Solanum cyanocarphium Blume, Bijdr. Fl. Ned. Ind. 13: 700. 1826.

Aubriot, Xavier & Knapp, Sandra, 2022, A revision of the " spiny solanums " of Tropical Asia (Solanum, the Leptostemonum Clade, Solanaceae), PhytoKeys 198, pp. 1-270 : 1

10. Solanum cyanocarphium Blume, Bijdr. Fl. Ned. Ind. 13: 700. 1826.

Figs 1D , 4E , 16

Solanum sarmentosum Nees, Trans. Linn. Soc. London 17(1): 58. 1834, nom. illeg., non Solanum sarmentosum Lam., 1794. Type. Malaysia. Penang: Sin. loc., 1822, N. Wallich s.n. [Wallich Catal. 2628f] (lectotype, designated here: GZU [GZU000255826]; isolectotype: BM [BM000886340], K-W [K001116668]).

Solanum bullatorugosum Dunal, Prodr. [A. P. de Candolle] 13(1): 236. 1852. Type. Indonesia. Java: Sin. loc., H. Zollinger 1018 (lectotype, designated here: G-DC [G00145849]; isolectotypes: G [G00301680], P [P00369075, P00369076]).

Solanum maingayi Kuntze, Revis. Gen. Pl. 454. 1891. Type. Malaysia. Malacca: Sin. loc., A.C. Maingay 1158 (lectotype designated by Turner 1998, p. 40, as “holotype”: K [K001080494]; isolectotype: LAE [acc. # 229591]).

Solanum sparsiflorum Elmer, Leafl. Philipp. Bot. 5: 1838. 1913, nom. illeg., non Solanum sparsiflorum Dammer, 1912. Type. Philippines. MIMAROPA: "Puerto Princesa (Mount Pulgar), Province of Palawan, Island of Palawan", May 1911, A.D.E. Elmer 13157 (lectotype, designated here: GH [00077851]; isolectotypes: BISH [BISH1005088], BM [BM000778206], CAL [acc. # 316497], E [E00273861], F [acc. # 384070, v0073463F], G [G00343321], HBG [HBG511492], K [K000195917], L [L0003665], LAE [acc. # 229593], LE, MO [acc. # 706769, MO-2289036], NY [00172293], P [P00379711], W [acc. # 1913-0005906], U [U0113978], US [00027804, acc. # 873055]).

Solanum thorelii Bonati, Bull. Soc. Bot. Genève, 1913, sér. 2, 5: 310. 1914. Type. Vietnam. Tây Ninh: “Caï Cong, environs de village", 1862, C. Thorel 1419 (lectotype, designated here: P [P00054148]; isolectotypes: P [P00054149, P00054150]).

Solanum sakhanii Hul, Fl. Photogr. Cambodge 522. 2013. Type. Cambodia. Sihanoukville: “Sihanoukville”, 3 May 2008, K.C. Cheng et al. CL929 (holotype: P [P00836398]; isotypes: P [P00836399, P00836400]).

Type.

Indonesia. Java: West Java ["in oryzetis siccis montium Seribu" Curug Seribu near Bogor; from protologue], C.L. Blume s.n. (lectotype, designated here: L [L0003630]) .

Description.

Herbs to small shrubs, creeping over the ground, to 1 m tall, armed. Stems decumbent, terete, black to dark brownish, prickly and sparsely stellate-pubescent; prickles to 5 mm long, to 2.5 mm in diameter at the base, straight or curved at the tip, awl-shaped to deltate, flattened, pale yellow in dry material, tan or purplish black in live plants, glabrescent; trichomes porrect-stellate, sessile to stalked, the stalks to 0.1 mm long, the rays 4-7, 0.1-0.4 mm long, the midpoints absent or to 0.4 mm long, sometimes purplish black in live plants; new growth moderately to densely stellate-pubescent, light green in dry material; bark of older stems brownish grey, sparsely stellate-pubescent. Sympodial units plurifoliate, the leaves usually not geminate. Leaves simple, more or less deeply lobed, the blades 4.5-9 cm long, 2-5 cm wide, ca. 2 times longer than wide, elliptic to ovate, chartaceous, slightly discolorous, moderately prickly with 3-7 prickles per leaf side, the prickles to 1 cm long, to 1 mm wide at the base, straight at the tip, awl-shaped, conical, pale yellow in dried material sometimes purplish black in live plants, glabrous; adaxial surface mid-green, moderately stellate-pubescent, the stellate trichomes porrect, sessile to stalked, the stalks to 0.1 mm long, the rays 3-5, 0.1-0.4 mm long, the midpoints to 1 mm long, 2-3 times longer than the rays; abaxial surface light green, moderately stellate-pubescent with trichomes like those of the adaxial surface; major veins 4-5 pairs, drying dark; base cuneate to truncate; margins shallowly to deeply lobed, the lobes 1-4 on each side, 0.5-1 cm long, deltate to oblong, apically rounded, the sinuses extending up to halfway to the midrib; apex rounded to acute; petiole 0.7-2.2 cm long, 1/8-1/5 of the leaf blade length, moderately stellate-pubescent, armed with 1-4 prickles like those of the blades. Inflorescences 1.5-3.5 cm long, apparently lateral, unbranched, with ca. 1-4 flowers, 1-2 flowers open at any one time, sparsely to moderately stellate-pubescent with trichomes like those of the stems but with longer midpoints, unarmed; peduncle 0-8 mm long, unarmed; pedicels 0.5-2 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, spreading to erect to somewhat nodding at anthesis, unarmed or with 1-11(-16) prickles, moderately stellate-pubescent with trichomes like those of the inflorescence axes, articulated at the base; pedicel scars spaced 0.5-6(-18) mm apart. Buds globose to oval, strongly included in the calyx lobes. Flowers 5-merous, apparently all perfect. Calyx with the tube 2-4 mm long, campanulate, the lobes 2-5 mm long, 1-1.5 mm wide, narrowly deltate, apically acute, densely prickly and stellate-pubescent abaxially with numerous prickles and trichomes like those of the pedicels but with shorter midpoints. Corolla 1-1.5 cm in diameter, white or purple, stellate, lobed ca. 1/2 of the way to the base, the lobes 3-6 mm long, 2-3 mm wide, deltate, spreading or somewhat campanulate and erect (not spreading and perpendicular to the pedicel) at anthesis, glabrous adaxially, moderately to densely stellate pubescent abaxially, the trichomes often purple-tinged. Stamens equal; anthers 3-5 mm long, ca. 0.5 mm wide, connivent, tapering, orange, glabrous, poricidal at the tips, the pores directed distally, not elongating to slits with drying; filament tube minute, glabrous; free portion of the filaments ca. 1 mm long, glabrous. Ovary conical, minutely glandular-puberulent; style ca. 5 mm long, slender, curved at the apex, glabrous; stigma capitate, the surface minutely papillate. Fruit a globose berry, 1-4 per infructescence, 1-1.6 cm in diameter, red when mature, the pericarp thin and smooth, glabrous; fruiting pedicels 2.5-4 cm long, 0.5-1 mm in diameter at the base, 1-1.5 mm in diameter at the apex, woody, deflexed and nodding, unarmed or with 1-10 prickles; fruiting calyx lobes elongating to 1.3 cm long, 1/2-3/4 the length of the mature fruit, the tips slightly reflexed, usually completely enclosing at least the lower half of the berry, with 6-17(-24) prickles, these often purplish black. Seeds 85-170 per berry, 1.75-2 mm long, 1.5-2 mm wide, flattened reniform, dull yellow, the surface minutely pitted, the testal cells sinuate in shape. Chromosome number: not known.

Distribution

(Fig. 17 ). Solanum cyanocarphium is widely distributed from southeastern Indochina to western Malay Archipelago (Borneo, Java, Sumatra and south Philippines).

Ecology and habitat.

Solanum cyanocarphium has been found growing on limestone, in riparian and secondary tropical forests as well as in open degraded vegetation; from 10 to 1,000 m elevation.

Common names and uses.

Cambodia. plone lane [Khmer] (Chassagne 138), trâp krab [Khmer] ( Hul and Dy Phon 2014); Indonesia. North Sumatra: tioeng (Toroes 1639), teroeng oetan (Toroes 2926); Malaysia/Singapore. tĕrong puyoh, tĕrong tikus, tĕrong pipit (all meaning "little brinjal", Burkill 1935); Vietnam. Dông Nai: blou xit [Mnong] (Pierre s.n.), cà co [Vietnamese] (Pierre s.n.).

Solanum cyanocarphium is said to have edible berries (Cuadra A1010) and the juice is drunk for fever ( Burkill 1935). Burkill (1935) also records its use to improve the appetite of elephants.

Preliminary conservation status

( IUCN 2019). Least Concern (LC); EOO 3,563,723 km2 (LC), AOO 92 km2 (EN). Historical collections of Solanum cyanocarphium suggest that this species is widely distributed but absence of recent collections from large areas of the historical range suggests it may merit conservation concern because of the increasing anthropogenic alteration of natural habitat throughout lowland and coastal tropical Asia ( Wikramanayake et al. 2002).

Discussion.

Solanum cyanocarphium is a weakly climbing shrub to vine that scrambles over vegetation with hooked prickles; most specimens are quite thin-stemmed and somewhat scrappy. It is superficially similar to S. procumbens but differs from it in the strongly accrescent and spiny fruiting calyx (not accrescent in S. procumbens ), larger prickles on stems and leaf blades, deeply lobed leaves (leaves usually entire in S. procumbens ) and larger and many seeded berries. The accrescent calyx in fruit is similar to that of S. involucratum , but that species is more robust and has pubescent, rather than glabrous berries. Hul and Dy Phon (2014) suggested that S. cyanocarphium was introduced to Indochina but described what we recognise as a synonym ( S. sakhanii ) as new; we consider S. cyanocarphium to be native across tropical Asia where it occurs.

Aubriot et al. (2016a) included S. cyanocarphium in a clade called ' S. cyanocarphium + S. sakhanii '; subsequent examination shows that the two taxa are synonyms (see above), not surprising given the very short branch lengths ( Aubriot et al. 2016a).

The Blume specimen in L (L0003630) we have selected as the lectotype of S. cyanocarphium is the only one we have found that corresponds to the protologue and is likely to have been the original material that Blume used.

Nees van Esenbeck (1834) only cited a Wallich catalogue number in the protologue of S. sarmentosum , but no herbarium. The Solanaceae from his own herbarium are now held at the herbarium (GZU) of the University of Graz ( Stafleu and Cowan 1981) and we have selected the specimen there (GZU000255826) as the lectotype, it has flower and fruit and an annotation " S. sarmentosum " in Nees van Esenbeck’s hand.

The protologue of S. bullatorugosum ( Dunal 1852) cites two specimens at G; we have selected the more complete sheet at G-DC (G00145849) with both flowers and fruits as the lectotype.

The collection cited in the protologue of S. sparsiflorum ( Elmer 1913) without specification of any herbarium (Elmer 13157) is represented in many collections; we have chosen that at GH (0007785) as the lectotype because it is the duplicate best matching the protologue and has both flower and mature fruit. Hul and Dy Phon (2014) cited a sheet in PNH as "holo-, PNH" but this does not constitute effective lectotypification.

Hul and Dy Phon (2014) cited a holotype at P for S. thorelii , but no herbarium was cited in the original protologue ( Bonati 1914). We select here the best preserved of the three duplicates of Thorel 1419 held at P (P00054148) as the lectotype.

Specimens examined.

See Suppl. materials 1-3.

Figures

Figure 1. Diversity of habits and habitats for Solanum subgenus Leptostemonum in tropical Asia A Solanum giganteum, a small tree of the tropical forest understory in India and Sri Lanka (field photograph, unvouchered, India) B S. trilobatum, a scandent species in its typical habitat, along the road in mangrove area (Meeboonya et al. RM 245, Thailand) C S. lasiocarpum, an erect shrub found in disturbed and human inhabited areas where it is often cultivated (Meeboonya et al. RM 272, Thailand) D S. cyanocarphium, a creeper with decumbent stems in forest understory (field photograph, unvouchered, Vietnam) E S. camranhense, a scandent shrub endemic to the coastal dunes of South Vietnam (field photograph, unvouchered, Vietnam) F S. insanum, the wild progenitor of the brinjal eggplant usually found in degraded areas (Meeboonya et al. RM 305, Thailand). Photograph credits: A S. More B, C, F X. Aubriot; D M. Nuraliev E S. Hul.

Figure 4. Diversity of fruits and seeds in tropical Asian spiny solanums A large oval fruit of Solanum insanum (Sampath Kumar et al. 126918, India) B condensed infructescence and small rounded fruits of Solanum hovei (field photograph, unvouchered, India) C large rounded hairy fruit of S. involucratum with strongly accrescent and spiny calyx (field photograph, unvouchered, Vietnam) D small rounded fruit of Solanum pubescens with elongated calyx lobes (Sampath Kumar et al. 126956, India) E rounded fruit of Solanum cyanocarphium with strongly accrescent and spiny calyx (field photograph, unvouchered, Vietnam) F small rounded fruit of S. trilobatum in transverse cut (Meeboonya et al. RM 242, Thailand) G rounded fruit and accrescent calyx of S. praetermissum in transverse cut (field photograph, unvouchered, Vietnam) H oval fruit of Solanum virginianum in transverse cut (field photograph, unvouchered, India). Photograph credits: A, B, D, F, H X. Aubriot C, E, G M. Nuraliev.

Figure 16. Solanum cyanocarphium Blume A herbarium specimen collected in Vietnam in 1929 (Lichy 18, P 00055844) B detail of the leaves and prickles (field photograph, unvouchered, Vietnam) C detail view of a flower (field photograph, unvouchered, Vietnam) D detail view of a fruit (field photograph, unvouchered, Vietnam). Photograph credits: A CC-BY, Museum national d'Histoire naturelle, Paris B-D M. Nuraliev.

Figure 17. Distribution of S. cyanocarphium.