Solanum houstonii Martyn, Gard. Dict. (Miller), ed. 9, no. 91. 1807, as ' houstoni'

Knapp, Sandra, Sagona, Eva, Carbonell, Anna K. Z. & Chiarini, Franco, 2017, A revision of the Solanum elaeagnifolium clade (Elaeagnifolium clade; subgenus Leptostemonum, Solanaceae), PhytoKeys 84, pp. 1-104 : 27-30

4. Solanum houstonii Martyn, Gard. Dict. (Miller), ed. 9, no. 91. 1807, as ' houstoni' Figures 2D, E , 3E, F , 4B , 10

Solanum carolinense Mill., Gard. Dict. ed. 8, no. 21. 1768, as ' carolinense ’’, non S. carolinense L. (1753). Type. Mexico. Sin. loc. [probably the Yucatán peninsula or Veracruz], W. Houstoun s.n. (lectotype, designated here: BM [BM000514926]).

Solanum tridynamum Dunal, Encycl. [J. Lamarck & al.] Suppl. 3: 776. 1814. Type. Mexico. Sin. loc. (no specimens cited; lectotype, designated here: Service du Patrimoine Historique de l’Université de Montpellier Node-Véran, Sol. Tab. 75, MPU310713).

Solanum amazonium Ker Gawl., Bot. Reg.1: tab. 71. 1815. Type. Cultivated in London at Chelsea Physic Garden, plants received from A.B. Lambert, said to be from Mexico (no specimens cited; lectotype, designated here: Ker, Botanical Register 1, tab. 71).

Nycterium amazonium (Ker Gawl.) Sims, Bot. Mag. 43: tab. 1801. 1816. Type. Based on Solanum amazonium Ker Gawl.

Solanum herbertianum Paxton, Paxton’s Mag. Bot. 5: 269. 1838. Type. Cultivated in England at the Epsom Nursery of unknown origin, no specimens cited; lectotype, designated here: Paxton, Paxton’s Magazine of Botany, un-numbered plate labelled " Solanum herbertiana ", facing page 269).

Solanum obtusilobum M.Martens & Galeotti, Bull. Acad. Roy. Sci. Bruxelles 12(1): 152. 1845. Type. Mexico. Puebla: Tehuacán [de las Granadas], 6000 ft, Aug 1840, H. Galeotti 1168 (lectotype, designated here: BR [BR00000523923]; isolectotypes: BR [BR00000523971], K [K000532440], K [K000063990 "Cordillera, Oaxaca"], P [P00366959 "Cordillera, Oaxaca"], W [W0003056 "Cordillera, Oaxaca"]).

Solanum tridynamum Dunal var. anoplocladum Dunal, Prodr. [A. P. de Candolle] 13(1): 334. 1852. Type. Cultivated in Geneva, Aug 1848, Anon. s.n. (holotype: G-DC [G00131315]).

Solanum tridynamum Dunal var. stylosum Dunal, Prodr. [A. P. de Candolle] 13(1): 334. 1852. Type. Mexico. " J. Pavón s.n. " [almost certainly collected by M. Sessé & J. Mociño, see Knapp 2008b] (holotype: G [G00343357]; probable isotypes: M. Sessé & J. Moçiño 1538 [MA-604686]; M. Sessé & J. Moçiño 5367 [MA-604687], 1538 bis [MA-604685]).

Solanum azureum Fernald, Proc. Amer. Acad. 35: 570. 1900. Type. Mexico. Sinaloa: Topalobampo [Topolobampo], 15-25 Sep 1897, E. Palmer 178 (lectotype, designated here: A [A00077437]; isotypes: ARIZ [ARIZ-BOT-0005048], GH [GH00077436], MICH [MICH1109885], P [P00324679], RSA [RSA0006300], S [S09-39899, S-G-5690], UC [UC00770350], US [US-315500]).

Type.

Based on Solanum carolinense Mill.

Description.

Shrub, 0.2-1.5 m tall. Stems erect, sparsely to densely armed, or unarmed; young stems densely stellate-pubescent; trichomes porrect, translucent and often reddish gold, sessile to subsessile, the rays 6 to 9, 0.1-0.3 mm long, the midpoints ca. 0.1 mm long, the prickles irregularly distributed throughout the plant, sometimes more dense on the calyx and pedicels, 1-10 mm long, 0.5-1.5 mm wide at base, straight, brown or reddish, sometimes yellow on the young stems, spaced 0.1-1 cm apart if dense, 1-10 cm apart if sparse; bark of older stems brown, glabrescent. Sympodial units usually difoliate, but not markedly geminate, sometimes plurifoliate. Leaves simple, variously lobed, 3-7(10) cm long, 1.5-3(5) cm wide, variable in shape, ovate to elliptic to broadly elliptic, 1.5-3 times longer than wide, mostly concolorous, drying yellowish green to green, densely stellate-pubescent adaxially and abaxially, the trichomes porrect, translucent, sessile to subsessile, the stalk up to 0.1 mm long, the rays 7 to 9, (0.1) 0.2-0.4 mm long, the midpoints 0.1-0.2 mm long; principal veins 3-6 pairs, raised abaxially, flat adaxially, spreading at ca. 45° to the midvein, the tertiary venation mostly not visible to the naked eye; base obtuse, rounded or truncate, sometimes oblique; margins variously lobed, rarely entire, the lobes 2-5 on each side, 0.1-1.5 cm long, usually rounded, rarely obtuse, the sinuses extending up to 1/3-1/2(2/3) of the distance to the midvein; apex rounded to acute; petiole 1-4 cm long, 1/2-1/10 of the leaf length, densely stellate-pubescent like the young stems. Inflorescence terminal or lateral, 4-7(15) cm long, usually unbranched but occasionally forked, with 4-9(26) flowers, usually with one hermaphrodite flower at the base, all distal flowers staminate; peduncle 0.5-2 cm long; rachis 1-6(9) cm long; pedicels 0.4-1 cm long, ca. 1 mm in diameter, filiform or apically dilated, densely or sparsely armed only in hermaphroditic flowers, unarmed in staminate flowers, articulated near the base; pedicel scars irregularly spaced 2-15 mm apart, prominent and brown. Buds strongly curved and zygomorphic, more so in staminate flowers; the corolla exserted ca. halfway from the calyx tube before anthesis. Flowers 5-merous, heterostylous, heterandrous and markedly dimorphic, the plants strongly andromonoecious, the basal flower long-styled and hermaphroditic, the distal flowers short-styled and staminate. Calyx tube 0.8-2 cm long, cup-shaped, the lobes 0.7-2.5 cm long, 1-2 mm wide, long-triangular with an elongate acumen, unarmed in staminate flowers, densely armed in hermaphroditic flowers, the prickles 0.5-1 cm long, 0.5-1 mm wide at base, yellow or reddish, straight, spaced 0.5-2 mm apart. Corolla 2-4 cm in diameter (slightly larger in staminate flowers), lilac or purple or occasionally white, the midvein paler and greenish yellow, drying yellow or brownish tan in herbarium specimens, stellate to broadly stellate, lobed ca. halfway to the base, the lobes 1-1.5 cm long, 0.8-1 cm wide, spreading or slightly reflexed at anthesis with abundant interpetalar tissue, the abaxial surfaces densely pubescent along the middle portions where exposed in bud, the adaxial surfaces glabrous or with a few trichomes along the margins, the tips somewhat cucullate. Stamens strongly unequal in both hermaphroditic and staminate flowers, with the 2 adaxial anthers shorter than the 3 abaxial anthers; filament tube minute; free portion of the filaments 1.5-2 mm long, glabrous; anthers tapering, yellow or occasionally purple (Sinaloa and Sonora), the surfaces smooth, poricidal at the tips, the pores about the same diameter as the anther apices, clearly delineated, directed distally, in hermaphroditic flowers three longer abaxial anthers ca. 9 mm long, two shorter adaxial ca. 6 mm long, in staminate flowers three curved abaxial anthers 15-20 mm long, two straight adaxial anthers ca. 10 mm long. Ovary in hermaphroditic flowers ca. 2 mm in diameter, conical. porrect-stellate pubescent in the distal third, the trichomes sessile to subsessile, with 7-10 rays, the rays 0.1-0.2 mm long, the midpoints 0.1-0.15 mm long; ovary in staminate flowers vestigal; style 20-30 mm long in long-styled flowers, in staminate flowers ca. 1.5 mm long; stigma capitate and slightly bilobed in hermaphroditic flowers, indistinctly bilobed and much smaller in staminate flowers. Fruit a globose dehiscent berry, (1-)2-2.5 cm in diameter, pale green with darker green mottled areas when young, dark brown or orange-brownish when dry, dehiscing irregularly in several parts, usually breaking open irregularly in the upper half of the fruit, the pericarp thin, smooth, glabrous; fruiting pedicels 0.5-1.5 long, 2-3 mm in diameter at base, 2-5 mm in diameter at apex, erect, herbaceous to woody; fruiting calyx (hermaphroditic flowers only) strongly accrescent, the lobes covering up to the total length of the fruit, sparsely or densely armed with prickles up to 8 mm long. Seeds ca. 20-40 per berry, 3-4 mm long, 3-4 mm wide, flattened reniform, dark brown to black, the surface minutely pitted, the testal cells pentagonal. Chromosome number: n=12 ( Averett and Powell 1972; Chiarini et al. in press).

Distribution

(Figure 12 ). Solanum houstonii is endemic to Mexico, but widespread in south central Mexico on both coasts, from the Yucatán Peninsula and Veracruz to Sinaloa and Sonora where it is sympatric with S. hindsianum ; from sea level to nearly 2000 m.

Ecology and habitat.

Solanum houstonii grows in a wide variety of dry and semi-deciduous forests, from Sinaloan thorn scrub to the humid semi-deciduous forests of the Caribbean coast (selva mediana subcaducifolia) and in open situations such as coastal dunes. Like other members of the group, it can form large colonies, especially in disturbed ground.

Common names and uses.

Mexico. Colima: mala mujer; Puebla: diente de burro; Sinaloa: melón de coyote, coyotillo; Sonora: sacamanteca; Veracruz: berenjena espinosa, necachane; Yucatán: berenjena de monte, k’omya’ axnik (Maya).

The leaves of S. houstonii are used for cleansing after birth and as a dieting aid (Nee & Taylor 29629) and for skin diseases and problems (Rivera 90).

Preliminary conservation status

( IUCN 2016). LC (Least Concern). EOO = 1,837,020 km2 (LC - Least Concern); AOO = 1,052 km2 (VU- Vulnerable). Based on its wide distribution S. houstonii is not of immediate conservation concern, but populations across Mexico exhibit significant heterogeneity in morphology that is presumably also reflected in genetic variability.

Discussion.

Solanum houstonii has the most pronounced flower dimorphism in group, with differences in size and shape of both corolla and anthers. Staminate flowers are larger and exhibit stronger anther zygomorphy, while hermaphrodites tend to be smaller and with anthers of more equal size. The anther dimorphism in hermaphroditic flowers of S. houstonii is similar to that seen in both flower morphs of S. hindsianum and S. homalospermum . On the coast of western Mexico S. houstonii grows in sympatry with S. hindsianum ; it differs from that species in its larger fruit, more dimorphic flowers, more prickly calyx of long-styled flowers, and usually lobed leaves. Populations from around the bay of Topolobambo in Sinaloa are unusual in having deeply pinnatifid leaves, and individuals from northwestern Mexico are polymorphic for purple or yellow anther coloration.

This species was long known as either S. amazonium or S. tridynamum ; Nee (1999) found that the name coined by Thomas Martyn ( Miller 1807) as a replacement for the illegitimate S. carolinense Mill. had priority, thus S. houstonii becomes the correct name for this distinctive and relatively widespread Mexican species. Martyn coined his new name for this plant in honour of its collector, William Houstoun (with the original spelling ' houstoni '), who sent specimens and seeds of plants collected in western Mexico to Philip Miller at the Chelsea Physic Garden in London ( Stearn 1974); many of Houstoun’s plants were grown in the gardens and glasshouses of Chelsea. We select here as the lectotype the single specimen in BM (BM000514926) with labels in Houstoun’s hand and that bears another in Miller’s hand (the upper label); both of these feature verbatim in the polynomial description of S. carolinense Mill. (see image in Global Plants, http://plants.jstor.org/ - search for S. houstonii ).

The protologue of S. tridynamum (Dunal 1814) cites no specimens, only two illustrations. One was an illustration in the collection of original drawings done for the Sessé and Mociño expedition to Mexico and Central America (see McVaugh 2000) and the other Dunal’s own, unpublished illustration copied from this original, now held in the collections at MPU. Dunal is likely to have seen the former illustration when Mociño was in Montpellier between 1812 and 1817 (see McVaugh 2000: 12); this set of original illustrations was then copied for Candolle, but no images of Solanum species are held in the set of copies at G. Two illustrations in the Torner Collection of the Hunt Botanical Institute (accession numbers 6331.0674 and 6331.1883) both clearly show the distinctive flower morphology of S. houstonii . One is a sketch (6331.1883) upon which the other (6331.0674) showing the whole plant is based. The illustration made by Node-Veran and cited by Dunal in the protologue (MPU310713; Fig. 11 ) is apparently copied from these originals, but we select it as the lectotype of S. tridynanum because it is the only one of these elements we can be sure was seen by Dunal at the time he described the species.

Solanum amazonium was described from specimens cultivated at the Chelsea Physic Garden sent by Aylmer Lambert ( Ker 1815); two sheets at Vienna (W-292940, W-292941) from the A.B. Lambert herbarium are possibly from the same introduction, but are not type material. The plate accompanying the description was prepared from live plants cultivated at Chelsea and is the only unambiguous authentic material that could be used as a lectotype. The protologue contains an excellent description of the anther dimorphism that differs in hermaphroditic and staminate flowers ( Ker 1815). Solanum herbertianum was similarly described from cultivated material ( Paxton 1838) of unknown origin that was grown in the Epsom Nursery in southern England. The plate is the only unambiguous material associated with the protologue and we select this here as the lectotype. The plant depicted is completely devoid of prickles (like that in Ker 1815) and may have come from the same stock. In contemporaneous horticultural collections in W there appear to have been two distinct morphs of S. houstonii in cultivation in Vienna, one with (e.g., W-0001800) and the other without prickles (e.g., W-0001799).

The lectotype we have selected for S. obtusilobum is one of two sheets held in BR, where Martens and Galeotti worked. We have selected the sheet (BR00000523923) that was labelled (but not published) as lectotype by M. Nee in 1986. The other duplicate of Galeotti 1168 in BR (BR00000523971) is clearly labelled as “2ème exemplaire" and is less complete. The protologue ( Martens and Galeotti 1845) cites Galeotti 1168 from "montagnes cactifères et arides près Tehuacan de las Granadas" in today’s state of Puebla. The sheet selected as the lectotype (BR00000523923) states only “Tehuacan”; other sheets labelled with the same number (Galeotti 1168) but with the locality "Cordillera de Oaxaca" are recognised here as isolectotypes, they are morphologically extremely similar to the sheets in BR.

Of the two varieties described in Dunal (1852) var. anoplocladum was based on plants cultivated in the gardens in Geneva and a specimen in the Candolle herbarium was cited, and var. stylosum was based on material from "Herb. Pavon" and is likely to be a collection made by Sessé and Mociño in Mexico (see Knapp 2008b for a discussion of the distribution of Sessé and Mociño’s collections by Pavón). This latter specimen (G00343357) could be related to the illustration used in the description of S. tridynamum , but does not exactly match it; it is probably from western populations with more deeply lobed leaves.

Of the two sheets of Palmer 178 held in the Harvard University herbaria (where Fernald worked) we select the A sheet (A00077437) as the lectotype of Solanum azureum because it is better preserved and has flowers. This plant comes from populations on the west coast of Mexico with deeply sinuate leaves and flowers with occasionally purple anthers.

Selected specimens examined.

MEXICO. Campeche: 3 km al S de Bolochen de Rejón, cerca de las Grutas de Xtacumbilxunan, por la carretera via ruinas a Campeche, 25 Jul 1986, Cabrera & de Cabrera 11770 (MEXU); Calkiní, 8 km al E de Tancuche, sobre el camino Calkini-Punta Arenas, 26 Jul 1987, Cabrera & de Cabrera 13920 (MEXU); 8 km al O de Nunkini, sobre el camino a Calkini-Punta Arenas, 19 Sep 1987, Cabrera & de Cabrera 14318 (MEXU); Ciudad del Carmen, Isla del Carmen, 12 km al SO de Puente de la Unidad, sobre la carretera Campeche-Ciudad del Carmen, 21 Sep 1987, Cabrera & de Cabrera 14535 (MEXU). Chihuahua: Urique, on road coming out of canyon from Urizue, 17 Aug 2005, Ford 11 (NY); Batopilas, La Bufa, SE of Creel, 12 Sep 1957, Knobloch 419 (BM); SW part of state, on road ca. 16.5 mi W of Ocampo, 1 Sep 1989, Mayfield et al. 301 (MEXU); SW Chihuahua, Aug 1885, Palmer 237 (BM, K, MEXU, NY). Colima: Coquimatlán, Coquimatlán, en un terreno solo cercano a zona habitacional, 1 May 2013, Gaitón Hinojosa et al. 2013- 05 (MEXU); 400 m al norte de El Trapiche, 3 Sep 1992, García Torres 78 (MEXU); Colima, 1 km antes de Cardona, 15 Mar 1994, García Torres 148 (MEXU); Armería, Rancho El Mirador, Valle de A, 19 Aug 1994, García Torres & Navarrete 183 (MEXU); Coquimatlán, Rancho Buenos Aires, 1 km del cruce Coquim.-El Poblado, 25 Jul 1995, García Torres 315 (MEXU); Colima, 2 km al SE de Colima, 20 Dec 1040, Langman 3190 (MEXU); Coquimatlán, Pueblo Juárez, 14 Jan 1985, Maillet 14 (MEXU); Comala, Ejido La Caja, cerca del Río Armería, 4 Aug 1992, Navarrete de la Paz 33 (MEXU); Colima, Jul 1897, Palmer 95 (LE, S); Manzanillo, 1 Dec 1890, Palmer 1035 (K, NY); Comala, Predio Cerro Alto y Potrero Nuevo 3 km al suroeste de Los Colomos, 28 Aug 1990, Román Miranda 1371 (MEXU); Armería, 1 km de la desv. carret. Colima-Manz, rumbo a Armería, 26 Feb 1994, Román Miranda 1795 (MEXU); Comala, 8 km al N de Comala, brecha a La Caja, 22 Aug 1984, Santana M. & Cervantes A. 546 (MEXU). Durango: San Dimas, La Desmontada, 7 km al S, por el camino a Mala Noche, 7 Mar 1990, González 2394 (MEXU); Tabahuelo (al catorce) 196 km al W de Tepehuanes, 31 Aug 1983, Torres C. et al. 3561 (MEXU, NY). Jalisco: Guadalajara, 5 miles S of Guadalajara, 12 Aug 1947, Barkley et al. 7505 (MEXU); Tecolotlán, 1.5 km al NE de Lindavista, km 59 carretera Guadalajara-Barra de Navidad, 10 Jul 2003, Carrillo-Reyes & Ortíz-Catedral 4139 (MEXU); Huejuquilla, Rancho Los Arroyos del Agua, 15 km al NW de Huejuquilla, 4 Aug 1990, Flores M. 2037 (MEXU); Villa Guerrero, 41 km al SW de Villa Guerrero, camino a Chimaltitlán, 18 Oct 1983, Lott et al. 2016 (BH, MEXU, NY); San Martín de Bolaños, 14 km al NE de San Martín de Bolaños, 1 km al SE de Chimaltitán, 21 Oct 1983, Lott et al. 2114 (MEXU); Amacueca, 2.5 km al NE de Cofradía del Rosario, 7 Nov 1992, Novoa L. & Acevedo R. 54 A (MEXU); San Martín de Bolaños, 5 km salida oeste a al Zualaga, 17 Feb 1992, Reynoso Dueñas et al. 634 (MEXU); Atoyac, camino a Atoyac, a partir de la carretera libre a Zacoalco-Sayula, 27 Jul 1991, Rodríguez C. 2107 (MEXU); Zapotitlán de Vadillo, 5 km al N de Zapotitlán de Vadillo, 26 Oct 1994, Sánchez & Santana M. 46 (MEXU); El Limón, Rancho El Recodo, 2 km al E de San Miguel Hidalgo, 1 Sep 1987, Santana Michel 2955 (MEXU); Ciudad Guzmán, Floripondio, km 13 carretera Autlán, 12 Mar 1994, Villareal de Puga et al. 16135 (MEXU); Tenamaxtlán, inmediaciones de la presa, camino a Tecolotlán y Tenamaxtlán, 24 Feb 1980, Villarreal de Puga 11507 (MEXU); Atoyac, Laguna de Sayula, entre el km 51-52 Autopista Guadalajara-Colima, 27 Mar 1993, Villegas F. & Macías R. 219 (MEXU); Amacueca, Laguna de Sayula, carretera Guadalajara-Sayula, 3 km después de la desviación a Tapalpa, 9 Jul 1993, Villegas F. & Ramírez D. 329 (MEXU); Autlán, ca. 1 mile south, 21 Aug 1949, Wilbur & Wilbur 2469 (DS, DUKE, MEXU). Nayarit: Nayar, recorrido entre el poblado y Aguamilpa y el Cerro del Colomo, 20 Sep 1991, Benítez-Paredes 3405 (MEXU); Nayar, 15 km al SW de Jesus Maria, camino a la Mesa del Nayar, 28 Jul 1990, Flores F. et al. 2122 (MEXU); Nayar, Santa Cruz, camino rumbo al rancho El Ocotillo [Santa Cruz de Guaybel], 11 Sep 1990, Rodríguez González 143 (MEXU); Tepic, km 20-40 terraceria hacia La Presa de Aguamilpa, 15 Jan 1988, Téllez V. 11198 (MEXU). Oaxaca: Tepelmeme Villa de Morelos, 45 miles S of Tehuacan, 26 Aug 1967, Clarke et al. s.n. (CAS); W slopes of Sierra Zongolica, 2km (by road) above Teotitlán del Camino on road to Huautla de Jiménez,, 8 Jul 1978, Cochrane et al. 8516 (BH, BM, CAS); along highway 190, 55 miles SE of Oaxaca, 2 Jul 1977, Croat 39945 (MEXU); carretera Oaxaca-Mitla km 39, 10 Feb 1989, Escalante Membrillo s.n. (MEXU); San Pedro Totolapa, Dto. Tlacolula, Valles Centrales, 3.5 km de Totolapan, hacia Tehuantepec, 14 Jul 1988, Flores Martínez 1328 (MEXU, NY); 2 km E of Tomallín, 18 Oct 1992, Newman et al. 435 (K); Mejia Station, 9 May 1894, Pringle 5850 (MEXU, MEXU, VT); Tlacolula, a 1 km al E de San Juan Guegoyache, o a 5 km al NE de Totolapán, 22 May 1982, Rico Arce et al. 378 (BM, MEXU); Valerio Trujano, Tomellín, Sep 1905, Rose & Painter 10073 (K, NY); beyond reservorio 2 km E of San Juan de los Cues, 6 Dec 1994, Way WS-023 (K). Puebla: Cerro Tres Cruces, 10 Dec 1967, Boege 689 (DUKE); Acatlán de Osorio, Route 190 near km 293, ca. 8 km SE of Acatlán, 12 Aug 1966, Cruden 1174 (MEXU); Coxcatlán, 1 km SE of Guadelupe Victoria, path to San José Tilapo, 13 Sep 2004, Guzmán C. 2121 (K); Acatlán de Osorio, Acatlán, 26 Jul 1942, Miranda 2119 (MEXU); Zapotitlán, Zapotitlán de las Salinas, 4 km al NE de Zapotitlán de las Salinas, sobre la Carretera a Tabuacán, 2 Jun 1975, Rzedowski 33231 (CAS, CORD); along Highway 190, 41.5 miles SE from Tehuitzingo, 2 Aug 1975, Torke et al. 381 (MEXU); Zapotitlán, 11 km SW of Tehuacan, on Hwy 125, 30 Jul 1974, Whalen 16 (BH). Querétaro: Cadereyta, entre Nizarrón [ Vizarrón de Montes] y Higuerillas, 23 Aug 1905, Altamirano 1678 (MEXU); Cadereyta, Vizarrón, ca. 10 km al N, 22 Jul 1991, Carranza 3292 (CAS); km 110 hacia al poblado de Camargo, carretera federal 120, 7 Jul 2000, Lira Charco et al. 1379 (MEXU); Peñamiller, Nacimiento, ca. 1.5 km de La Higuera rumbo al Balneario El Oasis, 12 Jun 2000, Ocampo et al. 874 (MEXU); Peñamiller, Mex. 120 entre Camargo y Peña Blanca, 28 Sep 2016, Ochoterena et al. 976 (MEXU); Tolimán, carretera San Pablo-Higuerillas, 17 Jun 1992, Orozco H. et al. 9957 (MEXU); Cadereyta, cerros calizos al este de Vizarrón, 31 Aug 1994, Orozco H. et al. 10757 (MEXU); Pinal de Amoles, orilla del Río Estorax, cerca de Bucarali, 15 Apr 1988, Rzedowski 46431 (MEXU); Peñamiller, 3.5 km al NE de Peñamiller, 25 May 1977, Zamudio 2097 (MEXU); Tolimán, 7 km al N de San Pablo Tolimán, sobre la brecha a Higuerillas, 12 Jul 1977, Zamudio 2212 (MEXU); Tolimán, 2 km al NE de "El Chilar", 16 Jun 1978, Zamudio 2868 (MEXU); Cadereyta, ca. 1 km al N de La Florida, 14 Jul 1997, Zamudio & Zamudio 10249 (MEXU). Quintana Roo: Isla Mujeres, a 1 km al N de El Faro de la punta sur, sobre la carretera perimetral, 8 Jan 1986, Cabrera & de Cabrera 10461 (CIQR, MEXU); Isla Mujeres, camino al Puerto de Abrigo, 1 Nov 1970, Cabrera et al. 17207 (CIQR, MEXU); Cozumel, Cobá, Ruinas de Cobá, 24 Nov 1980, Chan et al. 45 (CIQR, MEXU); Isla Mujeres, a small isle off the coast at Puerto Juarez, along gravel road between lighthouse and Garrafón beach on the S end of the isle, 11 Jan 1979, Doebley & Sager 203 (CAS, MEXU); Cozumel, Cozumel Island, coast of Yucatan, 11 Jun 1885, Gaumer 135 (K); José María Morelos, Chichankanab, Gaumer 1708 (BM, CAS, K); Cozumel, Cobá, parque arqueológico y natural Cobá, entre el camino y el tipo de pinturas, 21 May 1977, López Franco & Villers 1100 (MEXU); Cozumel, carretera Tulúm-Cobá, 10 km antes de llegar a Cobá, 19 Feb 1976, Moreno 494 (MEXU); Cozumel, Cobá, Grupo Coba, 20 Aug 1981, Rico-Gray & Chan 421 (MEXU); Isla Mujeres, a 6 km al SE de la cuidad de Isla Mujeres, a 200 m del Faro despues de Garrafón, 11 Feb 1980, Téllez V. & Cabrera 1576 (BM, MEXU); Cobá, orilla de Laguna Cobá, 18 Oct 1980, Téllez V. et al. 766 (MEXU); Solidaridad, Cobá, zona arqueologica, 9 Sep 1980, Ucán Ek 394 (CICY). San Luis Potosí: Tampamolon, 1 km N of Tampamolon, 12 Oct 1978, Alcorn 2028 (MEXU); Rioverde, ca. 27 km del entronque a la Presa El Realito, 19 Feb 2010, Guzmán C. 3279 (K). Sinaloa: Culiacán, La Lima, ca. 1 km al NE de La Lima, 12 Feb 1984, Aguayo C. & Gómez R. 2 (MEXU); Escuinapa, orilla de estero Navajas, ca. 8 km de Isla de Bosque, 5 Jan 1985, Aguiar H. 59 (MEXU); Badiraguato, Ojito de Agua, ca. 3 km de Huajote, carretera Badiraguta-Surutato, 7 Mar 1995, Armenta et al. 132 (MEXU); Cofradía, 7 Apr 1934, Bailey 21 (BH); Mazatlán, 7.4 km N of the Tropic of Cancer on road from Mazatlán to Culiacán, 31 Oct 1985, Bartholomew et al. 3643 (CAS, MEXU); Salvador Alvarado, cerros al SE de Tabullal a ca. 5 km, 15 Aug 1988, Bojórquez B. 721 (MEXU); Morcocito, Terreros, 3 miles N, ca. 48 miles N of Culiacán, 28 Jan 1962, Breedlove 1511 (DS, DUKE); Badiraguato, 28 km E of Badiraguato, 20 Aug 1986, Breedlove & Anderson 62857 (CAS); Cosalá, ca. 12 km al NO de Cosalá, 18 Oct 1986, Carrasco et al. 151 (MEXU); Agua Caliente de Zevada, por la casa de Silveiro Perez, Chapiro 41 (MEXU); Culiacán, km 27 carretera Culiacán-Guamuchil, Ejido La Campana, 29 Sep 1984, Chávez Montes & Niebla Armenta, 82 (MEXU); Calmoa, Río Fuerte, 21 Nov 1933, Gentry 930 (DS); Bahia Topolobampo, Cerros de Navachiste about Bahia Topolobampo, 26 Sep 1954, Gentry14362 (MEXU); Mazatlán, zona ganadera del poblado El Quelite, desviación km 33 carretera Mazatlán-Culiacán, 2 Nov 1994, González E. s.n. (MEXU); Elota, La Roble, Sind. La Cruz, González Ortega 56 (MEXU); Mazatlán, Urias, González Ortega 5609 (K, MEXU); Mazatlán, Mazatlán, Jul 1934, González Ortega 7249 (CAS); Elota, Mexican Highway 15, 7 miles N of the bridge over Río Piaxtla, ca. 87 km N of Mazatlán, 16 May 1973, Hansen et al. 1402 (BH, MEXU); Cosalá, Mineral de Nuestra Señora, 22 Jan 1988, Hernández A. & Hernández V. 567 (MEXU); Guasave, carretera 15 (libre) km 81, 1 km al S de Terrero, entre Mazatlán y Guamúchil, camino de terracería hacia El Progreso, menos de 1 km al E de la carretera 15, 30 Sep 2007, Igic & Vallejo-Marín 07s - 20 (MEXU); Culiacán, La Divisa km 6 al N de la carretera Culiacán-Sanalona, 25 Feb 1984, López Felix & Antio 16 (MEXU); Culiacán, 23 km al N de Culiacán camino a El Barco, 30 Jul 1983, Martínez S. et al. 4083 (MEXU, NY); Salvador Alvarado, along Hwy 15 about 59 miles N of Culiacán, 22 Dec 1971, Norris et al. 20104 (CAS, MEXU); Culiacán, carretera Culiacán-Guamuchil km 15, 12 Oct 1984, Ochoa Avalos & Bojórquez B. 188 (MEXU); Culiacán, El Chaparral, 30 Sep 1994, Pineda C. 477 (MEXU); Cosola, junction between Mazatlán and Culiacán, 22 Feb 1970, Roderick et al. 3136 (CAS); Culiacán, Costa Rica, km 5 carretera Costa Rica-C. Internacional, 18 Feb 1984, Rosas J. & Sánchez G. 1 (MEXU); Imala, 20 miles NE of Culiacán, about 1-2 miles beyond Cofradia along artificial lake, 24 Apr 1959, Thomas 7683 (CAS, DS, MEXU); Culiacán, La Loma de Rodriguera, camino a Las Guasimas, ribera del Río Humaya, 4 Feb 1984, Torres et al. 21 (MEXU); Culiacán, Dique La Primavera, ca. 8 km al SE de Culiacán, 14 Feb 1985, Vega Aviña & Bojórquez B. 1467 (MEXU); Mazatlán, El Venadillo, ca. 5 km al N de Mazatlán, 5 Aug 1985, Vega Aviña 1595 (MEXU); Ahome, San Miguel de Zapotitlán, al pie de Barobampo, 2 Mar 1999, Vega Aviña & Gutierrez 9884 (MEXU); E of Playa Escondida, near Mazatlán, 20 Dec 1974, Webster 19916 (MEXU); along Hwy 150, 20 miles N of Rosario, 27 Jul 1975, Whalen 182 (BH); Mocorito, Pericos, 8 miles S, 17 Mar 1955, Wiggins 13150 (DS, MEXU). Sonora: Alamos, 25 miles E of Navajos on road to Alamos, 19 Feb 1953, Blakley 3-1652 (DS); Cañada Tetabejo, Sierra Libre, al S de la Cuidad de Hermosillo, por la Carr. Fed. 15, 8 Aug 1995, Búrquez 95-154 (MEXU); Cañon el Abolio, Sierra Libre, al S de la Ciudad de Mermosillo, por la car retera Federal 15, 16 Jan 1996, Búrquez 96-15 (MEXU); Cañón Tepoca, km 177 a los lados de la Carr. Fed. 16, 7 Sep 1996, Búrquez et al. 96-960 (MEXU); Sahuaripa, región de Los Mulatos, Cerro La Estrella, 1 km al SE del pueblo Mulatos, 11 Sep 1996, Búrquez & Yetman 96-1127 (MEXU); Baviacora, 40 km al NE de Ures, camino a Moctezuma, 22 Jul 1978, Castellanos 206 (MEXU); Cumpas, Jécori, in ravine 1 mile N of Jécori, 20 Nov 1939, Drouet et al. 3695 (DS); Arivechi, 2 km al NW de Arivechi, carr. a Sahuaripa, 27 Sep 1996, Flores Martínez & Arvizo Y. 4751 (MEXU); Alamos, Río Mayo, San Bernardo, 10 Feb 1955, Gentry 1294 (K, MEXU); La Tinaja, 19 Nov 1890, Hartman 243 (K); Bacanora, 18 miles by road E of bridge of Río Yaqui at El Novillo, 27 Oct 1965, Hastings & Turner 65-197 (DS); Alamos area, SE side of Cerro Prieto, W of Alamos, 29 Dec 1989, Joyal 1339 (MEXU, NY); Las Cabras, ESE of Alamos, 13 May 1990, Joyal 1423 (MEXU); La Palmita, between Bacadehauchi and Nacori Chico, Tedehuachi drainage into Río Bavispe, 6 Feb 1991, Joyal 1520 (MEXU); road from Hermosillo to Sahuaripa, 3.8 mi E of Río Yaqui, 21 Feb 1987, Landrum et al. 5409 (MEXU, NY); Onavos, Rancho la Mula, 5 km W of Agua Amarilla, 28.2 km SW of Río Yaqui on Mex. 16 (km 195 E of Hermosillo), 27 Mar 1997, Reina G. & Van Devender 97-297 (CAS, MEXU); Huatabampo, ca. 6 km NW of Camahuiroa on road to Las Bocas, near Bachomojaqui, 24 Nov 1998, Reina G. et al. 98-2123 (MEXU, NY); Madgalena, Río Madgalena at Tubutama road, 23 Jul 2001, Reina G. & Van Devender 2001-598 (MEXU); Navajoa, 20 Jan 1931, Souviron & Erlanson 27 (BH); Alamos, estación de microondas, Cerro Prieto, 15 km al DE de Navojoa, carretera a Alamos, 1 Oct 1983, Tenorio L. & Torres C. 4644 (MEXU); San Javier, km 135 carretera 16 a 10 km al NW de al desviación al pueblo, 29 Aug 1996, Varela E. 96-184 (MEXU); San Javier, Cerro El Halcón ladera N a 3.5 km al NE del poblado camino brecha hacia el Cañón del Aliso, 11 Jan 1997, Varela & Cuamea 97-01 (MEXU); La Colorada, Colorado, 7 miles NE between Colorado and Mazatán, 6 Sep 1941, Wiggins & Rollins 326 (DS, NY); Mazatán, Mazatán, 5 miles S between Mazatán and Colorado, 7 Sep 1941, Wiggins & Rollins 382 (DS, NY); Cajeme, small valley in foothills 17 miles NE of Cajame on road to Tesopaco, 3 Mar 1933, Wiggins 6393 (DS); Ures, W side of mountains 6 miles N of Ures on road to Babiacari, 20 Sep 1934, Wiggins 7333 (DS). Tamaulipas: Aldama, 13 km NE of Aldama on road to Barra del Tordo, 22 Sep 1981, Fryxell 3706 (BH, MEXU, NY); Aldama, entre Rancho Nuevo y San Vicente, 28 Jul 1970, González-Medrano et al. 3122 (MEXU); Aldama, Ejido El Nacimiento, 5.5 km al S de Aldama, 31 Aug 1984, Hernández & Romo 1174 (MEXU); Soto de la Marina, Campamento Tortugero La Pesca, 4 Apr 2007, Martínez S. 39209 (MEXU); Aldama, 18 km E de la carretera Aldama-Barra del Tordo, 27 Jun 1992, Mora-López 189 (MEXU); Aldama, 17.2 km al NW de la desviación a Rancho Las Yucas, 26 Jun 1983, Torres C. 3141 (MEXU, NY); Aldama, 5 miles N or ca. 75 miles N of Tampico, 15 Jul 1984, Wilbur 35210 (DUKE). Veracruz: Actopán, alrededores de Laguna de la Mancha, 28 Nov 1975, Acosta & Dorantes 663 (MEXU, NY); Emiliano Zapata, 15 miles SE of Xalapa, 3 Aug 1947, Barkley et al. 2616 (CAS); Carrizal, 0.5 km de la desviación a Carrizal, por la carretera Jalapa [Xalapa]-Veracruz, 20 Jul 1975, Calzada 1837 (MEXU); Veracruz, Colonia Las Amapolas, en el km 100 de la carretera Xalapa Veracruz, 13 May 1977, Calzada 3182 (MEXU); Vega de Alatorre, Cañada de Mesillas entrada por Santa Ana, 24 Jul 1981, Calzada 7725 (MEXU); Actopán, Playa Paraíso, 0.5 km antes de la playa, 6 Sep 1977, Castillo Campos et al. 189 (MEXU); Coatepec, Cerro de las Palmas, 2 km antes de Jalcomulco en las faldas del Cerro de las Palmas, 20 May 1979, Castillo Campos & Tapia 700 (MEXU); Apazapan, 1 km E del pueblo, en las faldas (N) del Río Jalcomulco, 10 Oct 1991, Castillo Campos & Birke B. 6971 (MEXU); El Remuladera, carrretera Conejos a Huatusco, 13 Jul 1970, Chavelas P. ES- 971 (MEXU); Xalapa, km 13 carretera Jalapa[Xalapa]-Veracruz, 5 Dec 1985, Chehaiber & Espejo 201 (MEXU); entre rancho del Diamante y el rancho del Capulin, camino a Misantla, 3 Nov 2001, Cruz Durán et al. 5483 (MEXU); Actopán, La Mancha, carretera Cardel-Nautla, 1 Aug 1971, Dorantes 251 (CAS, MEXU); Emiliano Zapata, Barranca de San Antonio al E de Corral Falso, carretera Xalapa-Veracruz, 17 Aug 1971, Dorantes 260 (CAS, MEXU); Alto Lucero de Gutiérrez Barri, W de Laguna Salada, 26 Jun 1972, Dorantes 1033 (CAS, MEXU); Actopán, Sierra Manuel Diaz, 11 Jul 1972, Dorantes 1333 (CAS, MEXU); Alto Lucero, alrededores de Laguna Verde, 18 Nov 1975, Dorantes et al. 5209 (MEXU); Camarón de Tejeda, Matta de Agua, 13 Aug 1926, Fisher s.n. (DS); Veracruz, Villarin, 5 km al W del Puerto, 2 Jul 1990, García Bielma 536 (MEXU); a 4 km de Paso de Macho, hacia Camarón, 12 Jul 1970, González G. 133 (MEXU); La Antigua, Salmoral, camino a los cañales, a 1 km de Salmoral, 6 Apr 1988, González H. 267 (MEXU); Veracruz, Rancheria Neveria, carretera antigua nacional Xalapa-Veracruz km 90.5, 10 Sep 1981, Gutiérrez B. 723 (MEXU); Soledad de Doblado, Paso de Conejos, [Paso de los Cedros], 6 Aug 1966, Hernández Magaña 207 (DS, MEXU, NY); Colipa, 1841, Karwinski 1399 (LE); Cuitláhuac, along route 150 just NW of Cuitláhuac, 6 Jun 1960, King 2669 (DS, NY); Veracruz, along route 180, about 14 miles S of Veracruz, 7 Jun 1960, King 2701 (DS, NY); Puente Nacional, Jun 1838, Linden 243 (K); Emiliano Zapata, carretera Rinconada-Buenavista, 13 Oct 1999, Lizama M. 1479 (MEXU); hacia el Plan de Las Hayas km 739-761, 13 Aug 1969, Lot 454 (CAS, MEXU); Actopán, La Mancha, desviacion de la carretera J. Cardel a Nautla, cerca de Ursulo Galvan, 29 Nov 1969, Lot 483 (CAS, MEXU); Veracruz, Santa Rita, 600 m al NW de la estación de Santa Rita (agricultura prehispanica), 19 Oct 1987, Luna M. 36 (MEXU); Veracruz, 1 km de Santa Elena, 13 May 1988, Luna M. & Zola B. 262 (MEXU); Emiliano Zapata, Palmar, and vicinity, 3 Sep 1935, MacDaniels 443 (BH); 15 km S of Vega de la Torre, 18 Nov 1963, MacKee, H.S. 10974 (K); Emiliano Zapata, alrededores del poblado de la Balsa, 25 May 1982, Marquez, M. 1095 (MEXU); 10 km S of Nautla, 18 Nov 1963, McKee 10974 (MEXU); Paso de Ovejas, 1 km al N de Cantarranas, 25 Apr 1985, Medina A. & Acevedo R. 24 (MEXU); Totutla, along Hwy 144 W of Mata Oscura, km 36-37, 25 Jul 1965, Mertz 148 (MEXU); Puente Nacional, 6 km E of Banos de Carrizal (downstream), on N side of valley of Río Chico [upper part of Río de La Antigua], 8 km ESE of Emiliano Zapata [=Carrizal], 17 Apr 1983, Nee & Taylor 26632 (BH, BM, CAS, CORD, MEXU, NY); 20 km al N de Martinez de la Torre, rumbo a Papantla, 5 Sep 1967, Nevling & Gomez-Pompa 544 (MEXU); Juchique de Ferrer, 22 km de Palma Sola, rumbo a Plan de las Hayas, 16 Apr 1969, Nevling & Gomez-Pompa 1043 (MEXU); Cardel, a 500 m del puente del Río La Antigua, 27 Oct 1988, Orea L. 153 (MEXU); Actopán, 5 km antes de la desviación a san Nicolas, 5 Aug 1976, Ortega 427 (K, MEXU); Rancho La Palmilla, Oct 1929, Purpus s.n. (DS); near Rancho Camarones, Feb 1930, Purpus 11048 (K); Emiliano Zapata, Chavarillo, 22 Oct 1930, Reddick 157 (BH); Axocuapán, camino Coetzalán-Cueva del Abono, Ejido de Coetzalán, 14 Jan 1984, Robles H. 397 (MEXU); Acultzingo, Cumbres de Acultzingo 30km carretera Orizaba-Tehuacan, D-8-50, 30 Nov 1967, Rosas R. 876 (BM); Emiliano Zapata, La Laja, entre Corral Falso-Pinoltepec, a 900 m de al carretera Jalapa [Xalapa]-Veracruz, desv. a 16 km a SE de Jalapa [Xalapa], 20 Sep 1975, Sousa S. & Ramos 4836 (MEXU); La Laja-Pinoltepec, 6 Mar 1976, Trejo, L. 13 (MEXU); Ursulo Galvan, 31 Mar 1971, Ventura A. 3385 (DS); Dos Ríos, Palo Gacho, 17 Sep 1974, Ventura A. 10560 (MEXU); Dos Ríos, Carrizal, 25 Jul 1975, Ventura A. 11718 (MEXU); La Antigua, 700 m antes de la playa (agricultura prehispanica), carretera Cardel-Veracruz, camino de terraceria que sale a la playa a 500 m antes de la caseta de cobro, 21 Sep 1987, Zamora C. 598 (MEXU); Emiliano Zapata, carretera a Lencero, 28 Jul 1976, Zola B. 578 (MEXU); Paso de Ovejas, carretera El Faisán-La Víbora, 500 m de El Faisán, 16 Jun 1987, Zola B. & Mota 2135 (MEXU). Yucatán: a 14 km al O de Chemax, sobre la carretera Valladolid-Cancún, 16 Jul 1985, Cabrera & de Cabrera 8895 (CIQR, MEXU); Mayapan, alrededores de la zona arqueológica de Mayapan, a 1 km al S de Telchaquillo, carretera Tecch-Oxkutzoab, 21 Jul 1985, Cabrera & de Cabrera 9130 (CIQR, MEXU); a 2 km al S del crucero Las Coloradas-San Felipe, sobre la carretera Tizimín-Río Lagartos, 20 Dec 1985, Cabrera & de Cabrera 10064 (MEXU); 10 km al E de Telchak Puerto, sobre la carretera Pto. Progreso-Dzilam de Bravo, 23 Jan 1986, Cabrera & de Cabrera 10755 (MEXU, NY); a 4 km de Telchak Puerto sobre el camino a Telchak Pueblo, 18 Apr 1986, Cabrera & de Cabrera 11289 (CIQR, MEXU); 2 km al O de Ticuch, sobre la carretera 180 tramo Cancún-Valladolid, 21 Jul 1986, Cabrera & de Cabrera 11530 (MEXU); 3 km al E del crucero Río Lagartos-San Felipe, sobre el camino a Las Coloradas, 22 Jul 1986, Cabrera & de Cabrera 11566 (MEXU); Gruta de Blanacanche, 32 km al O de Valladolid, en los alrededores de la gruta, 23 Jul 1986, Cabrera & de Cabrera 11625 (MEXU); Progreso, 6 km al O del Puerto Progreso, sobre el camino a Yucalpeten, 24 Nov 1986, Cabrera & de Cabrera 12835 (CIQR, MEXU); Motul, 4 km al O de Motul, sobre el camino a Mérida, 21 Mar 1988, Cabrera & de Cabrera 15787 (CICY, MEXU); carretera Komchén-Cosgaya, 28 Apr 1992, Campos & Simá 2800 (MEXU); Progreso, km 10-16 carretera Sierra Papacal a Chuburná Puerto, 28 Apr 1992, Campos & Simá 2836 (MEXU); Muna, km 8 carretera a Opichén, 24 Sep 1984, Chan 3983 (CICY, MEXU); Río Lagartos, entronque de la carretera Río Lagartos a Las Coloradas, 13 Mar 1985, Chan 4835 (MEXU); Ticul, Hacienda Yokat, camino Muna-Ticul, 29 Apr 1985, Colunga 155 (MEXU); Cuzamá, 4 km al S de Chunkanán, en un cenote, 19 Nov 1995, Dorantes & Ek 34 (CICY); Hunucmá, 6 km de Hunucmá rumbo a Sisal, 10 Feb 1992, Durán et al. 1581 (MEXU); Mérida, junto a la discoteca Bin Bon Beach, 26 Jun 1992, Durán & Méndez 1626 B (CIQR, MEXU); San Felipe, 100 m al E del desvio de San Felipe a Río Lagartos, 1 Jul 1999, Durán et al. 3328 (MEXU); Tekon, 30 Sep 1955, Enríquez 98 (MEXU); Chuburná, 18 Jul 1956, Enríquez 686 (MEXU); 7 km S Xtampu to San Diego Guerrera, 18 Feb 1991, Escalante R. et al. 240 (K); Mérida, Dzilbilchaltún, 27 Sep 1982, Escalante 672 (CIQR, MEXU); Sinanché, a 13 km de Telchac Puerto camino a Dzilam, 17 Oct 1980, Espejel 99 (MEXU); Abalá, 2 km al W de Mucuyché, 31 Mar 1993, Feliciano Kú y Yam 254 (MEXU); Telchac Puerto, 10 km al W de Telchac Puerto, 9 Dec 1996, Feliciano Kú y Yam 626 (MEXU); Izamul, Gaumer 366 (BM, CAS, DS, E, G, K, LE, NY, W); Dzidzantún, Mina de Oro, May 1916, Gaumer and sons 23325 (K, NY); Yaxcabá, Stancabxonot, 1917, Gaumer 23622 (LE); Dzoncauich, a 10 km de Chacmay, 12 Apr 1983, Góngora 301 (MEXU); Tixkokob, cerca del poblado Ruinas de Ake, 28 Jun 1983, Góngora 694 (CICY, MEXU); Tinum, 1 km al E de Chichen Itza, 30 Aug 1991, Méndez et al. 261 (MEXU); Mérida, Sierra Papacal, 2 km al N, 22 Apr 1993, Méndez et al. 812 (MEXU); Mérida, ruinas de Dzibilchaltun [Chablekal], 4 Apr 1981, Narváez et al. 339 (CIQR); Sacalum, camino Sacalum-Mérida, 10 Jul 1981, Narváez & Puch 590 (CICY); Mérida, Opichén, Hcda. Xixim, 10 Feb 1983, Orellana 119 - BB (MEXU); Cantamayec, 6 km S Tixcalcupil, 5 Mar 1996, Ortíz M. s.n. (MEXU); entrada a la zona arqueológica de Mayapán, 11 Mar 1999, Peña-Chocarro & Tun 407 (BM, MEXU); Dzemul, 4 km al S de Dzemul, 4 Sep 1991, Pérez Lara 94 (MEXU x2); Buctzotz, a 15 km al E de Buctzotz, 5 Jun 1992, Pérez Lara 186 (MEXU); Abalá, 1 km al W de Hacienda Uayaicén, 30 Oct 1996, Pérez Lara 612 (MEXU); Tecóh, 1 km al S de Lepan, 2 Aug 1999, Pérez Lara 860 (MEXU); Hunucmá, 10 km al SE de Sisal, 9 Nov 1999, Pérez Lara 892 (MEXU); Abalá, 4 km al E de Tenozón, 14 Aug 1979, Pérez S. et al. 318 (MEXU); Oxkutzcab, salida de Oxkutzcab yendo a Lol-Tun, 7 Jul 1981, Puch & Narváez 447 (CIQR, MEXU); Temox, km 17 carretera Izamal-Temax, Dzoncauich, 25 Apr 1984, Puch & Ortiz 1319 (CICY); Hunucmá, al 8 km al S de Hunucmá, 19 Mar 1992, Reyes de los Santos 147 (MEXU); Tunkás, a 5 km al W de Tunkás, 23 Sep 1993, Reyes de los Santos 272 (MEXU); Timucuy, 2 km al E de Timucuy, 24 Aug 1994, Reyes de los Santos 367 (MEXU x 2); Yaxkukul, 1 km al N de Yaxkukul, 30 Aug 1995, Reyes de los Santos 477 (MEXU); Panabá, 2 km al S de Panaba, 25 Apr 1996, Reyes de los Santos 545 (MEXU); Chicxulub Pueblo, 5 km al N de Chicxulub, 2 Dec 1998, Reyes de los Santos, E. 788 (MEXU); Peto, 1 km al N de Peto, 30 Sep 1999, Reyes de los Santos 871 (MEXU); a 1.5 km al S de Conkal, 7 Jun 1984, Rivera 90 (CIQR); Xpacamul, Maya, Mérida, 29 Jul 1865, Schott 310 (BM); Tinum, Chichen Itzá, 8 Jun 1932, Steere 1065 (BM, MEXU); Dzemul, km 6 de al carretera Dzemul-Xtampú, 4 km S del entronque a las ruinas de Xtampú, 3 Dec 2004, Tapia et al. 1533 (MEXU); Baca, 8, 5 km al N de Baca, camino a ranchos ganaderos, 10 Dec 1998, Tun & González-Iturbe 443 (MEXU); Tepakan, carrretera Izamal sobre km 6 rumbo a Tepakan, 18 Jan 1982, Ucán Ek 1826 (MEXU); Valladolid, Pixoy, calle N de la plaza principal, 12 Oct 1983, Ucán Ek et al. 3021 (CIQR, MEXU); Mérida, Parque Científico, a 5 km de Sierra Papacal, 5 km al oeste, 14 Aug 2013, Uh Cauich et al. 4 (MEXU); Yaxcabá, Tixcacaltuyub, 27 Jun 1980, Vargas 68 (MEXU); Chac-May, 5 Dec 1985, Waizel Bucay s.n. (MEXU); 10 km N of Sinanche, 1 km E into Rancho Chumhabin [Chum Habin], 15 Feb 1994, Way 107 (CICY, K); Chicxulub Pueblo, ca. 4 mi SE of Chicxulub Puerto, 30 Jul 1972, Webster et al. 17551 (MEXU); N slopes of hills S of Ticul, 22 Feb 1982, White & Mott 96 (MEXU). Zacatecas: Moyahua, Los Otates, 20 Aug 1992, Enríquez E. 114 (MEXU); Apozol, desviación presa Achoquén (projecto Cañon de Juchipila), 3 Aug 1995, Enríquez E. 465 (MEXU); Juchipila, Apozol, 20 km al N de Juchipilas, 16 Aug 1984, Hernández Magaña et al. 9027 (MEXU).

Figures

Figure 2. Flowers of members of the Elaeagnifolium clade. A Solanum elaeagnifolium, with divergent anthers of approximately equal size and shape (Argentina, Mendoza, Knapp et al. 10470) B Solanum homalospermum (Argentina, Catamarca, Chiarini et al. 505) C Solanum hindsianum (cultivated in Arizona) D Solanum houstonii, hermaphroditic flower and strongly curved buds (Mexico, Yucatan, Pena-Chocarro et al. 407) E Solanum houstonii, staminate flower (Mexico, Queretaro, Ochoterena et al. 976) F Solanum mortonii (cultivated in Cordoba; from Barboza et al. 644). Photographs A, D, E by S. Knapp; B, F by F. Chiarini; C by Eugene Sturla.

Figure 3. Fruits of members of the Elaeagnifolium clade. A Solanum elaeagnifolium, immature berries with green mottling (Argentina, Cordoba, Barboza et al. 3434) B Solanum elaeagnifolium, mature fruit with sticky tan seeds (Argentina, Cordoba, Barboza et al. 3435) C Solanum mortonii, immature fruit exerted from the calyx (Argentina, Catamarca, Barboza et al. 3439) D Solanum mortonii, mature fruit exerted from the calyx (Argentina, Catamarca, Barboza et al. 3438) E Solanum houstonii, immature fruit enclosed in accrescent calyx (Mexico, Queretero, Ochoterena et al. 976) F Solanum houstonii, mature fruit enclosed in accrescent calyx with black seeds (Mexico, Queretero, Ochoterena et al. 976). Photographs by S. Knapp.

Figure 4. Habitats of members of the Elaeagnifolium clade. A Solanum elaeagnifolium, old stems with persistent berries and young new stems from underground rhizomes (Argentina, Mendoza, Knapp et al. 10470) B Solanum houstonii on rocky slope (Mexico, Queretaro, Ochoterena et al. 976) C Solanum mortonii, plant dug up showing extensive underground stems (Argentina, Catamarca, Barboza et al. 3437) D Solanum mortonii, large population on rocky slopes (Argentina, Catamarca, Barboza et al. 3437). Photographs by S. Knapp

Figure 10. Solanum houstonii Martyn. A Whole plant B Leaf with prickly petiole and stellate trichomes C Lobed leaf D Prickly stem with stellate trichomes E Bud F Staminate flower G Ovary and style of staminate flower H Anthers of staminate flower I Hermaphroditic flower spread J Gynoecium of hermaphroditic flower K Anther of hermaphroditic flower L Fruits M Seed. Drawn by Bobbi Angell A, E from Fryxell 3706 (NY 00751643) B from Duran et al. 4737 (NY 00751647) C from Reina G. 98 - 2082 (NY 00751617) D from Palmer 1243 (NY 00751622) F-H from Reina G. 98 - 2123 (NY 00751616) I-K from Cabrera 10755 (NY 00751572) L from Cabrera 11000 (NY 00751567).

Figure 11. Lectotype of S. tridynamum Dunal. Unpublished drawing by T. F. Node-Veran (1773 - 1852) based on an illustration of Sesse and Mocino (MPU 310713). Reproduced with permission of the Universite de Montpellier - Herbier MPU (Service de Patrimoine Historique); copyright Universite de Montpellier - Herbier MPU (SPH).

Figure 12. Distribution of S. houstonii and S. mortonii.