Solanum capsicoides
Not known.
A lowland species of humid or seasonally humid warm subtropical or tropical areas nearly or quite free from frost. Throughout its now extensive range (Nee, 1979) an often abundant weed in unshaded disturbed artificial weedy habitats such as roadsides, waste places, pastures, old coffee plantations, stream banks, beaches, cultivated land (presumably the edges), open woods and around dwellings. Probably preferring sandy habitats but also recording from wet alluvial clay and limestone derived soils. Native along the Atlantic coast of Brazil; introduced in the Guianas, Costa Rica, Nicaragua and Honduras, common on the Caribbean islands, common in Florida, sporadic in nearby states; sporadically introduced in Hawaii, Africa, India and Sri Lanka, more commonly in southern China, Java, Sumatra, and Australia. It can be expected in to eventually occur as a cultivated plant or as naturalized in any tropical or subtropical area of the world. At elevations up to 1500 m, the great majority of collections from below 1000 m with no geographic trends apparent.
Sporadically introduced and perhaps naturalized in western Africa (Sierra Leone, São Tomé and Principe, and Ghana) and a few collections recorded from East Africa (Ethiopia and Tanzania); native to the Atlantic coast of southern South America (Nee 1979), but now widespread as an introduction, often an abundant weed in unshaded disturbed habitats and around dwellings.
Solanum capsicoides shares the much-flattened seeds forming a wing around the embryo with several other species in an unnamed subsection (Nee, 1991, 1999). A key to distinguish the species is given by Nee (1991). It belongs to the Leptostemonum clade of Solanum (Bohs, 2005). Within Leptostemonum, it belongs to the Acanthophora clade, a monophyletic group that includes most of the species traditionally recognized in Solanum section Acanthophora Dunal (the S. mammosum species group of Whalen, 1984; Levin et al., 2006). Within the Acanthophora clade, S. capsicoides is sister to a clade that consists of S. acerifolium, S. atropurpureum, and S. tenuispinum, and the four species together form a well-supported monophyletic group (Levin et al., 2005).
D’Arcy, W.G. 1974. Solanum and its close relatives in Florida. Ann. Missouri Bot. Gard. 61: 819-867.
Edmonds, J. M. 2012. Solanum spp. 1-25, 46-51, 63-64, 67-72. In: J.M. Edmonds, Solanaceae, Flora of Tropical East Africa. Kew: Royal Botanic Gardens, Kew.
Nee, M. 1979. Patterns in biogeography in Solanum, section Acanthophora. Pp. 569–580 in J. G. Hawkes, R. N. Lester & A. D. Skelding (eds.), The Biology and Taxonomy of the Solanaceae. Academic Press, London.
Whalen, M.D. 1984. Conspectus of species groups in Solanum subgenus Leptostemonum. Gentes Herbarum 12 (4): 179-282.
Nee, M. 1991. Synopsis of Solanum section Acanthophora: a group of interest for glycoalkaloids. Pp. 257–266 In: J.G. Hawkes, R.N. Lester, M. Nee, and N. Estrada-R. (eds.). Solanaceae III: Taxonomy, Chemistry, Evolution. Richmond, Surrey, UK: Royal Botanic Gardens, Kew and Linnean Society of London.
Nee, M. 1999. Synopsis of Solanum in the New World. Pp. 285–333 in M. Nee, D. E. Symon, R. N. Lester & J. P. Jessop (eds.), Solanaceae IV: Advances in Biology and Utilization. Royal Botanic Gardens, Kew.
Levin, R.A., K. Watson & L. Bohs 2005. A four-gene study of evolutionary relationships in Solanum section Acanthophora.Amer. J. Bot. 92(4): 603–612.
Bohs, L. 2005. Major clades in Solanum based on ndhF sequences.Pp. 27-49 in R. C. Keating, V. C. Hollowell, & T. B. Croat (eds.), A festschrift for William G. D’Arcy: the legacy of a taxonomist. Monographs in Systematic Botany from the Missouri Botanical Garden, Vol. 104. Missouri Botanical Garden Press, St. Louis.
Levin, R.A., N.R. Myers, & L. Bohs 2006. Phylogenetic relationships among the "spiny" solanums (Solanum subgenus Leptostemonum). Amer. J. Bot. 93: 157-169.
chloroplast ndhF sequence: GenBank AF500808 (voucher: Bohs 2451, UT). http://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?db=nucleotide&val=27463399 chloroplast trnT-F sequence: GenBank AY266251 (voucher: Bohs 2451, UT). http://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?db=nucleotide&val=33355752 chloroplast trnS-G sequence: GenBank AY555460 (voucher: Bohs 2451, UT). http://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?db=nucleotide&val=49065900 nuclear ITS sequence: GenBank AY561265 (voucher: Bohs 2451, UT). http://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?db=nucleotide&val=49458083 nuclear waxy (GBSSI) sequence: GenBank AY562954 (voucher: Bohs 2451, UT). http://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?db=nucleotide&val=45826390
Local Names in Africa. Sierra Leone: Mano bonjema, Mano sakrim; Coakroach berry (Burkill 1985).
Allioni's name for the species was validly published and the description alone is ample enough for definitive determination, even without the photograph of the type at TO kindly furnished to Nee by Dr. B. L. Burtt of the Royal Botanic Garden, Edinburgh. The quite fitting epithet "capsicoides" is found on several old herbarium specimens in European herbaria, including on the type of S. ciliatum Lam., but it was never investigated nor taken up by later botanists. .
Solanum capsicoides has some peculiarities which would seem to adapt it to the naturally disturbed habitats of the sandy restingas. It is a a lower growing, more spreading species than its erect and single-stemmed relatives of mesic and wooded habitats such as S. acerifolium or S. atropurpureum.Winged seeds are probably an adaptation to local wind or water dispersal after passing of the seeds by birds in these relatives with small juicy berries. The seeds have become much broader in S. capsicoides and it is significant that they not only are blown around more easily than those of S. atropurpureum but that they also float for a few days. There seems no longer to be any provision for animal dispersal of the fruits of S. capsicoides since they regularly are found on herbarium specimens in the mature state; they are nearly dry at maturity and split irregularly to permit the seeds to escape in a not very effective shaker mechanism. The development (or retention?) of the striking bright orange-red color of the fruits in this context is difficult to explain.
Despite the considerable confusion in nomenclature involving this species, it is one of the most easily recognized and least variable in sect. Acanthophora. The large bright orange-red fruit and large straw colored seeds maintain their bright colors in all but the most poorly preserved herbarium specimens. The leaves are distinctive, appearing glabrous but for the sparse silky hairs; it is the only species in the section, and to my knowledge in subgen. Leptostemonum, without stellae on any part of the plant. The calyx with stout spines from enlarged, almost bulbous bases is distinctive in the section. It could be confused with extremes of S. atropurpureum which have shallowly lobed leaves, but this latter species has much smaller yellow fruits and yellowish rather than white corollas as well as stellae (even if only a few and confined to the base of the sinuses) on the underside of the leaf. Solanum aculeatissimum with which it has often been unnecessarily confused, is easily distinguished by the non-winged seeds, more copious glandular pubescence, and the presence of stellae on the underside of the leaf.
It is found sporadically in both western and eastern tropical Africa, and may be increasing in distribution; it is apparently often introduced from cultivation. Solanum capsicoides can be easily distinguished from other similar species by its large (to 3.5 cm in diameter) bright orange fruit with winged seeds; S. aculeatissimum and S. viarum have smaller fruits that are yellowish green at maturity and have unwinged seeds. Solanum mammosum has larger, irregularly shaped fruits usually (in Africa) with 4-5 protuberances at the base, unwinged seeds and purple flowers. Both S. capsicoides and S. mammosum have a spongy fruit mesocarp.
Solanum capsicoides was long known as S. ciliatum. Nee (1979) details the history of usage of these names. Edmonds (2012) cited the specimen we designate here as the lectotype of S. ciliatum as the holotype; we feel that lectotypification is more appropriate in this case, as the sheet is not annotated by Lamarck.
We have not seen the holotype (cited as in TO by Nee 1979) of S. capsicoides, and have designated lectotypes only in unambiguous situations (e.g., S. arrebenta and S. sinuatifolium for which no specimens are extant), where we have seen specimens and in situations where we have needed to correct previous incorrect citations of holotypes (e.g., Edmonds 2012). We are here following Nee (1979) in assigning S. arrebenta and S. sinuatifolium to the synonymy of S. capsicoides, but the plates in Flora Fluminensis are very schematic and could be of any member of the Acanthophora clade.