Phaseolus diversifolius Pittier, Bol. Tecn. Minist. Agric.

Phaseolus diversifolius Pittier, Bol. Tecn. Minist. Agric. View in CoL 5: 56 (1944). TYPE: VENEZUELA. Anzoategui, sabanas de Guaraguara, cerca de Santame, H. Pittier 14302 (holotype: not traced), non P. diversifolius Pers., Syn. Pl. View in CoL 2(2): 296 (1807).

Perennial scrambling or climbing vine up to 5 m or more, foliage and reproductive parts covered with long yellow (rarely brown) hairs, and minute glandular hairs intermingled, pubescent or glabrescent. Stems herbaceous, fibrous (sturdy), older stems woody, with nitrogen fixing nodules, often with adventitious roots with nodules, sparsely to densely pilose with yellowish-fulvous, straight hairs. Leaves with stipules ovate-lanceolate, often apiculate, unequally bilobed, upper portion 4–7 3 1–2 mm wide, obscurely veined; lower portion ca. 2 mm long, sometimes adnate to the stem, densely strigose; stipels oblong to triangular, 1–2 mm long, shorter than petiolules, 2-veined, covered with minute glandular trichomes; petioles 4–6 cm long, covered with hirsute, retrorse hairs, rachis considerably shorter, 5–10 mm long, with antrorsely appressed hairs, canaliculate; leaflets entire or lobed, with sinuous margins, oblong to ovate or widely ovate, sometimes rhombic or linear-lanceolate, usually obtuse or acute or even acuminate at apex, often with raised veins below, chartaceous, densely strigose to softly pilose on both surfaces, usually more densely so beneath; terminal leaflet 4–11.5 3 2.5–5 cm, lateral leaflets 3–9.5 3 2–4.5 cm. Inflorescences 10–50 cm long, peduncles up to 45 cm long, covered with straight, retrorse hairs, densely strigose distally; rachis 3–5 cm long, with (3–)5–8(–9) swollen nodes, these oblong, 2–4 mm long, with ca. 10 secretory orifices in two parallel rows, flowers clustered distally; primary bracts caducous, secondary bracts lanceolate, 8–10 mm long, caducous; bracteoles mostly persistent at anthesis, lanceolate, 6–15 mm long; pedicels shorter than calyx tube, 3–5 mm long, longer and twisting in fruit, covered with retrorse, straight hairs, antrorse at base of calyx; calyx campanulate, densely hirsute-strigose, tube longer than teeth, 5.5–10 3 3.5–8 mm, upper teeth deeply divided and broadly rounded at apex forming a lip; 5–6 3 5–6 mm, sinuses 3 mm, lower lip with lateral teeth oblong and rounded, 5–6 3 2–3 mm, and with the middle lower tooth 4–5 3 2 mm. Flowers golden yellow, (1.8–) 2.5–3.5 cm long, standard petal asymmetric, broadly ovate, 2.5–3 3 2.7–3 cm, bilobed at apex, two parallel, thick 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, 2.5–3.5 3 ca. 1.5 cm, with a prominent auricle at base, ca. 2 mm long, claw ca. 4 mm long; keel distinctly beaked, curved through 360, forming a complete coil, ca. 3 cm above the wing, with transverse pockets above the claws, claws 4–5 mm long, fused to staminal tube; androecium 3.5–4 cm long, staminal tube ca. 3 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, interstitium granular; ovary straight, with a basal nectary disk ca. 1 mm long, ovules 16–18, style with a tenuous lower part, upper portion thickened, cylindrical, curved, pollen brush 7–10 mm long, with long spreading hairs, produced beyond the stigma to form a short hook; stigma transversally-ovate, laterally placed. Fruit patent or rarely ascending, linear-oblong, cylindrical, truncate at the apex, valves coriaceous to woody, turning dark brown or black at maturity, densely pilose with a mixture of white and yellow hairs becoming more brownish at the sutures and base, 6–9 cm long, 7–11 mm wide, elastically dehiscent, beak 2–3 mm long, seeds arranged obliquely. Seeds oblong, subglobose, 3–4 3 3–4 mm, surface smooth, testa brown, hilum oblong, ca. 2 mm long, covered by an epihilum, rim-aril raised, with a white, excentric hippocrepiform aril consistently present towards the lens. Chromosome number: 2 n 5 20 ( Marechal 1969, as P. pilosus Kunth ; Mercado-Ruaro and Delgado-Salinas 1996).

Illustrations and Photographs — Bentham (1859) as Phaseolus lasiocarpus , or in http://floraBrasiliensis.cria.org.br; Micheli (1883) as Phaseolus balansae ; Pittier (1944) as Phaseolus lasiocarpus ; Lackey and D’ Arcy (1980) as Vigna lasiocarpa ; Lewis and Owen (1989) photographs of flower and fruit, as V. lasiocarpa ; Pott and Pott (2000) photograph of V. lasiocarpa .

Distribution and Habitat —The species occurs in southern Mexico (Campeche, Chiapas, Oaxaca, Tabasco, Veracruz, and Yucatan); Central America ( Belize, Costa Rica, El Salvador, Honduras, Nicaragua, and Panama); South America ( Colombia, Venezuela, Guyana, French Guiana, Suriname, Bolivia, Brazil, Ecuador, Peru, Paraguay, and Argentina) and in the Dominican Republic ( Fig. 3 View FIG ). Mainly in seasonal or permanently flooded plains or savannas or swampy grasslands, reported growing in sugar plantation ditches and on bank edges of rice fields; mostly from sea level to ca. 600 m, but ascending in Mexico and Colombia to 1200 m. In South America, in the Solim ~ oes and Amazon River basins, and in the Paraguay-Parana fluvial system. An ecological study of aquatic macrophyte diversity in the Pantanal Matogrossense National Park found Vigna lasiocarpa to be one of the most frequent species and distributed among ca. 60% of the 17 sampling sites (Pott et al. 2011). Flowering and fruiting occur throughout the year.

Vernacular Names —“Frijolillo” (H. Vibrans 8246), “Feij ~ aozinho-do-brejo” (Pott and Pott 2000).

Etymology —The species epithet alludes to the hairiness of the fruits (Lasio woolly or hairy and Carpus fruit).

Uses —The species is relished by livestock, and its seeds are eaten by birds (Pott and Pott 2000).

Representative Specimens Examined — See Appendix 1 for complete list. Argentina. — Corrientes: Depto. Santo Tome, Estancia Garruchos, 14 Feb 1960, T. M. Pedersen 5446 ( RBGE, US). Belize. — STANN CREEK DISTRICT: Carib Switch, Stann Creek-Middlesex Road, 13 Jan 1953, P. H. Gentle 7839 ( MEXU). Bolivia. — EL BENI: Trinidad, 236 m, O. Braun 75 ( US). Brazil. — ACRE: Rio Branco, Amazonia, Baixo Rio Branco , 1 Jan 1948, R. de Lemos Froes 23013 ( NY). Colombia. — CAUCA: east of Aganche , 1200–1500 m, 14 Jul 1922, F. W. Pennell 8328 ( NY). Costa Rica. — ALAJUELA: Alfaro Ruiz , entre Tapezco de Arriba y Tapezco de Abajo, 1900 m, 12 Nov 1964, A. Jimenez M. 1145 ( NY). Dominican Republic. — SAN CRISTÓBAL: Station Hatillo , 1 Nov 1977, A. H. Liogier 9073 ( NY). Ecuador. — GUAYAS: Milagro , 50 m, 30 Jun 1923, A. S. Hitchcock 20275 ( NY US). El Salvador. — AHUACHAPÁN: A.P. Santa Rita, ruta 7, La Laguna , 13 48 9 0 99 N, 90 4 9 0 99 W, 20 m, 12 Apr 2004, J. M. Rosales 2355 ( MEXU). Guyana. Demerara-Mahaica Region , east coast Demerara , Cane Grove Conservancy , Flagstaff , 6 37 9 60”N, 57 55 9 60 99 E, 19 Feb 1989, L. J. Gillespie 706 ( US). Honduras. — CORTÉS: en bananales de La Lima, 60 m, Jun 1977, R. H. Stover 4 ( MEXU). Mexico. — CAMPECHE: Municipio Champoton, Aguada Paraıso, 18 55 9 60 99 N, 90 21 9 0 99 W, 12 m, 15 Oct 1981, C. Chan 887 ( XAL). Nicaragua. — MANAGUA: Banco de Germoplasma , Hacienda Experimental de la Universidad Centroamericana, 15 Nov 1984, D. Soza 247 ( MO). Panama. — CHIRIQUÍ: vicinity of San Felix, eastern Chiriquı , 1 Jan 1912, H. Pittier 5225 ( US). Paraguay. — ALTO PARANÁ: in regione fluminis Alto Parana, 1 Oct 1909, K. Fiebrig 6177 ( BM, SI, US). Peru. — LORETO: Mishuyacu, near Iquitos, 100 m, 1 Feb 1930, C. Klug 1022 ( NY, US). Suriname. — DISTRICT NICKERIE: Nickerie, 28 Jun 1951, A. T. Semple 358 ( US). Venezuela. — AMAZONAS: at Cerro Yavita , Rıo Atabapo , Rıo Orinoco ; 19 Oct 1950, B. Maguire 29294A ( NY) .

Notes —Verdcourt (1970) designated Vigna lasiocarpa as the type species of V. sect. Lasiospron and, in addition, gave reasons for selecting the species Phaseolus lasiocarpus opposed to P. pilosus or P. hirsutus . Unfortunately, no lectotype was designated at that time between the two cited syntypes under P. lasiocarpus . Both syntypes were studied by us, and the Martius collection at M has been designated as lectotype. Regarding the original material of P. hirsutus at M, all syntypes were found except the one of Poeppig. The lectotype here designated is the Martius specimen from Rio Negro, which bears a description of this species in Martius’ handwriting. Recently, Cremers and Hoft (1998), following Grisebach (1860), listed Dolichos jacquinii DC. ( De Candolle 1825) in synonymy under V. lasiocarpa ; however, no new combination was proposed by those authors. De Candolle published Dolichos jacquinii for a plant described by Jacquin (1788) from the Caribbean jungles under the epithet D. lignosus L. ( Linnaeus 1753). No type specimen of the binomial has been found at G-DC or at BM-Banks herbarium, and only the species description can be used to interpret the name. Jacquin described a plant with white flowers on peduncles shorter than the leaves, and long straight pods with 18 seeds. Although Vigna lasiocarpa sometimes has pods containing 18 seeds, its inflorescences always exceed the length of the leaves and the species never has been reported to have white flowers. Thus, from its description Dolichos jacquinii is not identifiable as V. lasiocarpa , and the decision here is to exclude this name.

Phaseolus diversifolius Pittier was proposed as a synonym of V. lasiocarpa by Aymard (1999). We have not seen Pittier’ s plant and, thus, cannot comment upon its identity. The protologue of Phaseolus diversifolius is not in Latin and the name has been shown to be a later homonym of P. diversifolius Pers. (Persoon 1807) . Therefore, in accordance with Article 53 of the International Code of Nomenclature for algae, fungi, and plants (Turland et al. 2018), this species was not validly published, and the name is unavailable for use.

Vigna lasiocarpa is the most vigorous and conspicuous of all the species of V. subg. Lasiospron . It is most similar in overall morphology to its sister species V. longifolia . Vigna lasiocarpa differs from V. longifolia in being more often pubescent (especially on the leaves), having longer and wider calyces with the upper teeth more deeply divided and broadly rounded at the apex to form a lip, and having longer and thicker fruits. Both species have distinctive arillate seeds. Vigna lasiocarpa is the only species in the subgenus, up to now, that has been reported as a dysploid.