! [microfiche: IDC 110 I, 3-4]).

Bryonia alceifolia [sphalm. alceaefolia ] Willd. in Rottler, Neue Schriften d. Ges. Naturf. Freunde Berlin 4: 223. 1803. Coccinia cordifolia (L.) Cogn. var. alceifolia [sphalm. alceaefolia ] (Willd.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 531. 1881.

Coccinia grandis Type: India. [Tamil Nadu]: Tiruchinapally [Tiruchirappalli], male and female, fl, Nov 1793, Anon. [J.G. Klein, B. Heyne or J.P. Rottler] in Herb. J.G. Klein 177 (Lectotype, designated here: B-W! [B-W 18065], isolectotype: K!).

Momordica covel Dennst., Schlüssel Hortus malab.: 23. 1818. Cucumis pavel Kostel., Allg. med.-pharm. Fl. 2: 738. 1833. nom. illeg. [nom. superfl.]

Coccinia grandis Type: Covel Rheede, Hort. malab. 8: 27, t. 14. 1688 (Holotype: drawing in l.c.).

Momordica bicolor Blume, Bijdr. fl. Ned. Ind.: 928. 1825-26.

Coccinia grandis Type: Indonesia. [Java], Kuripan, in calcareis [on calcareous ground?], K.L. Blume 1012 (Holotype: L! [L 0587745]).

Momordica bicolor var. a Blume, Bijdr. fl. Ned. Ind.: 928. 1825-26. nom. inval. Indonesia. Maluku Province, Timor, A. Zippelius s.n. (L! [L 0587743]).

Momordica bicolor var. b Blume, Bijdr. fl. Ned. Ind.: 928. 1825-26. nom. inval. [Indonesia]. [Java], Parang Mts, calcareous mountains. K.L. Blume 1016 (L! [L 0587744]).

Bryonia moimoi Ser. in DC. Prodr. 3: 305. 1828. Moï-moï Adans., Hist. nat. Sénégal: 159, Paris. 1757. nom. inval. Coccinia moimoi (Ser.) M.Roem., Syn. Pepon.: 93. 1846. Type: Sri Lanka. Bryonia folio anguloso acuto glabro Burm., Thes. zeylan.: 48, t. 19, fig. 1. 1737 (Holotype: drawing in l.c.).

Momordica monadelpha Roxb., Fl. ind. [2nd ed.], 3: 708. 1832. nom. illeg. [nom. superfl.] pro parte. Bryonia foliis cordatis oblongis angulatis dentatis glabris. Fl. Zeyl. 356. 1747. nom. inval. Type: Sri Lanka. P. Hermann Mus. Zeyl. 2: 37 (Type: BM [BM000621642], BM [BM000621643]) [these two specimens are Cucumis maderaspatanus L.].

Coccinia grandis Type: Sri Lanka. P. Hermann Mus. Zeyl. 5: 225 (Lectotype, designated here:: BM [BM000595000]).

Coccinia grandis Type: Sri Lanka. P. Hermann Mus. Zeyl. 5: 321 (Type: BM [BM000621089]). Type: Sri Lanka. Bryonia folio anguloso acuto glabro Burm., Thes. zeylan.: 48, t. 19, fig. 1. 1737. (Type: drawing in l.c.). Type: Sri Lanka. Bryonia foliis subrotundis angulosis, momordicae facies Burm., Thes. zeylan.: 49, t. 19, fig. 2. 1737 (Type: drawing in l.c.). Type: Sri Lanka. Vitis alba indica Rumphius [G. E. Rumpf], Herb. Amboin. 5: 448, t. 166, fig. 1. 1747 (Type: drawing in l.c.).

Coccinia loureiriana M.Roem., Syn. Pepon.: 93. 1846. Bryonia grandis Lour., Fl. cochinch. 1(2): 595. 1790. nom. illeg. and Fl. cochinch. 2: 731. 1793. nom. illeg. Type: Sri Lanka. Bryonia foliis subrotundis angulosis, momordicae facies Burm., Thes. zeylan.: 49, t. 19, fig. 2. 1737. (Type: drawing in l.c.). Type: Sri Lanka. Vitis alba indica Rumphius [G. E. Rumpf], Herb. Amboin. 5: 448, t. 166, fig. 1. 1747 (Lectotype, designated here: drawing in l.c.).

Coccinia wightiana M.Roem., Syn. Pepon.: 93. 1846. Coccinia cordifolia (L.) Cogn. var. wightiana (M.Roem.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 531. 1881. Coccinia grandis var. wightiana (M.Roem.) Greb. in R. Mansfeld & J. Schultze-Motel, Verz. Landwirtsch. u. Gaertn. Kulturpfl. 2: 929. 1986. Coccinia indica Wight et Arn. β, Prodr. fl. Ind. orient.: 347. 1834.

Coccinia grandis Type: India. [Tamil Nadu]: [= Wall. Cat 6711a], [J.G. Klein, B. Heyne or J.P. Rottler] in Herb. Madras s.n. (Syntype: E! [E00174668, digital image: E, JPS]).

Coccinia grandis Type: India. [Tamil Nadu]: [Chennai, Saidapet], Nopalry [= in Wall. Cat. 6711b or e], female, fl, R. Wight 1124 (Lectotype, designated here: E! [E00174667, digital image: E, JPS]).

Coccinia grandis Type: India. [Tamil Nadu]: Negapatam [Nagapattinam], female, fl, fr, R. Wight 1124 (Syntype: E! [E00174666, digital image: E, JPS]).

Coccinia grandis Type: unknown. R.Wight 1124 (Syntypes: E! [E00174664], NY! [00172358, digital image: NY]).

Coccinia grandis var. quinqueangularis Miq., Fl. Ned. Ind. 1(4): 673. 1856.

Coccinia grandis Type: [Indonesia]. [Central Java]: near Soerakarta [Surakarta], T. Horsfield s.n. (Holotype: U!).

Coccinia schimperi Naudin, Ann. Sci. Nat. bot., ser. 4, 8: 366. 1857. Cephalandra schimperi (Naudin) Naudin, Ann. Sci. Nat. Bot. ser. 5, 5: 16. 1866.

Coccinia grandis Type: Ethiopia. In Semen [Semien Mts], female, fl, 1854, G.H.W. Schimper Herb. Abyss. 1215 (Lectotype, designated here: P! [the specimen with thick branch and fruit], isolectotype: P!).

Coccinia grandis Type: Ethiopia. Biria Dekeno et Dschadscha, 5000', female, fl, 1853, G.H.W. Schimper Herb. Abyss. 1215 (Syntype: P!).

Coccinia grandis Type: Ethiopia. Dschadscha, 5000 ’-5500’, male and female, fl, fr, 13 and 21 Jul 1853, G.H.W. Schimper Herb. Abyss. 1215 (Syntype: P!).

Coccinia grandis Type: Ethiopia. Dschadscha, 5000', male and female, fl, fr, 13 and 21 Jul 1853, G.H.W. Schimper Herb. Abyss. 1215 (Syntype: P!).

Coccinia grandis Type: Ethiopia. without details, G.H.W. Schimper Herb. Abyss. 1215 (Syntypes: BR! [BR0000005113958, digital image: BR, JPS], BR! [BR0000005114641, digital image: BR], BR! [BR0000008251053, digital image: BR, JPS], BR [BR0000008351050], G-DC!).

Coccinia grandis Type: Ethiopia. Dschadscha, female, 1853, G.H.W. Schimper Herb. Abyss. 1215 (Syntype: BR! [BR0000005111923, digital image: BR]).

Coccinia palmatisecta Kotschy, Sitzungsber. K. Akad. Wiss. Math.-Naturwiss. Cl. Abt. 1, 51: 360-361. 1865.

Coccinia grandis Type: [South Sudan], Kyk [Ciec (a Dinka subtribe) realm, S of confluence of Bahr al-Ghazal and White Nile], male, fl, no date given, M.L. Hansal s.n. (Lectotype, designated here: W! [K neg. 4837]).

Coccinia grandis Type: [South Sudan]. marshes in Noer [Nuer] realm [S to E of Malakal], no detailed location given, F. Binder s.n. (Syntype: W! [K neg. 4838]).

Cephalandra indica var. palmata C.B.Clarke in Hook.f., Fl. Brit. Ind. 2(6): 621. 1879.

Bryonia alceifolia [sphalm. alceaefolia ] Willd. in Rottler, Neue Schriften d. Ges. Naturf. Freunde Berlin 4: 223. 1803. Coccinia wightiana M.Roem., Syn. Pepon.: 93. 1846.

Coccinia grandis Type: India, [Tamil Nadu]: Tiruchinapally [Tiruchirappalli], male and female, fl, Nov 1793, Anon. [J.G. Klein, B. Heyne or J.P. Rottler] in Herb. J.G. Klein 177 (Syntypes: B-W! [B-W 18065], K!)

Coccinia grandis Type: India. [Tamil Nadu]: [= Wall. Cat 6711a], [J.G. Klein, B. Heyne or J.P. Rottler] in Herb. Madras s.n. (Lectotype, designated here: E! [E00174668, digital image: E, JPS]).

Coccinia grandis Type: India. [Tamil Nadu], [Chennai, Saidapet], Nopalry [= in Wall. Cat. 6711b and e], female, fl, R. Wight 1124 (Syntype: E! [E00174667, digital image: E, JPS]).

Coccinia grandis Type: India. [Tamil Nadu]: Negapatam [Nagapattinam], female, fl, fr, R. Wight 1124 (Syntype: E! [E00174666, digital image: E, JPS]).

Coccinia grandis Type: unknown. R.Wight 1124 (Syntypes: E! [E00174664], NY! [00172358, digital image: NY]).

Coccinia cordifolia (L.) Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 529. 1881. pro parte majore, non Bryonia cordifolia L.

Coccinia cordifolia (L.) Cogn. var. genuina Cogn. in A.DC. & C.DC., Monogr. Phan. 3: 531. 1881. nom. invalid.

Description.

Perennial climber or creeper. Stems up to 5 m, glabrous, when older often white pustulate. Petioles 0.5-5.5 cm, glabrous, rarely some trichomes on adaxial side. Leaves 3-11 × 3-13 cm, cordate to 3-lobate or 5-edged to 5-lobate, sometimes lobulate. Lobes triangulate, ovoid, oblong, to obovoid. Leaf margin dentate, teeth usually with yellowish-reddish to brownish gland (Fig. 7a), becoming black when dried. Margin rarely with short (<1 mm), whitish trichomes. Apex obtuse to acute with final tooth. Upper leaf surface glabrous, more or less dense hyaline to white pustulate. Lower leaf surface glabrous, with glands that are usually framed with lighter color between major nerves, nerves sometimes with white pustules. Probracts <1.5 mm or missing. Tendrils simple. Male flowers 1(-3) solitary, rarely in short racemes. Peduncle 0.3-1.5 cm, glabrous, pedicels of flowers in racemes up to 3.2 cm, glabrous, pedicels of solitary flowers up to 4.5 cm, glabrous. Bracts inconspicuous (<1 mm), often absent. Perianth tube glabrous. Calyx lobes 1.2-3.5 mm long, lineal, spreading to reflexed, often with reddish to brownish gland on lower side at the acute tip (Fig. 7a). Corolla 1.7-4 cm long, yellowish buff (mostly African collections) to snow-white (esp. outside Africa). Corolla lobes 0.7-1.7 cm. Filament column and anther head pale greenish, pollen sacs yellow. Female flowers 1(-2) solitary. Pedicel up to 1 cm, glabrous. Hypanthium glabrous, calyx and corolla like in males. Ovary glabrous. Style columnar, yellowish-green. Stigmas 2-lobed, greenish. Fruit globose to ellipsoid, cultivated also shortly cylindrical, 3-4.5 × 1.5 cm, glabrous, unripe greenish with few pale spots and/or lines, becoming scarlet red when ripe. Seeds 5-7 × 2.5-3.5 × 1.2 mm (L/W/H), asymmetrically obovate, face flat (Fig. 14f).

Phenology.

Flowering time: All over the year, but not at the end of the dry season or in cold seasons. Seems to need 1-2 weeks of at least 10 hours daylight with sunny weather for flower induction (pers. observ. from greenhouse cultivation).

Distribution.

Figs 26, 27. Benin, N Cameroon, S Chad, D. R. Congo (in Great African Rift valley), Djibouti, Egypt (along Nile, Elba Mts), Eritrea, Ethiopia, Kenya, Mali (except N), S Mauritania, Niger, Senegal, Somalia, South Sudan, S and E Sudan, N Tanzania (Arusha, Dar es Salaam, Manyara, Morogoro, Mwanza, Pwani, Tanga, Zanzibar), Uganda, mountains and oases of the southern and western Arabian Peninsula, tropical and subtropical India, subtropical Nepal, Pakistan, Sri Lanka, South East Asia, S China incl. Hainan. Occurrence likely due to recent introduction in: Australia (Northern Territory, Queensland, Western Australia), R. China, Maldives, Mauritius, Mozambique, USA (Florida, Guam, Hawaii, Wake Island), many tropical Pacific islands, Caribbean area, Central and tropical South America. 0 to 1600 m. Black clay soil, black cotton soil, sand, on limestone. There seems to be a certain salt tolerance ( Bharathi 2007). Grasslands, bushlands, (semi-arid) savannas, thickets, along rivers, ruderal sites, seemingly avoiding closed canopies (woodlands and forests). Coccinia grandis is, especially in tropical Asia, often reported from sand or from calcareous grounds (karst areas), which are well-drained. Although precipitation in South East Asia is much higher than in Africa, the quick loss of surface water allows Coccinia grandis to survive there.

Use.

Fruits (raw and cooked) and shoots (cooked) edible. The Luo eat the leaves as spinach ( Orech et al. 2005). The sap is used against diabetes ( Ramachandran and Subramaniam 1983) due to its hypoglycemic activity ( Chopra and Bose 1925; Shibib et al. 1993). It is used in traditional Indian medicine in India for various diseases, and seems to have a general antibacterial effect (see also Use, economic potential, and phytochemistry).

Vernacular names.

Agau or Chomir [most likely: Khamir/Xamtanga language of Agaw language family]: amballa gosa (G.H.W. Schimper 365, Schweinfurth 1893); Arabic: mogad ( Getahun 1974b); Bagirmi: na odio (A.J.B. Chevalier 9527); Bengali: tala-kucha ( Nadkarni and Nadkarni 1976), tela kucha [ tēlākucā] (J. Sinclair 4414); Canarese [Kannada]: tonde-konde, tonde-kayee [toṇḍe kāyi] ( Nadkarni and Nadkarni 1976); Chinese: hong gua [ Hóng guā] ( Wu et al. 2011); Dassanetch [Daasanach]: dali (C.J. Carr & C. Metolo 867); Galinya [Oromo]: raho (T. Ebba 624; Getahun 1974b); Gujarati: gholi ( Nadkarni and Nadkarni 1976); Hindi: kan-duriki-bel ( Nadkarni and Nadkarni 1976), kundru [kundrū] ( Bharathi 2007); Karen [in Mae Hong Son Province, Thailand]: khae-do ( de Wilde and Duyfjes 2008); Khmer: ( sloek’) bah ( Kéraudren-Aymonin 1975b); Kizigua: lukewja (R.E.S. Tanner 2691), pondwa (R.E.S. Tanner 2030); Lao: ( khùa 'phăk) tăm ling, tăm nhing ( Kéraudren-Aymonin 1975b), phak tam nin (J.-M. Dubost 200); Luo: nyathud gush (S.H. Padwa 71); Mahrati [Marathi]: ran-tondula, tondli ( Nadkarni and Nadkarni 1976); Malayalam: kova ( Nadkarni and Nadkarni 1976), kovakka ( Bharathi 2007); Masai: olaposhi (C.H. Pecler 3); Nepali: gol kankri ( Aryal 2007); Ngadha (Baba-Ngadha): 'uta kala (J.A.J. Verheijen 5415); Ngadha (Mataloko-Ngadha): uta kobho (J.A.J. Verheijen 5415); Persian [Farsi]: kabare-hindi ( Nadkarni and Nadkarni 1976); Punjabi: kanduri ( Nadkarni and Nadkarni 1976); Sanskrit: vimboshta, vimbaja, bimba, tundika [tu ṇḍikā] ( Nadkarni and Nadkarni 1976); Serer: ayandyaloup (J.-F. Ezanno Cucurbitaceae 9); Sinhala: kowakka (D. Philcox et al. 10454, D. Philcox et al. 10458); Somali: masskar (T. Ebba 804); Songhai: lombaria (A.J.B. Chevalier 2996); Sunda: aroi papassang ( Miquel 1855); Tamasheq [Touareg]: seffel (H. Lhote 2). Tamil: kovai [kowai] ( Nadkarni and Nadkarni 1976), kovakkai ( Bharathi 2007); Telugu: dondakaya [doṇḍa kāya] ( Bharathi 2007), dondatiga, kakidonda ( Nadkarni and Nadkarni 1976); Thai: in Central Region: phak tamlueng, in Northern Region: phak khaep ( de Wilde and Duyfjes 2008); Tigrinya: asumbeh (G.A. Schweinfurth & D. Riva 1007); Vietnamese: northern: hoa bát, southern: ( dây) bai bác, bình bát, ma'nh bát, lá bát ( Kéraudren-Aymonin 1975b); Wolof: barbouf (J. Trochain 663).

Remarks.

With the exception of South and South East Asia, Coccinia grandis is easily recognizable, especially by the lack of an obvious indumentum and the pale framed (in living state) glands in the axils of the nerves at the base of the lower leaf lamina. In NE Africa, collections with finely dissected leaves can be similar to Coccinia ogadensis . When compared to collections from South Africa, Coccinia sessilifolia var. variifolia and some forms of Coccinia mackenii are also similar, but Coccinia sessilifolia var. variifolia is glaucous, Coccinia mackenii plants have bifid tendrils, and both South African species lack colored leaf teeth and have erect to spreading calyx lobes instead of spreading to reflexed calyx lobes. In South and South East Asia, some vegetatively similar Gymnopetalum species (e.g., Gymnopetalum chinense ) can be mistaken for Coccinia grandis , as in both taxa the leaf shape is 5-edged to cordate, and glands on the lower leaf lamina can be found. However, Gymnopetalum species are rather densely beset with trichomes, have ribbed fruits and are monoecious, whereas Coccinia grandis is glabrous, has smooth fruits, and is dioecious, at least in wild populations.

Asian and (at least most of the) African Coccinia grandis differ genetically (e.g., in a short sequence in the 5'-beginning of the LEAFY -like 2nd intron) and in petal color (white in Asian, buff in African individuals). Hence, the distribution in Asia (at least India) is not due to human impact. Whether Coccinia grandis is introduced or native to Malesia, northern Australia and southern China and Taiwan is not known. Crossing experiments by Naudin (1862) indicate full compatibility between the African and Asian morphs though.

Taxonomic remarks.

Up to the 21st century (e.g., Bulbul et al. 2011; Hussain et al. 2011), there has been quite a lot of confusion about the valid name of the species that is now called Coccinia grandis . Wight and Arnott (1834) established the genus Coccinia , with the name based on the scarlet-red fruits of a species that Wight had collected several times during his 13-year stay in South India ( Stafleu and Cowan 1988). The name of the only species they described in their new genus, Coccinia indica Wight & Arn., is illegitimate since they included Bryonia grandis L., of which they ought to have adopted the epithet (Art. 52.1 ICN). The illegitimacy of Coccinia indica , however, does not affect the legitimacy of the genus name Coccinia (Art. 42.2), although there was also some confusion about which genus name to use. In 1845, Voigt published the correct combination Coccinia grandis (L.) Voigt with a description exactly matching Bryonia grandis .

Although Coccinia indica is not valid, and the problem seemed to have been solved, a third species name was brought into discussion by Cogniaux. He thought that Bryonia cordifolia L. and Bryonia grandis L. referred to the same species as Linnaeus cited Rumphius’ Vitis alba indica ( Coccinia grandis ) under Bryonia cordifolia ( Linnaeus 1763). Hence, Cogniaux created a more broadly circumscribed Coccinia cordifolia (L.) Cogn. (1878). However, Linnaeus described Bryonia cordifolia beforehand (1753) citing the description from his Flora Zeylanica number 354 (1747), and only in the 2nd and 3rd edition of his Species Plantarum did Linnaeus synonymize Rumphius’ Vitis alba indica (1747). Cogniaux therefore erred, when he stated that Linnaeus had based his Bryonia cordifolia on Rumphius’ figure, and the epithet cordifolia is hence misapplied in Coccinia .

There are four specimens in Linnaeus’ herbarium, which belong to Coccinia grandis but it is unclear which are type material. The number 1153.2 is designated (by Linnaeus?) as Bryonia grandis on the sheet and is therefore the best choice for lectotypification as is has been done by Nazimuddin and Naqvi (1984). The specimen 1153.13 bears the note " Bryonia foliis subrotundis angulosis, momordicae facie" on the specimen flip side referring to one of the citations to drawings in the protologue. Hence, this specimen is also original material. The other two specimens (1153.3 and 1153.12) do not bear indications that Linnaeus referred to them as Bryonia grandis , but there is no contraindication either, so they might be original material, too.

The drawing to Bryonia foliis subrotundis, angulosis, momordicae facie Burm. (Thes. zeylan.: 49, t. 19, fig. 1. 1737), which is original material of Bryonia grandis L. appears to be a product of artistic freedom. The calyx lobes of the three uppermost and the lowermost flowers are reflexed and match Coccinia / Bryonia grandis well, whereas the calyx of the other two flowers appears to consist of almost free elliptic petals, quite like in Momordica foetida (except for the soft spines that are missing in the drawing). The addition "momordica facies" seems to relate to this. Strangely, the drawing of Bryonia folio anguloso acuto glabro Burm. (Thes. zeylan.: 48, t. 19, fig. 1. 1737) matches the current definition of Bryonia grandis also well, but has not been cited by Linnaeus. Eventually, the synonyms that Burman used, which also include Bryonia cordifolia L. ( Cucumis maderaspatanus L.), made it hard to interpret the species and also lead to the confusion of Cogniaux.

The Forsskål 660 specimen (C10002122) has a hand-written field label by Forsskål (according to notes in JStor Plant Science) on the flip side, stating "Cucumis incerta. Arakis, Mour." (incerta meaning “uncertain”) and two hand-written identifications "Cucumis inedulis Fl. Yemen CXXII nr. 580" and "c. s. Arakis cent VI nr. 61 p. 169". The location in the text is the same as on the field label: Môr /Mour [Mawr, a small town about 30 km E of al-Luhayyah]. The former identification is a nom. nud. with the number 580 on page CXXII of Forsskål’s Flora Ægyptiaco-Arabica (1775). The second identification links to a Cucumis sativus variety that is validly described in that book on page 169. The description matches well Coccinia grandis except for the tuberculate ovary. Additionally the collection is supposed to be from Loheja [al-Luhayyah], but might only indicate the region, in which Mawr is localized. However, the Arabic name of Cucumis inedulis and Cucumis sativus var. arakis are both: Arakis [3raqīs], so they can be cross-referenced. The description of the variety also mentions that the plant is not edible, just as the supposed species epithet.

The genus name Turia has been created by Forsskål in his Flora Ægyptiaco-Arabica (1775). There is a debate, however, whether it is validly published ( Friis 1984). The descriptions of this part of Forsskål’s book (page number in Latin) must be used in consideration with the corresponding parts in the lists of local floras (Roman page numbers), which is on page CXXI in the case of Turia . The first species there is Turia sativa (no. 550), which is called “turia” in Arabic and is cultivated according to the epithet. This matches exactly the first description in the descriptive part of the book (p. 165). Forsskål lists five Turia species in the floristic part and describes five species in the descriptive part. Therefore, the genus Turia lacks a description ( Jeffrey 1962), because Forsskål does not mention any character to be typical for the genus. The name Turia was legitimately described by Gmelin in an extension of Linnaeus’ Systema naturae (1791), and so was the name Turia moghadd .

The typification of Turia moghadd is not straight-forward, because Forsskål added little, if any marks on the sheets ( Friis 1983). The flip side of the Forsskål 663 specimen has three notes: "Cucumis glandulosus", a second one with a different handwriting: " Bryonia Turia 35 Forsk" and a third one with another handwriting "Turia gijef Forsk Cent. 6 no. 38" with all words except for “Turia” crossed out. It can be hypothesized that these are different trials to identify the specimen, but not by Forsskål himself. Aside from these, the specimen lacks any written marks, but it matches exactly the description of Turia moghadd , as do the specimens Forsskål 662 and 666, which are syntypes. The Forsskål 663 specimen is chosen to be the lectotype, because the original description mentions the occurrence of fruits and this specimen is the only female of the three.

Willdenow describes Bryonia alceifolia in a travel report by J. P. Rottler from 1799, but he only mentions that he separates Rottler’s Bryonia epigaea from another new species, viz. Bryonia alceifolia , so the type was not necessarily collected in 1799. Willdenow knew Bryonia alceifolia from Klein’s specimens in his herbarium. Rottler was missionary in Tranquebar, the same place in which J. G. Klein was surgeon ( Jensen 2005). Both are known to have collected together (with B. Heyne). The lectotype collection is in the describer’s herbarium under the Klein number 177, but it is not certain, whether Klein collected the specimen himself, of if it was by Rottler or even Heyne. A duplicate with the same label data as in B-W is deposited in K. For Willdenow described the species, the specimen in his herbarium was chosen to be lectotype.

The name Momordica monadelpha Roxb. is superfluous, because Roxburgh synonymized Bryonia moimoi Ser. in total by citing the only element of that name and Bryonia grandis . The other elements of Momordica monadelpha are also interesting, though. Roxburgh cited " Bryonia foliis subrotundis " with the citation of Burman’s Thes. zeylan.: t. 19, fig. 1. 1737 and fig. 2, Vitis alba indica , all Coccinia grandis , and Herman’s Musæum Zeylanicum 356. The latter one consists of two specimens (2: 37), which are Cucumis maderaspatanus L., however, there are also two drawings (5: 225 and 5: 321) with the number 356. Both drawings represent Coccinia grandis because of the fruit size, fruit shape, and the flower morphology (calyx lobe length and position, corolla size), rather than Cucumis maderaspatanus L., Diplocyclos palmatus L. or Cayaponia laciniosa (L.) C.Jeffrey. That the drawing 5: 225 (BM000595000) shows a plant with male and female flowers on one individual might be explained best by artistic freedom.

The name Coccinia indica var. palmata C.B.Clarke is valid and legitimate but not obvious to typifiy. Despite Coccinia indica being illegitimate as a nomen superfluum, the variety is legitimate and validly described. Clarke cites Bryonia alceifolia Willd. and the Coccinia indica protologue with page 348 although Coccinia indica was described on page 347 ( Wight and Arnott 1834). Page 348, however, only comprises the β variety, which itself was the basis for Coccinia wightiana M. Roem. This unnamed variety consists of several elements, of which the literature citations of Bryonia palmata are mentioned with a question mark and are thus not eligible for typification (Art. 52.2 N1). The element " Bryonia palmata Linn.? herb. Madr.!" relates to a collection in Herbarium Madras that was identified a Bryonia palmata , but Wight and Arnott appear to have been in doubt whether the name was used sensu Linnaeus, hence the question mark after the name. This specimen (or a duplicate) is found in E and represents a deeply lobed Coccinia grandis just as the protologue of the β variety says. Clarke obviously takes the epithet of the variety from this misidentification and not from Bryonia palmata L., which is why the name was typified with this collection. That Clarke meant a deeply lobed Coccinia grandis is evident since he also cites Bryonia alceifolia , which is also deeply lobed. It is thus not a new combination and status change but a new variety.

The name Coccinia cordifolia var. genuina Cogn. cannot be regarded as intended to represent a new variety, because Cogniaux divides all specimens he had seen into three varieties. Hence, the epithet " genuina " and his " genuina " variety is just synonymous with the autonym.

Specimens examined.

(Selection, in total: 1066). Australia. Queensland: Weipa, Awonga Court, 12°37'28"S, 141°52'39"E, B.M. Waterhouse 7428 (BRI, CANB [CANB 682857]). Bangladesh. Sylhet: Sillet, F. de Silva in Wall. list 6700g (M [ “439”], M [ “440”]). Barbados. St. Michael: Bridgetown, old railway yard, Sep 1940, E.G.B. Gooding s.n. (BAR00002591). Brazil. Minas Gerais: No detailed location, <1840, P.C.D. Clausen s.n. (G-DC). Cameroon. North Region: Pitowa [Pitoa], c. 17 km NE of Garoua, W.J.J.O. de Wilde et al. 4932 (MO, WAG [WAG0225471]). Chad. Chari-Baguirmi: Baguirmi et région du Lac Fittri, Tjecna, A.J.B. Chevalier 9527 (G, P [P05620497]). P. R. China. Hainan: Ch’ang-kiang District [Changjiang country], [Bawangling Nature Reserve], Ka Chik Shan [Qicha hill], S.K. Lau 3008 (GH n.v., P [P06394498], S). Hong Kong: near Ouang-nei-Tchong [Wang nai chung] - Happy Valley, E.M. Bodinier 1367 (E). Yunnan: Nan-chiao [Meng che], C.W. Wang 76988 (A n.v., IBSC [0207527], KUN n.v., PE [01178321], PE [01178322]). R. China. Pingtung County: Hsinpi [Sinpi], Y.-P. Cheng 4603 (BR [BR0000005164189], TAIF). Cuba. Havana: Santiago de las Vegas, A. O’Donovan 5106 (G, S [S08-12050]). D. R. Congo. Orientale: Mahagi territory, Mahagi-Port, J. Lebrun 3800 (BR (2), EA, WAG [WAG0225470]). East Timor. Dili: just W of Dili, Tasutolu area, 8°33'57"S, 125°30'05"E, L.D. Cowie 10658 (L). Egypt. Red Sea Governorate: [ Hala’ib triangle], Gebel Shendib, G.W. Murray 3857 (K [K000037307]). Eritrea. Gash-Barka: Mai Mentai [at Sciagolgol River, SE of Agordat], N. Beccari 118 (FT). Northern Red Sea Region: [NE of Ghimda/Gimda], Pianura Sabarguma, A. Pappi 3970 (G, MO, P [P05620558], S [S08-12133], W). Ethiopia. Oromia: 85 km NE of Nazareth, along road to Awash, c. 5 km W of Metahara, 8°55'N, 39°55'E, J.J.F.E. de Wilde 6870 (BR, MO, WAG [WAG0225448], WAG [WAG0225449], WAG [WAG0225450]). Tigray: near Djeladjeranné, 23 Apr 1841, G.H.W. Schimper 1570 (P [P05620544], specimens from BM, G, MO, S [S08-12147], and W [W 0011063] might also be from this location, for details see under Cucurbita schimperiana ). Guyana. Demerara-Mahaica: Georgetown, W. Hahn 4810 (MO, US). India. West Bengal: Sibpur [Shibpur in Howrah city] near Calcutta [Kolkata], J.G. Hallier s.n. (M [ “999”]). Indonesia. East Java: Pasoeroean [Pasuruan], 17 Jul 1928, J.J. Ochse s.n. (B, BR [BR0000009938076], L [L 0587578], P [P06394462], U). Mali. Kayes: Bafoulabé Cercle, Bamafélé arondissement, 3.2 km ESE of Manantali, 13°10.847'N, 10°26.260'W, C.S. Duvall 130 (MO (2)). Mozambique. Maputo: Vila Luísa [Marracuene], along Incomáti river, A. Balsinhas 1273 (COI). Nepal. Madhya Pashchim: [Rapti], Tulsipur, O. Polunin et al. 5906 (E). Nicaragua. Chontales: St Tomas [Santo Tomàs], 1841, E. von Friedrichsthal s.n. (W). Niger. Agadez: [ Aïr Mountains], Mt Bagwezan [Mt Bagzan, Idoukal-n-Taghès], 31 May 1920, A. Buchanan s.n. (MO [acc. no. 1667128]). Pakistan. Sindh: 7 mls [c. 11.2 km] from Hyderabad to Tando M.[Mohammad] Khan, S. ul-Abedin 3839 (B, KUH n.v., MO). Senegal. Dakar: Jof [Yoff], J.-G. Adam 1806 (MO (2), [non P [P00694028/P05590411], which is a Citrullus sp.]). Singapore. South East District: Potong Pasir off Serangoon Road, J. Sinclair 5434 (E, L [L 0587600], P [P0639450]). Somalia. Togdheer: [c. 54 km SSW of Berbera], foot of Sheikh Pass, P.R.O. Bally 11827 (G (3), K, PRE). Sri Lanka. North Central Province: Anuradhapura, F.R. Fosberg & N. Balakrishnan 53431 (MO, PDA n.v.); ibid., W. Forstner s.n. W13705 (W). Sudan. North Kurdufan: Chursi [Khursi], K.G.T. Kotschy Iter nub. 308 (BM, BR (2), E, G (2), G-DC, GOET, K, L, M [ “400”], M [ “401”], M [ “782”], MO, P [P05620478], P [P05620484], P [P05620578], PRC, S [S08-12152], TUB [TUB004721], TUB [TUB004722], W [W 0011066], W (2), WAG [WAG0234161]). Tanzania. Tanga: Tanga town, near public beach, 200 m E of 05°03'19.2"S, 39°07'31.3"E, N. Holstein et al. 94 (B, DSM, M, WAG), and 95 (DSM, M). Zanzibar Urban/West: Chukwani, H.G. Faulkner 2782 (BR, COI, K n.v., P [P05620529]). Thailand. Bangkok: Bangkok, R. Zimmermann 66 (B (2), BR [BR0000009938878], G (2), H, L [L 0587657], M, MO, P [P06394565], PR [PR 801377], PRC, S, U, W). USA. Florida: along Indian River [lagoon], Coco [Cocoa], 9 May 1918, J.K. Small s.n. (FLAS [FLAS 29152], G, MO [acc. no. 1161454], US). Hawaii: O’ahu Island, Dole St, behind U Hawaii Manoa dorms, B. Kennedy 47 (MO, US [00674433]). Midway Atoll: Sand Island, 4208 Commodore Ave., F. Starr & K. Martz 080601-01 (BISH, PTBG [PTBG1000013099]). Vietnam. Da Nang: Tourane [Da Nang] and vicinity, no detailed location, J. Clemens & M.S. Clemens 3257 (A n.v., G, MO, P [P06394668], U, W, Z). Yemen. Al-Hudaydah: Mor [Mawr], 15°40'N, 28-30 Mar 1825, F.W. Hemprich s.n. (P [P06394550], S [S08-12138]).