Solanum aligerum Schltdl.

Knapp, Sandra, 2013, A revision of the Dulcamaroid Clade of Solanum L. (Solanaceae), PhytoKeys 22, pp. 1-432 : 37-47

2. Solanum aligerum Schltdl. , Linnaea 19: 301. 1847. Figure 10

Solanum pterocladum van Heurck & Müll.-Arg., Observ. Bot. 44. 1870. Type: Bolivia. La Paz: Prov. Larecaja, Sorata, 3000 m, May 1858, G. Mandon 415 (lectotype, designated here: G [G00016904]; isolectotypes: AWH [n.v.], F [F-760412, F-876198, F-1588536], G-DC [Morton neg. 8550], GH [GH00077740], BM [BM000815930], K [K000590235], NY [NY00172146], P [P00445070, P00445071, P00445072], S [S04-2969]).

Solanum manicatum Bitter, Bot. Jahrb. Syst. 50, Beibl. 111: 63. 1913. Type: Peru. Ayacucho: Prov. Huanta, road to Tambo above Osno, Río Apurímac, 2600-2700 m, A. Weberbauer 5643 (holotype: B [F neg. 2620], destroyed; lectotype, designated here: MOL; isolectotypes: F [F-647977, frag.], G [G00016963], GH [GH00077711]).

Solanum dotanum C.V. Morton & Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1079. 1938. Type: Costa Rica. San José: Laguna de La Chonta, NE of Santa María de Dota, 2000-2100 m, 18 Dec 1925, P. Standley 42265 (holotype: US [US-1252694]).

Solanum grossum C.V. Morton, Revis. Argentine Sp. Solanum 178. 1976. Type: Argentina. Tucumán: Dpto. Chicligasta, Estancia Las Pavas, 2000 m, 22 Nov 1926, Solanum Venturi 4632 (holotype: US [US-1548932, barcode US0027006]; isotypes: F, GH [GH00077675, GH00077676], NY [NY00172009, NY0017210], SI, US [US-1343321, barcode US01014178]).

Type

. Mexico. Michoacán: Angangueo, Oct, C. Schiede s.n. (holotype: B, destroyed, no duplicates found). Mexico. Michoácan: Wooded slopes 8-10 miles NW and WNW of Ciudad Hidalgo, among mountains west of Cerro San Andrés and 6-7 miles N of village of San Pedro Aguaro, in steep ravine along brook, 2850-3000 m, 19 48'N, 100 40'W, 18 Mar 1949, R. McVaugh & R.L. Wilbur 9917 (neotype, designated here: MEXU [MEXU-92021]; isoneotypes: BM [BM000578989], NY [NY00961953], US [US-2452307]).

Description.

Small tree or shrub, 1.5-5 m tall, the branches arching. Stems glabrous to densely pubescent with simple or dendritic uniseriate trichomes 0.5-1 mm long, glabrescent, often strongly winged from the decurrent leaf bases, the wings to 0.5 cm wide; new growth pubescent with simple or dendritic uniseriate trichomes 0.5-1 mm. Bark of older stems dark reddish brown, shiny. Sympodial units plurifoliate. Leaves simple, (3.5-)6-15 cm long, 1.2-5 cm wide, narrowly elliptic to lanceolate, membranous to slightly fleshy, the upper surfaces glabrous or sparsely (to occasionally densely) pubescent with simple or less often dendritic trichomes on the veins and lamina, the lower surfaces with dense tufts of dendritic trichomes 0.5-1 mm long in the vein axils, these trichomes occasionally extending along the midrib or to the lamina and the leaves more densely pubescent; primary veins 10-15(-20) pairs, the midrib keeled above, the veins often drying yellowish brown; base attenuate, often winged onto the stem; margins entire, sometimes revolute; apex acuminate; petioles 0.7-2 cm long, glabrous or pubescent like the stems, never twining. Inflorescences terminal or lateral, 4-15 cm long, many times branched, with 10-60 flowers, glabrous to pubescent, the trichomes simple or dendritic, ca. 0.5 mm long, denser at branch tips and at flower insertion points; peduncle 1.5-3.5 cm long; pedicels 1-1.2 cm long, ca. 0.5 mm in diameter at the base, ca. 1 mm in diameter at the apex, slender, erect to nodding, glabrous, articulated at the base and inserted into a short sleeve; pedicel scars mostly clustered at the tips of inflorescence branches on small platforms or irregularly spaced 2-5 mm apart more basally. Buds pointed ovoid or turbinate with an apical nipple, the corolla strongly exserted from the calyx tube before anthesis. Flowers all perfect, 5-merous. Calyx tube 1.5-2.5 mm long, cup-shaped, narrowing abruptly to the pedicel, the lobes 1.5-2 mm long, quadrate with an apiculate tip to 0.5 mm long, the tips densely pubescent with simple uniseriate trichomes, these extending adaxially, the adaxial surface densely papillate. Corolla 1.8-2 cm in diameter, white, often tinged with purple, stellate, lobed 2/3 to 3/4 of the way to the base, the lobes ca. 7 mm long, 5 mm wide, spreading, minutely pubescent abaxially, the tips cucullate, densely pubescent, the sinuses broad and thin, glabrous adaxially. Filament tube minute, the free portion of the filaments 0.5-1 mm long, glabrous; anthers 4.5-5.5 mm long, 1-1.5 mm wide, ellipsoid, loosely connivent, poricidal at the tips, the pores lengthening to slits with age. Ovary glabrous; style 6.5-7 mm long, glabrous; stigma minutely capitate to slightly clavate, the surface minutely papillose. Fruit a globose berry, 1-1.2 cm in diameter, black or greenish black when ripe, the pericarp thin and shiny, glabrous; fruiting pedicels 1.5-1.7 cm long, ca. 1.5 mm in diameter at the base, woody, more or less nodding with the weight of fruits. Seeds 20-40 per berry, 2.5-3 mm long, 2-2.5 mm wide, flattened reniform, reddish brown or pale yellow, the surfaces minutely pitted, the testal cells square, the cell walls disintegrating and the seeds appearing hairy. Chromosome number: not known.

Distribution

( Figure 11 ). Common from Mexico to Argentina, from 1500-3300 m. A gap in the distribution of Solanum aligerum occurs from Panama to Peru (see discussion).

Ecology.

Solanum aligerum has been collected most commonly in pine-oak forests in Mexico and adjacent Central America, and in cloud forests in the Andes.

Common names.

Guatemala: seconillo (Steyermark 34672).

Conservation status.

Least Concern (LC); EOO>50,000 km2 (LC) and AOO>5,000 km2 (LC). See Moat (2007) for explanation of measurements.

Discussion.

Solanum aligerum has the widest latitudinal range of all the New World members of the Dulcamaroid clade, occurring all down the cordilleras of both continents, but with a distinct gap between western Panama and Peru (see below). In spite of this huge range, the species is remarkably uniform in general aspect, but varies considerably in pubescence. In Bolivia, several populations have been found with plants that are uniformly and densely pubescent with dendritic trichomes over all plant parts; at one time I thought these were a new species and annotated some material as such, but in studying the Dulcamaroid clade in detail it has become apparent that these plants are merely pubescence extremes in this widespread species. They agree with other Bolivian material of Solanum aligerum in all other respects. It would be interesting to investigate if this pubescence variation is due to some habitat variation not apparent from herbarium labels.

Solanum aligerum is extremely similar to Solanum pubigerum , with which it broadly overlaps in central Mexico and Central America. The two species can be very difficult to distinguish, but Solanum aligerum has dendritic trichomes concentrated in the vein axils or over the entire lamina, while Solanum pubigerum has simple trichomes found all along the midrib on the leaf undersides. Flowers are larger in Solanum aligerum with the calyx lobes quadrate rather than deltate (this can be difficult to see), and the berries of Solanum aligerum are also larger and usually black or blackish green, rather than red, when ripe. The stems of S. aligerum are often (but not always) prominently winged, while those of Solanum pubigerum are never so. In general the two species appear to not occupy the same forest types where their ranges overlap.

The absence of Solanum aligerum in Colombia and Ecuador could indicate the northern and southern populations are in the process of differentation. This disjunction is unusual, but has also been found in the potato species Solanum morelliforme Bitter that is disjunct between Guatemala and Bolivia ( Simon et al. 2011). Solanum morelliforme populations from south of the disjunction were morphologically (as here in Solanum aligerum ) and molecularly identical. Without the locality being revealed, it is impossible to distinguish a plant of Solanum aligerum as coming from any particular area, so I am confident that these all represent a single taxon. Molecular studies at the population and geographical level may reveal some differentiating characters.

The holotype of Solanum aligerum was destroyed in Berlin and no duplicates have been traced. I have used a widely distributed collection (McVaugh & Wilbur 9917) from the same region of Mexico as a neotype; this typification also honors Rogers McVaugh, whose many years of collecting and study have so improved our knowledge of the flora of Mexico. Many syntypes exist for Solanum pterocladum , as Mandon’s collections were widely distributed and no herbarium was cited in the original publication. I have selected that in G as a lectotype, as that is where Mueller worked.

Specimens examined.

Argentina. Catamarca: Andalgalá, sobre la ruta no. 65, muy cerca del limite ca. de Tucumán, 4 Nov 1967, Hunziker 19509 (CORD); Andalgalá, Esquina Grande, 29 Apr 1915, Jorgensen 88791 (SI, US); Andalgalá, Valle del Río Chacra, 1650 m, 12 Feb 1950, Sleumer 18 (B). Jujuy: Ledesma, Abra de las Cañas, Parque Nacional Calilegua, 1750 m, 27 Nov 2004, Barboza et al. 1070 (CORD); Ledesma, Abra Colorado, 23 km de la entrada del Parque Nacional Calilegua, 16 Feb 2012, Barboza et al. 3549 (BM, CORD); Valle Grande, camino a Valle Grande, Río Jordan, 1400 m, 18 Oct 1964, Cabrera & Fabris 16142 (SI); Capital, Lagunas de Yala, 11 Nov 1978, Cabrera et al. 29778 (MO, SI); Capital, Sierra de Zapia, Mina 9 de octubre, camino a la antena, 16 Nov 1980, Cabrera et al. 32061 (SI); Santa Bárbara, S de El Fuerte, Finca Confines, 17 Nov 1992, Kiesling et al. 8332 (MO, SI); Palpalá, above Quebrada "Los Tomates", on road to Cerro Zapla, 1850 m, 13 Apr 2000, Nee et al. 50757 (NY); Santa Bárbara, Centinela, 11 Dec 1983, Rotman et al. 930 (SI). Salta: Parque Nacional El Rey, Cerro Chañar, 20 Nov 1981, Brown et al. 1638 (NY); Cafayate,1400 m, 24 Nov 1949, Araque M. & Barkley 19Ar 353 (F); Santa Victoria, Los Toldos, Rio Toldos, frente a Quebrada del Astillero, 3-5 km al E del pueblo, 30 Oct 1987, Novara 7100 (CORD, G); Santa Victoria, Los Toldos, cerros al E del pueblo, margen S del Río Toldos, 1700 m, 2 Nov 1989, Novara et al. 9107 (G); Santa Victoria, Camino al Lipeo, a 15 km de Los Toldos, 1800 m, 8 Nov 1975, Schiavone et al. 11901 C. (F, US); Capital, Quebrada de San Lorenzo, Oct 1926, Venturi 5080 (NY, US); Guachipas, Alemania, 1500 m, 16 Dec 1929, Venturi 9934 (US). Tucumán: Tafí, Apeadero Muñoz, 13 Feb 2012, Barboza et al. 3498 (BM, CORD); Trancas, Quebrada de Rearte, 1800 m, 15 Jan 1946, Bellonio 253 (LE, W); Cuesta del Clavillo, algo más abajo de La Banderita, 1750 m, 10 Nov 1952, Hunziker 10005 (CORD); Monteros, camino a Tafí del Valle, km 41-42, 12 Feb 1986, Hunziker 24873 (CORD); Chichigasta, Las Pavas, Oct 1911, Jorgensen 36423 (SI); Chichigasta, Cuesta Rio Cochuna, 1200 m, 28 Jan 1937, O’Donell 18158 (F); Chichigasta, Las Lenguas, cuesta del Rio Cochuma, 1400 m, 3 Nov 1930, Schreiter 6470 (B, SI, US); Tafí, Garabatal, Siambón, 1600 m, 25 Nov 1930, Schreiter 6552 (SI, US); Tafí, La Hoyada, Casa Arce, 1460 m, Nov 1932, Schreiter 8812 (CORD, GH); Tafí, Cumbre de Garabatal (Porteruelo), 1750 m, 23 Oct 1939, Schreiter 9809 (F, GH); Burruyacú, Cerro del Campo, 1800 m, 14 Dec 1928, Venturi 7755 (US); Tafí, cerca de Taficillo, 1200 m, 5 Oct 1929, Venturi 9609 (LE, NY, US).

Bolivia. Chuquisaca: Punilla, Guerrabamba, 3200 m, Feb 1949, Cárdenas 4136 (US); Azurday, La Angostura, ca. 15 km de Azurduy hacia el cañon de la Angostura, 2500 m, 11 Jan 2004, Huaylla & Guachalla 676 (K); Azurday, Tarvita, 2 km SW on Tarabuco-Azurday road, 2750 m, 25 Sep 1991, Kessler 3235 (GOET); Oropeza, entre el km 11 y Ravelo, 3300 m, 19 Nov 1993, Kiesling & Metzing 8425 (NY); Oropeza, Maragua, cercanias de Sucre, 3062 m, 4 Aug 2007, Velayos et al. 11136 (MA); ca. 8 km from Punilla towards Ravelo, 3300 m, 2 Oct 1994, Wood 8708 (K, NY); Oropeza, 2 km de la cruce a Potolo y Punilla, 2994 m, 29 Mar 2005, Wood et al. 22011 (K). Cochabamba: Carrasco, Montepunco, entrando por los Yungas, 2700 m, 18 Oct 1984, Beck 8931 (F, NY); Carrasco, ca. 75.5 km E of Epizana on Carretera Fundamental 4, 2439 m, 21 Nov 1976, Davidson 5095 (MO, US); Carrasco, Serrania Siberia, 20-35 km W of Comarapa (Prov. Santa Cruz) on the old Cochabamba-Santa Cruz Road (Hwy 4), 2000 m, 14 Jan 1990, Dorr & Barnett 7009 (MO, NY, US); Carrasco, Aguarrica, a 43 km de Epizana por la carretera que conduce a Santa Cruz, cerca de Aguarrica, 3030 m, 26 Dec 1982, Fernández Casas 7793 (MA, NY); Ayopaya, 10 km Cocapata-Cotacajes, 2750 m, 8 May 1997, Kessler et al. 9340 (NY); Carrasco, 5 km (by air and road) SE of bridge at Lopez Mendoza, 19 km by road NW of Epizana, on road from Comaropa to Cochabamba, 2900 m, 11 Feb 1987, Nee & Solomon 34093 (CORD, MO, NY, US); Carrasco, 5 km (by air and road) SE of bridge at Lopez Mendoza, 19 km by road NW of Epizana, on road from Comaropa to Cochabamba, 2900 m, 11 Feb 1987, Nee & Solomon 34093 (NY); Carrasco, narrow canyon of Rio Monte Puncu, 5 km NE of Monte Puncu, 10 km (by air) NW of Epizana, 2700 m, 10 Mar 1988, Nee & Solomon 36607 (CORD, NY, US); Chapare, trail leading NW from the road to tablas Monte along Río Corani Mayu, 2280 m, 2 May 1994, Ritter 919 (GH, MO); Chapare, 23.8 km N of Colomi (junction of the road to Candelaria) on road to Chapare, then 2.2 km NW (left) on side road. Upper Rio Cayani, 2700 m, 19 Oct 1985, Solomon 14386 (G, MO, NY); Chapare, km 104 del camino a Chapare, 3100 m, 2 Dec 1966, Steinbach 563 (F, MO, NY, S, US); Carrasco, Totora-Duraznillo, 2700 m, 30 Mar 1920, Steinbach 3826 (NY); Chapare, Sacaba, Cerro de Incachaca, 2500 m, 4 Sep 1921, Steinbach 5752 (F, G, MO, NY); Pojos, 2600 m, 5 Nov 1928, Steinbach 8621 (BM, E, MO, NY, S); Totora-Pocona, 2700 m, 7 Nov 1928, Steinbach 8685 (BM); Chapare, Incachaca-La Aduana (Sanctutonis), 2700 m, 8 Mar 1929, Steinbach 9550 (BM, E, F, G, MO, NY, S); Chapare, Durazno, Quebrada de Coraní, 2800 m, 18 Jun 1928, Steinbach 9854 (BM, E, F, G, MO, NY, S); Ayopaya, Independencia, c. 1 km before La Mina, 38 km north of Independencia along road to Sailapata, 2856 m, 15 May 2002, Wood et al. 8479 (BOLV, K); Chapare, descent from the dam towards Villa Tunari on road from Cochabamba to the Chapare, 3000 m, 25 Jun 1994, Wood 8545 (K, LPB). La Paz: Larecaja, Laripata, 2995 m, 13 Aug 2007, Aedo et al. 14644 (MA); Larecaja, Sorata, 2439 m, Feb 1886, Rusby 781 (NY); Murillo, Zongo, 2760 m, 15 Aug 2007, Aedo et al. 14734 (MA); Inquisivi, Rio Calachaca Jahuira-down river 1 km from Aguas Calientes de Calachaca, 10 km NE of Choquetanga, 3200 m, 17 Jul 1991, Lewis 39251 (G, MO); Larecaja, along road to Consata, ca. 4-15 km above (N of) Sorata, 2896 m, 24 May 1990, Luteyn & Dorr 13794 (NY, NY, US); Sud Yungas, La Paz Calacoto 69 km hacia el Este, pasando el Nevado Illimani, Estacion general Ikiko, 3100 m, 31 Dec 1980, Beck 3904 (F); Inquisivi, 3 km (by air) and 8 km (by road) SW of Inquisivi, above road from Quime to Inquisivi, 2950 m, 12 Mar 1988, Nee & Solomon 36672 (CORD, F, G, MEXU, MO, NY, US); Inquisivi, 6.5 km SE of Inquisivi, 1.2 km SW of Machacamarca, 3000 m, 18 Mar 1988, Nee 36718 (CORD, MO, NY, US); Larecaja, above Rio Lakhathiya, just above junction with Rio Tusca Jahuira, 19 May 2001, Nee et al. 51833 (NY); Inquisivi, comunidada Coquetanga-Aguas Calientes-Calachaca, cuenca del rio Calachaca-Jahura, 9 km de Choquetanga, 3400 m, 20 Jun 1994, Salinas 3275 (K, NY); Sud Yungas, Mina Chojilla camino de acceso de vehiculos a Kacapi, 2700 m, 4 Jul 2000, Siñani RS 230 (NY); Pedro Domingo Murillo, Rio Zongo Valley, 22.5 km below dam at Lago Zongo, 3000 m, 9 Oct 1982, Solomon 8431 (K, MO, NY); Inquisivi, Camino de Quime a La Paz a unos 2-3 km arriba de Quime, 3221 m, 16 Mar 2003, Wood & Ortuño 19411 (K, LPB). Santa Cruz: Florida, Comarapa Road, 28 km from Comarapa, 2650 m, 15 Aug 1991, Acevedo et al. 4623 (NY, US); Caballero, Siberia, 2-2.5 km debajo de la escuela de Siberia, camino al valle de Saipina, 2850 m, 30 Nov 2003, Jordan & Vargas 253 (MO); Vallegrande, on road from Guadalupe to Pucará, 2725 m, 4 May 2001, Nee et al. 51736 (BM, NY); Caballero, loc 2-3 km S de Siberia sobre el camino a Larkapampa, 3000 m, 25 Jul 1996, Saldías & Fernández 4607 (NY); Caballero, Parque Nacional Amboró, Siberia, 25 km al NW sobre la carretera Comarapa-Cochabamba, 3000 m, 5 May 1993, Vargas C. et al. 2302 (NY); Caballero, Parque Nacional Amboró, proximadades de Cerro Bravo a 10 km al N de Comarapa, alrededores de la Parcela permanente, 2400 m, 18 Oct 1993, Vargas C. & Jardim 3002 (NY); Caballero, Siberia, ca. 1-2 km arriba de la comunidad de Siberia, sobre un camino vecinal, entrando hacia el Parque Nacional Amboró, 3001 m, 12 Apr 2003, Wood et al. 19700 (BOLV, K);. Tarija: route Villa Montes to Tarija, 15 km apres Entre Rios, 1850 m, 4 Nov 1993, Billiet & Jadin 6067 (K, MO); Aniceto Arce, 39.9 km S of jct of road to Entre Rios, on road to Padcaya, 2100 m, 29 Apr 1983, Solomon 10249 (MO, NY).

Costa Rica. Cartago: Volcán Irazu and Volcán Turrialba, along road from Pacayas to Hacienda Central, 31 Mar 1982, Barringer et al. 2233 (F); SE slope of Cerro de la Muerte, Cordillera del Talamanca, along Interamerican Hwy 63 miles from downtown San Jose, 2700 m, 23 May 1976, Croat 35427 (MO); Cantón Paraíso, Reserva Forestal Los Santos, Cueva del Savegre, carretera Interamericana de la Georgina hasta la entrada a San Gerardo de Dota, 3180 m, 18 Mar 1997, González & Hammel 1840 (G, MO); Cantón de El Guarco, Cordillera de Talamanca. Carretera Interamericana, entre E1 Empalme y La Chonta, 2400 m, 10 May 1995, Hammel 19815 (BM, MO); Irazú, 2134 m, 24 Jun 1974, Kuntze 2300 (NY); Macho Gap Camp, 39 kilometers S. of Cartago and 3 miles N of Copey, 2500 m, 19 Feb 1943, Little 6022 (F, US);. Heredia: Cantón de Barva, P.N. Braulio Carrillo, Cordillera Central, potreros La Georgina, 2600 m, 29 Oct 1993, Fernández & García 1503 (MO); Puntarenas: Monteverde, at edge of Continental Divide and of Pacific side of slope, 1300 m, 18 Aug 1976, Solomon 5365 (MO); N. San Isidro del General, 12 Aug 1971, Spellman et al. 653 (MO); San José: northern Cordillera Talamanca, region of Cerro de la Muerte, on carretera Nacional 34.5 km N. of San Isidro and 2.6 km S of La Georgina Inn, 2652 m, 4 Apr 1978, Davidson 7235 (F, MO, NY); Cantón de Pérez Zeledón, Cordillera de Talamanca. Estación Cuericí, 2600 m, 29 Aug 1995, Gamboa 281 (BM, MO); Villa Mills, Cerro de la Muerte, Carretera Panamericana Sur, 2800 m, 26 Feb 1965, Jiménez M. 2989 (F, NY); Z.P. Cerros de Escazú, Cedral, falda norte del cerro Rabo de Mico, cuenca del Río Poás, 1600 m, 9 Oct 1991, Morales 168 (MO); Santa María de Dota, Laguna de la Chonta, northeast of Santa María, 2000 m, 18 Dec 1925, Standley 42265 (F, US); near Finca La Cima, above Los Lotes, North of El Copey, 2100 m, 21 Dec 1925, Standley 42764 (US); San Gerardo de Dota, 8 to 10 km down from the Interamerican Highway, 18 Sep 1975, Utley & Utley 3083 (F, MEXU, MO, NY).

El Salvador. Santa Ana: Near summit of Cerro Monte Cristo, 2134 m, 18 Jan 1959, Allen 7172 (F, NY, US); Montecristo, P.N. Montecristo, plan de los helechos, 2051 m, 13 Mar 2002, Carballo et al. 281 (LAGU); Cordillera Miramundo, mountain of Montecristo, 2000 m, 27 Jan 1966, Molina R. et al. 16706 (F, NY, US); P.N. Montecristo, al final del atajo para el Trifinio, 2000 m, 24 Jan 2002, Monterrosa et al. 215 (B, BM, LAGU).

Guatemala. Chimaltenango: Chichavae, 2400 m, 5 Dec 1933, Skutch 738 (US). Huehuetenango: San Juan Ixcoy, Sierra de los Cuchumatanes, along road to Huehuetenango, 5 miles S of San Juan Ixcoy, 4 Feb 1965, Breedlove 8542 (F); San Mateo Ixtatán, Sierra de los Cuchumatanes, 4 miles east of San Mateo Ixtatan on road to Barillas, 7 Feb 1965, Breedlove 8744 (F); Cerro Cananá, between Nucapuxlac and Cananá, Sierra de los Cuchumatanes, 2500 m, 18 Jul 1942, Steyermark 49046 (F); wet cloud forest at Cruz de Limon, between San Mateo Ixtatan and Nuca’, Sierra de los Cuchamatanes, 2600 m, 31 Jul 1942, Steyermark 49870 (F, NY). Quetzaltenango: highway km 172 junction Quetzaltenango, Huehuetenango and Totonicapan, 10 Jan 1974, Molina R. et al. 30200 (F, MO); Volcán Zunil, 2500 m, 22 Jan 1940, Steyermark 34672 (F). Quiché: sin. loc., 1942, Aguilar s.n. (F). Totonicapán: Sierra Madre Mountains south of Totonicapán, near Mirador (km 170), 2800 m, 20 Dec 1972, Williams et al. 41447 (BM). Zacapa: Jones, 2020 m, 5 Apr 2000, CDC-CECON 1535 (BM).

Honduras. Francisco Morazán: along road to Parque Nacional La Tigra, 22-25 km NE of Tegucigalpa, 1850 m, 1 Feb 1987, Croat & Hannon 64047A (MO, NY); Tegucigalpa, 11 km NE of Tegucigalpa. La Tigra National Park Summit, 2000 m, 26 May 1992, D’Arcy 18006 (MO); La Tigra, 25 km NE de Tegucigalpa, 2105 m, 24 May 1986, Manzano 177 (MA); Montaña La Tigra S.O. de San Juancito, 2000 m, 23 Apr 1964, Molina R. 13744 (F, NY, US); Cerro Uyuca, 8 km O. del Zamorano, 1100 m, 27 Apr 1984, Soihet M. 142 (MO).

Mexico. Chiapas: Chamula, Yalal Chin, 1829 m, 3 Apr 1965, Breedlove 9540 (MEXU); San Cristóbal de las Casas, SW slope of Zontehuiz, 2774 m, 21 Jun 1965, Breedlove 10435 (F, MEXU, US); Chenalho, in paraje Los Angeles Chiste, 1700 m, 22 Oct 1976, Breedlove 40863 (MEXU, MO); La Independencia, 6-10 km north-northeast of La Soledad along logging road from Las Margaritas to Campo Alegre, 1600 m, 26 Nov 1980, Breedlove & Almeda 47819 (MEXU, MO, NY); Zinacantán, on NW side of Muk’ta vits (Cerro Huitepec), 2743 m, 18 Feb 1966, Laughlin 125 (MEXU, US). San Cristóbal de las Casas, Santa Cruz en San Felipe, 15 Nov 1986, Mendez Ton 9743 (MEXU, NY); Pueblo Nuevo Solistahuacán, on ridge north of Clinica Hierba Buena near Pueblo Nuevo Solistahuacán, 1981 m, 25 Jan 1965, Raven & Breedlove 19978 (F); Tenejapa, at Colonia Ach’lum, 2652 m, 27 Dec 1965, Shilom Ton 454 (F, MEXU, NY, US); San Andres, NW of San Cristobal, 579 m, 26 Feb 1931, Souviron & Erlanson 81 (BH, US); Tenejapa, a 20 km al SW de Tenejapa camino a San Cristóbal de las Casas, 25 Sep 1983, Téllez V. & Pankhurst 7262 (MEXU, MO); Mazapa de Madero, Granada de Tlalcanaque, 10 km al Noreste, 2500 m, 22 Feb 1987, Ventura & López 4352 (MO, NY). Colima: Nevado de Colima, Nevado de Zapotlan, a few miles south of Ciudad Guzman (Zapotlan), 3000 m, 2 Jul 1956, Gregory & Eiten 292 (P); Guerrero: Taxco de Alarcón, Taxco. 19.9 km al NO, 2360 m, 28 May 1998, Cruz Durán 2260 (MEXU). Chilpancingo de los Bravo, a 6 km al W de Omiltemi, camino a la Soledad La Joyas, 27 Mar 1982, Martínez S. & Téllez V. 272 (MO); Chilpancingo, laderas superiores del Cerro Alquitran, cerca de Mazatlan, 2600 m, 11 Feb 1970, Rzedowski 27036 (F, MO x2); Chichihualco, a 8 km al SO de Filo de Caballo, 2435 m, 21 Apr 1985, Soto N. & Aureoles C. 8345 (MEXU). Hidalgo: Lolotla, carr. Pachuca-Huejutla, 3.5 km al N de Ixtlahuaco, 1670 m, 5 Apr 1986, Baker et al. 054 (NY); 17 mi N of Jacala, ca 56 mi from Zimapan, 1 Aug 1972, Dunn 19059 (MO); Zacualtipán, 1 km al SSE de Zacualtipan, 2400 m, 19 May 1976, Flores F. 234 (MO, US); Zacualtipán, Zacualtipán, 1 km al SSE, 2400 m, 19 May 1976, Flores M. 234 (CORD, MEXU); Tenango de Doria, Cirio, 8 km al este de Tenango de Doria, 1700 m, 27 Mar 1980, Hernández M. & Hernández V. 4189 (MEXU); Tianguistengo, 10 km al oeste de Tianguistengo (Tepeoco), 2100 m, Apr 1981, Hernández M. et al. 5806 (MEXU, MO); Metztitlán, noreste de Hidalgo, Rincón de los Ahuajes, 3.5 km de la desviación a Zoquizoquipan, 6 Jul 1992, López García 113 (MEXU); Zacualtipán, Paraje Cumbre de Tlahuelompa, 2 km al SW del ejido de Tlahuelompa, 18 Dec 1992, López García 392 (MEXU); El Alcalaque, 2 km al NW del pobaldo Eloxochitlán, 29 Dec 1992, López G. 433 (MEXU); Tlanchinol, a 4 km al E de Tlanchinol, camino a Apantlasol, 3 Sep 1997, Martínez S. 28444 (MEXU, NY); Palo Hueco, km 301 on highway between Jacala and Santa Ana, 1524 m, 28 Apr 1947, Moore 2672 (BH); Barranca below Trinidad Iron Works, 1524 m, 10 May 1904, Pringle 8835 (BH, BM, F, GOET, K, LE, MEXU, MO, US); Molango, Ismolinta, 2 km S de Ismolinta rumbo a Eloxochitlán, 1580 m, 19 Jun 1995, Sousa Peña 577 (MEXU); Zacualtipán, vereda entre El Reparo y Zahuastipan, 3 km de la Majonera, 29 Jun 1988, Vázquez et al. 4626 (F, NY); Zoquizoquipan, 1940 m, 2 Sep 1962, Vela G. 977 (MEXU). Jalisco: Yolox, Sierra de Juárez, Carretera Ciudad de Oaxaca. a Tuxtepec, cerca de Cerro Pelon, a 300 m. antes de San Pedro Yolox, 2850 m, 18 Jan 1989, Chazaro et al. 5815 (MEXU); Cuatitlán, 19-20 km al NE de Cuatitlan, 500 m al W de Llanos de San Miguel, 2200 m, 29 Apr 1988, Cuevas & Guzmán 2871 (MEXU); along lumber road, 0.3 km E of fork Cerro La Cumbre/ Rincón de Manantlán, at crossing of first small stream, N of Sierra de Manantlán Central, 17.7 km S of El Chante, 2250 m, 6 Jan 1980, Iltis et al. 2336 (F x2, MEXU, US); northwestern slopes of Nevado de Colima, above Jazmín, 2-3 km above settlement of El Isote, 2600 m, 26 Mar 1949, McVaugh & Wilbur 10066 (BM); Sierra de Manantlán (15-20 miles southest of Autlán), about 2 miles from Aserradero San Miguel Uno, west and south of divide towards Manzanillo, 2250 m, 4 Nov 1952, McVaugh & Sooby 13923 (BM); Venustiano Carranza, Puerto El Floripondio, camino a la Est. de Microondas Las Viboras, 2450 m, 22 Aug 1987, Rodríguez C. & Suárez J. 939 (MEXU). Michoacán: Zinapécuaro, Los Azufres, alrededores de la laguna larga, 31 May 1988, Díaz Barriga 4680 (MEXU); Zinapécuaro, Cañada La Hierbabuena, al SW de la Presa Laguna Larga, 2750 m, 17 Nov 1988, Jasso 532 (MEXU, MO); Melchor Ocampo, Cerro Camacho, Los Remedios, a 4 km al W de Ocampo, 2320 m, 26 Apr 1982, Martínez S. et al. 386 (BH, MEXU); Tzitzio, En Mil Cumbres a 31 km al SW de Cd Hidalgo, carretera a Morelia, 2500 m, 11 Oct 1983, Martínez S. et al. 4741 (MEXU, NY). Morelos: Tepoztlán, Santo Domingo Ocotitlán, en el cerro al N de la población, 2200 m, 26 Apr 1984, Gutiérrez 203 (MEXU). México: Temasceltepec, Rincón, 3 Oct 1932, Hinton 365 (BM, K, US); Temascaltepec, Comunidad, 2480 m, 5 Oct 1933, Hinton 3852 (BM, F, K, MO, US); Oaxaca: Comaltepec, immediately to the right of Hwy. 175, just beyond first major switchback on decent from Mirador below Cerro Humo Chico, in quebrada of Río Cerro Pelón, 2740 m, 1 Nov 1993, Boyle & Massart 2512 (BM, MEXU); Sierra Juárez, 2.5 miles by road. SW of summit at Cerro Pilon, on hwy 175 between Ixtlan de Juarez and Valle Nacional, 2550 m, 19 Jul 1972, Breckon & Breckon 1364 (F); San Martín Peras, Dto. Juxtlahuaca, a 2.7 km adelante de la desviacion de San Martin Peras, carretera para Coycoyan de las Flores, 2490 m, 20 Jun 1993, Calzada 18460 (MEXU, NY); Comaltepec, 46.5 km al NE de Istlán, carretera Oaxaca-Tuxtepec, 2650 m, 17 Dec 1987, Campos V. & Torres 886 (MEXU); Mixistlán, 30 km al S de Totontepec, Distr. Mixe, 2600 m, 10 Nov 1983, García Mendoza et al. 1287 (MO, NY); Patio de Arena, vicinity of Cerro Zempoaltepetl, ca. 5 km E of summit, 2800 m, 9 Aug 1950, Hallberg 879 (MEXU); Ixtlán, Sierra de Juárez, parte alta y cima de Cerro Pelon, 2900 m, 5 Aug 1985, Lorence et al. 4754 (MEXU); Yolox, distrito de Ixtlan, 8 km E of Yolox on road between Yolox and highway 175, 2900 m, 13 Apr 1981, Martín 531 (MO); Ixtepeji, La Botuda, Dto. Ixtlán, 10 May 2001, Santiago 49 (MEXU); Totontepec, Villa de Morelos, distr.: Mixe Tsa, 19 Aug 1991, Rivera Reyes 2814 (MEXU, NY); Santa María Teopoxco, Sierra Mazateca, near Plan de Guadalupe, 39 km by road (16 km by air) west o Huatla on road to Teotitlan, 2200 m, 17 Jan 1984, Solheim & Reisfield 1400 (NY); Puerto de la Soledad, 3 km al S de Huautla, sobre el camino a Teotitlán del Camino-Huautla, 10 Sep 1981, Téllez V. & Elisens 4698 (MEXU); Totontepec, 13 km al O de Totontepec, carretera a Villa Alta, Dto. Mixe, 28 Mar 1986, Torres C. & Ramírez 8450 (MEXU, MO, US); San Juan Tepeuxila, San Juan Tepeuxila, Dto. Cuicatlan, arroyo la primera toma, hacia Llano Chiflido, por Arroyo Paloma, 2369 m, 18 May 2002, Torres Colin et al. 16202 (MEXU). Puebla: Tlatlauqui vertiente norte, 1800 m, 31 Apr 1971, Boege 1738 (MEXU); Equimita, km 35 de la carretera que va hacia Cuetzalan, 22 Jun 1976, Inzunza 114 (MO); Huauchinango,1951 m, 7 Oct 1944, Sharp 441218 (MEXU, US); Coxcatlán, L y Griega, desviación a Coyomeapan, de La Brecha a Zoquitlán, 2540 m, 28 Sep 1984, Tenorio L. & Romero de T. 7502 (MEXU); Tezuitlán, Coaxisco 12 km al NW de Tezuitlan, 2030 m, 12 Apr 1985, Tenorio L. et al. 8630 (MEXU); Pahuatlán, Acahuales, 1900 m, 22 May 1986, Tenorio L. & Romero de T. 11383 (MEXU); Tezuitlán, Río Frío, 12 km al N de Teziutlan, carr. a Nautla, 1500 m, 7 Jul 1987, Tenorio L. et al. 14020 (MEXU, NY). Querétaro: Pinal de Amoles, El Ranchito, 1 km de El Rnachito camino a San Pedro Escanela, 1980 m, 10 Nov 1988, Carranza 1251 (MEXU). San Luis Potosí: 22 km W of Santa Catarina on highway 86 at km 49, 2200 m, 29 Sep 1965, Roe & Roe 2191 (BM). Veracruz: Xico, Tonalasco, 25 Jun 1986, Arriaga C. 389 (MEXU); Huayacocotla, Vereda Tzimentey, cerca del limite SE de la Reserca propuesta, 1750 m, 25 Apr 1981, Bastelleros & Ballesteros 421 (MEXU); Acultzingo, La Laguna, carretera Puerto del Aire, poblado, 2320 m, 16 Dec 1977, Calzada & Delgado 4181 (MEXU); Huayacocotla, Agua de la Calabaza, carretera Huayacocotla-Chicontepec, 1800 m, 20 Jul 1979, Calzada 5464 (F); Coatepec, Cerro Huilotepec entre Mesa de los Laureles y Tierra Grande, 2500 m, 27 Jul 1987, Cházaro B. et al. 5275 (MEXU, NY); Las Minas, al SE de Rinconada por el Cerro La Tolva, 2300 m, 11 Aug 1988, Duran E. & Burgos 570 (MEXU); Huayacocotla, vicinity of a large shrine and "Bienvenidos a Huayacocotla" arch over the highway, SW entrance to Huayacocotla on road from Palo Bendito, 2150 m, 21 Jul 1982, Diggs & Nee 2958 (BH, CORD, F); Huayacocotla, Agua de la Calabaza, 16 km NE de Huayacocotla, carretera a Zilacatipan, 1750 m, 24 Apr 1981, Juárez G. & Vasquez B. 68 (F); Huayacocotla, Xalapa, INIREB, along Huayacocotla-Zontecomatlan road, between Barro Colorado and Tepozanes, 2 km by road NE of Agua de la Calabaza and 5 km by road SW of Zilacatipan, 1800 m, 27 Apr 1983, Nee 26894 (F, MO, NY); Puerto del Aire, just W and high above Acultzingo, top of very steep pass on Highway 150 ( Tehuacán to Orizaba) near boundary between states of Veracruz and Puebla. Cumbres de Acultzingo, 2409 m, 27 Sep 1962, Ugent & Flores C. 2466 (BM); entre Los Ocotes y Helechales, 1 Jun 1980, Vargas et al. 306 (MEXU); Jalacingo, Ocotepec, cerca de cerro, 1750 m, 4 Mar 1970, Ventura A. 1008 (F, MO, NY); Acajete, Acajete, 1850 m, 1 Apr 1974, Ventura A. 9810 (MEXU, MO); Atzalán, La Florida, 1650 m, 16 Jun 1976, Ventura A. 12871 (F, MEXU); Altotonga, San Miguel Tlalpoalan, 1950 m, 2 Jun 1981, Ventura A. 18538 (MEXU); Acajete, La Joya, 2100 m, 1 Apr 1982, Ventura A. 19577 (MEXU, MO); Las Vigas, Rancho San Miguel, barranca El Cable, 2400 m, 11 Sep 1989, Zamora C. 1096 (MEXU);

Peru. Cusco: Urubamba, Ollantaytambo, Dist. Ollantaytambo, Abra de Malaga, 3600 m, 2 Jun 2002, Galiano et al. 4225 (MO); Hacienda Huyucalla, valle del Paucartambo, 3400 m, Jul 1930, Herrera 2986 (US); Paucartambo, 10 km from Puesto de Vigilancia (PN Manu) towards Paucartambo, 3313 m, 15 Mar 2012, Knapp et al. 10436 (BM, USM); Quispicanchis, near Marcapata, 24 Jul 1942, Metcalf 30729 (G, MO, US); Quispicanchis, entre Abra Walla Walla y Marcapata a 210 km de Cusco, 2800 m, 21 Apr 1988, Núñez V. et al. 8994 (F, MO x2); Pautarcambo to Tres Cruces, Cerro de Cusilluyoc, 3200 m, 2 May 1925, Pennell 14148 (F, NY); Calca, Dist. Lares, Mantto, 2100 m, 23 Jan 2003, Valenzuela et al. 1236 (MO); La Convención, Espiritupampa, Dist. Vilcabamba, 3300 m, 14 Oct 2003, Suclli et al. 1277 (BM, NY); La Convención, San Luis, valley of Lucumayu River, on road from Ollantaytambu to Quillabamba, environs of San Luis (former resturan) above Alfamayo, along roadside with 2 km of San Luis back up towards Abra de Malaga, 2800 m, 3 Apr 1997, Tupayachi & Emshwiller 3424 (BH); Paucartambo, Challabamba, Pesto Grande, 2900 m, 20 Apr 1986, Zimmerer 80 (NY). Junín: Tarma, Carpapata, al este de Carpapata, 8 Dec 2005, Marcelo Peña et al. 1920 (MOL). Puno: Carabaya, Ollachea to San Goban road, 2.5 km north of Ollachea, 15 Aug 1980, Boeke 3049 (F, MO, NY, US); Ollachea to San Gabon, 17 Jul 1978, Dillon et al. 1107 (BM, MO, NY).

Figures

Figure 10. Solanum aligerum Schltdl. (A drawn from Martinez S. 4741 B-D drawn from Torres B. 2022 E drawn from Nee 26870). Illustration by Bobbi Angell.

Figure 11. Distribution of Solanum aligerum Schltdl.