Chamaecrista absus (Linnaeus) Irwin & Barneby (1982: 664)

Souza, Alessandro Oliveira De, Cavalcante, Raphael Guarda & Silva, Marcos José Da, 2022, Taxonomic revision of Chamaecrista sect. Absus subsect. Absus (Leguminosae, Caesalpinioideae) with adjustments in the new classification, Phytotaxa 565 (1), pp. 1-82 : 6-12

1. Chamaecrista absus (Linnaeus) Irwin & Barneby (1982: 664)

≡ Cassia absus Linnaeus (1753: 537) ≡ Grimaldia absus (Linnaeus) Link (1831: 141) , non (Linnaeus) Britton & Rose (1930a: 299)

≡ Senna absus (Linnaeus) Roxburgh (1832: 340) .

Type :— s. loc., 1670–1677, (fl.), Ceylon s. n. (holotype BM000621504 !). (Figs. 1 and 2)

= Grimaldia opifera Schrank (1805: 187) . nom. nud.

= Cassia babylonica Schrank (1805: 312) . nom nud.

= Cassia thonningii De Candolle (1825: 500) . Type :—” in Guinea. C. viscosa Vahl ! herb, non Kunth.”, s. dat., s. col., s. n. (holotype G-DC, isotypes S, P-JUSS14532!).

= Senna exigua Roxburgh (1832: 339) ≡ Cassia exigua Roxburgh (1814: 31) . nom nud.

= Chamaecrista absus var. meonandra ( Irwin & Barneby 1978: 282) Irwin & Barneby (1982: 664) ≡ Cassia absus var. meonandra Irwin & Barneby (1978: 282) . Type:— MEXICO. Jalisco: Tequila, 29 September 1893, (fl., fr.), C.G. Pringle 4605 (holotype NY00003618!, isotypes BM000952123!, BR0000005117703!, BR0000005118038!, G00371053!, GH00053038!, F0057448F!, K000478168!, LE, MEXU00246986!, MEXU01167196!, MICH1107177!, MO-125068!, NDG24325!, NY003618!, P02436018!, P02436017!, PUL00000268! S, UC, US 00001376!) Syn. nov.

Herbs or subshrubs monocarpic, 0.1–1 m tall, erect, less frequent decumbent. Root nodules present. Branches straight or curved, yellowish-green, setose-viscous and puberulent including stipules, petiole and rachis; non-exfoliating bark. Stipules 1.3–5.5 × 0.4–1 mm long, lanceolate.Adult leaves alternate-distichous, the distal young leaves may be spiraled, regularly distributed along the entire stem, plagiotropic; petiole 1–5 cm long; rachis 0.4–1.3(–1.5) cm long; leaflets 2 pairs, 0.8–4.6 × 0.5–2.8 cm, the distal pair slightly larger than the proximal pair, elliptical, oboval or largely oboval, apex obtuse or rounded or rarely acute, base rounded, oblique on the distal pair, papyraceous, slightly discolored, dark green on the adaxial surface and light green on the abaxial surface, villosus on both sides, setulose-viscous on the margins and on the abaxial veins, sometimes glabrous or glabrescent; margin entire, flat or slightly revolute, ciliated or not; secondary veins 3–6 pairs, conspicuous and imprinted on both sides. Leafy appendix inter-petiolule, filiform, linguliform or triangular, deciduous in some adult leaves. Racemes 2–13(–25) cm long, sessile, 8–20(–40) flowers, erect with straight or slightly sinuous axis, congested or lax, terminal or opposite to leaves when branch grows after flowering, setulose-viscous and villosule, including pedicel, outer surface of bracts and bracteoles. Bracts 1–2.5 × 0.4– 1 mm, oval or lanceolate, persistent. Bracteoles 1–4 × 0.5–1 mm, triangular or lanceolate, deciduous on adult flowers. Buds 3.5–4.7 × 2.5–3.5 mm, ellipsoid or ovoid, apex obtuse. Flowers 5–9 × 3–5 mm; pedicel 1.5–4(–7) mm long, straight and ascending even when fruited; sepals 3–5 × 1–2.2 mm, oblong-elliptic or lanceolate, apex obtuse, setuloseviscous and puberulent, rarely glabrous, yellowish-green, with red spots externally or not; 5 flat, oboval or spatulate petals, the inner one of the same size as the others, it may be slightly falcate, but never curled and nor encircling the stamens, 4.2–7.2 × 1.5–3.5 mm, orange, red or yellow; stamens (2)3–6(7), 2–4.5 mm long, anthers not barbulate; ovary 1.8–2.5 × 1.5 mm, and setulose-viscous on both sides or rarely glabrescent, densely villosule at sutures; style 1.3–3 mm long, straight or slightly curved at apex, glabrous. Legumes 2–5 × 0.5–0.7 cm, linear-oblong or oblong, setuloseviscous and puberulent or only villosule and setulose along the sutures, dark brown. Seeds 5–8, 3.4–5 × 2.5–4 mm, obovoid, rhomboid or piriform, blackened.

Etymology:— “ absus ” from Latin means clothing, alluding to the trichomes present throughout the plant.

Geographical distribution and habitat:— Chamaecrista absus is the species with the largest area of occurrence of the entire genus, has a pantropical distribution and can be found in four continents: Africa, America, Asia and Oceania (Fig. 37A). In America it is reported from southern Arizona in the United States to Paraguay, being more frequent in Mexico and Central American countries. In Brazil it is found as native in the coastal regions of Bahia, Ceará and Rio Grande do Norte in restingas and degraded and/or anthropized environments, or as cultivated/spontaneous in the rest of the country. In Africa it occurs in practically every continent, except in the North region, growing in savannah areas, forest edges or pasture areas. In Asia it is native to Southeast Asia, occurring in India, Malaysia, Indonesia and Southeast China ( Wang et al. 2014). In Oceania it is recorded for Papua New Guinea and the northern portion of Australia. Found between 100 and 1650 m altitude. The low representation of Ch. absus in South America led Irwin & Barneby in 1978 to consider the species as introduced in this region, as well as in other areas of the world (e.g. Madagascar) although the place of origin is still unknown.

The species’ great ability to establish itself in different environments is probably due to its nodulation ability. Chamaecrista absus exhibits a pioneer plant behavior, with rapid and annual growth and occurs frequently in environments that have been recently degraded or disturbed, which is not common in plants of the section Absus , but quite frequent in species of the section Chamaecrista (e.g. Ch. nictitans ( Linnaeus 1753: 380) Moench (1794: 272) and Ch. rotundifolia ( Persoon 1805: 456) Greene (1899: 31)) which grow in similar environments and are capable of nodulating as well as Ch. absus .

Flowering and fruiting:— Chamaecrista absus is a species of annual growth with autogamous flowers, it has a very fast flowering period compared to the other species studied, and can be found with flowers and fruits at the same time all year round, flowering being more common in times of greater rainfall.

Conservation status:—The conservation of the species is given as Least Concern (LC) due to its wide distribution, for demonstrating excellent ability to colonize different environments and for being present in different environmental protection/conservation units around the world.

Taxonomic history and adjustments:— Cassia absus was described by Linnaeus in 1753 in Species Plantarum. In 1805, Schrank established the genus Grimaldia citing Grimaldia opifera as a type species. About 26 years later, Link (1831) combined C. absus into Grimaldia naming Grimaldia absus (Linnaeus) Link. In 1832 Roxburgh in the work “Flora of the plants of India ” combines C. absus in the genus Senna , naming Senna absus . In 1930, Britton & Rose performed the combination of Senna absus on the genus Grimaldia , treating it as “ Grimaldia absus (Linnaeus) Britton & Rose ”, however, this is a later homonym and should be disregarded, since the species was combined. first in Grimaldia by Link in 1831. In this same work Britton & Rose synonymized Grimaldia opifera in Grimaldia absus .

In 1978, Irwin & Barneby in a revision of Cassiinae transferred the species of the genus Grimaldia to Cassia subgen. Lasiohegma . Due to the many unique features that Grimaldia absus has (e.g. leaf phyllotaxis, presence of a foliate leafy appendix inter-petiolule, reduced flowers and androecium), Irwin & Barneby (l.c.) chose to consider the species in a monotypic section (= Cassia sect. Grimaldia ) while the other species belonging to the genus Grimaldia were allocated in the section Absus series Absoideae . In the samestudy, the authors described C. absus var. meonandra from Mexico and distinguished from the typical variety by the androecium commonly with (2)3–4 stamens (vs. (2–)5– 6(7) in the typical variety).

In 1978, Irwin & Barneby in a revision of Cassiinae transferred the species of the genus Grimaldia to Cassia subgen. Lasiohegma . Due to the many unique features that G. absus has (e.g. leaf phyllotaxis, presence of a leafy appendix inter-petiolule, reduced flowers and androecium), Irwin & Barneby (l.c.) chose to consider the species in a monotypic section (= Cassia sect. Grimaldia ) while the other species belonging to the ancient genus Grimaldia were allocated in the series Absoideae of the section Absus . This year, the authors described C. absus var. meonandra from Mexico and distinguished from the typical variety by the androecium commonly with (2)3–4 stamens vs. (2–)5–6(7) in the typical variety.

Analyzing the original works, we found that Cassia babylonica , Grimaldia opifera and Senna exigua are nomina nuda, as they were created without description, diagnosis or indication of a reference (e.g. illustration or specimen in herbarium); and we corroborate with the synonym Cassia thonningii (valid name) as it is just a variation of the species. Finally, analyzing collections from Central and South America, Africa, Asia and Australia, we concluded that Ch. absus var. meonandra does not hold up as a distinct taxon based solely on the variation of its stamens. The androecium with 5–7 stamens seems to be the most common variation for the species, as also noted by Irwin & Barneby (1978). However, it is not difficult to find flowers with androecium with 2–4 stamens, and this condition is not unique to the populations of Mexico as postulated by Irwin & Barneby (l.c.) when establishing the meonandra variety. Specimens from Africa and Asia have been found with flowers with the following stamen variations: 2–4, 3–5, 5–6, and 5–7. Thus, the only diagnostic feature of the meonandra variety is superimposed and continuous and must be considered a common variation of a polymorphic and widely distributed species. In view of this, we admit here Ch. absus as a species without infraspecific categories with the synonymization of Ch. absus var. meonandra .

Characterization and morphological relationships:—Although it has a wide distribution and certain polymorphism, Ch. absus has conserved diagnostic characters and is easily recognized as it is the only species in the section Absus to have: adult leaves alternate with a filiform, lingual or triangular leafy appendix inter-petiolule; flowers conspicuously reduced (5–9 mm long); androecium with (2)3–5(–7) stamens; fillets similar in size to anthers and style as short as the ovary (1.3–3 mm long); in addition to the presence of root nodules. Such characters are constant in all analyzed morphotypes and together are determinant to the recognition of the species. Other relevant characteristics of the species, but not exclusive, are the sessile and terminal inflorescences or opposite the leaves; flowers short pedicelled with 5 flat petals, the inner one slightly falcate and not curled or encircling the stamens, and the color of the petals that vary from orange, red or yellow.

Due to the presence of leafy appendix inter-petiolule, Ch. absus has already been confused, allocated and described in the genus Senna , as such structures were wrongly interpreted as convex nectaries ( Roxburgh 1832). Irwin & Barneby (1978) already mentioned the uncertainty about the origin and function of the leafy appendix of Ch. absus . Morphologically, the flat, foliate and non-secretory structure of the appendix is more similar to stipules than nectaries. However, a deeper micromorphological analysis (ontogenetic and anatomical) is necessary to elucidate the nature of this structure in terms of its origin and function.

Despite so many peculiarities, Chamaecrista absus is vegetatively similar to the species of the subsect. Absus , sometimes being confused in herbarium collections with Ch. hispidula and Ch. fagonioides . The similarity of these taxa is attributed to the leaves always with two pairs of papyraceous or membranous leaflets with the distal pair slightly larger than the proximal pair and with an oblique base, while the proximal pair has an obtuse or rounded base in addition to similar shapes and dimensions and branches with glandular trichomes. However, the diagnostic features of Ch. absus listed in the preceding paragraphs make it unmistakable with any species of the genus.

Specimens examined: — ANGOLA. Zoana , s. dat., J. Gossweiler 170 ( K) ; Huila: Chicunge , 1400m, 27 March 1962, (fl., fr.), Teixeira & Almeida 5543 ( L / WAG) . AUSTRALIA. Queensland: Cook district, 4.6Km south of Batavia on the peninsula Development Road, 19 April 1990, (fl., fr.), J.R. Clarkson & V.J. Neldner 8251 ( L) ; Rockhampton , 1864-66, (fl., fr.), A. Dietrich 1461, 2227 ( L / WAG) ; The Northern Territrory: Darwin, Nightcliff , 12°22’0”S, 130°52’59”E, (fl., fr.), R.L. Specht 176 ( L) . BENIN. 03 June 1999, (fl.), V. Adjakidje et al. 2704 ( K) ; Antacora: Natitingou , 02 September 1998, (fl., fr.), N. Sokpon 191 ( L / WAG) . Borgou: Nikki, Tanakpé , 10°00’00”N, 03°27’00”E, 14 August 1998, (fr.), B. Sinsin 1944 ( L / WAG) ; Parakou, 09°00’00”N, 02°00’00”E, 10 August 1999, (fl., fr.), A. Akoegninou 2801 ( L / WAG) ; Zou: Dassa, 07°49’00”N, 02°20’00”E, 07 June 1998, (fl.), V. Adjakidjé 1627 ( L / WAG) . BOLIVIA. s. loc. ( Yungas ), 20 April 1894, (fl., fr.), M. Bang 2139 ( NY) ; Milunguayensis Moro-Yungas, December 1917, (fl.), O. Buchtien 777 ( NY) . BOTSWANA. Ngamiland: Mwakupan at lake Ngami, 128km S of Maun , 20°30’0”S, 22°45’0”E, 10 March 1969, (fl.), R.J. Hoogh 156 ( L / WAG) . BRAZIL. Bahia: Caém, ca. 1 km da BR 324 na estrada para Caém , 11°11’00’’S, 40°25’00’’W, 440m, 22August 1993, (fl., fr.), L.P. Queiroz & N.S. Nascimento 3555 ( HUEFS) . Ceará: Alcântaras, Serra da Meruoca, 19 May 1977, (fr.), A. Fernandes s. n. ( EAC3212 , NY959821) ; Fortaleza, Campus Esc. Agronomia , 07 May 1973, (fl.), A. Fernandes s. n.( EAC2300 ) ; ib., Campus do Pici, 12 June 1974, (fl., fr.), A. G. Fernandes s. n.(NY959820) ; ib., 05 June 1986, (fl., fr.), E. Nunes s. n. ( EAC14661 ) ; Meruoca, Serra da Meruoca , Sítio São Jerônimo , 16 July 1957, (fr.), A. Fernandes s. n. ( EAC1733 ) ; ib., sítio Santo Antônio , 05 June 1961, (fr.), A. Fernandes s. n. EAC2104 ) ; Santa Quitéria, próximo a Serra do Quati , 07 June 1984, (fl., fr.), A. Fernandes et al. s. n. ( EAC12704 , NY959818 ) ; Sobral, Fazenda Macapá , 24 May 1976, (fl.), A. Fernandes s. n. ( EAC2775 , NY959819 ) ; ib., BR- 222 km 9, Sobral-Fortaleza, 13 June 1979, (fl., fr.), L. Coradin et al. 1921 ( CEN, NY) . Mato Grosso do Sul: Miranda, 17 March 2003, (fl., fr.), G.G. Hatschbach 74839 ( K) . Minas Gerais: Bambuí , 11 April 2001, (fl., fr.), J. F. Macedo 3985 ( PAMG) ; Belo Horizonte, Serra do Curral , 25 April 1991, (fr.), Pio 151 ( PAMG) . Rio Grande do Norte, Jucurutu, RPPN Stoessel de Brito , 01 June 2008, (fl., fr.), A. A. Roque 583 ( UFRN) ; Serra Negra do Norte, Estação Ecológica do Seridó ( ESEC), 06°34’04’’S, 37°15’20’’W, 02 May 2005, (fl.), R. T. Queiroz 296 ( UFRN) ; Serrinha dos Pintos, Sítio Serrinha do Canto , 17 April 2005, (fl.), R. T. Queiroz & F. R. Queiroz 322 ( UFRN) ; ib., 06°10’05’’S, 37°57’27’’W, 305m, 21 June 2006, (fl.), R. T. Queiroz 1023 ( UFRN) . BURKINA FASO. Bobo-Dioulasso: 10km E. of Bobo-Dioulasso, 11°10’00”N, 04°11’00”W, 17 October 1967, (fl., fr.), C. Geerling & J. Bokdam 1303 ( L / WAG) . BURUNDI. Bubanza: ferme de Randa, 11 February 1967, (fl., fr.), M. Reekmans 4772 ( L / WAG) . Bujumbura: plaine Rusizi km: I 0, jachere sur sables, 03°19’00”S, 29°19’00”E, 20 March 1979, (fr.), M. Reekmans 7714 ( L / WAG) . Bururi: Resha prov. Bururi, 03°53’00”S, 29°23’00”E, 19 January 1980, fl. (fr.), M. Reekmans 8567 ( L / WAG) . CAMEROOM. North Region; près Sangbe, (50km SSW de Tibati), feuille IGN 1 /200.000 Tibati , 18 September 1963, (fl., fr.), R. G. Letouzey 5808 ( L / WAG) . CHAD. Baguirmi et région Du Lac Fittri Kolkeli et Moïte, 12°37’0”N, 16°37’0”E, 06 September 1903, (fr.), A. J. B. Chevalier 9769 ( L) . CHINA. Hainan: Lingshui Xian, Xincun Zhen, Jiusuo Shan, on grassland, 50 m, 21 February 2013, (fl., fr.), Q. L. Wang 476 ( ATCH) . COLOMBIA. Cauca: Patía, Granja de la Universidad de Nariño, 10 November 1986, (fl.), O. Salazar de Benavides 7250 ( NY) . CÔTE D’IVOIRE. Adiopodoumé, 03 November 1954, (fl., fr.), J. F. Lelieveld 5 ( L / WAG) ; Bavé , W. of Bavé, 5 km E. of Comoe River, 09°37’00”N, 04°16’00”W, 04 August 1967, (fl., fr.), G. J. Amshoff 470 ( L / WAG) ; Dabou, E. of Dabou, 05°20’00”N 04°22’00”W, 30 August 1967, (fl., fr.), C. Geerlin & J. Bokdam 791 ( L / WAG) ; Ouangofetini, 10 km E. of Ouangofetini, 09°37’00”N, 03°58’00”W, 21 June 1968, (fl., fr.), C. Geerling & J. Bokdam 2907 ( L / WAG) ; Seye, SW of Seye, 09°24’00”S, 03°26’00”W, 01 July 1967, (fl., fr.), G. J. Amshoff 1972 ( L / WAG) . COSTA RICA. s. loc. Sommels des collines de Nicoya, December 1899, (fl., fr.), J. F. A. Tonduz 13571 ( NY) . CURAÇAO. Near Carmabi Experimental Station, s. dat., (fl.), P. A. W. J. Wilde 102 ( L) . DEMOCRATIC REPUBLIC OF THE CONGO. Dungu, Paro nat. De la Geramba , March 1951, (fl.), D. Saeger 1113 ( L / WAG) ; Kasenga, Haut-Shaba , vallés du Luapula, près du village Tanganika , Près de la chute Mufunfu , 12 March 1971, (fr.), U. A. M. Druk 445/30 ( L / WAG) ; Léopoldville, Madimba, Kisantu, 24 February 1949, (fr.), Calles 2037 ( L / WAG) ; Mbanza-Ngungu, Kibulu , 05 February 1979, (fl., fr.), L. Pauwels 6156 ( L / WAG) ; Ndjili, Ndjili terr. Ngafula , 25 March 1976, (fr.), L. Pauwels 5533 ( L / WAG) . ECUADOR. Guayas: Guayaquil, Cerro Santa Ana, 24 March 1955, (fl., fr.), E. Asplund 15884 ( NY) ; Isla Puná, Puná Nueva to Concorida, 02°44’00”S, 79°55’00”W, 05 July 1987, (fl., fr.), J. E. Madsen 63579 ( NY) . ETHIOPIA. s. loc. 1837, (fr.), Kotschy 460 ( K). Tigray: Adwa, 20 km along road Adwa to Asmara, 14°19’00”N, 38°52’00”E, 1650m, 12 September 1970, (fl., fr.), J. J. F. E. De Wilde 7110 ( L / WAG) . GABON. Nyanga; Monts Doudou, au Sud-Ouest du village Moukoualou, qui se trouve à 6km Sud de Mourindi ( Brigade de Faune de Moukalaba ), 2°36’0”S, 10°46’0”E, 21 March 2000, (fl., fr.), J. M. Moussavou 881 ( L / WAG) . GHANA. path between Pepease and Nkwantanang, Kowahu district , 14 April 1954, (fl., fr.), J. K. A. Morton 775 ( L / WAG) ; between Adidome and Ho, 2/3 of the way, 10 May 1956, (fl.), J. K. A. Morton 2056 ( L / WAG) . GRENADA. s. loc., Lowcher-Tempé road, near the Boramie Station , 11 December 1904, (fl., fr.), W. E. Broadway 1745 ( NY) . GUATEMALA. Chiquimula: Vicinity of Ataluapa, 11 December 1969, (fl., fr.), A. M. Rositto 25380 ( NY) . Guatemala: along river Villalobos, 12 January 1966, (fr.), A. M. Rositto 16052 ( NY) . GUINEA. Boffa; près de Tugnifily, 06 February 1979, (fl., fr.), S. Lisowski 51420 ( L / WAG) . GUINEA-BISSAU. Pessubé, 11 December 1937, (fl., fr.), J. V. G. Espirito Santo 1078 ( LQWAG) . INDIA. Monghyr, s. dat., (fl.), s. col. ( K001120513 ) ; Himachal Pradesh: pine forest 7 km to Bharwain, 11 October 1977, (fl.), L. J.G. van der Maesen 2869 ( L / WAG) ; Dehradun, 4, 5 km to Sahabradhara , 16 October 1977, (fr.), L. J.G. van der Maesen 2986 ( L / WAG) ; Madhya Pradesh: Chhindwara, 27 September 1979, (fr.), P. Remanandan 4288 ( L / WAG) . Surguja: Khuri , 12 October 1947, (fr.), W. N. Koelz 19258 ( L) ; Uttar Pradesh: Allahabad, 09 October 1978, fl. (fr.), Bumisra 813 ( L) ; Majhola (Pilibhit), 23 September 1972, (fl.), C. L. Malhotra 50995 ( L) . Uttarakhand: Dehra Dun, Rispana, 08 August 1934, (fl.), C. R. Babu 33302 ( L) . INDONESIA. Ned. Indie. Semongkrong, 01 November 1921, (fl., fr.), J. Jeswiet 1057 ( L / WAG) ; Java: Timur, Baluran, 01 March 1950, (fl.), F. W. Rappard 137 ( L / WAG) ; Lesser sunda islands: Island Bali ( North ), Pura Pulaki, 31 March 1964, (fl., fr.), A. Dilmy 1042 ( L) . JAMAICA. s. loc., Kings House Grounds, 25 November 1897, (fr.), W. H. Harris 6707 ( NY) . Saint Andrew: East of the Cane River above coastal highway bridge, 25 October 1964, (fl., fr.), G. R. Proctor 25597 ( NY) . KENYA. 4/7 Kora Game Reserve , Tana River District, 07 December 1984, (fl.), G. M. Mungai 84/274 ( L / WAG) . MALAWI. Shire Highlands, Zambesia, s. dat., Buchan 363 ( K) . Rumphi: Rumphi, Chelinda river 3000ft., 04 May 1974, (fr.), J. Pawek 8601 ( L / WAG) ; Binga, Manzituba Marsh, Chizarira Game Reserve , 25 February 1972, (fl., fr.), P. J. Thomson 549 ( L / WAG) . MALAYSIA. Penang: Tanjong Bunga, Seashore below Sandycroft Soldiers’ Leave Home, 31 October 1951, (fl.), J. Sincalir 39389 ( L) . MEXICO. Baja California Sur: La Paz , Los Limpios , Sierra La Laguna , al E de Todos Santos, BCS, 18 October 1985, (fl., fr.), P. Tenorio L. 10617 ( NY) ; Chiapas: 20 km al S de Ocozocuautla, camino a Villa Flores, en el Cerro Brujo, 25 October 1985, (fr.), E. M. M. Salas 14310 ( NY) ; Coahiula: Artega, a 7 km al N de Arteaga, camino a Nueva Italia , 19 November 1983, (fr.), E. Martínez S. & F. Barrie 5398 ( MEXU, XAL) ; Jalisco: Falda del cerro El Huehuentan, Nacastillo, Jal, 03 November 1978, (fl., fr.), J. A. S. Magallanes 1205 ( NY) ; Guadalajara: Río Caliente , La Primavera, Zapopan, Jal, 01 August 1967, (fl.), C. L. Díaz Luna 328 ( NY) ; Hostotipaquillo: 2 mi W of Barranquitos , ca. 10 mi E of Plan de Barrancas, on road to Tequila, 23 August 1957, (fl.), H. S. Irwin 1273 ( NY) ; La Huerta , Ca. 8 mi SW of La Resolana, road to La Huerta, 16 November 1960, (fr.), R. McVaugh 21109 ( NY) ; Tala, a largo del Arroyo que parte del Rancho em Bosque Escuela, hasta el Arroyo Caliente, La Primavera, 15 October 1988, (fr.), A. Rodriguez C. & J. J. Reynoso D. 1585 ( XAL) ; Tequila , July 1886, (fl., fr.), E. Palmer 412 ( NY) . Michoacán de Ocampo: Huétamo, En Las Trincheras, 12 km al N de Huétamo , 24 October 1984, (fr.), J. C. S. Núñez 6666 ( NY) ; La Huacana , a 6 km al SW de La Huacana, 27 October 1981, (fl., fr.), J. C. S. Núñez 3566 ( XAL) ; Nocupétaro, 7 km al N de Nocupétaro, carr. a Villa Madero, 29 September 1982, (fl., fr.), J. C. S. Nuñez 4649 ( NY) ; Morelos: Emiliano Zapata, Cerro Gordo , Mpio. Dos Ríos, 14 October 1970, (fl.), F. Ventura A. 2610 ( NY) ; Nayarit: Km 31, 4-5 m N of Compostela, Nay , 22 September 1960, (fl., fr.), R. McVaugh 19361 ( NY) . Sinaloa: Mazatlán, 01 January 1895, (fl., fr.), F. H. Lamb 375 ( NY) ; Sinaloa, Quebrado de Manzana, Sierra Surotato, 10 September 1941, (fl., fr.), H. S. Gentry 6536 ( NY) . Sonora: Imuris, 17.1 km northeast of Imuris on MEX 2 , 09 September 2002, (fl., fr.), S. C. Doan 1205 ( NY) ; Yecora. Arroyo Hondo, 11.5 km E of El Kipor, 4 km W of Chihuahua border on Mex. 16, 28°26’30”N, 108°32’30”W, 11 September 1996, (fl., fr.), T. R. Van Devender 96-613 ( NY) . Veracruz: Actopan, Trapiche, 06 October 1971, (fl., fr.), F. Ventura A. 4361 ( XAL) ; Coatepec, 4 km al este de Tuzamapan, 02 November 1975, (fr.), M. Morales et al. 15 ( XAL) ; Cosamaloapan, Ciudad Alemán, 16 November 1968, (fl., fr.), G. M. Calderon 1159 ( NY) ; Dos Rios, Corral Falso , 03 October 1974, (fl., fr.), A. Delgado 1978 ( XAL) ; Zacualpan, Corral de las Piedras, October 1906, (fl., fr.), C. A. Purpus 2339 ( NY) . MOZAMBIQUE. Lourenco Marqueen Forto Menrique, andados ca. 1, 5 m para a Bela Vista, 25 March 1975, (fl., fr.), A. Marques 2673 ( L / WAG) . NAMIBIA. Caprivi; near Ngoma, 17°52’59”S, 24°43’0”E, 10 February 1988, (fl.), G. L. Maggs 199 ( L / WAG) . NETHERLANDS ANTILLES. Bonaire, 1909, (fl., fr.), I. Boldingh 7303 ( NY) . NIGERIA. s. loc., Idah [Zuoora], November 1841, (fl., fr.), Vogel 13 ( K) . Adamawa: Yola, government reservation on Jimeta-Yola Road, 23 October 1971, (fr.), M. G. Latilo 63475 ( L / WAG) . PAPUA NEW GUINEA. 10 June 1928, (fr.), J. Jeswiet 80 ( L / WAG) ; Central District: New Guinea; bulogo Creek c. 18 miles N. of Port oresby, 11 April 1967, (fl., fr.), R. Pullen 6665 ( L) . PARAGUAY. Assunção, 05 May 1889, fl. T. Morong 700 ( NY) ; Cordillera: 5 km SE of Emboscada. “ Isla Alta ”, 25°08’00”S, 57°15’00”W, 26 May 1990, (fr.), E. M. Zardini 20648, 20737 ( NY) . PERU. Lambayeque: Portachuelo de Olmos, 01 June 1972, (fl., fr.), E. C. Valenzuela 5209 ( NY) ; Piura: Província Chulucanas, Cerro Vicus, 52 km de la carretera Piura-Morropon, 24 May 1992, (fl.), I. M. S. Veja 6206 ( NY) . RWANDA. Kibungu, plain Kilali, 03 February 1958, (fl., fr.), G. M.D.J. Troupin 5904 ( L / WAG) . SENEGAL. Kolda; Diaroume, 13 October 1961, (fl., fr.), J. G. Adam 18419 ( L / WAG) . SIERRA LEONE. between Mussaia and Kabala N. P., 27 November 1965, (fr.), J. K. Morton 2869 ( L / WAG) . SOUTH AFRICA. Waterberg Dist. 6m. E. Hermanusdoorns: Gravelly loam soil in Mogol River Valley , 13 March 1946, (fl., fr.), L. E. W. Codd 1000 ( L / WAG) ; Regio TVL. 8 miles from Nylstroom on road to Naboomspruit at bridge over a branch of the Nyl river , 15 March 1971, (fl., fr.), L. A. Coetzer 18 ( L / WAG) . SRI LANKA. Hambantota: Hambantota, Ruhuna National Park , Block I Karaugaswala, 17 January 1969, (fl., fr.), N. Wirawan 810 ( L) . TANZANIA. Dodoma, Ikowa reservoir, 06°12’00”S, 36°14’00”E, 980m, 21 February 1975, (fl.), I. Backéus 1141 ( L / WAG) ; Morogoro, Mikumi National Park, 07°15’00”S, 37°10’00”E, 14 April 1984, (fl., fr.), G. Nevers & S. Charnley 3281 ( L / WAG) . UGANDA: North Buganda; Mukono, Kyagwe, Bule-Buyaga. VR-65-30, 0°16’59”S, 32°42’0”E, 20 March 1992, (fr.), P. K. Rwaburindore 3378 ( L / WAG) . UNITED STATES. Arizona: Montezuma, Pima Co. North side of Montezuma Bluff, Baboquivari Canyon, Baboquivari Mts. , 28 September 1931, (fl., fr.), M. F. Gilman B 234 ( NY) . VENEZUELA. Distrito Capital: Caracas, Around Caracas. Sabana Grande, 11 December 1921, (fr.), H. F. Pittier 9927 ( NY) . ZAMBIA. Lusaka: Chalimbana Research Station; Lusaka E. 21km, 27 January 1993, (fl., fr.), M. G. Bingham 8807 ( L / WAG) ; Namwala: Kafue National Park. Musa river , 1 mile from Musa-Kafve, 22 March 1964, (fr.), B. L. Mitchell 25/7 ( L / WAG) . ZIMBABWE. Makuti, ( Top of Escarpment ), 22 February 1985, (fl.), R.D.A. Bayliss 10206 ( L / WAG)

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