Rhynchosia edulis Griseb., Abh. Bohm. Ges. Wiss. Goet.

Bezerra, Luísa Maria De Paula Alves, Cândido, Elisa Silva, Vargas, Wanderleia De, Torke, Benjamin Marland, Lewis, Gwilym Peter & Perez, Ana Paula Fortuna, 2024, Taxonomic revision of Rhynchosia Lour. (Leguminosae, Papilionoideae, Phaseoleae) in South America, Phytotaxa 643 (1), pp. 1-67 : 26-27

9. Rhynchosia edulis Griseb., Abh. Bohm. Ges. Wiss. Goet. 19: 123. 1874.

Eriosema edule (Griseb.) Burkart. Darwiniana. 6(2): 261. 1943. Type:— ARGENTINA, “ In Flusssande des Río Primero bei Córdoba, Avarillo del Campo”, November 1870 (fl./fr.), P. G. Lorentz 351 (Lectotype, designated here: GOET 008904 [digital image!]; isolectotype: SI 002515 [digital image!]) . Iconography: Rodriguésia 70: 9. 2019. ( Fig. 6 D–G )

Rhynchosia melanosticta Griseb. Abh. Königl. Ges. Wiss. Göttingen. 19: 124. 1874. Dolicholus melanostictus (Griseb.) Kuntze. Revis. Gen. Pl. 3(3): 62. 1898. Type: ARGENTINA, “ Tucuman, in fruticetis montium pr. Siambon”, s.d., Lorenz s.n. (Holotype: GOET 008903 [digital image!]).

Eriosema volubile Micheli. Mém. Soc. Phys. Genève. 28(7): 36. 1883. Type : PARAGUAY, “L’Assomption sur le bord des chemins”, May 1874 (fl./fr.), Balansa 1553 (Lectotype, designated inadvertently [as holotype] by Grear [1978]: G 00379283 [digital image!]; isolectotypes: F 911437 [fragment] [digital image!], P 02748529 [digital image!]).

Eriosema volubile fo. pubedensiore (Micheli) Chodat & Hassl. Bull. Herb. Boissier. 2(4): 905. 1904. Type: PARAGUAY, “in dumeto pr. Velenzuela”, 1900 (fl.) Hassler 6951 (Lectotype, first-step designated by Grear [1978]; second-step designated here: G 00379282 [digital image!]; isolectotypes: BM 000538488 [digital image!], G 00379281 [digital image!], NY 00007877 [!]) .

Rhynchosia nigropunctata S. Watson. Proc. Amer. Acad. Arts 22: 408. 1887. Dolicholus nigropunctatus (S. Watson) Rose. Contr. U.S. Natl. Herb. 10(3): 101. 1906. Type: MÉXICO, “ Jalisco, Tequila”, 1886 (fl./fr.), E. Palmer 402 (Lectotype, designated inadvertently [as holotype] by Grear [1978]: GH 00063808 [digital image!]; isolectotypes: US 00004690 [digital image!]).

Rhynchosia rupicola Brandegee. Zoë. 5(10B): 202–203. 1905. Type: MÉXICO, Sinaloa, Cerro Colorado , November 1902 (fl./fr.), Brandegee s.n. (Lectotype, designated inadvertently [as holotype] by Grear [1978]: UC 83531 [digital image!]; isolectotype: GH 00063816 [digital image!]).

Dolicholus apoloensis Rusby. Bull. New York Bot. Gard. 6(22): 515–516. 1910. Rhynchosia apoloensis (Rusby) J.F. Macbr. Publ. Field Mus. Nat. Hist., Bot. Ser. 4(4): 91. 1925. Type : BOLIVIA, Apolo, February 1902, (fl.) Williams 97 (Lectotype, designated inadvertently [as holotype] by Grear [1978]: NY 00007793 [!]; isolectotypes: BM 000931749 [digital image!]).

Dolicholus ixodes Standl. Contr. U.S. Natl. Herb. 18(3): 107–108. 1916. Rhynchosia ixodes (Standl.) Standl. Publ. Field Mus. Nat. Hist. Bot. Ser. 4(8): 214. 1929. Type : PANAMÁ, Vicinity of Penomé, 22.III.1908, (fl.) R. S. Williams 119 (Holotype: US 00004680 digital image; isotype: NY 00007784[!]).

Eriosema nigropunctatum Brandegee. Univ. Calif. Publ. Bot. 7(10): 327. 1920. Type : MÉXICO, Vera Cruz, “Palmilla near Zacuapan”, IX.1919, (fl./fr.) C. A. Purpus 8386 (Holotype: UC 200794 [digital image!]; isotypes: GH 00066302 [digital image!], MO 859773 [digital image!], NY 00007842 [!]).

Rhynchosia rariflora Standl. Publ. Field Mus. Nat. Hist., Bot. Ser. 17(3): 264–265. 1937. Type : México, Guaseremos, Río Mayo “valley margins”, September 1936 (fl./fr.), H. S. Gentry 2883 (Holotype: F 863596 [digital image!]; isotypes: ARIZ 04033 [digital image!]; GH00063815 [digital image!], MEXU 00246959 [digital image!], MO 125148 [digital image!], S 6-5331 [digital image!], US 00004825 [digital image!]).

Rhynchosia pinetorum Standl. Publ. Field Mus. Nat. Hist., Bot. Ser. 17(4): 370. 1938. Type: HONDURAS, Siquatepeque , Dpto. Comayagua, July 1936 (fl./fr.), T. G. Yuncker et al. 5791 (Holotype: F 857675 [digital image!]; isotypes: GH00063810 [digital image!], K 000082211 [digital image!], MO 1116261 [digital image!], NY 00026930 [!]).

Vine herbaceous, prostrate to twining, stems branched, pilose to pubescent, indumentum white to yellow, with yellow vesicular glands and bulbous-based trichomes. Stipules 2–8.3 × 0.5–1.5 mm, free, persistent, lanceolate to triangular, externally pilose to pubescent with brown to black vesicular glands and bulbous-based trichomes. Leaves trifoliolate; petiole 1–5.2 cm long; leaf rachis 0.5–2.2 cm long; stipels persistent; leaflets 1.2–6.8 × 1–4.4 cm, ovate, deltoid, or rhombic, surface not bullate, acute to attenuate at apex, truncate to rounded at base, margin entire, pilose to pubescent, with brown to black vesicular glands on the abaxial surfaces. Inflorescences axillary, paniculate, 7–21 cm long, many-flowered, exceeding the length of the subtending leaf; bracts caducous; pedicel 0.5–4 mm long. Flowers 5–8 mm long excluding the pedicel; calyx 3–5 mm long, pubescent with brown to black vesicular glands and bulbous-based trichomes, lobes lanceolate, not exceeding the corolla in length; standard petal 5.7–9 × 4–7.5 mm, oblong to obovate, rounded at apex, pilose to pubescent, with brown to black vesicular glands at the apex and bulbous-based trichomes, claw 0.5–1.5 mm long, auricle ca. 0.5 mm long; wing petals 5–9 × 1–1.5 mm, oblong, claw 0.5–2 mm long, auricle 0.5–1 mm long; keel petals 4.5–10 × 1.5–3 mm, falcate, claw 1–2.5 mm long; stamens 6–10 mm long; ovary 2.9–3.4 mm long, pubescent, with yellow vesicular glands. Fruits 1.5–2.5 × 0.5–0.7 cm, brown, without constrictions between the seeds, oblong to narrowly obovate in outline, pilose, with yellow to black vesicular glands and bulbous-based trichomes. Seeds 3–4 × 3–4 mm, suborbicular, unicolored, brown to black.

Nomenclatural and taxonomic notes: — Grisebach (1874) described the species R. edulis and R. melanosticta in Plantae Lorentzianae.According to their protologues, these species differ by characters related to habit and inflorescence type, R. edulis being an herb, with erect stems and peduncles equaling the petioles, and R. melanosticta with twining stems and peduncles twice the petiole length. When analysing the type material of both species, we conclude that there is no morphological justification for maintaining the two names as separate species. We, therefore, follow Grear’s (1978) decision in placing R. melanosticta as a synonym of R. edulis , since the name R. edulis was published one page earlier in the original publication.

In the protologue of R. edulis, Grisebach (1874) mentioned a collection from “Cordoba, in arenosis ad fl. Rio Primiero”. Grear (1978) erroneously cited as the holotype the collection Lorentz & Hieronymus 1202 from Salta, Argentina, deposited in B. The collection that best matches the description of the type location given in the protologue is Lorentz 351. There are two duplicates of this collection, one of them deposited in GOET and the other in SI. As the author did not specify a particular specimen as type, we designate here the specimen in GOET (GOET 008904) as the lectotype of R. edulis (see Turland et al. 2018, Art. 9.11), since it contains most of the types of Grisebach.

In the protologue of Eriosema volubile fo. pubedensiore the authors cited two collections: “ Valenzuela, Jan. n. 6951” and “in dumeto pr. Bellavista Apa, Dec., n. 8068”. Grear (1978) unintentionally lectotypified the collection Hassler 6951 deposited in G when he cited it as the holotype. However, there are two duplicates of the collection in G (G 00379282 and G 00379281) and Grear (1978) did not specify a particular specimen as type. His type designation should be treated as a first-step lectotypification and we designate here the specimen with barcode number G 00379282 as the required second-step lectotype of E. volubile fo. pubedensiore (see Turland et al., 2018, Art. 9.10, Ex. 11; Prado et al. 2020).

Rhynchosia edulis has an extensive geographical range and occurs in many different habitats. It also displays extensive variation in vegetative characters, which explains the assignment of different names to the taxon by different taxonomists ( Grear 1978). The species resembles R. minima vegetatively and in flower characters ( Fig. 6 G ); their habit and leaflet shape are quite similar ( Fig. 6 D ), and they both have a wide distribution in South America. However, R. edulis is characterized by having bulbous-based trichomes all over the plant, and especially abundant on the fruit ( Fig. 6 E–F ) (vs. absent in R. minima ), dark vesicular glands only on the abaxial surface of the leaflets (vs. on both surfaces) and inflorescences paniculate (vs. racemose).

Reproductive phenology: —The species flowers throughout the year; and fruits from January to June and August to December.

Distribution and habitat: — Rhynchosia edulis occurs in the southern United States, Mexico, some countries of Central America ( Costa Rica, El Salvador, Honduras, Nicaragua) and in South America. It has a wide distribution in South America, occurring in Argentina, Bolivia, Brazil, Colombia, Paraguay, Peru and Venezuela ( Fig. 7 ). It can be found in pastures, grasslands, disturbed areas, along roadsides, in cerrado vegetation and forest edges.

Conservation status: —Least Concern (LC). Rhynchosia edulis is one of the most widespread species of Rhynchosia in South America. It is classified as LC based on its vast EOO of 21,818,239 km 2 (GeoCAT, accessed in May 2022).

Selected Specimens Examined:— ARGENTINA. Cordoba: Dep. Capital, Villa Warcalde, recta Martinoli, antes de La Carolina , 11 February 1960 (fr.), L. A. Espinar 726 ( NY) ; ibid., Villa Belgrano , 28 April 1963 (fl./fr.), L. A. Espinar 1624 ( NY) . Dep. Colón, Sierra Chica, Casa Bamba , 27 November 1898 (fl./fr.), F. Kurtz 10297 ( MBM) . Corrientes: Dep. Santo Tomé, Ruta 38, 26 km NW de Ruta 14, 1 December 1981 (fl.), S. G. Tressens et al. 1394 ( MBM) . Formosa: Dep. Pilcomayo, Sol de Mayo , 14 km SW de Laguna Blanca, 25 October 1995 (fl./fr.), A. Schnini & G. Cuadrado 30331 ( NY) . Misiones: Candelaria, Santa Ana , 14 November 1945 (fl.), J. E. Montes 1457 ( NY) . BOLIVIA. Beni: Ballivian , Carmen Florida, Río Beni, 7 km upstream from Rurrenabaque, Tacana Indian Village , 16 September 1989 (fl.), D. E. Williams 998 ( NY) ; Itenez, Magdalena , 11 April 1979 (fl.), A. Krapovickas & A. Schinini 34801 ( NY) . Cochabamba: Narciso Campero Leyes, Comunidad Sauce Phujyu , 22 February November (fl./fr.), R. G. Pardo 18 ( NY) ; Santibanez, 6 m NE of El Convento , 13 March 2003 (fl./fr.), L. Rico et al. 1561 ( NY) . La Paz: Sud Yungas, Chicaloma , 06 February 1996 (fl.), Beck St. 22592 ( NY) . Santa Cruz: Andres Ibañes, alrededores de Tierras Nuevas–El Palmar , 8 km al SW de la Ciudad de Santa Cruz de la Sierra, 13 November 1994 (fl./fr.), I. G. Vargas 3648 ( NY) ; Serranía del Mutún , 09 March 2008 (fl.), D. Villarroel et al. 2033 ( UB) ; Santiesteban , 12.2 km N of Mineros, 20 February 1994 (fl./fr.), M. Nee 45080 ( NY) ; Velasco , barbecho al borde del camino, 1 km al S de San Ignacio, 4 April 1986 (fl.), R. Seidel 165 ( NY) ; Warnes, Santa Rita del Monte, from La Angostura to Bermejo , 24 February 2003 (fl./fr.), L. Rico et al. 1363 ( NY) . Tarija: Cercado, cerca Vivero CODETAR/Coimata, 19 December 1985 (fl.), E. Bastian 189 ( NY) . BRAZIL. Bahia: Barra da Estiva, Side road ca. 2km. from Estiva, about 12 km. N. of Senhor do Bonfim on the BA 130 to Juazeiro, 27 February 1974 (fl./fr.), R. M. Harley et al. 16514 ( NY) ; Lauro de Freitas, Caji , em frente ao supermar, 15 July 1997 (fl.), R. Soeiro s/nº ( RB) . Ceará: S. loc., s.d. (fl./fr.), F. Alemão & M. Cysneiros 408a ( R) . Espírito Santo: Vitória, Vila Velha , 6 March 1946 (fl.), Brade et al. 18086 ( RB) . Goiás: 25km S de Niquelândia margem da mata de Galeria , 24 January 1972 (fr.), H. S. Irwin et al. 34941 ( UB) ; Niquelandia, CODEMIN, cerca de 500m do cruzamento para Rosariana, na bica do lado esquerdo, 15 April 1996 (fr.), R. Marquete et al. 2537 ( NY, RB) ; Estr. Jataí/Caiaponia a 115 km de Itajaí, alt 740 m, 20 June 1978 (fr.), G. J. Shepherd et al. 7474 ( UEC) ; Gallery forest and adjacent cerrado, ca. 17km S. of Niquelândia , elev. ca. 750m, 23 January 1972 (fr.), H. S. Irwin et al. 34902 ( UB) ; Posse, Rodovia Brasília-Fortaleza , 220km de Formosa, 08 January 1965 (fl.), R. P. Belém & J. M. Mendes 108 ( IPA, NY) . Mato Grosso: Cáceres, Faz. Descalvados , 5 November 1978 (fl./fr.), A. C. Allem 2428 ( NY) ; Poconé, Carretera MTII ( Brozo-Poconé ) 56km of nordeste de Poconé, 31 August 1977 (fl./fr.), R. Shultz-Kraft et al. s.n. ( NY) . Mato Grosso Do Sul: Corumbá, Estrada Miranda , 23 July 1977 (fl.), P. E. Gibbs et al. 5389 ( UEC) ; ibid., Fazenda São Bento , Dist. Nabileque , 16 November 1977 (fl.), A. Allem & G. Vieira 1252 ( CEN) ; ibid., Serra Grande , Planalto Residual do Urucum , interior de Floresta Estacional Residual, 21 December 2004 (fr.), R. R. Silva & J. S. Velasquez 1307 ( UEC) ; ibid., Serra Santa Cruz , Planalto Residual do Urucum . Borda de Floresta Estacional Semidecidual , estrada para mina 7, 10 November 2003 (fr.), R. R. Silva & R. Silva 920 ( UEC) ; Estrada Campo Grande Aquidauana , km 110, Fazenda Leão December 1976 (fl.), G. J. Shepherd et al. 391 ( UEC) ; Estrada para Rochedo , entrada para a fazenda Novo Mundo , 25 January 1979 (fl./fr.), E. P. Heringer et al 788 ( UEC) ; Ladário, Serra do Rabichão , Planalto Residual do Urucum , acesso Fazenda Uruba , 10 December 2003 (fr.), R. R. Silva & M. V. Silva 641 ( UEC) ; ibid., 26 February 2004 (fl.), R. R. Silva & M. V. Silva 749 ( UEC) ; Maracaju , estrada BR 267, 23 March 2002 (fl./fr.), A. Sciamarelli et al. 1549 ( UEC) ; Paranaíba , cultura de milho, 1 March 1981 (fr.), L. Oliveira 34 ( UEC) ; Sete Quedas , 20 March 1982 (fl./fr.), M. R. Melo 375 ( UB) . Minas Gerais: Diamantina, ca. 20km E. of Diamantina , cerrado, 22 March 1970 (fr.), H. S. Irwin et al. s.n. ( UB) ; Joaquim Felício, 3km N. of Joaquim Felício , 5 March 1970 (fl./fr.), H. S. Irwin et al. s.n. ( UB) ; Ouro Preto, Distrito de Cachoeira do Campo , 16 May 2017 (fl./fr.), L. M. P. A. Bezerra 85 ( BOTU) ; Paraopeba, Faz. do Zé Alfredo , 23 June 1957 (fl.), E. P. Heringer 5652 ( UB) ; Ritápolis, Ibitutinga , Estrada secundária à BR-494, junto à antiga estação de trem, 1 February 2013 (fl.), M. Sobral et al. 15394 ( RB) ; Rod. Araxá/Uberaba , km 381 (cerrado), alt. 1000 m., 22 February 1978 (fl./fr.), J. Semir et al. 685 ( UEC) ; Rod. BR 116 entre Caratinga e Governador Valadares , km 477, 08 March 1977 (fl./fr.), G. Shepherd et al. 4368 ( UEC) ; São João Del Rei , 5 km da cidade (aterro), 07 April 1978 (fl./fr.), G. Bufarah et al. IZ-397 ( UEC) ; Tiradentes , campo rupestre próximo à cidade, 06 December 1983 (fl./fr.), H. F. Leitão Filho et al. 15275 ( UEC) ; Uberlândia, Rodovia Uberlândia a Catalão , 06 June 1978 (fr.), P. R. Salgado & D. Bianchine IZ-527 ( UEC) . Paraná: Laranjeiras do Sul, Km 127 , 12 February 1969 (fl.), G. Hatschbach 21128 ( NY) ; Ponta Grossa, Parque Vila Velha , s.d. (fl.), F. Trzeciak 57 ( UEC) ; Sengés, Rio Cajuru , margens, 04 December 1988 (fl./fr.), M. Siveira et al. 81 ( UEC) . Rio Grande do Sul: Torres, Faxinal , 28 March 1998 (fl./fr.), A. Flores 197 ( ICN) . Santa Catarina: Laguna, Praia do Sol , 17 March 2005 (fl./fr.), G. Hatschbach et al. 79242 ( MBM) . São Paulo: Estrada de Salto de Pirapora a Pilar do Sul , próximo a Fazenda Palmeiras , 03 December 1998 (fr.), A. M. G. A. Tozzi & J. L. A. Moreira 348 ( UEC) ; Iperó, Fazenda Ipanema , porteira da Fepasa, alt. 520 m, 04 December 1998 (fr.), A. M. G. A. Tozzi & J. L. A. Moreira 386 ( UEC) ; Matão, cultivada no Instituto de Pesquisas IRI , procedente dos arredores de Matão, s.d. (fl.), Iri 1036 ( UB) ; Pereira Barreto, Ilha Solteira, Perimetral , 28 November 1990 (fl.), R. S. Saito 5 ( UEC) ; Santo Antônio da Alegria , 10 November 1994 (fr.), A. M. G. A. Tozzi & J. C. Galvão 94-218 ( UEC) ; São José do Rio Preto / Mirassol , 20º48’36’’S 49º22’50’’W, 22 September 1995 (fl./fr.), A. A. Rezende 184 ( UEC) ; ibid., 23 January 1996 (fl.), A. A. Rezende 296 ( UEC) ; Sorocaba, nativo à margem da Rodovia Castelo Branco , 11 February 1976 (fl.), H. F. Leitão Filho et al. 1664 ( UEC) ; Taquarivaí, estrada para Itapeva próximo ao Rio Apiaí Mirim , beira da mata mesófila, 07 December 1993 (fr.), V. C. Souza et al 4894 ( UEC) ; Votorantim, estrada do Carafá , 727, entrada da fazenda João Urquiza, 16 December 2012 (fr.), V. C. Souza 35111 ( RB) ; ibid., Serra de São Francisco , Represa de Itupararanga, Cerrado , 12 January 1984 (fl.), V. F. Ferreira 3197 ( RB) . Tocantins: Aurora do Tocantins , Rodovia Campos Belos a Taguatinga, próxima ao Rio Sobrado , 11 February 1994 (fl.), G. Hatschbach & J. M. Silva 60401 ( NY) . COLOMBIA. Cundinamarca: Bogotá, Honda 80 km NO de Bogotá, 3 April 1981 (fl./fr.), R. Schulize-Kraft, R. Reid & G. Keller Grein 56-1 ( NY) ; Hacienda el Cucharo , between Tacaima and Pubenza, 7 May 1944 (fl.), E. P. Killip et al. 38320 ( US) ; Sucre: Archipielago de San Bernardo, Isla Mucura , December 1980 (fl.), G. Moreno & R. Lopez 20 ( NY) . Tolima: Carretera Mariquita–Ibagué , 58 km S de Mariquita, 3 April 1981 (fl./fr.), R. Schulize-Kraft, R. Reid & G. Keller Grein 60-2 ( NY) . COSTA RICA. Las Cañas, Hacienda La Pacifica, Jaragua Garden , 20 November 1962 (fl.), A. T. Semple L100 ( US) . EL SALVADOR. San Salvador, Cerro de San Jacinto , August 1922 (fr.), S. Calderón 1067 ( US) . HONDURAS. El Paraiso; Along Rio Lizapa , between Galeras and Lizapa Grande, 24 June 1947 (fl./fr.), R. A. Molina 163 ( US) . MÉXICO. Chiapas, Slopes on the bank of the Rio Lagas four miles SW of Soyala along the road to the Pan American Highway , 26 June 1964 (fl.), D. E Breedlove 6568 ( US) . NICARAGUA. Managua, steep northeast slope among rocky outcrops; 43 miles towards Managua from Nicaraguan-Honduran border by the Inter-American Highway through Choluteca, Honduras and Somoto, 1 August 1946 (fl./fr.), T. Morley 752 ( US) . PARAGUAY. Amambay: cerca de Bellavista, malezas junto al río Apa, February 1982 (fl./fr.), J. F. Casas & J. Molero FC6273 ( NY) ; Cordillera; Dep. Cordillera, Caaceepe, ( Barrio Buena Vista ) on selva degradada, 2 February 1987 (fl./fr.), E. Bordas 4149 ( NY) ; Estero de Ypoá, 8 km S of Puerto Guyrati, around Arroyo Surubiy , 24 February 1993 (fl.), E. Zardini & T. Tilleria 35251 ( NY) . PERU. San Martin: entre Banos i Bellavista, 12 September 1948 (fl./fr.), R. A. Ferreyra 4836 ( US) ; camino al Km 9 , banda de Shilcayo, 12 January 1982 (fr.), L. Ramirez 28 ( NY) . UNITED STATES. Arizona: Sycamore Canyon (near Ruby), Santa Cruz Co., 24 September 1939 (fr.), T. H. Kearney 14458 ( US) . VENEZUELA. Aragua: Maracay, Girardot, parcela ubicada em la Av: circunvalación, urbanización El Castaño, 28 October 1985 (fr.), H. Rodriguez 1806 ( NY) . Bolivar: 35 km SE de Upata, em a vía a Guasipati , 13 October 1979 (fr.), N. Xena 473 ( NY) . Miranda: en campos de Los Chorros , 20 December 1937 (fr.), F. Tamayo 333 ( US) . Monagas: Carretera Maturín–Barcelona, 5 km al Oeste de Maturín, 28 February 1978 (fl./fr.), R. Shultze-Kraft, D. Escobar & E. Garcia s.n. ( NY) . Trujillo: Carretera 7: Agua Viva–Boconó , 68 km al N de Boconó, 30 April 1981 (fl./fr.), R. Shultze-Kraft & A. Flores ( NY) .

Figures

FIGURE 6. Rhynchosia species. A–C. R. diversifolia. D–G. R. edulis. H–I. R. erythrinoides. (A, B, E by T.C. Monteiro); (C by L.M.P.A. Bezerra); (D, F, G by D. S. Gissi) and (H, I by N. R. Marcial).

FIGURE 7. Geographical distribution of Rhynchosia diversifolia, R. edulis and R. erythrinoides in South America.