Solanum lasiocarpum [Dunal ]

Solanum lasiocarpum

Citation author:

Dunal

Citation:

Histoire des Solanum 223. 1813.

Type:

Lectotype: Tab. 35 in Rheede, Hortus Indicus Malabaricus 2. 1680, designated by Whalen et al., 1981.

Last edited by:

Bohs, L.

Written by:

M.D. Whalen, D.E. Costich and C.B. Heiser, Jr.

Habit:

Erect or spreadingly branched, lignescent perennials, 1-2 m tall. Stems invested with a mixture of stipitate and essentially sessile stellate trichomes, the multiseriate stalks of stipitate stellae variable in length, the longest 05-2.5 mm, with lateral rays 5-8, 0.4-1.5 mm long. the midpoint length variable, longer than the rays in some forms, manifestly shorter in others; cauline prickles often short, 0.2-0.5 (-0.8) mm long, laterally compressed, often broad-based and elongate-deltoid in outline, usually spaced at least by their length, often becoming sparse on later growth.

Sympodial structure:

Sympodial units 2-foliate, geminate.

Leaves:

Leaves simple, the blades 20-35 x 15-30 cm, ca. 1-1.4 times as long as wide, broadly ovate or ovate, thin-textured, pubescent adaxially, sericeous or farinaceous in appearance, the hairs variable, with stellae sessile, the largest with midpoints 0.8-2.5 mm long and with 2-5 variously reduced lateral rays 0.1-0.5 mm long, also with scattered stellae bearing well-developed, ascending rays, the largest stellae sometimes grading into much smaller ones with short midpoints, abaxially felty-pubescent with short-stalked and sessile stellae; armed with prickles similar to those of the stem, these scattered along petiole, midrib and principal lateral veins; major lateral veins 4-5 on each side; base truncate or obtuse; margin coarsely dentate with 3-6 deltoid or narrowly deltoid, acute- or round-tipped lobes at lateral vein terminations, the interlobal sinuses entire or with 1-2 smaller tooth-like lobes; apex acute; petioles 3-14 cm, 1/5-1/2 the length of the blades, pubescent with with a mixture of stipitate and essentially sessile stellate trichomes like those of the stem.

Inflorescences:

Inflorescences usually short, 0.4-0.9 cm, extra-axillary, often very close to a leaf pair, unbranched, with 6-16 flowers, the distal ones female-sterile, the axes stellate-pubescent, frequently prickly; peduncle 0.1-0.4 cm; rachis 0.3-0.6 cm; pedicels 4-9 mm in flower, 10-15 mm in fruit, spaced 0.25-1.25 mm apart, articulated at the base.

Flowers:

Flowers with the calyx broadly campanulate, 6-9 mm wide, 6-9 mm long, the tube 2.5-4.5 mm, the lobes 3-5 x 2.5-5 mm, deltoid or broadly ovate-apiculate, abaxially densely sericeous or occasionally felty, the stellae with delicate, elongate midpoints, rays and stalks. Corolla 2.5-3.5 cm in diameter, 7-16 mm long, stellate, thin-textured, white, the tube 5-8 mm, the lobes 6-9 x 3-6 mm, ovate-lanceolate or lanceolate, stellate-pubescent abaxially, glabrous adaxially. Stamens with filaments ca. 0.1-0.2 mm; anthers 6-8.5 x 1.5-2.2 mm at base, linear-lanceolate, attenuate, connivent, yellow, the pores minute and directed distally. Ovary pubescent, the hairs appearing simple but with underdeveloped rays at base; style 5-10 x 0.25-0.6 mm, cylindrical, glabrous; stigma capitate.

Fruits:

Fruits 1-5 per inflorescence, 2.5-3.5 cm in diameter, globose, orange when ripe, densely spreading-hirsute with sessile stellate hairs with elongate midpoints 1.5-4 mm long; the lateral rays 5-15, short, 0.1-0.4 mm long, inserted at trichome base, these persistent at maturity.

Seeds:

Seeds numerous, 2.2-3.5 x 1.75-2.5 mm, tan, lenticular, broadly reniform-ovate in outline, the surfaces minutely pitted.

Chromosome number:

2n = ploidy missing =24 voucher missing =

Distribution:

Low to middle elevations, 0-1000 m, in forest openings, disturbed sites and second growth thickets; extending from tropical India east through Indochina, extreme southern China, Malaysia and Indonesia to the Philippines and New Guinea.

Phylogeny:

Solanum lasiocarpum belongs to the Leptostemonum clade of Solanum (Bohs, 2005). Within Leptostemonum, it belongs to the Lasiocarpa clade, a monophyletic group that includes most of the species traditionally recognized in Solanum section Lasiocarpa Dunal (Whalen et al., 1981; the S. quitoense species group of Whalen, 1984; Levin et al., 2006). Within this clade, chloroplast sequences from the trnT-F region indicate that S. lasiocarpum belongs to a clade that also includes S. felinum, S. hyporhodium, S. candidum, S. pseudolulo, S. quitoense, S. repandum, and S. vestissimum (Bohs, 2004). Resolution is poor to non-existent among the species of this latter clade. However, it should be noted that the Asian species S. repandum and S. lasiocarpum belong to the same clade as S. candidum in cpDNA analyses. This result is consistent with the hypothesis of Heiser (1986, 1987, 1996) that S. candidum may be one of the closest relatives of the disjunct Asian species of Solanum section Lasiocarpa.

Commentary:

The following commentary is taken verbatim from Whalen et al. (1981):

Solanum lasiocarpum is closely related to the Neotropical S. candidum. The species are so similar that we are not completely at ease retaining them. The smaller prickles, smaller leaves, shorter inflorescences, smaller more sericeous calyces, narrower anthers and smaller fruits that distinguish S. lasiocarpum from S. candidum constitute a long list of differences, but each is quantitative, and in every case, there is some overlap. Until field studies of S. lasiocarpum can be undertaken, the retention of these species is the conservative treatment. If they are combined, the name S. lasiocarpum, based on an Asian type, will have to be applied to the American plants, as well.

Solanum lasiocarpum must be of ultimate American origin, and its present distribution presents a puzzling historical problem, as does that of S. repandum. The great variability of S. lasiocarpum and its morphological divergence from its American counterpart make recent human introduction an unsatisfactory explanation. If dispersal was by birds or early man, it is surprising that the species is not found on intervening islands of Oceania.

The resurrection of a little known epithet for this species is unfortunate, since the name S. ferox has been consistently applied for over a century. Linnaeus gave no reference to a specimen or plate in his protologues of S. ferox, and typification of that name is still problematical. It is clear, however, that the name does not belong to the species treated here, because Linnaeus described a prickly calyx completely enclosing the berry. More likely the name refers to the plant currently called S. involucratum Blume. The latter interpretation was followed by Dunal and other early nineteenth century authors. Misapplication of the name to the species here called S. lasiocarpum began with Nees in 1837 who was followed by Bentham and Hooker in floristic works (see lists of synonymy). Several recent authors have become aware of this misapplication and have taken up the name S. stramonifolium for S. lasiocarpum. This use of the name S. stramonifolium began with Dunal, who must have been confused by Jacquin’s erroneous citation, “In Indiae Orientalis.” In fact, S. stramonifolium is clearly typified and is a species of northern South America already treated in this monograph.

Typification of S. lasiocarpum is straightforward. In Dunal’s 1816 and 1852 works, he cited two specimens, one of which is currently housed at P. However, these specimens had not yet been seen by him at the time of initial publication in 1813 (“v.s. 1814”), and cannot be accepted as types. Dunal’s original concept of the species was based entirely on two pre-Linnaean illustrations (R. Morison, Plantarum Historiae Oxioniensis, sect. 13, tab. 2, Fig. 12 and Rheede, Hortus Indicus Malabaricus 2, tab. 35 ). The Rheede plate has been chosen as lectotype because of its greater clarity and detail.

Commentary added by Bohs (9/2005):

Molecular phylogenies of section Lasiocarpa remove some of the problems posed by Whalen et al. (1981) for the distribution of S. lasiocarpum. Although trees based on chloroplast vs. nuclear sequence data differ in the relationships hypothesized for some Lasiocarpa species (e.g., S. hirtum), both types of data consistently place the neotropical S. candidum together in a clade with the Asian species S. repandum and S. lasiocarpum (Bohs, 2004; Bohs, unpubl. data). Whalen et al. (1981) state that it is surprising that, given that its ancestor must have been dispersed from the New World, S. lasiocarpum is not found on outlying islands of Oceania. Solanum repandum occupies a more or less intermediate range on these outlying islands, so this problem seems to be solved. However, it still remains to be determined how and when the ancestor of S. lasiocarpum and S. repandum arrived in the South Pacific and how the morphological differences between the two Asian species evolved. Poor resolution and conflicting gene topologies for species of section Lasiocarpa suggest that complex patterns of hybridization and introgression may be involved in many species in the section. Perhaps the same processes have contributed to the confusing morphological character suites seen in S. repandum and S. lasiocarpum.

The application of the name S. ferox L. has not been critically examined at this time. According to Heiser (1996), S. ferox is the correct name for the species called S. lasiocarpum and S. repandum in Whalen et al. (1981). More research is needed to clarify the application of this name and to designate a lectotype or neotype for S. ferox L.

Heiser recognizes two varieties of S. ferox. Solanum ferox var. ferox includes plants with prickly calyces that completely enclose the fruit. Solanum ferox var. lasiocarpum (Dunal) Miq. includes plants in which the calyx is not spiny and does not enclose the fruit. This latter variety encompasses domesticates as well as possibly wild plants and includes the species recognized as S. lasiocarpum and S. repandum in Whalen et al. (1981). Domesticaed forms lack prickles, have large fruits, and in some cases have both simple and stellate-haired races. These have been called S. ferox var. repandum (G. Forst.) Bitter or recognized as cultivate group Domesticum within S. ferox.

A summary of Heiser’s classification from Heiser (1996) follows. An alternative scheme divides S. ferox into a number of cultivar-groups with informal names (Hasan & Jansen, 1994). It is clear that there is more taxonomic and nomenclatural work to be done on the Asian species of Solanum section Lasiocarpa.

Solanum ferox L., Sp. Pl. ed. 2: 267. 1762. Type: unclear. Non S. ferox Burm. f (1978), S. ferox Dunal in DC. (1852), S. ferox Miq. (1861).

Solanum ferox var. ferox

syn. S. involucratum Blume, Bijdr. 701. 1826. Type: Dutch East Indies, Java, or Malaya. Location of type unclear. Non S. involucratum Kurz (1877).

Includes plants where the prickly calyx completely encloses the fruits.

Solanum ferox var. lasiocarpum (Dunal) Miquel. Flora van Nederlandsch Indië, vol. 3. 1856.

syns. S. lasiocarpum Dunal, Histoire des Solanum 223. 1813. Lectotype: Tab. 35 in Rheede, Hortus Indicus Malabaricus 2. 1680, designated by Whalen et al., 1981.

S. repandum G. Forst., Florulae Insularum Australium Prodromus 18. 1786. Type: Society Islands. Forster, not found (possibly K, see Symon, 1985). A further search should be conducted before a neotype is selected.

S. ferox var. repandum (G. Forst.) Bitter, Bot. Jahrb. Syst. 55: 84. 1919. Based on S. repandum G. Forst.

S. lasiocarpum var. domesticum Heiser in D’Arcy (ed.), Solanaceae: Biol. & Syst. 413. 1986. Type: Cultivated at Bloomington, Indiana, from seeds originally purchased in market in Bangkok, Thailand, 21 Sept 1980, Heiser 8008 (holotype, IND).

Includes plants with non-prickly calyces that do not enclose the fruits.

References:

Whalen, M.D., D.E. Costich & C.B. Heiser, Jr. 1981. Taxonomy of Solanum section Lasiocarpa.
Gentes Herb. 12: 41-129.

Whalen, M.D. 1984. Conspectus of species groups in Solanum subgenus Leptostemonum.
Gentes Herbarum 12 (4): 179-282.

Symon, D.E. 1985. The Solanaceae of New Guinea.
J. Adelaide Bot. Gard. 8: 1-171.

Heiser, C.B., Jr. 1986. A new domesticated variety and relationships of Solanum lasiocarpum.
Pp. 412-415 in Solanaceae: biology and systematics, ed. W. G. D’Arcy. New York: Columbia University Press.

Heiser, C.B., Jr. 1987. Origins of Solanum lasiocarpum and S. repandum.
Amer. J. Bot. 74: 1045-1048.

Hasan, M., & P. Jansen 1994. Solanum.
Pp. 249-252 in Prosea: plant resources of South-east Asia 8, Vegetables. Prosea, Bogor.

Heiser, C.B., Jr. 1996. Reappraisal of Solanum ferox, S. lasiocarpum, and S. repandum.
Solanaceae Newsletter 4(2): 44-50.

Bohs, L. 2004. A chloroplast DNA phylogeny of Solanum section Lasiocarpa (Solanaceae).
Syst. Bot. 29: 177-187.

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.

Genetics:

nuclear ITS sequence: GenBank AY263457 (voucher: Heiser 8008, IND) nuclear waxy (GBSSI) sequence: GenBank AY996433 (voucher: Ansyar 9605, IND) chloroplast trnS-G sequence: GenBank AY998432 (voucher: Ansyar 9605, IND) chloroplast trnT-F sequence: GenBank AY266256 (voucher: Ansyar 9605, IND) http://www.ncbi.nlm.nih.gov/entrez/viewer.fcgi?db=nucleotide&val=33355757

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Solanaceae

Solanum lasiocarpum Dunal

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