Vigna trichocarpa, (C. Wright) A. Delgado, Monogr. Syst. Bot.

Delgado-Salinas, Alfonso, Lavin, Matt, Snak, Cristiane & Lewis, Gwilym P., 2022, Systematics of Vigna subgenus Lasiospron (Leguminosae: Papilionoideae: Phaseolinae), Systematic Botany (Basel, Switzerland) 47 (1), pp. 97-124 : 114-117

5. VIGNA TRICHOCARPA (C. Wright) A. Delgado, Monogr. Syst. Bot. Missouri Bot. Gard. 45: 1254 (1993).

Phaseolus trichocarpus C. Wright , in F. A. Sauvalle, An. Acad. Ci. Habana, Rev. Ci. 5: 337 (1869) and in Fl. Cuba: 30 (1873).

TYPE: CUBA. [Pinar del Rıo Province], “ en la orilla de las lagunas dentro del agua. Vuelta Abajo ”, no date, C. Wright s.n. (holotype: GH! ; isotypes: BM!, G-2!, K-3! , MO! , NY- 3! , P! , US! , WU! ). (See notes). Figure 9 .

Phaseolus ovatus Benth. var. glabratus Benth. , in Mart. Flora Brazil. 15(1): 188 (1859). TYPE: BRAZIL. “prope Bahiam, in humidis cum forma pilosa”, W. D. Salzmann s.n. (holotype: M, not seen; isotypes: K!, MO!, RBGE!). non sensu Chodat & Hassler, Bull. Herb. Boiss. Ser. 2, 4: 909. (1904).

Phaseolus schottii var. campestris forma transiens Hassl., Candollea 1: 464. 1923. TYPE: BRAZIL. Bahia in pratis humedis, W. D. Salzmann s.n. (holotype: G-DEL, not seen; isotypes: K!, MO!, RBGE!), synon. nov. (See notes).

Phaseolus lanceolatus Bello, Anales Soc. Esp. Hist. Nat. 10: 262 (1881). TYPE: PUERTO RICO. Without locality, Bello s.n. (holotype: B, destroyed). Neotype designated by Santiago-Valentın et al. Taxon 64: 333, 2015: Puerto Rico, Pueblo Viejo, 19 July 1914, J.A. Stevenson 2097 [NY, barcode 01007063].

Phaseolus schottii var. campestris forma guyanensis Hassl., Candollea 1: 464. 1923. TYPE: FRENCH GUIANA. Without locality, Poiteau s.n. (G, not seen) synon. nov. (See notes).

Phaseolus productus Ducke, Arch. Jard. Bot. Rıo de Janeiro 4: 99 (1925). TYPE: [ BRAZIL. Para State], “in campis inundatis Jutahy inter Almeirim et Prainha inter frutices ad ripam fluminis”, 16 Apr 1923, Ducke s.n. (lectotype: RB!, No. 17.285, designated here).

Vigna paludosa Milne-Redh., Kew Bulletin 2: 27 (1947). TYPE: SIERRA LEONE. Rokupr, common among grass in swamps, 29 Jan 1935, Deighton 2953 (holotype: K!).

Perennials or annuals (?), scrambling or climbing vines up to 3 m, with foliage and reproductive parts covered with minute glandular hairs, and sparse or dense pubescence. Stems hollow, slightly woody at base, often with adventitious roots, sparsely to densely strigose, with yellow, retrorse hairs. Leaves with stipules ovate to narrowly-lanceolate, spurred, upper portion 8–12 ˟ 1–2 mm, 5–6-veined, not reflexed, persistent; lower portion 3–4 mm long, sparsely pilose; stipels oblong or triangular, ca. 1.5 mm long, subequal in length to petiolules, glabrous except for minute glandular trichomes; petioles 3.5–8 cm long, covered with retrorse hairs, rachis considerably shorter, ca. 1 cm long, with some antrorsely appressed hairs on the adaxial side, canaliculate; leaflets entire, ovate, narrowly ovate, to lanceolate, acute or acuminate at apex, with raised veins below, membranaceous; sparsely to densely strigose, terminal leaflet 3.5–11 ˟ 1.5–3 cm, lateral leaflets 7–9 ˟ 2–3.5 cm. Inflorescences up to 22 cm long, peduncles 16–20 cm long, covered with short retrorse hairs, densely strigose distally; rachis 1.5–2 cm long, with 1–4(–5) nodes, the nodes oblong, spatuliform, 2–3 mm long, orifice distribution scalariform, with transverse separations, flowers clustered distally; primary bracts, caducous, secondary bracts ca. 6 mm long, caducous; bracteoles mostly persistent at anthesis, 5–8 mm long, longer than calyx tube; pedicels shorter than calyx tube, 1–2 mm long, longer and twisting in fruit, covered distally with retrorse, straight hairs; c alyx campanulate, sparsely strigose at the base, glabrous distally, 1.5–2 ˟ ca. 2 mm, upper teeth slightly divided, narrow and acute, not forming a lip, teeth triangular, subequal, 0.5 mm long, lower tooth slightly longer than lateral teeth. Flowers golden yellow, 8–12 mm long, standard petal asymmetric, broadly ovate, ca. 10 ˟ ca. 10 mm, bilobed at apex, with two parallel callosities on the lamina above the point of folding, and two fleshy auricles above a short claw; wing petals longer than keel, with an obovate lamina, ca. 1.2 cm long, 5–6 mm wide, with an auricle at base, claw ca. 1.5 mm long; keel distinctly beaked, coiled through ca. 360, almost forming a complete circle; ca. 7 mm above the wing, with transverse pockets above claws, the claws ca. 2 mm long, fused to the staminal tube; androecium ca. 1.5 cm long, vexillary stamen with a basal appendage; anthers oblong-ovate, ca. 1 mm long, basifixed to sub-basifixed to filaments; pollen grains triporate, with a coarsely reticulate exine; ovary straight, with a basal nectary disc ca. 0.5 mm long, ovules 7–8 per ovary, style with a tenuous lower part, upper portion thickened, cylindrical, curved, pollen brush 2–3 mm long, with long spreading hairs, produced beyond the stigma to form a conical appendage; stigma transversally-ovate, sub-apically placed. Fruit ascending, oblong, flattened, valves thin-walled, not constricted between the seeds, turning dark brown or black at maturity, sparsely strigose, with yellow, straight hairs, (3–) 3.5–4 cm long, 6–7(–8) mm wide, beak 2 mm long, straight, elastically dehiscent. Seeds D- shaped, 2–3.5 ˟ 3–4 mm, surface smooth, testa light to dark brown, hilum oblong, as long as seed width, rim-aril distinctly raised, covered by an epihilum, lacking a cartilaginous aril. Seedlings with hypogeal germination, eophylls without stipules. Figure 9 .

Illustrations — Berhaut (1976) as Vigna paludosa Milne-Redh. Maxted et al. (2004) and Acevedo-Rodrıguez (2003) as Vigna longifolia .

Distribution and Habitat —Southern Mexico to Brazil, including the Greater Antilles, and likely native in west Africa ( Fig. 3 ). In general, in seasonal or permanently flooded plains or marshy environments, sometimes reported in coastal vegetation and sprawling in floating plant islands in rivers and lakes of South America, or growing in old rice fields; altitude 0 to 300 m. Flowering and fruiting have been recorded throughout the year, except in June. In South America, in the Solim oes ~ and Amazon River basins and northern Restinga Atlantic Forest and in the Paraguay-Parana fluvial system and the southern Restinga Atlantic Forest.

Etymology —The specific epithet refers to the plant’ s hairy fruits (Trichos hairs and Carpos fruit).

Vernacular Names —“Feij ~ aozinho amarelo” ( Brazil). “Habichuela cimarrona” ( Puerto Rico).

Representative Specimens Examined — See Appendix 1 for complete list. Belize. — STANN CREEK DISTRICT: ca. one mile, WSW of Hopkins, 17 Apr 1976, G. R. Proctor 35795 ( IJ, MO). Bolivia. — SANTA CRUZ: Velasco Province, Reserva Ecologica El Refugio, 14 45 ' 47 '' S, 61 52 ' 51 '' W, 100 m, 15 Oct 1994, T. J. Killeen 6839 ( MEXU). Brazil. — AMAZONAS: District Careiro, Lago Redondo, 3 Jan 1964, G. Marlier 14375 ( US). Colombia. — CHOCÓ: Rıo Atrato , 2–5 hr below Rıo Sucio , above Loma Teguerre, 16 May 1967, J. A. Duke 10988 ( MO, NY). Costa Rica. — CARTAGO: Instituto Interamericano de Ciencias Agrıcolas , 609 m, 1 Feb 1959, A. T. Semple 1 ( US). Dominican Republic. — SAMANÁ: Sanchez District , seccion La Majagua, Yaqueson (Jackson), entre cano ~ La Bestia y cano ~ Punta Arena, 19 16 ' 0 '' N, 69 31 ' 60 '' W, 30 Mar 1996, B. Peguero 134 ( MEXU). French Guiana. 10 km west of Mana , Savane de Criques Jacques , near St. Laurent , 19 Dec 1954, R. S. Cowan 38884 ( NY, US); St. Laurent do Maroni, 24 Feb 1914, R. Benoist 75 (P). Guatemala. — PETÉN: Lake Zotz, 18 May 1933, C. L. Lundell 3299 ( MICH). Guyana. Siparuni-Potaro Region , Essequibo R., near Iwokrama Rainforest Reserve , 4 16 ' 60”N, 58 30 ' 0 '' W, 65 m, 20 Mar 1996, D. Clarke 1356 ( NY). Honduras. — CORTÉS: Agua Azul tract No. B. North shore of Lake Yojoa near the canyon, 16 Aug 1951, P. Kamb 2092 ( BM). Jamaica. — SLIPE DISTRICT: without locality, 22 Jul 1973, G. R. Proctor 33461 ( IJ). Mexico. — TABASCO: ejido San Ramon, campo petrolero San Ramon, 17 Jun 1996, G. Ortız 2051 ( MEXU). Nicaragua. — RÍO SAN JUAN: delta of Rıo San Juan , 23 Mar 1961, G. S. Bunting 829 (F, US). Panama. — CANAL ZONE: bridge over inlet of Limon Bay , S of Ft. Sherman, 22 Aug 1960, J. E. Ebinger 975 ( MO). Peru. — LORETO: Province Maynas, Urco-Cocha, comunidad de Vargea, 10 Aug 1998, A. Zamora U. 11 ( HUT). Puerto Rico. Guaynabo, 3 Jul 1924, H. H. Whetzel 1 ( BH); Pueblo Viejo, no date, J. A. Stevenson 2097 ( NY). Suriname. — COMMEWIJNE DISTRICT: Charlottenburg, no date Wullschlaegel s.n. ( W). Senegal. — REGIONE DE SÉDHIOU: Ile du Diable , Silinki , Sorange , Forest de Bondie , 2 Mar 1964, R. P. Berhaut 7165 (M-2). Sierra Leone. Madina (Buya-Romende), 11 Aug 1953 H. D. Jordan 907 (K).

Notes — Howard (1988), using the sparse information available for Wright’ s collections in Cuba , listed among nine collections of Phaseolus , two collections named as P. trichocarpus : C. Wright s.n., and C. Wright 2341. The latter was collected in San Mateo (Pinar del Rıo), whereas the former lacking a collection number was given no exact collection locality. Furthermore, Howard considered as the holotype the specimen at GH of Wright 2341, although Wright’ s protologue mentions a plant listed by Grisebach as P. ovatus non Benth., “growing in the water, next to the shore of the lagoons in Vuelta Abajo.” Vuelta Abajo is also located in the Cuban province of Pinar del Rıo, and was several times visited by Wright in 1862 (21, 22 July, 4 August, 4 November). Delgado-Salinas (1993) transferred Phaseolus trichocarpus C. Wright to Vigna , and designated as lectotype the same collection C. Wright 2341, pointing out that the holotype might possibly be found in HAB. Numerous duplicates of C. Wright 2341 as well of C. Wright s.n. were distributed to different herbaria (BM, G-2, K-3, MO, NY-3, P, US, WU). In some herbaria (e.g. MO), both collections were mounted on the same sheet. The specimen at US even has a different collection locality: La Habana, Playa Santa Ana, with the collection year given as 1860, whereas duplicates at NY, US, and WU show 1865 as the year of collection.

Bello’ s type material of Phaseolus lanceolatus is no longer extant at B. With no original material, Bello’ s description, particularly of the stipules and the collecting locality ( Puerto Rico), leaves little doubt as to the plant’ s identity. A neotype was designated by Santiago-Valentın et al. (2015).

As mentioned earlier, we did not locate the type specimen of P. schottii var. campestris forma guyanensis and, therefore, we have accepted Amshoff’ s (1939) taxonomic decision in placing it under Phaseolus trichocarpus (= Vigna trichocarpa ). Regarding the forma transiens of Hassler, Salzmann’ s collection from Bahia, Brazil “in humidis” with Bentham’ s handwriting on the specimen was designated as lectotype. It is important to mention that different collections of Phaseolus were made by Salzmann in Bahia and those have been considered in different publications under different names: Bahia in humidis “ Phaseolus luteus , ” 1831, Salzmann s.n. (RBGE); Bahia, in collibus humidis, Phaseolus luteus, Salzmann s.n. (MO), and also Bahia in humidis, 1830, Salzmann 181 (G); a mixed collection at G of Salzmann 182 / Salzmann 183 (MO), are all here identified as Vigna trichocarpa . In addition, Bahia “ hirsutis, ” Salzmann s.n. (P), and Bahia “ Phaseolus luteus ” Salzm. , foliis hirsutis, Salzmann s.n. (P) are also assignable to Vigna trichocarpa , and it is possible that at least some of these specimens (especially those without a number) are from the same gathering.

Vigna trichocarpa is by far the most widely distributed and morphologically distinctive species of the subgenus, especially in fruit. The ascending fruits that parallel the inflorescence axis characterize V. trichocarpa . In addition, the seeds of V. trichocarpa hang from longer funicles relative to the shorter funicles of other species of V. subg. Lasiospron . The stipules of Vigna trichocarpa are distinctive in having the lower portion forming a spur with entire margins. Such a stipule is found otherwise among V. subg. Lasiospron species in V. schottii .

Figures

FIG. 9. Illustration of Vigna trichocarpa. A. Habit, natural size. A1. Portion of petiole. A2. Portion of stem. A3. Leaf showing narrow leaflets. B. Stipules (˟ 3). C. Variation in leaflet shape outlines (˟ 1). D. Extrafloral nectaries on inflorescence node (˟ 6). E. Flower bud, including bracteoles (˟ 3). F. Calyx opened out, external surface (˟ 6). G. Standard petal outer surface (˟ 3). H. Wing petal (˟ 6). H1. Wing petal from bud (˟ 3). I. Keel petals (˟ 6). J. Staminal tube with free stamen (˟ 6). K. Gynoecium (˟ 6). L. Fruits, natural size. M. Seed, hilar view (˟ 9). Drawn from Jansen Jacobs 4681 (habit and flower); Berg et al. 19796 (fruit and seed); Prance et al. 8048; Berg et al. 19796; Glaziou 9392 (leaflet outlines). Illustration by Pat Halliday.

FIG. 3. Geographic distributions of the six species of Vigna subg. Lasiospron, which are ordered, top to bottom, by mean latitude. Filled circles indicate species with amphi-Atlantic distributions.