Solanum pentaphyllum, Bitter, Repert. (Bitter)

Tepe, EJ & Bohs, L, 2011, A revision of Solanum section Herpystichum, Systematic Botany 36 (4), pp. 1068-1087 : 1082-1083

8. SOLANUM PENTAPHYLLUM Bitter,

Repert. Spec. Nov. Regni Veg.12: 70. 1913. Bassoviapentaphylla (Bitter) Pittier, Cat. Fl. Venez. 2: 356. 1947.

—TYPE: COLOMBIA. Tolima: Ibagué, 1845 (fr), J. Goudot s. n. (holotype: G–G00104280 [scan!]; photos of holotype [Morton neg. 8546]: F–1589922!, MO– 1781232!, NY!; isotype: G–G00096093 [scan!]; photos of isotype [F neg. 23140]: F–758058!, MO–1691340!, NY!).

Solanum pentaphyllum Bitter var. caraboboanum Bitter, Repert. Spec. Nov. Regni Veg. 12: 70. 1913. Bassoviapentaphylla (Bitter) Pittiervar. caraboboana (Bitter)Pittier, Cat.Fl.Venez. 2: 356. 1947.

— TYPE. VENEZUELA. Carabobo: Valencia, Oct 1843 (fl), N. Funck 791 (holotype: G–G00104281 [scan!]; possible isotypes: BR–BR00000988419!, P–P00578798 [scan!]).

Herbaceous vine, terrestrial, trailing over ground or fallen logs. Stems slender, herbaceous, sparsely to densely pubescent, the trichomes 1–2 mm long. Sympodial units plurifoliate. Leaves 5-pinnate, the blades 1.2–7 × 1.5–8 cm, about as long as wide, membranaceous to chartaceous, sparsely sandpunctate, sparsely pubescent adaxially with widely scattered, short, wide-diameter trichomes 0.3–1.5 × 0.1–0.2 mm, with thinner trichomes occasionally present on midvein adaxially, glabrous to pubescent only on the veins abaxially, the margins entire, the rachis sparsely to densely pubescent with hairs like those of the stem, the pubescence especially dense in the adaxial groove; lateral leaflets (the most distal pair) 0.8–4 × 0.4–2 cm, oblique, elliptical on lower side, obovate on upper side giving the leaflets a falcate appearance, the base oblique, acute to obtuse to slightly cordate on the proximal side, the apex obtuse to rounded to minutely acuminate, the basal pair slightly smaller than the upper pair, the upper pair more strongly oblique, the petiolules 0.5–2 mm on upper pair of leaflets, 1.5–6 mm on lower pair, glabrous to densely pubescent, especially in the adaxial groove; apical leaflet 0.8–4.5 × 0.6–2.4 cm, obovate, rhomboid, the base acute to cuneate, the apex obtuse to shortly acuminate, sessile to shortly petiolulate, the petiolule 0.5–2.5 mm, glabrous to densely pubescent, especially along adaxial groove; petioles 1.5–5.5 cm, sparsely sand-punctate, sparsely to densely pubescent, especially in adaxial groove. Internodes 2.5–9 cm. Inflorescences 3–7 cm long, unbranched, extra-axillary, with 1–3 flowers, the axes sparsely pubescent; peduncle 2–6 cm, slender; rachis 0.5–1 cm; pedicels 5–10 mm in flower, 15–22 mm in fruit, slender, sparsely pubescent, spaced 2–10 mm apart. Calyx 1.5–2 mm long, the tube 0.5–1 mm long, the lobes 0.5–0.8 × 1–1.2 mm, deltoid, acute to acuminate and slightly thickened at tips, sparselypubescentwithshorttrichomes (<0.1 mm), themargins ciliate; fruiting calyx slightly accrescent, the lobes 1.5–3 × ca. 1.5–1.8 mm. Corolla 1–1.5 cmindiameter, ca. 0.5 mm long, stellate, membranous, white, the tube 1–3 mm, the lobes 3–5 × 1.5–2 mm, lanceolate, acute at tips, glabrous abaxially and adaxially, the margins ciliate. Stamens with filaments 0.6– 1 mm, glabrous; anthers 2–2.5 × 0.8–1.2 mm. Ovary glabrous; style 3–5 × ca. 0.2 mm, clavate, papillose in lower half; stigma capitate. Fruits 2.2–3 × 1–2.5 cm, ovoid-rhomboid, flattened, the apex acute, green to purplish brown, glabrous. Seeds 2.2– 2.5 × 1.8–2 mm, ca. 0.5 mm thick, lenticular, orangish tan in color, the surface sparsely granular. Figure 2A .

Habitat and Distribution— Solanum pentaphyllum occurs in Colombia and Venezuela. Two collections are also known from Costa Rica. It is usually terrestrial, but occasionally grows over fallen logs; 600–1,700 m in elevation ( Fig. 8 ).

Phenology— Flowering specimens have been collected year round; fruiting specimens have been collected from Feb. to Apr., and in Aug.

Etymology— This species takes its name, pentaphyllum , from its 5-foliate compound leaves.

Notes— Solanum pentaphyllum is one of the ground-trailing species, and can easily be differentiated from the others based on its 5-foliate leaves. The leaves from several populations in Antioquia, Colombia are smaller than the average and are reddish below. The two collections known from Costa Rica are indistinguishable from South American collections. We obtained trnT–trnF and trnS–trnG sequences from one of the Costa Rican collections (Haber & Zuchowski 9863, NY), and it was well supported as sister to the South American collection of S. pentaphyllum rather than to Costa Rican S. phaseoloides ( Fig. 4 ). Based on available collections, S. pentaphyllum is apparently absent from Panama, thus the Costa Rican collections represent a significant disjunction from the species’ primary range in Colombia and Venezuela. The labels of two collections state that the deep green to purplish-brown fruits dig themselves into the ground during development (Pittier 8031, GH; Tamayo 2224, G). From these labels, it is unclear whether the fruits are actively pressed in to the ground by the plant and are truly geocarpic (this seems unlikely because the infructescences of this species are thin) or if they become buried during development as they lie on the ground. This intriguing phenomenon merits further research.

The protologue of S. pentaphyllum var. caraboboanum states that this variety differs from variety pentaphyllum in that it is smaller and less pubescent than var. pentaphyllum , and has a distribution in northern Venezuela. These morphological characters hold true for some specimens, but the same variation can be found among specimens throughout the range of the species. We were unable to identify any consistent morphological character to reliably identify variety caraboboanum , and could not separate the varieties unless their provenance was known. As a result, S. pentaphyllum var. caraboboanum is not recognized in this treatment.

The holotype of S. pentaphyllum at Gis clearly identifiable because the locality information on Goudot’s label matches the protologue. An isotype is also housed at G. Several of the photos of this specimen indicate Brazil as the country of origin; however, the locality on Goudot’s label, Ibagué, is unequivocally Colombian. Furthermore, Goudot lived in Colombia and most of his collections are from there ( Palmer 1918). He also collected in northern Venezuela from the upper Orinoco to Puerto Cabello on the Caribbean coast, but he does not seem to have collected in Brazil.

The holotype of S. pentaphyllum var. caraboboanum is also at G. Possible isotypes, held at BR and P, are also labeled Funck 791, but the localities of these two specimens differ from the holotype.

Additional Specimens Examined— COSTA RICA. Alajuela: Cantón de San Ramón, Cordillera de Tilarán, Bajo Los Rodríguez, Río La Esperanza, Quebrada Mirasol, Finca Araya, 10°18′30″N 84°35′0″W, 500–600 m, 10 Mar 1993 (fl), Bello et al. 4862 ( INB); Reserva Biológica Monteverde, Finca Villalobos, 8 km Sof Volcán Arenal, 10°23’N 84°43’W, 1,000 –1,200 m, 21 Apr 1990 (fl), W. Haber & W. Zuckowski 9863 (INB, NY).

COLOMBIA. Antioquia: Frontino, Corregimiento Nutibara, Región Murí, camino hacia La Blanquita, 6°45’N 76°25’W, 1,440 m, 10 Jul 1986 (fl), P. Acevedo-Rodríguez et al. 1203 ( NY); Frontino, Corregimiento La Blanquita, región de Murrí, vía Nutibarra-La Blanquita, 6°45’N 76°25’W, 1,350 –1,450 m, 10 Jun 1988 (fl) R. Callejas Posada et al. 6525 (MO, NY); Frontino, Region of Murrí, road between Nutibara and La Blanquita, 06°40’N 76°26’W, 1,460 m, 10 Feb 1989 (fl, fr), J. M. MacDougal et al. 3876 ( NY); Parque Nacional Natural Las Orquideas, Sector de Calles, Río Calles, 6°31’N 76°19’W, 1,300 –1,320 m, 29 Mar 1991 (fl), J. G. Ramírez & E. Muñoz 4062 ( MO). Chocó: Slopes of Serranía del Darién Eof Ungía, 8°01’N 77°05’W, 300–1,300 m, 19 Jun 1976 (st), A. Gentry et al. 16770 ( MO). El Valle:Río Dagua Valley, La Margarita, 3°30’N 75°52’W, 760 m, 4 Apr 1939 (fl, fr), E. P. Killip 34895 (COL, F, GH, NY, US). Los Llanos: Intendencia, El Meta, near Villavicencio, 3°16’N 73°05’W, 600 m, 20 Jan 1939 (fl), O. Haught 2551 (COL, F, MA, US). Meta: Trayecto desde la Vereda Aguas Claras (escuela) hasta el puente colgante del Río Ariari, 760–800 m, 25 Oct 1995 (fl), J. L. Fernández 12883 ( COL); Cubarral, Vda. Aguas Claras alrededores de la escuela Aguas Claras, 3°47′42”N 73°54′37″W, 855 m, 19 Nov 1995 (fl), M. E. Morales 590 ( COL).

VENEZUELA. Aragua: Colonia Tovar, between Maracai and Chorona, 10°24’N 67°17’W, 5,500 ft, 1856 (fl, fr), A. Fendler 2092 (GH, GOET); between Portachuelo and Ocumare, 10°34’N 69°23’W, 20 Jan 1924 (fl), H. Pittier 11379 (G, NY, US); Toma de Rancho Grande, 10°27’N 68°0’W (fl), H. Pittier 15311 ( US); Alto de Choroni, 10°29’N 67°36’W, 1,600 m, 16 Apr 1967 (fl), B. Trujillo 7632 ( WIS). Barinas: Bolivar, Altimira, La Gallineta Caserío El Celoso near feldspar mine, 8°50’N 70°35’W, 1,500 –1,700 m, 6 Jun 1988 (fr), L. J. Dorr et al. 5444 (MO, NY); Bolivar, near feldspar mine, between La Soledad and Santo Domingo, 8°51’N 71°35’W, 1,300 m, 24 Nov 1984 (fl, fr), H. Van der Werff & F. Ortega 6142 (MO, NY). Carabobo: Montagne La Soledad, 2,000 ft, Dec 1840 (?) (st), J. Linden 1599 (G); Hacienda de Cura, near San Joaquín, 10°06’N 67°46’W, 1,300 m, 15 Aug 1918 (fl, fr), H. Pittier 8031 (GH, US); Soto dela selva pluvial de Borburrata, 10°21’N 68°03’W, 600 m, Feb 1942 (fl, fr), F. Tamayo 2224 (G, US). Distrito Capital: Along old road between Portachuelo and Peñita (Petaquire) and Carayaca, between Colonia Tovar-Junquito road and Hacienda El Limon, 10°28’N 67°11’W, 25 May 1963 (fl), J. A. Steyermark 91446 (F, NY, US, VEN). Lara: Jiménez, Between Alto del Viento to Cerro Pando, 09°39’N 69°34’W, 1,400 m, 26 Oct 1982 (fl), G. Davidse & A. C. González 21180 ( NY); Iribarren, Vecinidades de la Laguna Negra, Loma de Los Naranjos, Montaña de Macanillal y Fila de San Estaban, 9°52’N 69°18’W, 1,300 –1,500 m, 24 Mar 1975 (infl), J. A. Steyermark et al. 111628 ( VEN); Jiménez, Montaña Oscura, en La Briza, 9°52’N 69°21’W, 1,630 m, 8 Aug 1970 (fl, fr) J. A. Steyermark et al. 103578 (M, US, VEN). Trujillo: 13 km ESE of Bocono, 1 km Wof Guarameca, 9°11’N 70°09’W, 1,600 m, 16 Mar 1982 (infl), R. L. Liesner et al. 12907 (MO, NY). Yaracuy: El Amparo camino abierto hasta la fila, 8°21’N 71°39’W, 1,300 m, 19 Mar 1973 (fl), E. Diederichs 108 ( MO); Cerro la Chapa, 10°12’N 68°24’W, 1,250 m, 12 Apr 1999 (fl), J. R. Grant et al. 99–03335 ( US); Nirgua, Serranía Santa María - Cerro la Chapa, 10°12’N 68°35’W, 1,200 –1,350 m, 31 Apr 1994 (fl), W. Meier et al. 3919 ( MO).

Figures

FIG.2. Flowers, fruits, and habit of Solanum sect.Herpystichum. A. Solanum pentaphyllum, habit and flower (from O. Haught 2551, F). B. S. dalibardiforme, habit and flower (from K. von Sneidern 3121, US). C. S. dolichorhachis, habit, flower, and immature fruit (from F. H. Eggers 14641, A).

FIG.8. Distribution of Solanum pacificum and S. pentaphyllum.

FIG.4. Summary phylogeny of Solanum sect. Herpystichum based on based on plastid psbA-trnH, trnT-trnF, and trnS-trnG, and nuclear waxy, ITS, cos1C, cos5, cos9B, cos10B, and cos11 sequences (E. J. Tepe et al. 2011). For clarity, replicate accessions of all species except for S. evolvulifolium and S. pentaphyllum have been omitted. Provenance is indicated for species with multiple accessions, species with compound leaves are underlined, and the asterisk indicates an especially robust specimen of S. evolvulifolium. The topology is based on Bayesian analyses, and branch support values are Bayesian posterior probabilities/maximum parsimony bootstrap.