Research Article |
Corresponding author: Xavier Aubriot ( x.aubriot@gmail.com ) Academic editor: Eric Tepe
© 2016 Xavier Aubriot, Caroline Loup, Sandra Knapp.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Aubriot X, Loup C, Knapp S (2016) Confirming the identity of two enigmatic “spiny solanums” (Solanum subgenus Leptostemonum, Solanaceae) collected by Jean-Baptiste Leschenault in Java. PhytoKeys 70: 97-110. https://doi.org/10.3897/phytokeys.70.9758
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Taxonomic revision of the tropical Asian species of Solanum revealed two names, Solanum poka Dunal and Solanum graciliflorum Dunal, whose identities were uncertain and whose application has always been tentative. Material collected in Java at the beginning of the 19th century by Jean-Baptiste Leschenault de la Tour and used to describe these taxa has not been found, despite extensive searches in European herbaria. We here stabilise use of these names by comparing herbarium specimens and drawings of original material made by the artist Toussaint François Node-Véran. Detailed descriptions with synonymy, preliminary conservation assessments and specimen citations are provided for both species. Lectotypes are designated for all names (including synonyms) and epitypes designated for S. poka and S. graciliflorum to stabilise usage.
Exploration, Jean-Baptiste Leschenault de la Tour, Indonesia, Montpellier, Nicolas Baudin, Toussaint François Node-Véran, typification
In 1800, shortly after he became First Consul of the Republic of France, Napoléon Bonaparte approved an expedition along the “coasts of New Holland” (Australia). The expedition, led by Nicolas Baudin, has been cited as one of the most ambitious and the most enriching for collections of natural history of the great scientific expeditions of the early 19th century (
The Baudin expedition lasted four years (1800-1804) and its explicit purpose was “observation and research relating to Geography and Natural History”. The crew included 24 scientists and artists, among them were three botanists and five gardeners that had been carefully selected by Antoine-Laurent de Jussieu, then director of the Muséum National d’Histoire Naturelle (
In the course of preparing a monographic revision of the spiny solanums from tropical Asia (see
Given that no plant specimens corresponding to the protologues have been found, despite extensive searches, we consider the unpublished Node-Véran drawings the most appropriate and only extant possibilities for lectotypifying both S. graciliflorum and S. poka. These two names have long been treated as confusing, or ignored; they have rarely been used (see below in each species treatment), and few herbarium specimens we have seen have been annotated with either name. Most specimens of the taxa we here recognise as S. graciliflorum and S. poka have been annotated incorrectly as widespread weedy taxa (e.g., S. torvum Sw.) or with names we here consider synonyms (e.g., S. athroanthum Dunal); this reflects the limited taxonomic work previously done on tropical Asian Solanum, whose taxonomy has not been revised in detail since
Searches for type specimens of S. graciliflorum and S. poka were made using the resources available in Global Plants (http://plants.jstor.org/) and physically in the herbaria where duplicates could possibly be kept (A,
Solanum athroanthum Dunal, Prodr. [A. P. de Candolle] 13(1): 208. 1852.
Type. Indonesia. Java: [Prov. Banjinwanyne] “in sylvis prope Sukaradja” [Sukaraja], 1846, H. Zollinger 2907 (lectotype, designated here: G-DC [G003043306]; isolectotypes: G-DC [G00301684],
Based on an unpublished illustration of Leschenault collection kept in the Node-Véran collection in Montpellier (lectotype, designated here: Service du Patrimoine Historique de l’Université de Montpellier Node-Véran, Sol. Tab. 47 [MPU028534]); Indonesia. East Java: Blambangan [Sumberwaru, Badjulmati], T. Horsfield s.n. (epitype, designated here:
Scandent shrub to 2 m, armed. Young stems terete, brownish grey, very sparsely stellate-pubescent and prickly, the stellate trichomes porrect, sessile to subsessile, the rays (4-)5–8, 0.1–0.25 mm long, the midpoints to 0.15 mm long, the prickles to 7 mm long, to 8 mm wide at base, curved, deltate, laterally flattened, pale yellow, glabrous; bark of older stems dark brownish grey, glabrescent. Sympodial units difoliate, the leaves geminate, usually similar in size. Leaves simple, the blades (4.5-)7–11 cm long, (1.5-)3–5 cm wide, ca. 2 times longer than wide, elliptic to ovate, chartaceous, slightly discolourous; adaxial and abaxial surfaces sparsely to very sparsely stellate-pubescent and usually with at least some prickles, the stellate trichomes porrect, sessile to subsessile, the rays 6–8, 0.1–0.25 mm long, the midpoint to 0.25 mm long, usually as long as the rays, the prickles 0–10(-12) per leaf side, mostly inserted on the midvein, to 9 mm long, to 2 mm wide at base, straight or slightly curved at the tip, awl-shaped, conical, pale yellow, glabrous; major veins 3–4 pairs drying dark; base attenuate to truncate; margins shallowly to deeply lobed, the lobes 1–3 on each side, 0.5–2.5 cm long, broadly deltate, apically rounded, the sinuses extending up to 2/3 of the distance to the midvein; apex rounded to acute; petiole 0.5–1.8 cm long, 1/10–1/6 of the leaf blade length, sparsely stellate-pubescent with porrect, subsessile trichomes denser at the very base, with 0–2 prickles like those of the blades. Inflorescences leaf-opposed or apparently lateral and borne between leaf pairs, 2–4 cm long, unbranched to up to 6 times branched, with 15–50+ flowers; axes sparsely to very sparsely stellate-pubescent, unarmed; peduncle 1–2(-2.5) cm long, with 0–1 prickles like those of the leaves and stems; pedicels 4–7 mm long, erect, articulated at the base, very sparsely stellate-pubescent, unarmed; pedicel scars spaced 1–5 mm apart. Flowers 5-merous, apparently all perfect. Calyx 1.75–2 mm long, campanulate, pubescent with sessile porrect stellate trichomes like those of the stems, unarmed, the lobes 0.25–0.5 mm long, deltate, apically acute. Corolla 0.5–1 cm in diameter, white to pale lilac, stellate, lobed nearly to the base, the lobes 4–5 mm long, ca. 1 mm wide, narrowly deltate to linear, reflexed at anthesis, densely stellate-pubescent abaxially, the trichomes porrect, sessile, the rays 4–6, 0.1–0.2 mm long, the midpoints the same size than the rays or to 0.25 mm long. Stamens slightly unequal; filament tube < 0.5 mm long; free portion of the filaments almost equal, 0.5–1.25 mm long; anthers unequal, three of the five 4.5–5 mm long and two 3–4 mm long, all 0.5–0.75 mm wide, glabrous, connivent, tapering, poricidal at the tips, the pores not lengthening to slits with age. Ovary conical, minutely glandular-puberulent; style ca. 5.5 mm long, slender, curved at the apex, glabrous; stigma capitate, minutely papillate. Fruit a globose berry, 6–50+ per infrutescence, 3–5 mm in diameter, the pericarp shiny, red when mature, glabrous; fruiting pedicels 0.8–1.2 cm long, ca. 0.5 mm in diameter at the base, tapering to a slightly enlarged apex, woody, spreading, unarmed; fruiting calyx lobes slightly expanding to 1.5 mm long, ca. 1/5 the length of the mature fruit, deltate to lanceolate, unarmed. Seeds 6–9 per berry, 3.5–4 mm long, 3–3.5 mm wide, flattened-reniform, orange-brown, the surface minutely pitted, the testal cells pentagonal in outline.
A Lectotype of Solanum graciliflorum, illustration Sol. Tab. 47 [MPU028534] made by T.F. Node-Véran (1773–1852). Reproduced with permission of the Université de Montpellier – Herbier
The few known collections were flowering and fruiting between May and August.
(Fig.
Data Deficient (DD); known only from seven collections, several of which are of uncertain localities. Solanum graciliflorum has not been re-collected since the first half of the 20th century, indicating it is certainly of conservation concern. Recollection of this species and exploration of the type locality are priorities.
Indonesia. Bali: Perepat Agoeng, 21 Jul 1934, de Voogd 2177 (A); Gorontalo: North Celebes, Jun 1875, Riedel s.n. (K); Java: sin loc., 1802, Horsfield 15 (K); West Java, Bogor, Anonymous s.n. (K); Malaku: “Malay Archipelago, Dawalore [Ambon, Dawa-lour]”, Aug 1883, Riedel s.n. (K).
Solanum graciliflorum is a poorly known species represented by very few collections that presents a combination of morphological features that makes it readily recognisable among tropical Asian spiny solanums. It is superficially similar to S. cyanocarphium Blume, a sympatric species that is distributed across the Sunda Shelf region, and to S. retrorsum Elmer, that occurs mainly in the Philippines. Solanum graciliflorum can be distinguished from both of them by its much sparser indumentum, stout, deltate stem prickles (rather than slender and awl-shaped), and tiny, delicate flowers (hence the species epithet) that are clustered in dense, many-flowered inflorescences. Molecular data show that S. cyanocarphium and S. graciliflorum are not closely related; S. graciliflorum is nested within the Sahul-Pacific clade while S. cyanocarphium is an unresolved species of uncertain affinities (see
Solanum graciliflorum is the type of section Graciliflorum (Dunal) Seithe, a section partly based on the informal grouping made in
In the protologue Dunal referred to an illustration made by Node-Véran, ‘Dun. Suppl. 7. Sol. Mss. tab. 4.’, an orthographic error for ‘Dun. Suppl. Sol. Mss. tab. 47.’ according to the sequence of figure numbers and to the caption on the illustration in Montpellier. We were unable to find any herbarium material matching the illustration in either P or
Solanum torvum var. scabrescens Miq. Fl. Ned. Ind. 2: 648. 1861.
Type. Indonesia. Sumatra: sin. loc., F.W. Junghuhn s.n. (holotype: L [L0403917])
Based on an unpublished illustration of Leschenault collection kept in the Node-Véran collection in Montpellier (lectotype, designated here: Service du Patrimoine Historique de l’Université de Montpellier, Node-Véran, Sol. Tab. 55 [MPU028527]); Indonesia. Java: sin. loc., T. Horsfield s.n. (epitype, designated here:
Shrubs to 3 m, armed. Young stems terete, black to dark brownish, moderately stellate-pubescent, usually densely prickly distally, sometimes unarmed, the stellate trichomes porrect, sessile or variously stalked, the stalks to 0.2 mm long, the rays (4-)5–8, 0.1–0.25 mm long, the midpoints reduced to globular glands; prickles to 3.5 mm long, to 2.5 mm wide at base, straight, awl-shaped to deltate, conical, pale yellow, glabrescent; bark of older stems brownish gray, sparsely stellate-pubescent. Sympodial units difoliate, the leaves geminate. Leaves simple, the blades 11–24 cm long, 4–13 cm wide, ca. 1.5–3 times longer than wide, elliptic to broadly ovate, chartaceous, slightly discolorous; adaxial surface moderately stellate-pubescent with porrect, sessile and less often variously stalked trichomes, the stalks to 0.1 mm long, the rays 4–8, 0.1–0.4 mm long, the midpoints to 0.25 mm long; abaxial surface moderately stellate-pubescent with trichomes like those of the adaxial surface, but more often stalked; prickles 0–6 per leaf side, to 6 mm long, to 1.5 mm wide at base, straight or slightly curved at the tip, awl-shaped, conical, pale yellow, glabrous; major veins 6–8 pairs drying yellow; base shortly attenuate to truncate; margins entire or shallowly to deeply lobed, the lobes 1–5 on each side, 0.5–5 cm long, rounded to apically acute, the sinuses extending up to 2/3 of the distance to the midvein, deltate; apex acute; petiole 1.5–4 cm long, 1/10–1/5 of the leaf blade length, densely stellate-pubescent with porrect, sessile trichomes like those of the blades, with 0–5 prickles like those of the stems. Inflorescences apparently lateral or leaf opposed, 2–5 cm long, unbranched to up to 2 times branched, with ca. 5–20 flowers, moderately to densely stellate-pubescent, unarmed; peduncle 0.5–1.5 cm long, with 0–1 prickles; pedicels 0.5–1.2 cm long, erect, articulated at the base, densely stellate-pubescent, unarmed; pedicel scars spaced 2–4 mm apart. Flowers 5-merous, apparently all perfect. Calyx 4–7 mm long, campanulate, moderately stellate-pubescent, densely stellate-pubescent on the midvein, unarmed, the lobes 3–5 mm long, the lower part deltate and abruptly constricting to an elongate acumen, the acumen 3/4 the total lobe length, the abaxial surface more or less strongly keeled along the midvein. Corolla 1–2 cm in diameter, white, lobed for ca. 1/2–2/3 of the way to the base, the lobes 5–8 mm long, 2–3.5 mm wide, deltate, spreading at anthesis, densely stellate-pubescent abaxially on parts exposed in bud. Stamens equal; filament tube < 0.5 mm long; free portion of the filaments 0.75–1.5 mm long; anthers 5–6.5 mm long, ca. 0.75 mm wide, connivent, tapering, poricidal at the tips, the pores not lengthening to slits with age. Ovary conical, minutely glandular-puberulent; style 0.6–1 cm long, slender, curved at the apex, with few scattered hairs at the tip; stigma capitate, minutely papillate, stellate-pubescent. Fruit a globose berry, 8–18 per infrutescence, 0.8–1.5 cm in diameter, the pericarp smooth, bluish green when young turning to dark greyish yellow, glabrous; fruiting pedicels 1.2–2.5 cm long, ca. 1–1.5 mm in diameter at the base, ca. 2–3 mm in diameter at the apex, woody, erect, unarmed; fruiting calyx lobes not expanding. Seeds 100–200 per berry, ca. 1.75–2 mm long, 1.5–1.75 mm wide, flattened reniform, pale yellowish, the surface minutely pitted, the testal cells sinuate in outline.
Flowering and fruiting throughout the year.
(Fig.
Least Concern (LC); EOO > 100,000 km2 and AOO > 10,000 m2 (see
Indonesia. Central Sulawesi: Banggai regency, Luwuk District, Bunta Subdistrict, Sumber Agung, Gunung Hek, Sungai Hek, Cabang Tiga, 980 m, 27 Feb 2004, Hendrian et al. 964 (E, L); Sigi Regency, near the river S of Tongoa, 650 m, 17 Mar 1981, Johansson et al. 419 (K, L); Java: sin. loc., Horsfield s.n. (
Solanum poka was long ignored after its first publication (
Based on morphology, S. poka belongs to the Torva clade (sensu
In the protologue Dunal referred to an illustration made by Node-Véran, ‘Dun. Suppl. 7. Sol. Mss. tab. 55’, but cited no herbarium material. Similarly to the situation of S. graciliflorum, we were unable to find any herbarium material matching the illustration in either P or
We have only seen three specimens of S. poka from Java, the cited type locality, all collected by Thomas Horsfield, an American physician who collected on Java contemporaneously with Leschenault in the early part of the 19th century (
We thank the curators of A,
List of collections used in the study with full locality and descriptive notes
Data type: List of collections in xls format
Explanation note: Details of the herbarium material examined for this manuscript (including full locality and descriptive notes).