Ayers, Johnson & Ayers, 2022

3b. Diastatea tenera subsp. durangensis E.P.Johnson & T.J. Ayers , subsp. nov. TYPE: MÉXICO. Durango: Mpio. Suchil. Laguna de Mogotes, alrededores, 8 km de El Aleman, Reserva de la Biosfera La Michilıa, [23.47, -104.26], 17 Nov 1986, F. Acevedo 366 (holotype: CIIDIR; isotypes: ASC!, IBUG, MEXU).

Stems 9.5–37 cm tall, simple or a few weak side branches. Leaves, the lowermost deltoid, 2–4 mm long, 1.2–1.8 mm; middle to upper leaves linear, 6–24 mm long, 0.5–1.6 mm wide. Inflorescence with few to 12 flowers; bracts 5.4–20 mm long, 0.4–0.9 mm wide. Flowers with pedicels 3.7–17.5 mm long in flower; calyx lobes subequal, narrowly triangular or linear, 1.1–2.5 mm long in flower, margins with 1–2 pair of teeth at or below the middle, glabrous or ciliolate; corolla purplish-blue or purple, the nectar guides pale yellow, gradually brighter toward throat, with projections at throat; tube 8.8–13.5 mm long from calyx sinus to lateral corolla sinus, 0.8–1.6 mm wide at middle; dorsal lobes obovate, apex acute to mucronate, with narrowed claw attaching distal portion to deltoid base, 6.1–9.0 mm long, 1.8–4.0 mm wide; ventral lobes broadly oblanceolate with mucronate tip, 7.1–11.3 mm long, 2.2–4.3 mm wide; filaments 9.5–13 mm long; anthers 1.7–2.0 mm long. Fruits narrowly ellipsoid, with an unequal base

mirroring gibbous corolla, 5.2–10 mm long, 2.2–3.6 mm wide. Figures 4D View FIG , 8 View FIG , S 4B View FIG .

Chromosome Number —Unknown.

Etymology —The new subspecific epithet reflects that the center of distribution of this taxon is in the Mexican state of Durango.

Distribution —Mexican states of Durango, Guanajuato, Jalisco, and Zacatecas ( Fig. 7 View FIG ).

Notes —Exceptionally large-flowered individuals of D. tenera have been found in Durango. They appear to be the end point of a cline stretching from Puebla north to Durango, in which the flowers of D. tenera increase dramatically in size. The populations in Zacatecas and Guanajuato have the long corolla tubes of D. tenera subsp. durangensis but their lower corolla lobes are slightly shorter than the populations in Durango and Jalisco.

Because of the distinct difference in size between the individuals found mainly in Durango and the rest of D. tenera , and the fact that the collections appear to have a distinctive range adjacent to the central Mexican Plateau, they are described here as a new subspecies. We chose subspecific rank over varietal rank because varietal rank in the Campanulaceae has been dismissed by Lammers (2006). Future examination of the subspecies should look at habitat and pollinator differences as possible drivers of floral differentiation.

Additional Specimens Examined — Mexico. — DURANGO: San Juan de Michis, Mpio. de Suchil , 25 Oct 1984, Acevedo 150 (TEX, WIS); Mpio. Suchil. San Juan de Michis , 3.5 km al S del poblado, Reserva de la Biosfera La Michilıa, [23.47, -104.26], 21 Oct 1985, Gonzalez & Acevedo 3546 (ASC); Mpio. Suchil, Sierra de Michis, San Agustın, Reserva de la Biosfera La Michilıa , [23, -104], 12 Oct 1984, Jimenez 186 (ASC); Mpio. Suchil, El Aleman, 8 km cerca del Arroyo Las Joaquinas, Reserva de la Biosfera La Michilıa, 18 Nov 1986, Acevedo 376 (ASC); Mpio. Suchil, Reserva de la Biosfera La Michilıa, Portrero Mesa Larga, [23.5, -104.3], 17 Nov 1993, Garcıa A. & Alvarado 1911 (ASC); Mpio. Suchil, Portrero Los Anegados, 3 km de el Aleman, Reserva de la Biosfera La Michilıa, [23.47, -104.26], 11 Oct 1986, Acevedo 306 (ASC); Mpio. Suchil. Potrero El Taray, Reserva de la Biosfera La Michilıa, [23.41, -104.25], 12 Sep 1983, Acevedo s. n. (ASC); Mpio. Suchil. Reserva de la Biosfera La Michilıa, camino Suchil-Charcos, km 51, entrada al Rancho La Pe na ~, [23.4, -104.2], 11 Oct 1992, Garcıa A. 1700 (ASC). — GUANAJUATO: 8 km northeast of Santa Rosa, 10 Nov 1970, McVaugh 24150 (MEXU); Mpio. San Jose, Ca nada ~ Las Peras 3 km al SE de El Zorrillo, 6 Nov 2003, Gutierrez G. et al. 787 (MEXU). — JALISCO: Rancho Viejo, about 10 km east-northeast of Huejuquilla el Alto, 31 Oct 1963, Feddema 2310 (NY, WIS). — ZACATECAS: Plus or minus 35 km al W de Jalpa, sobre la carretera a Tlaltenango; 27 km del entronque con la carretera Jalpa-Juchipila, 23 Oct 1970, Rzedowski & McVaugh 1039 (MEXU, MO, NY, LL).

DIASTATEA Scheidw. subg. DIASTATEA

Middle and upper cauline leaves ovate to lanceolate and sometimes narrowly elliptic, glabrous to pubescent. Corolla tube cylindric, without gibbous projection at base.

Distribution —The species within this subgenus are found in the following Mexican states and Central and South American countries: Chiapas, Colima, Durango, Guanajuato, Guerrero, Hidalgo, Jalisco, Mexico, Michoacan, Morelos, Nuevo Leon, Oaxaca, Puebla, and within the Distrito Federal; Costa Rica, El Salvador, Guatemala, Honduras, and Panama; Argentina, Bolivia, Colombia, Ecuador, Peru, and Venezuela.

4. DIASTATEA MICRANTHA (Kunth) McVaugh, Bull. Torrey Bot. View in CoL Club 67: 143. 1940. Lobelia micrantha Kunth , Nov. Gen. Sp. [H.B.K.] 3: 316. 1818. Rapuntium micranthum (Kunth) C. Presl, Prodr. Monogr. Lobel. View in CoL : 25. 1836. Dortmanna micrantha (Kunth) Kuntze, Revis. Gen. Pl. View in CoL 2: 972. 1891 as ‘ Dortmannia .’ Laurentia micrantha (Kunth) Zahlbr., Bull. Torrey Bot. Club 24(8): 386. 1897; non Laurentia micrantha (E.Mey.) A.DC., Prodr. [A. P. de Candolle] 7(2): 411. 1839. TYPE: ECUADOR. “in Regni Quitensi, prope pagum Puembo, alt. 1300 hex. Floret Julio.” A. Humboldt & A. Bonpland s. n. (holotype: P-image! isotypes: B-W [presumed destroyed], F-image! [photo of isotype at B, photo 9110]).

Lobelia subtilis Kunth , Nov. Gen. Sp. [H.B.K.] 3: 317. 1819. Rapuntium subtile (Kunth) C. Presl, Prodr. Monogr. Lobel. View in CoL : 25. 1836. TYPE: ECUADOR. Quito, Puembo. A. Humboldt & A. Bonpland s. n. (holotype: P-image!; isotype: P-image!).

Lobelia ruderalis Willd. ex Roem. & Schult., Syst. Veg. , ed. 15 bis [Roemer & Schultes] 5: 56. 1819. TYPE: PERU. “Habitat in Peru.” A. Humboldt & A. Bonpland s. n. (holotype: B-Wimage!).

Lobelia draba Willd. ex Roem. & Schult., Syst. Veg. , ed. 15 bis [Roemer & Schultes] 5: 67. 1819. TYPE: no locality or collector on specimen (holotype: B-W-image!).

Lobelia parviflora M.Martens & Galeotti, Bull. Acad. Roy. Sci. Brux. 9(2): 41. 1842. Dortmanna parviflora (M.Martens & Galeotti) Kuntze, Revis. Gen. Pl. View in CoL 2: 973. 1891 as ‘ Dortmannia .’ TYPE: MÉXICO. Oaxaca, Yavezia, cordillera [from description: “On trouve cette espece au bord des ruisseaux peu ombrages de la cordillere orientale d'Oaxaca, pres de Yavezia, de Socorro et de Castrasana, de 6,500 –7,500 pieds d’ elevation”], Jan 1840, H. G. Galeotti 1970 (holotype: BR-image!; isotypes: BR-image!, K-image!, US-image!, W-image!).

Laurentia ovatifolia B.L.Rob., Proc. Amer. Acad. Arts 26: 166. 1891. Laurentia micrantha var. ovatifolia (B.L.Rob.) E.Wimm. in J.F.Macbr., Publ. Field Mus. Nat. Hist., Bot. Ser. 13, pt. 6: 476. 1937. Diastatea micrantha var. ovatifolia (B.L.Rob.) E.Wimm., Ann. Naturhist. Mus. Wien View in CoL 56: 332. 1948. LECTOTYPE, here designated from two syntypes: MÉXICO. San Luis Potosi: hills near Cardenas, 8 Sep 1890, C. G. Pringle 3302 (lectotype: GH!; isolectotypes: BR, COLO, PH, S, NDG, K, GOET, W-image!).

Laurentia michoacana B.L.Rob., Proc. Amer. Acad. Arts 26: 167. 1891, nomen nudum.

Laurentia pedunculata Brandegee, Univ. Calif. Publ. Bot. 6: 73. 1914. Laurentia micrantha var. longibracteata E.Wimm., Revista Sudamer. Bot. 2: 104. 1935. Diastatea micrantha var. longibracteata (E.Wimm.) E.Wimm., Ann. Naturhist. Mus. Wien View in CoL 56: 332. 1948. TYPE: MÉXICO. Finca Covadonga, Jul 1913, C. A. Purpus 6705 (holotype: UC-image!; isotypes: GH!, MO-image!, US-image!).

Diastatea serrata Standl. & L.O.Williams, Ceiba View in CoL 1: 91. 1950. TYPE: HONDURAS. Department of Morazan. Drainage of the Rıo Yeguare, at about Longitude 87 W and Latitude 14 N, Alt. 850 m, 26 Nov 1948, L. O. Williams 14786 (holotype: US-image!).

Stems 2.7–80 cm tall, simple to branched, 0.2–3.0 mm wide at base, minutely winged, pubescent with soft hairs evenly distributed around stem, often becoming less pubescent distally, rarely glabrous. Leaves pseudo-petiolate; pseudo-petioles 1–22 mm long; margins ciliolate and decurrent into narrow wings on stem; blades purple to green, sometimes green adaxially and purplish abaxially, base subcordate to narrowly cuneate, apex round to acuminate; lowermost blades round, ovate, lanceolate to narrowly elliptic, 2.5–11 mm long, 2–8 mm wide, the margins regularly serrate to erose, glabrous or with sparsely scattered stiff hairs adaxially and on veins abaxially; upper blades ovate to narrowly lanceolate, 4.5–60 mm long, 2.0– 26 mm wide, the margins regularly or irregularly serrate, ciliolate; pubescence on veins abaxially. Inflorescence weakly to strongly secund with few to 30 flowers per inflorescence; bracts linear to elliptic, 2–53 mm long, 0.2–12 mm wide, spreading. Flowers with pedicels 1.5–25(–57) mm long in flower, glabrous, spreading to ascending, often bent upward distally in fruit; hypanthium obconic in flower, 0.5–0.9 mm in diameter, 0.3–0.5 mm in length from calyx sinus to attachment point with pedicel, not much expanded in fruit, glabrous, calyx lobes subequal to equal, from one-third to two-thirds the length of the fruit, narrowly triangular or linear, 1.1–3.5(–5) mm long in flower, green, margins entire, glabrous or ciliate; corolla purplish-blue or white, if white, then lobes lavender or blue, nectar guides yellow sometimes with green raised projections at throat; tube cylindrical, 2.4–4.5 mm long from calyx sinus to lateral corolla sinus, 0.7–1.1 mm wide at middle; dorsal lobes linear to linear spatulate, 1–2 mm long, 0.2–0.3 mm wide; ventral lobes oblong, sometimes with a mucronate tip, 0.9–2.5(–3.5) mm long, 0.4–1.3(–1.6) mm wide; filaments 2.8–4.6 mm long; anthers 0.5–1(–1.5) mm long, dark purple, either with downy white covering or appearing glabrous, sometimes with short hairs near apex of dorsal anthers, plus two large and many small white trichomes at apex of ventral anther pair. Fruits narrowly to broadly ellipsoid sometimes slightly curved at tip, 3–6.5(–8) mm long, 1.4–2.6 mm wide, light brown or purplish-brown; seeds ovoid to ellipsoid, 0.4–0.5 mm long, orange to dark-orange. Figures 3F, S View FIG 4D View FIG .

Chromosome Number —2 n 5 14 ( Fig. S5C View FIG ).

Distribution —Mexican states of: Chiapas, Colima, Durango, Guanajuato, Guerrero, Hidalgo, Jalisco, Mexico, Michoacan, Morelos, Nuevo Leon, Oaxaca, Puebla and the Distrito Federal; Central and South American countries of: Costa Rica, El Salvador, Guatemala, Honduras, and Panama; Argentina, Bolivia, Colombia, Ecuador, Peru, and Venezuela ( Fig. 6 View FIG ).

Notes — Diastatea micrantha can be distinguished by its minute corolla tube and corolla lobes, and its tendency to self-pollinate. Eight collections from Oaxaca have unusually large flowers for D. micrantha . Because Oaxaca is the center of distribution for D. virgata , these large-flowered specimens warrant further examination and should be included in future morphological and molecular work.

The type of D. micrantha was originally collected in Ecuador, but its center of distribution is in the Trans-Mexican Volcanic Belt. Its broad geographic range and morphological variation have led to it being described as at least ten different species.

Lobelia xalapensis and L. diastateoides can be easily confused with D. micrantha because their ovaries in flower are essentially nearly superior. Lobelia xalapensis can be distinguished from D. micrantha by a shorter more ovoid fruit, corolla split dorsally to the base, and prickly pubescent pedicels. Lobelia diastateoides can be distinguished from D. micrantha by a perennial habit and corolla split nearly to the base.

McVaugh noted in his 1940b revision of Laurentia that Laurentia michoacana B.L.Rob. is a nomen nudum and we want to re-emphasize that here. B. L. Robinson (1891) makes it clear that Pringle 3337, despite being distributed under the name Laurentia michoacana B.L.Rob. , “should be for the present doubtfully referred to as Lobelia parviflora Mart. & Gal. , since it is highly probable that the type of this species will be found on investigation to be a Laurentia .” It was, in fact, found to be

a Laurentia and is now considered a heterotypic synonym of D. micrantha . Many herbaria hold Pringle 3337 as a type including the following: (P-image! labeled as a holotype; BR-image!, K, COLO, LL!, MEXU!, MSC, CM, PH, WIS! labeled as isotypes).

Additional Specimens Examined — Colombia. — DEPARTMENT OF CUNDI- NAMARCA: Sibate, 13–15 Oct 1917, Pennel 2508 (GH). — DEPARTMENT OF MAG- DALENA: Santa Marta, Nov 1898 –1901, H. H. Smith 1496 (LL). Ecuador. — PROVINCE OF CHIMBORAZO: Ca non ~ of the Rıo Chanchan, about 5 km north of Huigra, 19–28 May 1945, Camp E-3340 (NY). El Salvador. — DEPARTMENT OF SANTA ANA: Camino entre La Cumbre y el bosque nebuloso de El Olimpo, faldas bajas del Cerro El Aguila, 7 Dec 1993, Linares & Martınez 1197 (MEXU). Guatemala. — DEPT. BAJA VERAPAZ: Purulha, B. V., 21 Sep 1995, Veliz 95–5095 (MEXU, TEX). — DEPT. HUEHUETENANGO: Huehuetenango, Todos Santos Cuchumatan, Xemal , 25 Nov 1998, Veliz 98–6900 (MEXU); Huehuetenango, Chiantla, cerca del Mirador, 26 Sep 1996, Veliz 96–5802 (MEXU, TEX); Barranco in oak forest near Ocubila, 10 km west of Aguacatan, 27 Nov 1962, Williams & Williams 21773 (LL); Huehuetenango, Todos Santos Cuchumatan, La Torre , 25 Nov 1998, Veliz 98–6870 (MEXU, TEX). — DEPT. QUICHÉ: Quiche, Mts. east of Quiche, 20 Nov – 4 Dec 1940, Grant 689 (GH). — DEPT. SACATEPÉQUEZ: Sacatepequez, volcan de Acatenango, Barranca del durazno, 6 Oct 1999, Veliz 99–7351 (TEX); San Lucas Sacatepequez, Cerro Alux., 14 Oct 2005, MacDonald & Mendez 32 (TEX). — DEPT. SOLOLÁ: Ravines near Nahuala, Sierra Madre Mountains, 17 Dec 1962, Williams et al. 23197 (LL). — DEPT. TOTONICAPÁN: Bancos humedos del bosque nebuloso de Marıa Tecun, 21 Nov 1965, Molina R. 15922 (NY). Honduras. — DEPT. MORAZAN: Cerro de Uyuca, 23 Dec 1950, Sauer 1578 (WIS). Mexico. — CHIAPAS: Sierra de Tonala, Sep 1913, Purpus 6809 (GH); Mt. Ovando, near Escuintla, 14 Sep 1945, Matuda 6030 (LL); 11 km al NE de Tuxtla Gutierrez Chiapas, Ca non ~ del Sumidero, 1 Oct 1984, Torres C. et al. 6340 (ASC); Shishintonil, Mpio. Tenejapa, [16.85, -92.49], 18 Nov 1994, Chame & Luna 708 (MEXU); Pase por el camino de Nachig, Zinacantan, 15 Nov 1988, Santos & Martınez 494 (MEXU); San Juan Chamula , 30 Dec 1987, Ruız 456 (MEXU, TEX); Kulaktik, Mpio. Tenejapa, 25 Jan 1983, Ton 5365 (MEXU); Laguna Montebello, Montebello National Park, Muncipio of La Trinitaria, 16 Nov 1972, Breedlove 29538 (MEXU, TEX); 17 km al O de Comitan de Domınguez, por la carretera a San Cristobal de las Casas, 4 Sep 1984, Cabrera & Mendez 7563 (MEXU); 6–10 km north-northeast of La Soledad along logging road from Las Margaritas to Camp Alegre. Municipio of La Independencia, 24 Oct 1976, Breedlove 41024 (MEXU); Steep slope above Rancho Nuevo, 9 miles southeast of San Cristobal la Casas. Municipio of San Cristobal la Casas , 5 Nov 1965, Breedlove 14067 (MEXU, LL, WIS); Loc. a 10 km al este del entronque a Villahermosa, sobre la carretera Tuxtla Gutierrez-San Cristobal de las Casas, 3 Dec 1983, Cabrera & Cabrera 5948 (ASC, MEXU); Extensive marsh at south end of the Valley of San Cristobal, Municipio of San Cristobal Las Casas , 23 Oct 1971, Breedlove & Thorne 21295 (MEXU); San Juan Chamula , 2 Oct 1987, Ruız 210 (TEX); Slope along a small river in the barrio of Tih Ha, paraje of Mahbenchauk. Municipio of Tenejapa, 28 Oct 1966, Ton 1438 (MEXU); Paraje of Yehts 'Uk'um. Municipio of Tenejapa, 23 Nov 1964, Breedlove 7475 (TEX); Tujtiltik, 1 km abajo del pueblo cafetales de Cancuc, 22 Dec 1987, Cruz 336 (MEXU, TEX, WIS); Clınica Yerba Buena, 2 km northwest of Pueblo Nuevo Solistahuacan. Municipio of Puebla Nuevo Solistahuacan, 23–24 Jan 1965, Raven & Breedlove 19835 (TEX); San Juan Chamula , 12 Oct 1988, Ruız 1096 (MEXU, TEX, WIS). — COLIMA: Rancho El Jabalı, 22 km (airline) NNW of Colima in the SW foothills of the Volcan de Colima, Colima/Jalisco line passes through ranch, lakeside 3 km E of house on main road to Lagos Calabozo & Epazote (intersection of main road with road to Caballeriza and airstrip), [19.5, -103.7], 30 Oct 1990, Lott et al. 3032 (TEX); Rancho El Jabalı, 22 km (airline) N of Colima in the SW foothills of the Volcan de Colima, Colima/Jalisco line passes through ranch. Near Lago Epazote, [19.5, -103.7], 5 Oct 1991, Vazquez V. 1366 (MEXU, TEX); Rancho El Jabalı, 20 km (airline) N of Colima in the SW foothills of the Volcan de Colima, El Agostadero, ridgetop in hills above the shrine 4.9 km SE of Hac. San Antonio on the road to Comala, [19.4, -103.7], 7 Nov 1991, Sanders 11813 (MEXU). — DISTRITO FEDERAL: Barrancas Mixcoac, Oct 1930, Lyonnet 1389 (MEXU); El Mirador de Santa Ana, delegacion de Milpa Alta, 30 Oct 1976, Ventura A. 2361 (MEXU); San Francisco Tlalnerantla, Xochimilco, Sep 1984 , Trejo s. n. (MEXU); Sta. Rosa, en ladera humeda, asoleada, matorral denso, 4 Oct 1952, Matuda 26622 (MEXU); In crevices in pedregal near Contreras, 12 Nov 1944, Sharp 441602 (MEXU); Santa Cecilia, delegacion de Xochimilco, 5 Dec 1976, Ventura A. 2458 (MEXU); Pedregal de San Angel , 1 Nov 1965, Sosa s. n. (MEXU); Pedregal de San. Angel, Oct 1913, Reiche s. n. (MEXU); Pedregal, Nov 1932, Lyonnet 1026 (MEXU); Rancho El Conejo, delegacion de Chimilco, 17 Oct 1976, Ventura A. 2325 (MEXU); Pedregal de San Angel , 5 Nov 1980, Madero 456 (MEXU); Pedregal de Tlalpan, 9 Oct 1937, Lyonnet 1686 (MEXU); Pedregal de San Angel , 5 Nov 1980, Madero 600 (MEXU); Bosque de Tlalpan; delegacion Tlalpan, [19.29, -99.20], 30 Oct 2000, Ramos L. 707 (MEXU); Frente a la casita de las Ciencias, 23 Oct 1996, Mera et al. 88B (MEXU); Tlaltequehuacan, municipio de San Andres Chiautla , 8 Nov 1981, Ventura A. 3802 (MEXU); Cerro del Judıo, delegacion de Contreras, 8 Oct 1977, Ventura A. 3085 (MEXU); Cerro de San Miguel , Santa Ana, delegacion de Milpa Alta, 6 Oct 1976, Ventura A. 2260 (MEXU); Primer Dinamo, delegacion de Contreras, 2 Nov 1979, Ventura A. 3592 (MEXU); Mirador Santa Ana, delegacion de Milpa Alta, 13 Nov 1976, Ventura A. 2386 (MEXU). — DURANGO: Mpio. Suchil. Reserva de la Biosfera La Michilıa, Cerro El Almagre, Sierra de Urica, al E de San Juan de Michis , [23.4, -104.1], 16 Oct 1986, Gonzalez 2130 (ASC); Portero El Papalote de Chon Macias, San Juan de Michis, Mpio de Suchil , 29 Oct 1985, Alvarado 468 (MEXU). — GUANAJUATO: Steep rocky mountainsides ca 8 km northeast of Santa Rosa, in second-growth oak-forest, 10 Nov 1970, McVaugh 24139 (MEXU). — GUERRERO: Chialpa de Alvarez, 5.5 mi E of Chilapa on MEX 93, road to Atlixtac, [17.59503, -99.11725], 9 Nov 2006, Ayers et al. 1775 (ASC); Chilapa de Alvarez, 7 mi W of Chilapa on MEX 93 road to Chilpancingo, 205 mi E of Zoquiapa, [17.55865, -99.25012], 8 Nov 2006, Ayers et al. 1771 (ASC); A 22 km al E de Chilpancingo, carr. a Chilapa. Mpio. Chilpancingo, 5 Nov 1983, Tenerio L. & Ramamoorthy 4824 (MEXU); Moist slope near creek about 2 km below Omiltemi, 26 Oct 1944, Sharp 441520 (MEXU); Verde Rico, 3.85 km al SO, [17.70, -99.95], 1 Oct 1999, Soto 17325 (MEXU); Verde Rico, 5.03 km al N, [17.75, -99.88], 8 Dec 1998, Soto 12531 (MEXU); El Alquitran (la antena), municipio de Chilpancingo de los Bravo, 3 Nov 1991, Lozada 1642 (MEXU); San Gabriel , 3 km al E, camino a Tenoxcotitlan, [18.58, -99.65], 23 Oct 1997, Duran 1664 (MEXU); Microondas road up Cerro Alquitran, marked “El Tejocote” on Hwy 95 W of Mazatlan; ca 10 km from Hwy 95, 29 Sep 1983, Anderson 12872 (MEXU, NY). — JALISCO: Shaded bank near Guadalajara, 23 Sep 1889, Pringle 2985 (GH); Volcan Tequila, along road to microwave station, [20.8, -103.8], 23 Oct 1970, Webster & Breckon 15867 (MEXU, TEX); 26.3 km S of Talpa de Allende, on the way to La Cuesta, on road cut in edge of oak forest, 6 Nov 2002, Yahara et al. 2705 (MO); Mpio. Zapotitlan, Rancho El Jabalı, 22 km NNW de Colima, en las faldas del Cerro El Campanario, en las orillas del Rıo Cordoba, [19.46, -103.7] 3 Nov 1990, Vazquez V. & Phillips 250 (MEXU). — MÉXICO: Amatepec, just N of Amatepec, 14.2 mi S of MEX 134 at Tejupilco, 4.5 mi S of Rıo Bejucos bridge on highway, [18.76082, -100.1662], 6 Nov 2006, Ayers et al. 1766 (ASC); Carretera de Tejupilco a Amatepec. San Simon , approx. 15 km al S de Ixtapan y 12 km al N de Amatepec. Milpa cosechada dentro de la curva de la carretera, [18.7, -100.2], 8 Dec 1996, Vibrans 6459 (MEXU); Fraccionamiento La Herradura, municipio de Huixquilucan, 12 Nov 1967, Rzedowski 25121 (LL); Mpio. de Tlalmanalco, carretera que va de Chalco-Amecameca, 9 km al ESE de Chalco, colina que esta a un lado de la carretera, 22 Nov 1976, Garcıa P. 232 (MEXU, WIS); Mesa del venado, Mpio. Temascalcingo, [19.99, -100.09], 27 Sep 2004, Cruz 1240 (MEXU); Mpio. Tlalmanalco, Santo Tomas Atzingo, 1 km al NE del poblado, 2 km al E de la carretera Tlalmanalco-Amecameca, 20 Nov 1981, Garcıa P. 1568 (MEXU, TEX); Cajones, Distr.: Temascaltepec, 17 Nov 1935, Hinton et al. 8655 (TEX); Villa de Allende, 5 Oct 1952, Matuda 26805 (MEXU); 6 km al SE de Cuijingo, Mpio. de Tepetlixpa, 16–17 Nov 1980, Garcıa 911 (MEXU); El Remedio, 4 Nov 1951, Gold 8694 (MEXU); Cerro Pelon, camino hacia la colinia de la mariposa Monarca, [19.37, -100.27], 13 Nov 2005, Melgoza et al. 957 (MEXU); Temascaltepec, 2 mi W of Temascaltepec on road to Valle de Bravo, [19.05353, -100.05653], 7 Nov 2006, Ayers et al. 1769 (ASC); Toll road from Zitacuaro to Toluca, 3 mi W of San Francisco , 1.8 mi W of junction to Villa de Allende, [19.42343, -100.16782], 5 Nov 2006, Ayers et al. 1762 (ASC); Mpio. Ocuilan, Laguna Tonatihua, [19.1, -99.3], 6 Oct 1987, Robles 624 (MEXU); Camino a Llano redondo, ejido San Juan Xoconuxco , [19.36, -100.24], 26 Dec 2005, Tenorio et al. 1705 (MEXU).— MICHOACÁN: Cerca de la cima del Cerro Altamirano (llano), Mpio. Contepec, [19.97, -100.15], 22 Nov 2004, Manrıquez 4518 (MEXU); Hills of Patzcuaro, 8 Nov 1890, Pringle 3337 (MEXU, LL, WIS); Patzcuaro, 8.1 mi E of Tingambato on MEX 14, just W of junction to Pichataro and Paracho, [19.49632, -101.77302], 2 Nov 2006, Ayers et al. 1761 (ASC); 7 km al E de Villa Jimenez , sobre el camino a Copandaro, 5 Oct 1986, Rzedowski 40775 (MEXU); Lado Este del Cerro El Aguila, subiendo por San Nicolas Obispo. Municipio de Morelia , 9 Dec 2007, Tenorio et al. 2378 (MEXU); 1 km SE de Jesus del Monte, Municipio de Morelia, 30 Oct 1988, Medina G. 1425 (MEXU); Agua Verde 3 km E de Copandaro en el Lago Zirahuen. Mun. Villa Escalante, 14 Dec 1980, Gonzalez G. 667 (MEXU); 1.0 km W of Dos Aguas on road to Coalcoman, 18 Nov 1983, Barrie 601 (TEX); Salida a Santa Clara del Cobre, municipio de Patzcuaro, 7 Nov 1985, Escobedo 587 (MEXU); 2.5 km al SE de Ajuno, Carretera Patzcuaro-Uruapan; Mpio. de Patzcuaro, 3 Nov 1986, Mayo 578 (MEXU); Lado Sureste del Cerro El Aguila subiendo por Huatzanguio, municipio de Morelia, [19.62, -101.38], 10 Nov 2007, Garcıa 379 (MEXU); A 5 km al S de Uruapan, camino a Nueva Italia.Mpio. Uruapan, 15 Nov 1983, Martınez S. et al. 5322 (MEXU); Mun. Los Reyes, a 18.5 km al SE de Tinguindin, [19.66, -102.40], 19 Nov 2001, Martınez 34663 (MEXU); Sparsely forested slopes of Cerro Potrerillos, ca. 5 miles north of Cotija and 2 miles south of Jiquilpan, 5–9 Oct 1961, King & Soderstrom 4611 (TEX); Mpio. de Ciudad Hidalgo, Mil Cumbres, a 2.8 km al E de Mil Cumbres, 14 Nov 1988, Ramomoorthy 4519 (MEXU); 2 km al SE del Cerro del Estribo, municipio de Patzcuaro, 8 Dec 1986, Gardu no ~ 2437 (MEXU); Lado NW del Cerro San Miguel , municipio de Patzcuaro, 13 Nov 1985, Barriga 1748 (MEXU); Guanatini-Pamatacuaro; municipio Los Reyes, 16 Oct 1984, Labat 1320 (MEXU); Bonilla, municipio de Patzcuaro, 23 Oct 1991, Escobedo 2238 (MEXU); Patzcuaro, Cerro Estribo, [19.51463, -101.63725], 1 Nov 2006, Ayers et al. 1760 (ASC, MEXU); El Cerrito, 1.5 km al E de Jerahuaro, municipio de Zinapecuaro, 20 Oct 1989, Jasso 1710 (MEXU); Los Tres Cerritos, municipio de Zacapu, 19 Oct 1988, Nu nez ~ 355 (MEXU). — MORELOS: 2–3 km al N de Tepoztlan, sobre la ca nada ~ de la zona arqueologica de Tepoztlan, Mpio. Tepoztlan, [19.0, -99.10], 5 Oct 1986, Flores F. & Cabrera 162 (MEXU). — NUEVO LEÓN: Rancho San Luis Aculco, Chalco , 8 Nov 1980, Hinton 18109 (TEX). — OAXACA: La Sirena, Miahuatlan, 23 Oct 1995, Hinton 26399 (TEX); En Puente Jaltengo a 15 km al N de Candelaria Loxichia, camino Pochutla-Oaxaca, Martınez S. et al. 2470 (ASC, MEXU); Tlaxiaco, 22 Oct 1957, MacDougall s. n. (MEXU); Sto. Tomas Octepec, Tlaxiaca, 17 Nov 1972, MacDougall H518 (NY); Teposcolula, 1 Dec 1895, Robinson 1506 (GH); Santiago Juxtlahuaca, San Sebastian Tecomaxtlahuaca , km 8 de la carretera Tecomaxtlahuaca-San Martın Peras, [17.310, -98.098], 2 Nov 1994, Panero 5427 (NY, TEX); A 35.5 km al NE de Putla, camino a Tlaxiaco, Distr. de Putla, 27 Nov 1983, Martınez S. & Barrie 5808 (MEXU); Barranca de Pajaro on the road to Tlaxiaco, 27 Nov 1983, Barrie 754 (TEX); A 5 km al S de Tlaxiaco, Distrito de Tlaxiaco, 27 Oct 1980, Tellez 3869 (MEXU); Santiago Juxtlahuaca, San Martın Peras , km 16 de la desviacıon a Ca nada ~ de Lobos sobre el nuevo camino a campos de fresas, [17.329, -98.121], 24 Oct 1994, Panero et al. 5283 (TEX); Juxtlahuaca, Santiago Juxtlahuaca, 22 Oct 1997, [17.2674, -98.0], Calzada 22292 (MEXU). — PUEBLA: Parque Estatal General Lazaro Cardenas, [19.03, -98.11], 1 Nov 1994, Romero 584 (MEXU); Between Acatzingo and El Seco, 10 Oct 1944, Sharp 441275 (MEXU); Moist steep bank above river near La Vente, 15 Nov 1944, Sharp 441660 (MEXU); Parque Estatal General Lazaro Cardenas, [19.01, -98.11], 25 Oct 1994, Romero 556 (MEXU); Tepoxuchilt, 27 Oct 1910, Antonio s. n. (MEXU); Parque Estatal General Lazaro Cardenas, [19.03, -98.11], 1 Nov 1994, Romero 598 (MEXU); Malinche San Miguel Canoa , 20 Oct 1968, Boege 1011 (MEXU); Matamoros, 2 Oct 1942, Miranda 2254 (MEXU). Panama. — PROV. HERRERA: Roadside ca 3 km N of Las Minas, 19 Jan 1981, D'Arcy & Sytsma 14287 (MEXU). Peru. — PROV. CONTUMAZÁ: Dept. Cajamarca, 28 Jun 1992, Sagastegui et al. 14748 (ASC). — PROV. OTUZCO: Dpto. La Libertad, Huaranchal, 6 Jun 1958, Lopez et al. 2657 (NY). Venezuela. — ESTADO DE MIRANDA: Los Teques, 29 Nov 1938, Alston 5262 (LL).