| Literature DB >> 31379450 |
Sandra Knapp1, Maria S Vorontsova2, Tiina Särkinen3.
Abstract
Solanum L. (Solanaceae) is one of the largest genera of angiosperms and presents difficulties in identification due to lack of regional keys to all groups. Here we provide keys to all 135 species of Solanum native and naturalised in Africa (as defined by World Geographical Scheme for Recording Plant Distributions): continental Africa, Madagascar (incl. the Indian Ocean islands of Mauritius, La Réunion, the Comoros and the Seychelles), Macaronesia and the Cape Verde Islands. Some of these have previously been published in the context of monographic works, but here we include all taxa. The paper is designed to be used in conjunction with the web resource Solanaceae Source (www.solanaceaesource.org) and hyperlinks provide access to online descriptions, synonymy and images (where available) of each species. All taxa treated and specimens seen are included in searchable Suppl. material 1, 2.Entities:
Keywords: Solanum ; Africa; Aldabra; Azores; Canary Islands; Cape Verde; Comoros; La Réunion; Madagascar; Madeira; Mauritius; Seychelles; cultivated plants; identification; keys; weeds
Year: 2019 PMID: 31379450 PMCID: PMC6661264 DOI: 10.3897/phytokeys.127.34326
Source DB: PubMed Journal: PhytoKeys ISSN: 1314-2003 Impact factor: 1.635
The 135 species of (native, naturalised and widely cultivated) occurring in Africa as defined by Brummitt (2001), with their places of original publication and clade membership as currently understood (Major Clades sensu Bohs 2005; minor clades are divisions within these sensu Bohs 2007; Stern et al. 2011; Vorontsova et al. 2103; Särkinen et al. 2015, 2018; Aubriot et al. 2016; Tepe et al. 2016).
| Species | Place of original publication | Major Clade | Minor clade |
|---|---|---|---|
| Prodr. [A. P. de Candolle] 13(1): 366. 1852. |
| Old World-Africa | |
| Collectanea [Jacquin] 1: 100. 1787 [‘1786’]. |
|
| |
| Tent. Fl. Abyss. 2: 105. 1850 [1851]. |
| Old World-Africa | |
| Cent. Pl. 2: 10. 1756. |
| Old World-Africa | |
| Gard. Dict. ed. 8, no. 26. 1768. | African non-spiny (ANS) | ||
| Phytotaxa 10: 32. 2010. |
| Old World-Africa | |
| Fl. Bras. (Martius) 10: 68, fig. 5, 32–33. 1846. |
| Gardneri | |
| Kew Bull. 1894: 149. 1894. |
| Old World | |
| Gard. Dict. ed. 8, no. 5. 1768. | Morelloid | Black nightshade | |
| Tabl. Encycl. 2: 23. 1794. |
| Old World-Africa | |
| Beskr. Guin. Pl. 126 1827. |
| Old World-Africa | |
| Boll. Reale Orto Bot. Giardino Colon. Palermo 7: 188. 1908. |
| Old World-Africa | |
| Syll. Ratisb. 1: 200. 1824. |
|
| |
| Repert. Spec. Nov. Regni Veg. 11: 18. 1912. |
| Old World-Africa | |
| Fl. Madag., Fam. 176: 75. 1994. |
| Old World-Madagascar | |
| Anales Hist. Nat. 1: 44. 1799. | Cyphomandra | Pachyphylla | |
| Fl. Madag., Fam. 176: 77. 1994. | African non-spiny (ANS) | ||
| Prodr. [A. P. de Candolle] 13(1): 292. 1852. |
| Old World-Madagascar | |
| Prodr. [A. P. de Candolle] 13(1): 291. 1852. |
| Old World-Africa | |
| Tent. Fl. Abyss. 2: 102. 1850. |
| Old World-Africa | |
| Syst. ed. 10: 935. 1759. |
| Old World-Africa | |
| Auct. Syn. Meth. Stirp. Hort. Regii Taur. 64. 1773. |
|
| |
| Sitzungsber. Kaiserl. Akad. Wiss., Math.-Naturwiss. Cl. 38: 576. 1860. |
| Old World-Africa | |
| Prodr. [A. P. de Candolle] 13(1): 365. 1852. |
| Old World-Africa | |
| Tabl. Encycl. 2: 18. 1794. | Morelloid | Black nightshade | |
| Linnaea 19: 304. 1847. |
| Torva | |
| Fl. Aegypt.-Arab. 47. 1775. |
| Old World-Africa | |
| Fl. Aegypt.-Arab. 47. 1775. |
| [not assigned] | |
| Phytologia 34: 282. 1976. |
| Old World-Madagascar | |
| Pl. Bequaert. 1: 425. 1922. |
| Old World-Africa | |
| Boll. Soc. Bot. Ital. 1925: 107. 1925. |
| Old World-Africa | |
| Beskr. Guin. Pl. 126 [146]. 1827. |
| Old World-Africa | |
| Bot. Jahrb. Syst. 38: 57. 1905. |
| Old World-Africa | |
| Sp. Pl. 184. 1753. |
| ||
| Icon. 3: 22, tab. 243. 1795. |
| Elaeagnifolium | |
| Prodr. Fl. Nep. 96. 1825. | Brevantherum | ||
| Prodr. [A. P. de Candolle] 13(1): 201. 1852. |
| Old World-Madagascar | |
| Hist. Nat. |
| Old World-Africa | |
| Collectanea [Jacquin] 4: 125. 1791. |
| Old World-Africa | |
| Hist. Nat. |
| Old World-Africa | |
| Bot. Jahrb. Syst. 28: 473. 1900. |
| Old World-Africa | |
| Sp. Pl. 184. 1753. | African non-spiny (ANS) | ||
| Prodr. [A. P. de Candolle] 13(1): 284. 1852. |
| Old World-Africa | |
| Phytologia 34: 282. 1976. |
| Old World-Madagascar | |
| Plant Syst. Evol. 228: 44. 2001. |
| ||
| Bot. Jahrb. Syst. 38: 184. 1906. | African non-spiny (ANS) | ||
| Tabl. Encycl. 2: 23. 1794. |
| Old World-Africa | |
| Prodr. [A. P. de Candolle] 13(1): 85. 1852. | African non-spiny (ANS) | ||
| Notizbl. Bot. Gart. Berlin-Dahlem 12: 90. 1934. |
| Old World-Africa | |
| Sp. Pl. 188. 1753. |
| Old World-Africa | |
| Mant. 1: 46. 1767. |
| Old World-Tropical Asia | |
| Fl. Madag., Fam. 176: 97. 1994. | African non-spiny (ANS) | ||
| Bot. Jahrb. Syst. 54: 501. 1917. |
| Old World-Africa | |
| Hort. Kew. ed. 1, 1: 247. 1789. | Archaesolanum | ||
| Repert. Spec. Nov. Regni Veg. Beih. 16: 107. 1923. |
| Old World-Africa | |
| Candollea 50: 217. 1995. |
| Old World-Africa | |
| Enum. Pl. (Willdenow) 1: 238. 1809. |
| Old World-Africa | |
| Blyttia 24: 368. 1966. |
| Old World | |
| Kew Bull. 41: 435. 1986. |
| Old World-Africa | |
| Kew Bull. 52(3): 703. 1997. |
| Old World-Africa | |
| Sp. Pl. 185. 1753. | Potato | Tomato | |
| Tent. Fl. Abyss. 2: 106. 1850. |
| Old World-Africa | |
| Mant. Pl. Altera: 205. 1771. |
| Old World-Africa | |
| Bot. Jahrb. Syst. 38: 185. 1906. | African non-spiny (ANS) | ||
| Prodr. [A. P. de Candolle] 13(1): 99. 1852. | African non-spiny (ANS) | ||
| Ann. Missouri Bot. Gard. 73: 498. 1986. |
| Old World-Madagascar | |
| Syst. Bot. 35: 904. 2010. |
| Old World-Africa | |
| Sp. Pl. 187. 1753. |
|
| |
| Suppl. 147. 1781. |
| Old World-Africa | |
| Repert. Spec. Nov. Regni Veg. Beih. 16: 42. 1923. |
| Old World-Africa | |
| Delic. Fl. Faun. Insubr. 3: 16. 1788. | Brevantherum | ||
| Bull. Misc. Inform. Kew 1894: 128. 1894. |
| Old World-Africa | |
| Sp. Pl. 186. 1753. |
| Old World-Tropical Asia | |
| Syst. Nat., ed. 13[bis] 2(1): 385. 1791 | Morelloid | Black nightshade | |
| Hort. Kew, ed. 1, 1: 250. 1789. | Potato | Basarthrum | |
| J. Linn. Soc., Bot. 21: 426. 1885. |
| Old World-Madagascar | |
| Fl. Madag., Fam. 176: 115. 1994. | African non-spiny (ANS) | ||
| Phyt. Canar. 2. 3(3): 123. 1845. |
| ||
| Repert. Spec. Nov. Regni Veg. Beih. 16: 163. 1923. |
| Old World-Africa | |
| Sp. Pl. 186. 1753. | Morelloid | Black nightshade | |
| Repert. Spec. Nov. Regni Veg. 11: 208. 1912. | Morelloid | Black nightshade | |
| Res. Sci. Somalia Ital. 1: 128. 1916. |
| Old World-Africa | |
| Bull. Misc. Inform. Kew 1894: 127. 1894. |
| Old World-Africa | |
| Prodr. [A. P. de Candolle] 13(1): 250. 1852. |
|
| |
| Syst. Bot. 35: 903. 2010. |
| Old World-Africa | |
| Cent. Pl. 1: 8. 1755. | Potato | Tomato | |
| Syst. Bot. 35: 902. 2010. |
| Old World-Africa | |
| Fl. Trop. Afr. [Oliver et al.] 4, 2: 220. 1906. | Morelloid | Black nightshade | |
| Tabl. Encycl. 2: 21. 1794. |
| Old World-Madagascar | |
| Prodr. [A. P. de Candolle] 13(1): 50. 1852. | Morelloid | Black nightshade | |
| Encycl. [J. Lamarck & al.] Suppl. 3: 775. 1814. |
| Old World-Africa | |
| Tabl. Encycl. 2: 23. 1794. |
| Old World-Africa | |
| Flora 27: 784. 1844. |
| Erythrotrichum | |
| Prodr. [A. P. de Candolle] 13(1): 304. 1852. |
| Old World-Africa | |
| Uganda Prot. (H.H.Johnston) 1: 326. 1902. | African non-spiny (ANS) | ||
| J. E. Afr. Nat. Hist. 99: 230. (2010) 2011. |
| Old World-Africa | |
| Fl. Madag., Fam. 176: 123. 1994. | African non-spiny (ANS) | ||
| Fl. Bras. (Martius) 10: 18, tab. 1, fig. 1-8. 1846. | Morelloid | Black nightshade | |
| Gard. Dict. ed. 8, no. 6. 1768. | Morelloid | Black nightshade | |
| Tent. Fl. Abyss. 2: 98. 1850. |
| Old World-Africa | |
| Notizbl. Bot. Gart. Berlin-Dahlem 12: 92. 1934. |
| Old World-Africa | |
| Pflanzenw. Ost-Afrikas C (Engler): 352. 1895. |
| Old World-Africa | |
| Pflanzenw. Ost-Afrikas C (Engler): 33. 1895. |
| Old World-Africa | |
| Tabl. Encycl. 2: 25. 1794. |
| Sisymbriifolium | |
| Revis. Gen. Pl. 3(3): 227. 1898. |
| Old World-Africa | |
| Sert. Somal. 47. 1882. |
| Old World-Africa | |
| Abh. Königl. Akad. Wiss. Berlin 1894: 63. 1894. |
| Old World-Africa | |
| Prodr. [A. P. de Candolle] 13(1): 289. 1852. |
| Old World-Africa | |
| Linnaea 43: 327. 1882. |
| Old World-Africa | |
| Repert. Spec. Nov. Regni Veg. 10: 547. 1912. | Morelloid | Black nightshade | |
| Fl. Aegypt.-Arab. 45. 1775. | African non-spiny (ANS) | ||
| Naturw. Reise Mossambique (Peters) 1: 237. 1861. |
| Old World-Africa | |
| Fl. Trop. Afr. [Oliver et al.] 4, 2: 217. 1906. |
| Old World-Africa | |
| Ann. Missouri Bot. Gard. 76: 351. 1989. |
| Old World-Madagascar | |
| Sp. Pl. 188. 1753. |
| Old World-Africa | |
| Kew Bull., 52(3): 706. 1997. |
| Old World-Africa | |
| Prodr. [O. P. Swartz] 47. 1788. |
| Torva | |
| Fl. Madag., Fam. 176: 130. 1994. | African non-spiny (ANS) | ||
| Gen. N. Amer. Pl. 1: 128. 1818. | Morelloid | ||
| Prodr. [A. P. de Candolle] 13(1): 36. 1852. |
| ||
| Bot. Jahrb. Syst. 54: 435. 1917. | African non-spiny (ANS) | ||
| Sp. Pl. 185. 1753. | Potato | Petota | |
| PhytoKeys 16: 67. 2012. | Morelloid | Black nightshade | |
| PhytoKeys 8: 4. 2012. |
| Old World-Africa | |
| Repert. Spec. Nov. Regni Veg. Beih. 16: 40. 1923. |
| Old World-Africa | |
| Pflanzenw. Ost-Afrikas C (Engler): 353. 1895. |
| Old World-Africa | |
| Hort. Kew. ed. 1, 1: 252. 1789. |
| Old World | |
| Prodr. [A. P. de Candolle] 13(1): 240. 1852. |
|
| |
| Gard. Dict. ed. 8, no. 2. 1768. | Morelloid | Black nightshade | |
| Nov. Pl. Descr. Dec. 56. 1798. |
| Old World-Tropical Asia | |
| Sp. Pl. 187. 1753. |
| Old World-Tropical Asia | |
| Bot. Mag. 113: tab. 6914. 1887. | Wendlandii-Allophyllum | ||
| Bull. Jard. Bot. État Bruxelles 17: 82. 1943. |
| Old World-Africa | |
| Fl. Hongk. 243. 1861. |
| Androceras-Crinitum | |
| Linnaea 43: 326. 1882. |
| Old World-Africa |
Country distribution of species in Africa (as defined here); introduced (incl. cultivated) species in brackets (epithet); taxa not included in the keys because they are known from a singleton cultivated specimen, are in italic type. All records based on specimens examined by the authors with verified identities. The status of is not completely clear, but it is most likely to be introduced from the New World, so is treated as that here; , on the other hand, appears to have a worldwide distribution, so is treated as native. The occurrence of in Senegal is doubtful, the specimen is very old and the label may be in error. Cultivated plants are often not collected, so the absence of records of commonly cultivated crops (e.g. , , ) should not be interpreted as lack of occurrence, merely as lack of collections. was recorded from Eygpt by Fawzi and Habeeb (2016) with a verifiable photograph; this Mexican species is widely cultivated and easily naturalised and is likely to be spreading around the Mediterranean.
| Country | Species |
|---|---|
| Algeria | |
| Angola | |
| Azores | ( |
| Benin | |
| Botswana | |
| Burkina Faso | |
| Burundi | |
| Cabo Verde | ( |
| Cameroon | |
| Canary Islands (Spain) | |
| Central African Republic (CAR) | |
| Chad | |
| Comoros (incl. Mayotte) | |
| Democratic Republic of the Congo | |
| Republic of the Congo | |
| Cote d’Ivoire | ( |
| Djibouti |
|
| Egypt (incl. Hala’ib triangle)* | |
| Equatorial Guinea | ( |
| Eritrea | |
| Ethiopia | ( |
| Gabon | |
| Gambia | |
| Ghana | ( |
| Guinea | ( |
| Guinea-Bissau | |
| Kenya | |
| Lesotho | ( |
| Liberia | ( |
| Libya | |
| Madagascar | |
| Madeira (Portugal) | ( |
| Malawi | |
| Mali | |
| Mauritania | |
| Mauritius (incl. La Réunion) | |
| Morocco | |
| Mozambique | |
| Namibia | |
| Niger | |
| Nigeria | |
| Rwanda | |
| São Tome e Principe | |
| Senegal | |
| Seychelles | |
| Sierra Leone | |
| Somalia | |
| South Africa | |
| South Sudan | |
| Sudan (incl. Hala’ib triangle)* | |
| Swaziland | |
| Tanzania | |
| Togo | ( |
| Tunisia | |
| Uganda | |
| Western Sahara |
|
| Zambia | ( |
| Zimbabwe |
*Possession of the area known as the Hala’ib triangle is disputed between Egypt and Sudan, species occurring there are listed under both coutries.
Figure 1.Heat map of diversity in Africa. Darker degree squares indicate greater species richness. The middle to high elevation regions of eastern Africa (Kenya/Tanzania) have the highest high species diversity, followed by secondary areas of species richness in the Ethiopian plateaus, dry areas of central Madagascar, South Africa and the area around Mount Cameroon. We have not analysed how collecting effort has influenced these patterns, but it is likely to be important. As the Clade has the largest number of species in Africa, diversity in that clade drives species richness overall (see Vorontsova and Knapp 2016, figure 2). Map prepared by Sarah Ficinski.
Figure 2.A, B Hook.f. (Allophyllum-Wendlandii Clade) C Bitter (Morelloid Clade) D Mill. (Morelloid Clade) E Lam. ( Clade) F Dunal ( Clade) G Bitter ( Clade) H Bitter & Dammer ( Clade). Photos A, B, F, G H by M.S. Vorontosova C, D, E by S. Knapp.
Figure 3.A Forssk. (African non-spiny Clade) B Dunal (African non-spiny Clade) C Scop. (Brevantherum Clade) D Spreng. (Dulcamaroid Clade) E Dunal ( Clade) F L. ( Clade) G L. (Potato Clade) H Aiton (Archaeosolanum Clade). Photos A, C, D, E, F, G, H by S. Knapp B by M.S. Vorontsova.
| 1a | Trichomes of stems and leaves stellate or echinoid |
|
| 1b | Trichomes of stems and leaves simple (unbranched) or dendritically branched, never stellate or echinoid |
|
| 2a | Anthers ellipsoid in outline; inflorescences many times branched; branching dichasial; stems without prickles |
|
| 2b | Anthers tapering in outline; inflorescences branched or unbranched; branching monochasial; stems with or without prickles |
|
| 3a | Shrubs, small trees or woody vines |
|
| 3b | Herbs or if plants woody, only at the base; never true vines, occasionally scandent |
|
| 4a | Stems with hooked prickles; anthers tapering |
|
| 4b | Stems without prickles; anthers ellipsoid |
|
| 5a | Small trees with foetid cordate leaves; flowers waxy pink or greenish white; anther connectives enlarged; fruit a large turbinate berry with fleshy pulp; cultivated tree tomato |
|
| 5b | Shrubs or woody vines; leaves, flowers and anther connectives not as above |
|
| 6a | Small shrubs with paired (geminate) leaves; flowers nodding, white; fruit held on erect pedicels, orange; cultivated or occasionally naturalised in northern Africa |
|
| 6b | Woody vines or scandent shrubs; leaves, flowers and fruit not as above; continental Africa, Madagascar; or cultivated |
|
| 7a | Leaves lanceolate, with at least some branched trichomes on new growth; leaves of a pair more or less the same shape; inflorescence usually with 2 flowers; berry dark orange; cultivated “Jerusalem cherry” |
|
| 7b | Leaves ovate or elliptic, completely glabrous; leaves of a pair markedly different in shape; inflorescence with more than 2 flowers; berry pale orange; only recorded from Egypt |
|
| 8a | Base of pedicel enclosed in a small sleeve of rhachis tissue usually more than 0.5 mm long; Mediterranean northern Africa; Macaronesia; if in other parts of Africa, cultivated |
|
| 8b | Base of pedicel peg-like, sometimes enclosed in a small sleeve of rhachis tissue, if so the sleeve less than 0.5 mm long; Continental tropical Africa; Madagascar; native plants |
|
| 9a | Leaves pinnate or deeply pinnatifid |
|
| 9b | Leaves simple (at most the margins toothed) or at most ternate |
|
| 10a | Fleshy prostrate herbs; leaves pinnatifid, the leaflets not distinct; inflorescences unbranched |
|
| 10b | Spreading herbs or herbaceous scramblers, not fleshy; leaves pinnate with distinct leaflets; inflorescences branched or less often unbranched |
|
| 11a | Anthers dimorphic, of different sizes and two of the five with horn-like projections; Macaronesia and northern Africa |
|
| 11b | Anthers equal in size and shape, if unequal only slightly so; widespread or cultivated |
|
| 12a | Trichomes simple with a single long terminal cell (bayonet hairs); fruit a large greenish berry with purple stripes (more than 3 cm diameter), with abundant solid mesocarp; herbaceous vine |
|
| 12b | Trichomes simple or branched; fruit variously coloured (usually less than 1 cm in diameter), with juicy mesocarp; annual or short-lived perennial herbs |
|
| 1a | Leaves glabrous on both surfaces |
|
| 1b | Leaves with at least some pubescence on either surface (this sometimes sparse along veins and midrib) |
|
| 2a | Inflorescence few-flowered, unbranched (at most furcate in |
|
| 2b | Inflorescence many flowered, usually many times branched |
|
| 3a | Flowers appearing fasciculate and axillary; corolla usually somewhat campanulate; fruit orange; South Africa |
|
| 3b | Flowers not appearing fasciculate; corolla stellate, the petals spreading or reflexed; fruit colour green, black or not known, never orange; Madagascar |
|
| 4a | Leaves clustered on short shoots; calyx lobes deltate, not divided to base; dry forests |
|
| 4b | Leaves not clustered on short shoots; calyx lobes long triangular, divided to the base; wet forests |
|
| 5a | Flowers or fruits (or pedicel scars) in tightly packed groups on individual branches (these sometimes very short and the inflorescence appearing spicate) |
|
| 5b | Flowers spaced on the open inflorescence, often unevenly so |
|
| 6a | Leaves clustered on short shoots |
|
| 6b | Leaves spaced along the stem |
|
| 7a | Anthers opening by pores that elongate with age; mountains of continental Africa |
|
| 7b | Anthers opening by delineated pores that do not elongate with age; Madagascar |
|
| 8a | Leaves fleshy, thick and coriaceous, the venation not visible in dry specimens; fruit with thick pericarp (woody?) |
|
| 8b | Leaves membranous to coriaceous, not markedly thick and fleshy, the venation visible in dry specimens; fruit with thin pericarp, the seeds visible through the berry wall |
|
| 9a | Petioles with long, simple trichomes (these not extending to the lamina); seeds 4–8 per berry; inflorescence axis thin and delicate |
|
| 9b | Petioles glabrous or with minute dendritic trichomes; seeds 20–40 per berry; inflorescence axis robust | |
| 10a | Leaf trichomes simple (unbranched) |
|
| 10b | Leaf trichomes branched (dendritic to echinoid) |
|
| 11a | Inflorescence axis unbranched, the flowers closely spaced |
|
| 11b | Inflorescence axis branched, often many times so |
|
| 12a | Leaves clustered along stem; fruit orange; South Africa |
|
| 12b | Leaves spaced along shoots; fruit purple or black; Madagascar |
|
| 13a | Flowers or fruits (or pedicel scars) in tightly packed groups on individual branches (these sometimes very short and the inflorescence appearing spicate) |
|
| 13b | Flowers spaced on the open inflorescence, often unevenly so |
|
| 14a | Stems strongly quadrangular; at least some leaves with shallow lobes; plants of seashore and dune habitats |
|
| 14b | Stems terete; leaves not lobed; plants of forests and forest edges |
|
| 15a | Leaf pubescence very sparse, confined to the midrib or near the petiole; flowers not heterostylous; Madagascar |
|
| 15a | Leaf pubescence variable, not very sparse, along veins and lamina; flowers heterostylous; mountains of continental Africa |
|
| 16a | Abaxial leaf surfaces with tufts of trichomes in the vein axils (domatia) |
|
| 16b | Abaxial leaf surfaces with trichomes on lamina and/or along veins, not with prominent tufts in the vein axils (domatia) |
|
| 17a | Inflorescence many times branched, open and with many flowers (more than 20); calyx lobes broadly deltate; petioles to 4 cm long, thin and flexuous; Mayotte (Comoros) |
|
| 17b | Inflorescence furcate, more congested and with fewer flowers (fewer than 20); calyx lobes deltate; petioles to 2.5 cm long, thicker; Madagascar |
|
| 18a | Calyx lobes 0.8–2 mm long; inflorescences with 10–16 flowers |
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| 18b | Calyx lobes 4–6 mm long; inflorescences with 3–10 flowers |
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| 19a | Abaxial leaf surfaces evenly pubescent on veins and lamina |
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| 19b | Abaxial leaf surfaces pubescent only along the veins and midrib, the trichomes not extending to the lamina |
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| 20a | Anther pores lengthening to slits with age; flowers heterostylous; leaves evenly distributed along branches; mountains of continental Africa |
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| 20b | Anther pores not lengthening to slit with age; flowers not heterostylous; leaves usually at least somewhat clustered on short shoots; Madagascar |
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| 21a | Leaves densely pubescent with golden (when dry) loosely dendritic trichomes; flowers more than 2 cm in diameter; anthers 4–6 mm long; widespread in Madagascar |
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| 21b | Leaves sparsely pubescent with white (when dry) congested dendritic trichomes; flowers 2 cm in diameter or less; anthers 3.5–4 mm long; dry forests of southern Madagascar |
|
| 22a | Inflorescence unbranched, with few flowers; pedicels 1.8–4.5 cm long |
|
| 22b | Inflorescence many times branched, with many flowers; pedicels 0.8–1.2 cm long |
|
| 23a | Anther pores lengthening to slits with age; flowers heterostylous; pedicels with pubescence like the inflorescence rhachis; mountains of continental Africa |
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| 23b | Anther pores not lengthening to slit with age; flowers not heterostylous; pedicels always glabrous; Madagascar |
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| 1a | Leaves shallowly lobed, pubescent with long, tangled eglandular trichomes; anthers tapering, horned near the base, tightly connivent; seeds more than 5 mm long; fruit a dry berry; Mediterranean |
|
| 1b | Leaves simple or ternate, glabrous or pubescent, but the trichomes not long and tangled, glandular; anthers markedly horned, spreading; seeds less than or equal to 5 mm long; fruit a brightly coloured, juicy berry; laurisylva forest in Macaronesia |
|
| 2a | Leaves simple or ternate, the base truncate or cordate if leaves unlobed; anthers yellow, horned in lower third; berry bright red; Madeira |
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| 2b | Leaves simple, the base cordate; anthers black, horned about halfway up from the base; berry orange or red; Tenerife, Canary Islands |
|
| 1a | Buds turbinate and strongly pointed; petals strongly reflexed, with shiny green dots at the base of each; anthers tightly connivent with “glue”; fruit a shiny red berry, often ellipsoid; native plants in Mediterranean northern Africa |
|
| 1b | Buds rounded, often somewhat inflated; petals spreading, without shiny green dots; anthers not tightly connivent with “glue”; fruit red or black, globose; cultivated plants, occasionally naturalised throughout the region |
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| 2a | Flowers white; anthers on equal filaments; leaves with axillary tufts of trichomes on the lower surfaces (domatia), usually simple, occasionally pinnatifid; berry (very rarely) black |
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| 2b | Flowers purple; one filament slightly longer than the other 4; leaves completely glabrous, pinnatifid, rarely simple; berry bright shiny red |
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| 1a | Leaves shallowly to deeply pinnatifid |
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| 1b | Leaves entire to sinuate-dentate |
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| 2a | Glandular trichomes present (e.g. along stems, petioles and leaves), plants usually sticky to touch when fresh |
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| 2b | Glandular trichomes absent (e.g. along stems, petioles and leaves), plants not sticky to touch when fresh |
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| 3a | Anthers less than 1.8 mm long |
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| 3b | Anthers more than or equal to 1.8 mm long |
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| 4a | Inflorescences with 10–40 flowers; pedicels spaced 1–2 mm apart, sharply bent at the base (near articulation point) in flower and fruit |
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| 4b | Inflorescences with 2–5(-10) flowers; pedicels spaced 0–1 mm apart, nodding, erect or spreading in flower and fruit, reflexed and slightly curved in some species in fruit but never in flower |
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| 5a | Calyx lobes 1–1.5 mm long in flower; fruiting calyces not accrescent, the tube remaining 1–1.7 mm long and lobes 1–1.5 mm long; fruit black when ripe, not markedly shiny, with a glaucous cast |
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| 5b | Calyx lobes 1.5–2.5 mm long in flower; fruiting calyces accrescent, the tube 3–4 mm long and lobes 2.5–8.0 mm long; fruit green when ripe, shiny |
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| 6a | Leaf bases attenuate to cuneate; inflorescences mostly intermodal, with 4–8(-10) flowers; pedicels spaced 0.3–1 mm apart; calyx lobes 1.7–2.5 mm long; corollas with yellow-green central eye with black-purple V-shaped margins; anthers 1.0–1.4 mm long; berries dark green to green-brown marbled with white lines, becoming usually translucent and glossy, lower half of berries covered with enlarged calyces but berry mostly visible; seeds brown; stone cells (1-)2–3, these 0.5 mm in diameter; northern Africa |
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| 6b | Leaf bases truncate; inflorescences mostly leaf-opposed, with 2–5(-7) flowers; pedicels spaced 0(-1) mm apart; calyx lobes 1.5–2.0 mm long; corolla with yellow-green or translucent basal star without black-purple colouration; anthers 1.2–2.0 mm long; berries pale green, shiny becoming dull, opaque, usually completely enveloped by enlarged calyces; seeds pale yellow; stone cells 4–6, these (0.5-)0.8–1 mm in diameter; only known from South Africa |
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| 7a | Anthers more than or equal to 2.8 mm long |
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| 7b | Anthers less than 2.8 mm long |
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| 8a | Inflorescences with bracteoles present in most individuals; buds narrowly ellipsoid; corolla deeply stellate, the lobes narrowly lanceolate; berries with more than 30 stone cells |
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| 8b | Inflorescences never with bracteoles; buds globose, ovoid or narrowly ellipsoid; corolla rotate-stellate, the lobes long- triangular with rounded tips; berries with (0-)2–4 stone cells |
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| 9a | Calyx lobes appressed to spreading in fruit, never strongly reflexed |
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| 9b | Calyx lobes strongly reflexed in fruit |
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| 10a | Calyx accrescent in fruit, calyx tube 3–4 mm long and lobes 2.5–8 mm long |
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| 10b | Calyx not accrescent in fruit, calyx tube 1–2 mm long and lobes 1–1.5 mm long |
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| 11a | Buds ellipsoid; calyx tube 1.5–2.0 mm long, lobes 1–1.5 mm long, elongate-deltate with rounded tips; fruiting pedicels persist when fruits mature and fall off; Cameroon line (Cameroon and Equatorial Guinea), above 2,000 m elevation |
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| 11b | Buds subglobose; calyx tube 0.8–1.0 mm long, lobes 0.5–0.8 mm long, triangular with rounded to acute tips; fruiting pedicels generally do not persist and fall off with maturing fruits; in continental Africa only in South Africa and around the Mediterranean |
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| 12a | Leaves rhomboidal to lanceolate; filaments 1.2–1.5 mm long, anthers 1.3–1.8(-2) mm long; seeds 1.6–1.8 mm long and 1.3–1.5 mm wide |
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| 12b | Leaves broadly to narrowly ovate to elliptic; filaments 0.5–1.3 mm long; anthers 1.8–2.5 mm long; seeds 1.8–2.2 mm long and 1.5–1.7 mm wide |
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| 13a | Calyx with broad and relatively transparent sinuses, lobes elliptic to triangular, rounded at tip; free part of the filaments 1.0–1.3 mm long; mature berries slightly ellipsoid, shiny yellow, orange or red; stone cells always absent |
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| 13b | Calyx with narrow, sharp triangular sinuses, lobes deltate with acute or rounded tips; free part of the filaments 0.5–0.7 mm long; mature berries round, dull black or green; stone cells 0–4; in Africa only in South Africa and around the Mediterranean |
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| 14a | Anthers less than 1.8 mm long |
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| 14b | Anthers more than or equal to 1.8 mm long |
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| 15a | Pedicels spaced 1–2 mm apart, pedicels sharply bent at the base (near the articulation point) in flower and fruit |
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| 15b | Pedicels spaced 0–0.5 mm apart, pedicels nodding, erect or spreading in flower and fruit |
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| 16a | Leaves with entire margins, occasionally sinuate-dentate; calyx lobes 0.3–0.5 mm long in flower, 1(-2) mm in fruit; mature fruits black, the surface very shiny |
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| 16b | Leaves shallowly toothed, occasionally entire; calyx lobes 1.0–1.5 mm long in flower, 1.5–2 mm in fruit; mature fruits purple-black or green, the surface dull |
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| 17a | Anthers less than 2.8 mm long |
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| 17b | Anthers more than or equal to 2.8 mm long |
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| 18a | Berries without stone cells |
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| 18b | Berries with 2–22 stone cells |
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| 19a | Pedicels persisting and not dropping with mature fruits; calyx lobes in fruit mostly strongly reflexed |
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| 19b | Pedicels dropping with mature fruits; calyx lobes in fruit appressed to slightly spreading, rarely strongly reflexed |
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| 20a | Leaves rhomboidal to lanceolate; filaments 1.2–1.5 mm long, anthers 1.3–1.8(-2) mm long; seeds 1.6–1.8 mm long and 1.3–1.5 mm wide |
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| 20b | Leaves broadly to narrowly ovate to elliptic; filaments 0.5–1.3 mm long; anthers 1.8–2.5 mm long; seeds 1.8–2.2 mm long and 1.5–1.7 mm wide |
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| 21a | Calyx with broad and relatively transparent sinuses, lobes elliptic to triangular, rounded at tip; filaments 1.0–1.3 mm long; mature berries slightly ellipsoid, shiny yellow, orange or red; stone cells always absent |
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| 21b | Calyx with narrow, sharp triangular sinuses, lobes deltate with acute tips; filaments 0.5–0.7 mm long; mature berries round, dull black or green; stone cells generally absent (2–4 stone cells common in Asian material) |
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| 22a | Buds elongate-oblong; fruiting peduncles strongly deflexed at the base (bent downwards at junction with the stem) |
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| 22b | Buds ellipsoid to subglobose; fruiting peduncles straight or ascending |
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| 23a | Pedicels spaced 1–2 mm apart, sharply bent at the base (near the articulation point) in flower and fruit; seeds 1.5–2 mm long and 1–1.5 mm wide |
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| 23b | Pedicels spaced 0–0.7 mm apart, straight, spreading or reflexed in flower and fruit; seeds 1.8–2 mm long and 1.5–1.6 mm wide |
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| 24a | Prostrate herb; leaves narrowly elliptic to lanceolate, base strongly attenuate; inflorescences with 1–5 flowers; pedicels stout and spreading; calyx lobes linear-oblong with rounded apices; mountains of Ethiopia |
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| 24b | Upright or spreading herb; leaves broadly ovate to elliptic, base acuminate, acute, obtuse, truncate to abruptly attenuate; inflorescences with 2–40 flowers; pedicels thinner, spreading to strongly reflexed; calyx lobes triangular, broadly deltoid or ovate with acute to rounded apices |
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| 25a | Pedicels strongly bent downwards at the base (near articulation point) in flower and fruit |
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| 25b | Pedicels spreading, stout or pendent in flower, occasionally recurved in fruit but never strongly bent downwards at the base |
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| 26a | Inflorescences unbranched or more often branched, often with small leaves (bracteoles?); calyx lobes broadly deltate to mere enations of the rim; style exserted 1.0–1.5 mm beyond anther cone; mature berries 3–4(-5) mm in diameter, dull yellowish brown |
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| 26b | Inflorescences unbranched, never with small leaves; calyx lobes triangular; style exserted 0–1 mm beyond anther cone; mature berries 6–10 mm in diameter, dull black |
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| 27a | Inflorescences with bracteoles present in most individuals; buds narrowly elliptic; berries with more than 30 stone cells |
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| 27b | Inflorescences never with bracteoles; buds globose, ovoid or narrowly ellipsoid; berries with 0–14 stone cells |
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| 28a | Berries with 2–14 stone cells; leaf base strongly attenuate |
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| 28b | Berries without stone cells; leaf base not strongly attenuate |
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| 29a | Buds elongate-oblong; calyx lobes broadly deltate to triangular with acute tips; fruiting peduncles strongly bent at the base near junction with the stem; fruiting pedicels thin, reflexed and slightly recurved; seeds 1.2–1.4 mm long and 1.0–1.2 mm wide |
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| 29b | Buds globose-subglobose; calyx lobes broadly deltate with rounded tips; fruiting peduncles straight; fruiting pedicels stout, erect and spreading; seeds 2–2.8 mm long and 1.5–1.8 mm wide |
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| 1a | Flowers yellow; anthers tightly connivent and tapering to a beak-like tip; fruit a bright red berry; cultivated tomatoes |
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| 1b | Flowers white or purple; anthers ellipsoid, not tapering to a beak-like tip; fruit green or whitish green (often with purple stripes) |
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| 2a | Corolla lobes deltate to triangular; anther cone stout, the style included; berry usually more than 1 cm in diameter (often much larger), fewer than 20 per infructescence; leaflets with serrate margins; cultivated tomato |
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| 2b | Corolla lobes narrowly triangular; anther cone long and narrow, the style exserted; berry less than 1 cm in diameter, more than 20 per infructescence; leaflets with entire margins; cultivated currant tomato |
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| 3a | Leaves at most ternate, usually simple; fruit a berry more than 3 cm in diameter; plant not bearing underground tubers; cultivated pepino |
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| 3b | Leaves pinnate; fruit a berry less than 3 cm in diameter; plant bearing underground tubers; cultivated potato |
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| 1a | Young flower buds turbinate; calyx densely pubescent within over entire surface; young stems sulcate; axillary leaves absent |
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| 1b | Young flower buds oblong to orbicular; calyx lobes pubescent within only in distal quarter; young stems terete; axillary leaves common |
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| 1a | Young stems and petioles noticeably winged; mature fruit densely pubescent; invasive plant in Tanzanian highlands and South Africa |
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| 1b | Young stems and petioles not markedly winged (terete or slightly ridged); mature fruit glabrous; native or invasive, widespread |
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| 2a | Flowers with stamens of differing lengths (due either to unequal anthers or unequal filaments or both); arid eastern and north-eastern Africa |
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| 2b | Flowers with all stamens equal in length; widespread |
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| 3a | Corolla strongly zygomorphic, with the two lower lobes much larger; flowers often enantiostylous; Canary Islands |
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| 3b | Corolla only weakly zygomorphic, the lower lobes somewhat but not markedly larger; flowers not enantiostylous; continental Africa |
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| 4a | Leaves narrowly elliptic to lanceolate; calyx lobes linear and awn-like; corolla always 5-merous; ripe berry orange; Gran Canaria |
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| 4b | Leaves ovate; calyx lobes linear; corolla often 4-merous; ripe berry yellow or yellowish green; Tenerife and Gran Canaria |
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| 5a | Leaves orbicular to reniform, 1.2–2.5 cm long, wider than long; petioles longer than leaves; rare in north-eastern Somalia |
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| 5b | Leaves ovate to elliptic or lanceolate, 2–14 cm long, longer than wide; petioles shorter than leaves; arid eastern and north-eastern Africa |
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| 6a | Stem prickles dense, acicular, less than 0.5 mm wide at base, pale yellow; fruit fully concealed by the accrescent calyx |
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| 6b | Stem prickles absent or sparse, if present wider than 1 mm at base, yellow to orange or brown; fruit at least partly exposed |
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| 7a | Leaves subentire to lobed; anthers of equal length; seeds very dark brown to almost black |
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| 7b | Leaves entire; one anther much longer than the others; seeds dull yellow to orange-brown |
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| 8a | Flower one per inflorescence, peduncle and rhachis absent; corolla pentagonal, lobed for 1/4–1/3 of the way to the base, 0.9–1.3 cm in diameter; southern Africa |
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| 8b | Flower usually more than one per inflorescence, peduncle and/or rhachis present in at least some inflorescences; corolla usually stellate, lobed for more than 1/3 of the way to the base or, if lobed, for 1/4–1/3 of the way to the base then corolla of long-styled flowers broader than 1.3 cm in diameter; widespread |
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| 9a | Trichomes on young stems and adaxial (upper) surfaces of the leaves simple only, never stellate |
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| 9b | Trichomes on young stems and adaxial (upper) surfaces of the leaves stellate |
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| 10a | Flowers pale bluish purple; anthers 8–12.5 mm; fruit globose or extended into a “nipple” |
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| 10b | Flowers white or greenish white; anthers 5–7.5 mm; fruit globose |
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| 11a | No stellate hairs anywhere on the plant; fruit bright orange at maturity; seeds winged, 4–6 mm long |
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| 11b | Stellate hairs almost always present on the abaxial (lower) surface of the leaves; fruit yellow at maturity; seeds not winged, 1.8–2.8 mm long |
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| 12a | Leaf lobes 2–3 pairs, extending 1/3–1/2 distance to the midvein; calyx lobes 5–6.5 mm long, often caudate |
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| 12b | Leaf lobes 3–5 pairs, extending less than 1/3 of the distance to the midvein; calyx lobes 0.8–2 mm long, acute |
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| 13a | Leaves entire, 3–10 times longer than wide; shrubs erect, 1–6 m tall; stem trichomes with partly fused rays; southern Madagascar |
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| 13b | Leaves entire or lobed, 1–3(8) times longer than wide; shrubs erect, scandent or climbing, 0.2–6 m tall; if leaves entire and more than 3 times longer than wide, then shrub less than 1 m tall and not in southern Madagascar; stem trichomes with free rays |
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| 14a | Leaves 9–13(20) cm long; corolla 2–3.1 cm in diameter; juvenile branches with dark red prickles; south-eastern Madagascar |
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| 14b | Leaves 1.5–7 cm long; corolla 1–2 cm in diameter; juvenile branches with grey-brown or red-brown prickles; south-western Madagascar |
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| 15a | Leaf blades 3–7 cm long, concolorous to weakly discolorous, yellow-green |
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| 15b | Leaf blades 1.5–3(4) cm long, strongly discolorous, green-brown adaxially and glaucous abaxially |
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| 16a | Prickles and leaf venation noticeably dark orange to red, contrasting with the yellow-green to red-green leaf surface; southern Madagascar |
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| 16b | Prickles and leaf venation not a contrasting colour, yellow to green or red-brown; widespread or naturalised |
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| 17a | Mature fruit green, never developing to bright yellow, orange or red; plants weakly andromonoecious; fruits 1–1.5 cm in diameter |
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| 17b | Mature fruit yellow, orange or red; plants hermaphroditic or andromonoecious, fruits 0.5–6 cm in diameter, if andromonoecious, then fruits more than 1.5 cm in diameter |
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| 18a | Shrub to tree 1.5–9 m tall; young stems and leaves densely ferruginous pubescent; trichomes on the inflorescences and pedicels not glandular; prickles straight to slightly curved |
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| 18b | Shrub to 3 m; young stems and leaves pubescent green to brownish; trichomes on the inflorescences and pedicels glandular; prickles curved |
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| 19a | Mature fruit yellow or greenish yellow, 1.5–5(6) cm in diameter; corolla on long-styled flowers (1.3)2–6 cm in diameter; plants mostly andromonoecious |
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| 19b | Mature fruit orange to red, 0.5–1.2(1.7) cm in diameter; if mature fruit orange (1)1.5–2.5(5) cm in diameter, the plant cultivated; corolla on long-styled flowers 0.8–3 cm in diameter; plants mostly hermaphroditic |
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| 20a | Cultivated tree 5–10 m tall with copious flowers; corolla bright purple aging to white with both colours usually present in an inflorescence, 5.5–8 cm in diameter |
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| 20b | Wild plants or cultivated vegetables, less than 6 m tall; corolla white to mauve or purple with the colour constant within each individual, 1.6–6 cm in diameter |
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| 21a | Petiole usually decurrent, leaf bases attenuate (cuneate); trichomes on abaxial leaf surface (if present) stalked with 4(5) rays |
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| 21b | Petiole never decurrent, leaf bases cordate to cuneate; trichomes on abaxial leaf surface (if present) sessile or stalked with 6–16 rays |
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| 22a | Plant clearly stellate-pubescent and armed, drying yellow-green to red-brown; wild plant |
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| 22b | Plant usually glabrous and unarmed, drying a distinctive red-brown colour; cultivated plant |
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| 23a | Climbers or scandent plants |
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| 23b | Plants erect or rarely semi-scandent |
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| 24a | Prickles on young stems straight |
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| 24b | Prickles on young stems strongly curved |
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| 25a | Prickles pale straw-yellow, to 20 mm long; petiole with sessile stellate trichomes; corolla ca. 2.5 cm in diameter; style strongly curved; Mediterranean northern Africa (adventive from Asia) |
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| 25b | Prickles yellow (but not straw-coloured) or brown; petiole trichomes usually stalked; corolla 3.5–5 cm in diameter; style straight; southern Kenya |
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| 26a | Corolla white, 1.3–1.6 cm in diameter; seeds 5.5–6 mm long; Kenyan mountains |
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| 26b | Corolla mauve to purple, 3.5–6 cm in diameter; seeds 3–4 mm long; eastern and southern Africa, Madagascar |
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| 27a | Calyx inflated, fully covering the berry at maturity; young stems densely covered with straight prickles; northern Madagascar |
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| 27b | Calyx not inflated, the berry exposed at maturity; young stems prickly or unarmed; widespread |
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| 28a | Corolla lobed for more than half way to the base; shrubs or trees up to 6 m; variety of habitats |
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| 28b | Corolla lobed for half way to the base or less; small shrubs up to 2 m; weeds of open arid environments or cultivated crops; relatives of the eggplant |
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| 29a | Leaves 8–18 cm long, strongly discolorous; young fruits plain green; seeds 3.5–4.5 mm long; wet upland habitats |
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| 29a | Leaves 2–8 cm long, concolorous or sometimes discolorous; young fruits striped in different shades of green; seeds 2.2–3.5 mm long; arid upland or lowland habitats |
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| 30a | Fruit globose, never apiculate, 1.4–1.7 cm in diameter, 4–10 per infructescence; young stems with yellow (when dry) long-stalked trichomes, the stalks 1–3 mm |
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| 30b | Fruit turbinate or cone-shaped, sometimes globose, usually somewhat apiculate, 2.8–5 x 1.8–4.5 cm, 1–3(5) per infructescence; young stems usually lacking long-stalked yellow (when dry) trichomes |
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| 31a | Fruit distinctly turbinate or cone-shaped, 2.8–3.7 x 1.8–2.2 cm; leaves on fertile branches elliptic and subentire, 6–8 x 2.5–4 cm, ca. 2.5 times longer than wide; 2100–3000 m elevation |
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| 31b | Fruit globose, usually somewhat apiculate, 3–5 x 2–4.5 cm; leaves on fertile branches ovate(elliptic) and lobed(subentire), 8–15 x 6–12 cm, 1.5–2 times longer than wide; 1200–2100(3200) m elevation |
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| 32a | Prickles straight or occasionally curved; petioles 1/3–2/3 as long as the leaf blades |
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| 32b | Prickles on young stems strongly curved; petioles less than 1/3 of the leaf blade length |
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| 33a | Leaves entire, densely stellate-pubescent on both sides; eastern and north-eastern African highlands |
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| 33b | Leaves lobed, adaxially glabrescent; eastern and north-eastern Africa |
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| 34a | Leaves 2–4 cm long; curved prickles on young stems 5–10 mm long; eastern and north-eastern Africa |
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| 34b | Leaves 6–8 cm long; curved prickles on young stems 1–3 mm long; coastal Kenya, rare |
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| 35a | Fruit with soft pericarp, in a variety of shapes and colours, edible; common fasciation in the flowers (e.g. increase in the number of flower parts up to 8), inflated ovaries; cultivated species |
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| 35a | Fruit spherical, yellow, with comparatively hard pericarp, not edible; flowers 5-merous; wild plants |
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| 36a | Leaves lobed with primary and secondary lobes, the primary lobes extending 2/3–3/4 of the distance to the midvein and secondary lobes always present; southern Africa and northern African coasts around the Mediterranean |
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| 36b | Leaves entire or lobed, lobes extending up to 2/3 of the distance to the midvein, secondary lobes usually not present; widespread |
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| 37a | Leaf margins and venation nearly white and contrasting with greenish red-brown adaxial leaf surface; trichomes on the abaxial surface of the leaves with 10–17 rays; Ethiopian highlands |
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| 37b | Leaf margins and venation the same colour as the rest of the leaf blade; trichomes on the abaxial surface of the leaves with 5–12(15) rays; widespread |
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| 38a | Leaf lobes apically obtuse to acute, sometimes rounded, sometimes with secondary lobes; lobes 1/4–2/3 of the distance to the midvein; leaves and young stems glabrescent to moderately pubescent |
|
| 38b | Leaf lobes apically rounded, sometimes obtuse, never with secondary lobes; lobes up to 1/3(1/2) of the distance to the midvein; leaves and young stems usually densely pubescent |
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| 39a | Calyx lobes on long-styled flowers 7–10 mm long, ovate and foliaceous, apically obtuse; South Africa |
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| 39b | Calyx lobes on long-styled flowers 4–7 mm long, deltate or long-triangular apically acuminate; northern Africa and Cape Verde Islands |
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| 40a | Calyx lobes on long-styled flowers 4–7 mm long, deltate, ca. 1/6 as long as the fruit at maturity; continental Africa north of the equator |
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| 40b | Calyx lobes on long-styled flowers 6–7 mm long, long triangular, 1/2 to 1/3 as long as the fruit at maturity; Cape Verde Islands (Senegal?) |
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| 41a | Prickles straight; corolla on long-styled flowers 1.8–2.5 cm in diameter; anthers ca. 4.5 mm long; Madagascar; Mauritius, Réunion |
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| 41b | Prickles curved or straight; corolla on long-styled flowers 2.5–4.5 cm in diameter; anthers 5–9 mm long; widespread |
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| 42a | Leaves usually entire, sometimes lobed; trichomes on the abaxial leaf surface sessile or with stalks up to 0.1 mm long; fruits 1.5–3 cm diameter |
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| 42b | Leaves lobed; trichomes on the abaxial leaf surface with stalks up to 0.5(1) mm long; fruits 2.5–4.5 cm diameter |
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| 43a | Leaves velvety red-brown adaxially; calyx lobes on long-styled flowers ovate to oblong, foliaceous, 7–10 mm long |
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| 43b | Leaves yellow-green to green-brown adaxially; calyx lobes on long-styled flowers deltate, usually not foliaceous, 2.5–6 mm long |
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| 44a | Leaves concolorous to weakly discolorous, indumentum yellowish; leaves ca. 1.5 times longer than wide; young stems terete to angular; north-eastern Africa |
|
| 44b | Leaves strongly discolorous, indumentum dirty greenish brown adaxially and whitish yellow abaxially; leaves 1.5–2.5 times longer than wide; young stems with somewhat raised longitudinal ridges; southern Africa |
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| 45a | Leaves on fertile branches with distinct lobes, at least some of the lobes longer than 1/4 of the distance from the midvein to the leaf edge |
|
| 45b | Leaves on fertile branches entire or subentire or with some shallow lobes no longer than 1/4 of the distance from the midvein to the leaf edge |
|
| 46a | Leaves more than 3 times longer than wide |
|
| 46b | Leaves 1–3 times longer than wide |
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| 47a | Plant unarmed or prickles straight, orange to red; trichomes on the abaxial surfaces of the leaves with (9)12–14 rays |
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| 47b | Plant densely armed with curved broad-based pale-yellow prickles; trichomes on the abaxial surfaces of the leaves, if present, with 0–8 rays |
|
| 48a | Anthers 2.5–3.2 mm long; trichomes anywhere on the plant with 0–4 rays; South Africa |
|
| 48b | Anthers 5–6.5 mm long; trichomes anywhere on the plant with 5–8 rays; north-eastern Africa |
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| 49a | Leaves with at least some secondary lobing present |
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| 49b | Leaves lobed once only |
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| 50a | Flowers 3–7 per inflorescence, mauve to purple; anthers 5.5–7 mm long; South Africa |
|
| 50b | Flowers 6–50 per inflorescence, white; anthers 9–10 mm long; invasive in South Africa and Kenya |
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| 51a | No prickles visible anywhere on the plant |
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| 51b | At least some prickles visible on the plant |
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| 52a | Plant scandent or scrambling, sometimes erect; anthers (4)5–6.5 mm long; inland eastern Africa |
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| 52b | Plant erect; anthers 2.3–5(8.5) mm long; widespread |
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| 53a | Corolla 1.5–3 cm in diameter; mauve or purple; southern Africa, Indian Ocean islands |
|
| 53b | Corolla 0.9–1.5 cm in diameter; white (only occasionally pale violet); widespread on continent or cultivated |
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| 54a | Fruiting pedicels deflexed, curved; anthers 3.5–5.2 mm long; South Africa and Namibia |
|
| 54b | Fruiting pedicels straight and spreading; anthers 4.5–8.5 mm long; Mauritius and Réunion |
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| 55a | Inflorescences 2.5–6 cm long, with 5–22 flowers |
|
| 55b | Inflorescences 1–2.5 cm long, with 1–4(10) flowers |
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| 56a | Leaf blades 5–18 cm long, petioles 1–4 cm long; plant cultivated for leaves or fruits, widespread |
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| 56b | Leaf blades 3–8 cm long, petioles 0.5–1.5 cm long; wild plant in southern Africa |
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| 57a | Prickles on young stems predominantly curved |
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| 57b | Prickles on young stems predominantly straight, sometimes straight and reflexed |
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| 58a | Young prickles and indumentum red; scandent shrubs; Madagascar |
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| 58b | Young prickles and indumentum yellowish, green or brown; erect or scandent shrubs; widespread (mainland Africa, rarely Madagascar, Indian Ocean Islands) |
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| 59a | Young stems with a dense covering of long-stalked trichomes and prickles of different lengths; 1000–3000 m elevation |
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| 59b | Young stems with a sparse or dense covering of sessile or short-stalked trichomes of uniform lengths; 0–1000 m elevation |
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| 60a | Stem trichomes with stalks 0.1–0.4 mm long; anthers 6–10 mm long; coastal eastern Africa |
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| 60b | Stem trichomes sessile or the stalks to 0.1 mm long; anthers 3–8.5 mm long; widespread |
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| 61a | Plants of the Indian Ocean islands (Mauritius, Réunion, Aldabra) |
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| 61b | Plants of continental Africa |
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| 62a | Fruiting pedicels straight and spreading; corolla to 3 cm in diameter; petiole not winged from decurrent leaf base; Mauritius and Réunion |
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| 62b | Fruiting pedicels curved and deflexed; corolla to 2 cm in diameter; petiole slightly winged from decurrent leaf base; Aldabra (Seychelles) |
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| 63a | Calyx in flower 7–9 mm long, apically caudate; calyx in fruit 10–12 mm long; trichomes with midpoints much longer that the rays; Tanzanian highlands, rare |
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| 63b | Calyx in flower 2–6 mm long, apically acute to acuminate or rarely caudate; calyx in fruit less than 10 mm long; trichomes with the midpoints short than or equal to the rays; widespread |
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| 64a | Plant always erect; leaves discolorous; corolla 0.8–1.5 cm in diameter, lobed for 1/2–2/3(3/4) of the way to the base |
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| 64b | Plant often scandent or semi-scandent; leaves usually concolorous; corolla (1)1.5–2.5 cm in diameter, lobed for 2/3 of the way to the base or more |
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| 65b | Inflorescences 2.5–6 cm long, with 5–22 flowers; polymorphic weed across highland areas of Africa and Madagascar |
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| 65a | Inflorescences 1–2.5 cm long, with 1–4 flowers; southern Africa |
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| 66a | Prickles 3–5 mm long; leaf blades 1.5–2.5(9) cm long, leaf base narrow-cuneate to attenuate; South Africa and Namibia |
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| 66b | Prickles 1–4(7) mm long. Leaf blades (1)2.5–14 cm long, leaf base cuneate to truncate; widespread |
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| 67a | Calyx 2–4 mm long in flower |
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| 67b | Calyx 4–6 mm long in flower |
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| 68a | Prickles on young stems 1.5–2(3) mm long; leaves 2.5–5.5 cm long; seeds 2.5–3.5 mm long; inland eastern Africa |
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| 68b | Prickles on young stems 2–4 mm long. Leaves 3–14 cm long; seeds 1.8–2.5 mm long; coastal eastern Africa |
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| 69a | Leaves drying yellow-green; petiole 1/4–1/3 of the leaf length; eastern and north-eastern Africa |
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| 69b | Leaves drying dark red-brown; petiole 1/3–1/2 of the leaf length; southern Africa |
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| 70a | Inflorescence with more than 10 flowers |
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| 70b | Inflorescence with 10 or fewer flowers |
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| 71a | Leaves 1–4(6) cm long; corolla 1.3–2.4 cm in diameter; seeds almost black; north of the equator |
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| 71b | Leaves (5)7–25 cm long; corolla 0.7–1.5 cm in diameter; seeds yellow to orange-brown; widespread |
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| 72a | Inflorescences 1–2.5 cm long, peduncle 0–2 mm long; trichomes on the young stems with (5)8–20 rays; western Africa |
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| 72b | Inflorescences 2.5–8 cm long, peduncle 2–30 mm long; trichomes on the young stems with 6–8 rays; widespread |
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| 73a | Inflorescences branched many times (more than once); Kenya and Tanzania |
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| 73b | Inflorescences unbranched or forked; widespread |
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| 74a | Young prickles almost filiform and easily bendable like bristles, with no firm prickles anywhere on the plant; eastern African savannah |
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| 74b | Young prickles sturdy when touched; widespread |
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| 75a | Calyx lobes oblong, foliaceous, apically obtuse; lowland eastern Africa, rare |
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| 75b | Calyx lobes deltate, sometimes long-ovate or narrow-oblong, not clearly foliaceous, apically acute to acuminate; widespread |
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| 76a | Calyx 7–12 mm long in flower |
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| 76b | Calyx 2–7 mm long in flower |
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| 77a | Prickles on young branches 8–16 mm long; leaves elliptic; Ethiopian highlands |
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| 77b | Prickles on young branches 2–7 mm long; leaves ovate; Tanzania, South Africa and Madagascar |
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| 78a | Corolla lobed for 3/4 of the way to the base or more; trichomes on the young stems with midpoints (0.5)1.5–2.5(3) mm long; Tanzanian highlands |
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| 78b | Corolla lobed for 1/2–2/3 of the way to the base; trichomes on the young stems with midpoints 0.15–5 mm long; South Africa and Madagascar |
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| 79a | Young stems with a dense covering of long-stalked trichomes and prickles of different lengths; Madagascar |
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| 79b | Young stems with no bristles, only prickles and trichomes of uniform length; South Africa |
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| 80a | Young prickles and indumentum red; pedicels (1)1.5–4 cm long; Madagascar |
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| 80b | Young prickles and indumentum pale yellow to brown or purple, never red; pedicels 0.2–1(1.4) cm long; mainland Africa, rarely in Madagascar |
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| 81a | Young stems with a dense covering of long-stalked trichomes and prickles of different lengths; 1000–3000 m elevation |
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| 81b | Young stems with a sparse or dense covering of sessile or short-stalked trichomes of uniform lengths; 0–1000 m elevation |
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| 82a | Corolla lobed for 3/4 of the way to the base or deeper; plants scrambling or climbing, sometimes erect |
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| 82a | Corolla lobed for 1/2–2/3 of the way to the base; plants erect, sometimes scandent |
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| 83a | Prickles on young stems 2–4 mm long; eastern African coastal areas |
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| 83b | Prickles on young stems (3)4–13 mm long; arid environments across continental Africa |
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| 84a | Prickles reflexed; seeds nearly black |
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| 84b | Prickles perpendicular to the stem; seeds pale yellow to orange-brown |
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| 85a | Leaves 1–2 times longer than wide; leaf lobes to 2/3 of the distance to the midvein; leaves concolorous to strongly discolorous; north-eastern Africa |
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| 85b | Leaves 1.5–3 times longer than wide; leaf lobes to 1/2 of the distance to the midvein; leaves concolorous; South Africa |
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| 86a | Petiole 1/3–2/3 as long as the leaf blade; South Africa |
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| 86b | Petiole 1/6–1/3 as long as the leaf blade; widespread |
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| 87a | Leaf blades (5)11–25 cm long; inflorescence with 5–22 flowers; common variable highland weed widespread across Africa |
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| 87b | Leaf blades 1.7–9 cm long; inflorescence with 1–6 flowers; southern Africa |
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| 88a | Leaves elliptic; corolla 1.5–2.2 cm in diameter, mauve to purple |
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| 88b | Leaves ovate, sometimes oblong; corolla 0.9–1.3 cm in diameter, usually white |
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| 89a | At least some inflorescences branched several to many times; inflorescence with (10)20–150 flowers; erect shrubs to small trees |
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| 89b | Inflorescences unbranched or forked; inflorescence with 1–10(20) flowers; scrambling, scandent or erect shrubs |
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| 90a | Plants with at least some visible prickles or bristles |
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| 90b | Plants with no prickles or bristles visible |
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| 91a | Young stems with soft or thin bristles |
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| 91b | Young stems with firm deltate prickles |
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| 92a | Stem bristles soft and white, 0.4–0.7 mm wide at base; leaves ca. 2 times longer than wide; 1500 -1800 m elevation; Tanzania |
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| 92b | Stem bristles erect purple-black or brown, 0.2–0.4 mm wide at base; leaves 2.5–3.5 times longer than wide; 1800–2600 m elevation; eastern Africa |
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| 93a | Leaf blades 12–40 cm long, whitish grey underneath with trichomes, falling as white powder when touched; inflorescences with 30–150 flowers; anthers 2.5–3 mm long |
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| 93b | Leaf blades 4–10(16) cm long, yellow-green, brown or rarely whitish grey underneath, trichomes not falling when touched; inflorescences with 10–30 flowers; anthers 3.5–6 mm long |
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| 94a | Leaf base cuneate to acuminate; anthers 3.5–4 mm long; stem trichomes with midpoints 0.4–1(2) mm long; Democratic Republic of the Congo, Rwanda, Uganda, Tanzania; rare |
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| 94b | Leaf base rounded, rarely cuneate; anthers 4–6 mm long; stem trichomes with midpoints 0.05–0.2 mm long; widespread |
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| 95a | Leaves elliptic, with 8–12 pairs of primary veins, 2.5–3.5 times longer than wide |
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| 95b | Leaves ovate, sometimes elliptic, with 4–8 pairs of primary veins, 1.5–2.5(3) times longer than wide |
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| 96a | Leaves concolorous; calyx lobes 2–4(5) mm long, long-acuminate; Sudan, Eritrea, Ethiopia, Somalia, 1500–2800 m elevation |
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| 96b | Leaves discolorous; calyx lobes 1–2 mm long, obtuse to acute, sometimes acuminate; southern Ethiopia, Somalia and further south, 650–1900 m elevation |
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| 97a | Leaf base cuneate to acuminate; anthers 3.5–4 mm long; stem trichomes with midpoints 0.4–1(2) mm long; Democratic Republic of the Congo, Rwanda, Uganda, Tanzania; rare |
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| 97b | Leaf base rounded, rarely cuneate; anthers 4–6 mm long; stem trichomes with midpoints 0.05–0.2 mm long; widespread, common |
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| 98a | Plants with no prickles visible |
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| 98b | Plants with at least some visible prickles. |
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| 99a | Leaves 3–8 times longer than wide, narrow-elliptic or lanceolate, rarely narrow-ovate |
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| 99b | Leaves 1–3 times longer than wide, orbicular, ovate, obovate, elliptic or oblong |
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| 100a | Corolla 0.9–1.5 cm in diameter; trichomes on young stems with ca. 8 rays; native plant in eastern and north-eastern Africa, black cotton soils |
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| 100b | Corolla 2.5–3(5) cm in diameter; trichomes on young stems with 10–14(16) rays; weed in southern Africa and Mediterranean |
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| 101a | Leaves less than 1.5 times longer than wide, orbicular, ovate or obovate; arid north-eastern Africa |
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| 101b | Leaves more than 1.5 times longer than wide, ovate, elliptic or oblong; widespread |
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| 102a | Erect shrub to small tree 1.5–4 m tall; leaves and flowers readily deciduous, so stems often bare; seeds orange-brown |
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| 102a | Scandent to erect shrub up to 1 m tall; leaves and flowers not readily deciduous, stems not bare; seeds dark brown to almost black |
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| 103a | Leaf base attenuate; corolla lobed for ca. 3/4 of the way to the base; trichomes on the young stems with 10–20 rays |
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| 103a | Leaf base cuneate, rounded or truncate, rarely attenuate; corolla lobed for 1/2–2/3 of the way to the base; trichomes on the young stems with 6–9(12) rays |
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| 104a | Petioles 1/3–2/3 as long as the adult leaves; flowers 4–8 per inflorescence; seeds 2.8–3.5 mm long; Angola |
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| 104b | Petioles 1/6–1/4 as long as the adult leaves; flowers 8–15 per inflorescence; seeds 3.6–4.8 mm long; eastern and south-eastern Africa |
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| 105a | Calyx lobes 5–10 mm long in flower, foliaceous, ovate to elliptic; leaf apex rounded, sometimes obtuse; arid north-eastern Africa |
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| 105b | Calyx lobes 1–4.5 mm long in flower, not clearly foliaceous, deltate; leaf apex acute or obtuse, sometimes rounded; widespread |
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| 106a | Scandent shrub; inflorescences 3.5–7 cm long; young stems with stalked trichomes, the stalks 0.1–0.4 mm long; eastern African highlands |
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| 106b | Erect herb or shrub; inflorescences 1–3.5 cm long; young stems with sessile or stalked trichomes, the stalks up to 0.15 mm long; widespread |
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| 107a | Fruit (1)1.5–2.5(5) cm in diameter; plant cultivated for leaves or fruits; widespread and common in western and eastern Africa |
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| 107b | Fruit (0.7)0.75–1.1 cm in diameter; wild plant in southern Africa |
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| 108a | Leaves elliptic; corolla 1.5–2.2 cm in diameter, mauve to purple |
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| 108b | Leaves ovate, sometimes oblong; corolla 0.9–1.3 cm in diameter, usually white |
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| 109a | Prickles on young stems predominantly curved |
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| 109b | Prickles on young stems predominantly straight, sometimes straight and reflexed |
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| 110a | Most leaves on reproductive branches less than two times longer than wide |
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| 110b | Most leaves on reproductive branches two times longer than wide or longer |
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| 111a | Corolla lobed for ca. 3/4 of the way to the base or deeper; seeds dark brown to almost black; continental Africa north of the equator |
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| 111b | Corolla lobed for ca. 2/3 of the way to the base; seeds yellow to orange-brown; continental Africa south of the equator |
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| 112a | Inflorescences with 1(2) flowers; trichomes on young stem with 12–18 rays |
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| 112b | Inflorescences with (1)2–20 flowers; trichomes on young stem with 6–10 rays |
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| 113a | Leaves drying yellowish green; anthers 3–4 mm long; southern Africa |
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| 113b | Leaves drying reddish brown; anthers (4.5)5.5–8 mm long; Madagascar |
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| 114a | Young stems with a dense covering of long-stalked trichomes and prickles of different lengths; 1000–3000 m elevation |
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| 114b | Young stems with a sparse or dense covering of sessile or short-stalked trichomes of uniform lengths; 0–1000 m elevation |
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| 115a | Plant erect, 0.4–1.5 m tall; corolla 0.9–1.4 cm in diameter; anthers 2.5–3.5 mm long |
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| 115b | Plant scandent, sometimes erect; corolla (1.2)1.5–3 cm in diameter; anthers 4–8 mm long |
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| 116a | Inflorescence 3–4(6) cm long; trichomes on abaxial surfaces of the leaves with stalks 0.15–0.4 mm long; eastern Africa |
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| 116b | Inflorescence 1–2.5 cm long; trichomes on abaxial surfaces of the leaves with stalks 0–0.15 mm long; southern Africa |
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| 117a | Calyx 7–9 mm long at anthesis, the lobes caudate; trichomes on young stems with midpoints (0.5)1.5–2.5(3) mm long; Tanzanian highlands |
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| 117b | Calyx 2–6 mm long at anthesis, the lobes acute to acuminate; trichomes on young stems with midpoints 0–1.5 mm long; widespread |
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| 118a | Leaves 3–6 times longer than wide, the bases attenuate; arid eastern and north-eastern Africa |
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| 118b | Leaves 2–3 times longer than wide, the bases cuneate to cordate; eastern Africa and Madagascar |
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| 119a | Corolla (1.8)2–3 cm in diameter; anthers (5.5)6–10 mm long |
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| 119b | Corolla 1–2(2.4) cm in diameter; anthers 4–6(7) mm long |
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| 120a | Inflorescences with 1–2(5) flowers; young stems with trichomes sessile or stalked, the stalks up to 0.2 mm long; Madagascar |
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| 120b | Inflorescences with 3–10 flowers; young stems with trichomes always stalked, the stalks 0.1–0.4 mm long; eastern Africa |
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| 121a | Leaves 3–8 cm long, densely stellate-pubescent on both sides; eastern African coasts |
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| 121b | Leaves 6–13 cm long, adaxially glabrescent; eastern African mountains |
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| 122a | Leaves apically rounded, with thick-stalked trichomes on the adaxial surface; inflorescence with 1–2(3) flowers; eastern Africa |
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| 122a | Leaves apically acute to obtuse, sometimes rounded, without thick-stalked trichomes on the adaxial surface; inflorescence with (1)2–20 flowers |
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| 123a | Leaves with 2–3 pairs of primary veins; seeds dark brown to almost black; continental Africa north of the equator |
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| 123b | Leaves with 4–6 pairs of primary veins; seeds yellow to orange-brown; continental Africa south of the equator or near the equator |
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| 124a | Prickles usually under 2 mm long; leaves 2.5–5.5 cm long; trichomes on the leaves and stems with stalks ca. 0.1 mm long; inland eastern Africa |
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| 124b | Prickles over 2 mm long; leaves 3–14 cm long; trichomes on the leaves and stems sessile or with stalks under 0.1 mm long; coastal eastern Africa |
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| 125a | Leaves more than two times longer than wide |
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| 125b | Leaves less than two times longer than wide |
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| 126a | Corolla 2.5–3(5) cm in diameter; anthers 6–10 mm long; invasive in southern Africa and around the Mediterranean |
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| 126b | Corolla 0.9–2.3 cm in diameter; anthers 2.5–6 mm long; Ethiopia, Tanzania or further south |
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| 127a | Prickles on young stems 8–16 mm long; calyx 7–12 mm long at anthesis; Ethiopia |
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| 127b | Prickles on young stems 1–6 mm long; calyx 2.5–7 mm long at anthesis; Tanzania or further south |
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| 128a | Plant scrambling shrub; leaves almost glabrous, trichomes on the leaves with up to 4 rays; Tanzanian coastal forest, rare (possibly extinct) |
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| 128b | Plant erect; leaves stellate-pubescent, trichomes on the abaxial sides of the leaves with 17–15 rays; southern Africa |
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| 129a | Leaves elliptic; corolla 1.5–2.2 cm in diameter, mauve to purple |
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| 129b | Leaves ovate, sometimes oblong; corolla 0.9–1.3 cm in diameter, usually white |
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| 130a | Leaves on reproductive branches less than 2 cm long |
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| 130b | Leaves on reproductive branches more than 2 cm long |
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| 131a | Leaves and flowers predominantly on short shoots, few on main stems; corolla lobed for 1/2–2/3 of the way to the base; seeds yellow to orange-brown; southern Madagascar |
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| 131b | Leaves and flowers predominantly on main stems, few on short shoots; corolla lobed for ca. 3/4 of the way to the base; seeds dark brown to almost black; continental Africa north of the equator |
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| 132a | Calyx lobes fused in fruit, ca. 10 mm long, fully covering the mature fruit; leaves ovate |
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| 132b | Calyx lobes free in fruit, 4–5 mm long, the mature fruit at least partly exposed; leaves orbicular, elliptic or ovate |
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| 133a | Inflorescences with 1(2) flowers; trichomes on young stem with 12–18 rays |
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| 133b | Inflorescences with (1)2–20 flowers; trichomes on young stem with 6–10 rays |
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| 134a | Leaves elliptic |
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| 134a | Leaves ovate |
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| 135a | Leaves drying yellow-green; anthers 3.5–5.2 mm long; southern Africa |
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| 135b | Leaves drying reddish brown; anthers 5.5–8 mm long; Madagascar |
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| 136a | Anthers 3.2–3.5 mm long; southern Africa |
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| 136b | Anthers 4.5–8 mm long; widespread |
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| 137a | Calyx lobes 3–7 mm long at anthesis |
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| 137b | Calyx lobes 0.5–3 mm long at anthesis |
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| 138a | Leaves yellow-green; trichomes on the abaxial sides of the leaves with midpoints 1–2 mm long; Tanzania |
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| 138b | Leaves reddish; trichomes on the abaxial sides of the leaves with midpoints to 0.5 mm long; Madagascar |
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| 139a | Leaves 6–13 cm long; prickles on young stems 1–2 mm long; highland eastern Africa |
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| 139b | Leaves 1–5(6) cm long; prickles on young stems 2–10 mm long; continental Africa, north of the equator or Madagascar |
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| 140a | Pedicels 0.2–1 mm long; seeds almost black; continental Africa, north of the equator |
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| 140b | Pedicels 1.5–2(2.5) cm long; seeds dull yellow to orange-brown; Madagascar |
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