Reyna, Saarela & Peterson & Valdés-Reyna, 2014

Saarela, Jeffery M., Peterson, Paul M. & Valdés-Reyna, Jesus, 2014, A taxonomic revision of Bromus (Poaceae: Pooideae: Bromeae) in México and Central America, Phytotaxa 185 (1), pp. 1-147 : 10-24

1. Bromus anomalus Ruprecht ex Fournier (1886: 126) . Figs. 1–3 .

Bromus anomalus Rupr. in Galeotti (1842: 236), nom. nud. Bromopsis anomala (Rupr. ex E. Fourn.) Holub (1973: 167) . Zerna anomala (Rupr. ex E. Fourn.) Henrard (1940: 499) . Type:— MÉXICO. Estado de México: Teotihuacan, July–August 1865, Hahn s.n. (lectotype P-00748832!, designated by Soderstrom & Beaman 1968: 496). We agree with Soderstrom & Beaman that the lectotypes chosen by Shear (1901: 243) [Cerro San Felipe in prov. Oajacensi, 8000 ft, Nov– April 1840, H. Galeotti 5757 (lectotype P-00624352!, isotypes US-A865510! fragm. ex P, LE-00000752!)] and Wagnon (1952: 469)

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[Hidalgo: Real de Monte, H. Galeotii 5815 (P)] are ambiguous.

Bromus meyeri Swallen (1950: 29) . Type:— MÉXICO. Nuevo León: Dulces Nombres , and just Eof border into Tamaulipas, 24ºN, 99.30'–100.0' W , 1850 m, rhizomatous perennial to 2 ft tall, amongst low thorny shrubs on north hillside, 18 June 1948, F. G . Meyer & D. J . Rogers 2561 (holotype US-1962983 !, isotype MO-1598683 !) .

Plants perennial, not rhizomatous. Culms up to 110 cm tall, 2–4 mm wide at base, erect or ascending, bases often weakly to strongly decumbent, glabrous, scabrous or pubescent below inflorescences; nodes 2–5, glabrous to pubescent. Leaf sheaths glabrous, pubescent, or pilose, hairs up to 1.5 mm long, midrib distinctly narrowing below the collar; auricles usually present, sometimes absent; ligules 0.2–1.6 mm long, glabrous, erose-lacerate, ciliate; blades up to 35 cm × 2–4 mm, flat, adaxial and abaxial surfaces glabrous or weakly to moderately pubescent or pilose, hairs up to 1 mm long, margins serrulate. Panicles up to 18 cm long, open, nodding, branches ascending or spreading, scabrous, 1–5 spikelets per branch. Spikelets 1.5–2.8 cm long, 4–12-flowered, elliptic to lanceolate, terete to moderately laterally compressed; glumes usually pubescent, hairs up to 0.4 mm long, occassionally glabrous to puberulent, margins hyaline, midnerves glabrous or scabrous; lower glumes 4.9–7.5(–8.5) mm long, triangular or narrowly lanceolate, 1- or 3-nerved, green, purplish-green to purple along and between the nerves; upper glumes 6–10.5 mm long, obovate-lanceolate, 3-nerved, green, purplish-green to purple along and between the nerves, apices obtuse, acute or mucronate, mucros up to 0.6 mm long; lemmas 7–10 mm long, elliptic to lanceolate, rounded over the backs, apices acute, obtuse or truncate, 5–7-nerved, green, purplish-green to purple along and between the nerves, sometimes ± translucent between nerves, marginal nerves sometimes very faint, moderately to densely pubescent to pilose on the backs and margins, hairs up to 0.8 mm long, longest hairs usually along the margins, backs and margins occassionally glabrous to puberulent with hairs up to 0.2 mm long; awns 1.2–3 mm long, inserted <0.5 mm below the apex of the lemma, straight; paleas shorter and narrower than the lemmas, backs glabrous or pubescent, keels ciliate, cilia up to 0.2 mm long; anthers 1.5–3.2 mm long; caryopses 5−9 mm long, light brown. 2 n = 14 ( Wagnon 1952).

ATAXONOMIC REVISION OF BROMUS (POACEAE)

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Distribution: ―Native. In México B. anomalus is known from the states of Aguascalientes, Baja California Sur, Chiapas, Chihuahua, Coahuila, Distrito Federal, Durango, Guanajuato, Hidalgo, Jalisco, Estado de México, Michoácan, Morelos, Nuevo León, Oaxaca, Puebla, Querétaro, San Luis Potosí, Tamaulipas, Tlaxcala, Veracruz and Zacatecas ( Fig. 4 ). It is present in Guatemala in the states of Solola and Quezaltenango, and in Costa Rica in the states of Cartago and San José ( Fig. 4 ). Bromus anomalus extends north into western Texas ( Pavlick & Anderton 2007) and has also been reported for New México ( Allred 1993).

Ecology: —This common species occurs in the mountains in pine-oak forests on slopes, forest margins, along creeks and ravines, and on rock outcrops, often in calcareous soils; associated with Pinus durangensis Martínez (1942b: 23) , P. cembroides Zuccarini (1832: 392) , P. rudis Endlicher (1847: 151) , P. teocote Schlechtendal & Chamisso (1830: 76) , P. chihuahuana Engelm. in Wislizenus (1848: 103), P. johannis Robert-Passini (1978: 366) , P. ayacahuite C. Ehrenb. ex Schlechtendal (1838: 492) , Picea engelmannii Parry ex Engelmann (1863: 212) , Cupressus arizonica Greene (1882: 64) , Juniperus deppeana Steudel (1840:835) , J. flaccida Schlechtendal (1838: 495) , Arctostaphylos pungens Kunth in Humboldt et al. (1819: 278), Agave montana Villareal-Quintanilla (1996: 191) , Physocarpus Cambessèdes (1824: 239) Rafinesque (1838: 73) , Ceanothus Linnaeus (1753: 195) , Garrya macrophylla Bentham (1840: 50) , Quercus sideroxyla Bonpl. in Humboldt et al. (1809: 39), Q. emoryi Torr. in Emory (1848: 151), Arbutus xalapensis Kunth in Humboldt et al. (1819: 279), A. arizonica ( Gray 1886: 317) Sargent (1891: 317) , Cercocarpus montanus Rafinesque (1832: 146) and C. ledifolius Nutt. in Torrey & Gray (1840: 427). Elevation: 3440–3630 m ( Costa Rica), 2100–3200 ( Guatemala), (790–) 1000–3710 m, usually> 2000 m ( México); sites at elevations below 2000 m are mostly in Coahuila, Nuevo León and Tamaulipas.

Common Names: ― México brome, nodding brome (English); bromo dormilon (Spanish).

Comments: ― Bromus anomalus is the most common and widely distributed species of section Bromopsis in México, where it is used as fodder ( Saulés & Dávila Aranda 1992). The species was not reported for Guatemala by Swallen & McClure (1955) or Soderstrom & Beaman (1968), but was reported there by Pohl & Davidse (1994); they cited Smith 658, as we do here, along with another collection from Guatemala.

Bromus anomalus is the smallest species of sect. Bromopsis in Mexico. Larger individuals of B. anomalus may be confused with B. richardsonii . It can be distinguished from B. richardsonii by its awns 1.2–3 mm long [vs. 3–6.5 mm long], auricles usually present [vs. absent], lemmas 7–10 mm long [vs. (9–)10–13.5(–15) mm long] and blades 2–4 mm wide [vs. (3–)4–9(–13) mm wide]. Acollection from Nuevo León (Peterson et al. 16765) with broad leaves (up to 6.5 mm wide), long lemmas (up to 13 mm), awns up to 4 mm long, pubescent lemmas and glumes, and auricles better fits B. richardsonii , but we keep it here based on the presence of auricles, which we have not seen in material of B. richardsonii . The extreme lengths of the longer awns, longer lemmas and broader leaves observed in this specimen are not included in the key or description, because its placement in B. anomalus is uncertain.

On 6 October 2000 the second author was accompanied by two Tarahumara (Gustavo Arturo Palma Aguirre and Juvencio Antonio Bustillos Ramirez) on a short foray along the edge of the Barranca Río Verde (P.M. Peterson, J. Cayouette, G.A. Palma Aquirre & J.A. Bustillos Ramírez 15372) to collect B. anomalus seed (basiáhuari) used to begin the fermentation process to make a corn drink called tesquino ( Kennedy 1963). This differs from the species Saulés & Davila Aranda (1992) reported for this purpose ( B. carinatus ).

Taxonomic status of Bromus meyeri — Bromus anomalus is a morphologically variable taxon. Individuals typically have hairy lemmas throughout most of the species’ range. Individuals with lemmas glabrous or puberulent from the mountains of Nuevo León and Tamaulipas were described as B. meyeri by Swallen (1950), from a collection gathered at Dulces Nombres, Nuevo León. Aside from noting B. meyeri to be a taxon with auricles and stating, incorrectly, that B. latiglumis (Scribner ex Shear 1900: 40) Hitchcock (1906: 211) , a Canadian/ U.S.A. taxon that does not occur in México, is the only other known species of Bromus to have auricles at the mouths of the sheaths, he did not provide a diagnosis of his new species. Two years after its description, Wagnon (1952) treated B. meyeri as a synonym of B. anomalus , as have some recent authors ( Pavlick 1995, Clayton et al. 2002 onwards). Bromus meyeri was recognized by Soderstrom & Beaman (1968), who suggested it to be most closely related to B. texensis ( Shear 1900: 41) Hitchcock (1913: 381) , a rare species from southern Texas ( Pavlick & Anderton 2007); B. anomalus differs from B. texensis by its longer awns, fewer-flowered spikelets, shorter glumes and lower sheaths short-pilose (vs. villous). Soderstrom & Beaman (1968) did not mention its similarity to B. anomalus . In their key, B. anomalus and B. meyeri are separated by lemma pubescence: "lemmas glabrous, scabrous, or inconspicuously short hirsute" in the lead to B. meyeri , and "lemmas villous over the back or along the margins" in the lead to B. anomalus . Bromus meyeri has also been recognized by Beetle (1977, 1987), Espejo- Serna et al. (2000) and Pavlick et al. (2003), all without comment.

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ATAXONOMIC REVISION OF BROMUS (POACEAE)

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We have studied plants in the field, including in the vicinity of the type location, and herbarium, and we agree with Wagnon (1952) in treating B. meyeri as a synonym of B. anomalus . Although many collections from the Sierra Madre Oriental in Nuevo León have lemmas that are variously glabrous to puberulent (e.g., Peterson, Valdés- Reyna & Sosa Morales 16699, 16703, 16756, 16760, 16763, 16768, 16780, 16783, Peterson & Valdés-Reyna 15872, Peterson & Saarela 21100, Mueller & Mueller 428, 1168, Harvey 987) or have short pubescence (e.g., Peterson & Saarela 21073, 21102)—character states not found elsewhere throughout the range of B. anomalus —the plants otherwise fall within the range of variation of B. anomalus . Plants with densely pubscent lemmas are also present in the same areas (e.g., Peterson & Valdés-Reyna 15824, 15891; Peterson, Saarela & Flores Villegas 21373; Peterson, Valdés-Reyna & Sosa Morales 16753, 16765, 16778; Peterson & Valdés-Reyna, Peterson & Saarela 21103; Mueller 2290; Mueller & Mueller 424, 425, 1058, 1078). Other specimens are intermediate, with dense pubescence along the margins and lemma backs glabrous to puberulent (e.g., Peterson & Valdés-Reyna 15863, 15889). We consider plants previously recognized as B. meyeri to be a local form (in the nontaxonomic sense of the word) of B. anomalus , consistent with varying lemma pubescence found in several other Bromus species , including B. exaltatus and B. richardsonii .

Distinguishing Bromus anomalus and B. porteri ( Coulter 1885: 425) Nash (1895: 512) — Bromus anomalus and B. porteri have long been confused in North America. Bromus porteri has been variously recognized as a distinct species ( Wagnon 1952, Pavlick 1995, Pavlick & Anderton 2007, Saarela 2008, Saarela & Peterson 2012) or as a synonym of B. anomalus ( Kearney & Peebles 1960, Hitchcock et al. 1969, Jones et al. 1997, Douglas et al. 2001), and the characters used to distinguish these taxa have not been applied consistently. Hitchcock (1913) recognized only B. porteri in México. Wagnon (1952) distinguished B. anomalus from B. porteri by lower glumes 1- or 3-nerved [vs. consistently 3-nerved], auricles usually present on lower leaves [vs. absent] and the midrib of the culm leaves tapered just below the collar [vs. not tapered just below the collar]. By these criteria, he did not consider B. porteri to be present in México. By contrast, Soderstrom & Beaman (1968) recognized both B. anomalus and B. porteri in México, and their circumscription of the species was different than that of Wagnon (1952). They treated plants with 1-nerved lower glumes, awnless upper glumes and lemmas usually obtuse as B. anomalus , and plants with 3-nerved lower glumes, upper glumes awn-tipped and lemmas acute or bifid at the apex

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as B. porteri , further noting that some specimens have unusual features. Subsequent to the revision of Soderstrom & Beaman (1968), both taxa have been recognized in México ( Beetle 1977, 1987, Gould & Moran 1981, Rivera & Aranda 2004).

We find the character states used by Soderstrom & Beaman (1968) to be intermixed among specimens, and conclude that they cannot be used to distinguish two taxa. Our circumscription of taxa follows that of Wagnon (1952), which has been utilized in recent treatments of Bromus in North America ( Pavlick 1995, Pavlick & Anderton 2007, Saarela 2008, Saarela & Peterson 2012). By these criteria, we have seen no specimens of B. porteri in México. Bromus porteri is distributed in western North America from Manitoba to British Columbia and south to California and New México ( Pavlick & Anderton 2007).

Specimens Examined:― COSTA RICA. Cartago: Cantón de Turrialba, Cuenca del Matina , Estación Crestones, [9.4978°N, 83.5°W], 3630 m, 6 November 1996, Billen Gamboa R . 828, E. Alfaro & A . Picado (INB- 60061, MO-5175364 ) . San José: Cantón de Pérez Zeledón P. N . Chirripó, 9.4569°N, 83.5106°W, 3460 m, 7 December 1996, E . Alfaro 1028 (CR-221452, INB-60060); Cantón de Pérez Zeledón P. N . Chirripó, 9.4664°N, 83.4972°W, 3440 m, 17 July 1998, E . Alfaro 1734 (INB-60059, MO-5652818 ) . GUATEMALA. Solola: ca. 4 km Eof Godinez , [14.7°N, 91.1°W], 2100 m, 5 December 1963, L. O . Williams, A . Molina R. & T. P. Williams 25190 (F-1652130); cerca la estación de micro-onda de Guatel, Sierra de Chuatroj , Santa Catarina Ixtahuacán, Sololá, 14.74°N, 91.29°W, 3200 m, 17 August 1977, D. N . Smith 658 (F-185163, F-2069944). Quezeltenango: La Esperanza, [14.84°N, 91.52°W], 2400 m, September 1954, M . de Koninck 50 ( US-2151632 , US-2153251 ) . MÉXICO. Aguascalientes: Ladera Ndel Cerro San Juan , 5 km al Ede Tepezela, [22.2167°N, 102.1667°W], 2300 m, J. Rzedowski 25009 ( MICH-1119140 ); Mpio. San José de Gracía, 12 km al SW de La Congoja , 22.09°N, 102.63°W, 2700 m, 16–17 October 1973, J . Rzedowski & R . McVaugh 801 ( MICH-1119142 , NY, US); Sierra Fria Potosina, at 5.6 mi Sof Jtn of road Rancho Cienega de Gallardo , 22.0967°N, 102.7094°W, 2594 m, 30 September 2012, P. M . Peterson, K . Romaschenko, J . Rodriguez Avalos, M . Herrera-Simoni & K . Garcia Rodriguez 24840 (US). Baja California Sur: along trail to El Picacho de la Laguna, Sierra de la Laguna, Eof Todos Santos, 23.5333°N, 110.0833°W, 1830 m, 26 December 1947, A . Carter, A. M . Alexander & L . Kellogg 2395 ( MEXU, US- 1936958 ); La Chuparosa, 23.5667°N, 110°W, 17 October 1893, T. S . Brandegee 73 ( US-591101 ); La Chuparosa, 23.5667°N, 110°W, 2 October 1899, T. S . Brandegee 27 ( US-1009557 ); La Laguna, Sierra de la Laguna, Eof Todos Santos , 23.5667°N, 110°W, 1650 m, 25 December 1947, A . Carter, A. M . Alexander & L . Kellogg 2341 ( US- 1936947 ); La Paz, Sierra La Laguna , Valle, 23.5333°N, 109.9°W, 1800 m, 19 October 1998, M . Domínguez L . 2632 ( ARIZ-351918 , SD-143637 ) . Chiapas: ca. 15 mi SE of Teopisca , [16.45°N, 92.28°W], 2330 m, 21 August 1953, J. R . Reeder & C. G . Reeder 2031 ( RSA-POM-287190 ); Mpio. Zinacantán, along Mexican Hwy . 190 at paraje Granadilla, [16.5°N, 93°W], 4500 ft, 1 July 1965, D. E . Breedlove 10609 ( US-3007632 ) . Chihuahua: 19.6 km W of Balleza and 74.2 km Eof Guachochi, [27.19°N, 106.47°W], 2120 m, 18 September 1991, P. M . Peterson, C. R . Annable & J . Valdés-Reyna 10753 ( US); 30 km de Buenaventura, [29.93°N, 107.18°W], 5 September 1981, Ma. Elena Siqueiros 1401 ( MEXU); Sierra Madre Occidental, 20.3 mi Sof Creel on road towards Rocheachic , 27.5387°N, 107.51°W, 2510 m, 5 October 2000, P. M . Peterson & J . Cayouette 15367 ( CAN, MO, US); Sierra Madre Occidental, 9 mi Sof Guachochi at edge of Barranca Río Verde 1 km along trail descending into canyon, [26.69°N, 107.06°W], 2470 m, 6 October 2000, P. M . Peterson, J . Cayouette, G. A . Palma Aquirre & J. A . Bustillos Ramírez 15372 ( CAN, US); along drainage of Río Candasneno at Cascada de Basaseachic , ca. 1 mi Sof village of Basaseachic along trail leading to cascada, 28.1333°N, 108.25°W, ca. 2000 m, 14 October 1984, R. & M. Spellenberg 7930 ( NY) . Coahuila: 16 mi Sof Saltillo on MEX 57 , [25.22°N, 101.1°W], 6800 ft, 21 July 1970, L. H . Harvey 8501 ( MICH-1119120 ); 21 km SE of Saltillo on Hwy. 57 towards Matehuala, 25.247°N, 100.9061°W, 2460 m, 19 September 2001, P. M . Peterson & J . Valdés-Reyna 15806 ( ANSM, CAN, MO, US); 26 mi Eof Saltillo , 25.4167°N, 101.4°W, 2316 m, 3 September 1960, J. R . Reeder, C. G . Reeder & T. R . Soderstrom 3292 ( US- 2473591 ); 4 km Eof Los Lirios on road to Laguna de Sánchez, 50 km SE of Saltillo , [25.39°N, 100.41°W], 2600 m, 8 October 1988, P. M . Peterson & C. R . Annable 06254 ( US); Cañón de San Lorenzo, ca. 5 a 6 km al Oriente de Buenavista , paraje de “Los Aguajes”, 2000 m, 20 November 1974, J . Marroquín 3075 ( ANSM); El Carmen, 29.15°N, 102.71°W, 26 August 1936, E. G . Marsh, Jr. 628 ( GH); entrance to Monterreal , at 33 mi Eof Arteaga, 25.2365°N, 100.4426°W, 2619 m, 27 September 2001, P. M . Peterson & J . Valdés-Reyna 15942 ( ANSM, CAN, MO, US) & 15943 ( CAN, US); just above Monterreal on road to Mesa de las Tabla and 34.7 mi Eof Arteaga, 25.2291°N, 100.4331°W, 2733 m, 27 September 2001, P. M . Peterson & J . Valdés-Reyna 15947 ( ANSM, CAN, US); Maderas del Carmen , 11.2 mi NE of Los Pilares, 28.9367°N, 102.61°W, 1953 m, 21 September 2007, P. M . Peterson, J. M . Saarela, S . Lara Contreras & J . Reyna Álvarez 20954 ( CAN, MO, US); Maderas del Carmen , 13.7

ATAXONOMIC REVISION OF BROMUS (POACEAE)

Phytotaxa 185 (1) © 2014 Magnolia Press 15 mi NW of Pilares and 0.4 mi Sof El Cinco Junction, 28.9538°N, 102.5853°W, 2365 m, 7 September 2005, P . M . Peterson & J . Valdés-Reyna 18900 ( CAN, MO, US); Maderas del Carmen , 13.8 mi NE of Los Pilares, 28.9536°N, 102.5854°W, 2335 m, 21 September 2007, P . M . Peterson, J . M . Saarela, S . Lara Contreras & J . Reyna Álvarez 20978 ( CAN, MO, US); Maderas del Carmen , 28.9899°N, 102.61°W, 2280 m, 8 September 2005, P . M . Peterson & J . Valdés-Reyna 18917 ( ANSM, CAN, MO, US); Maderas del Carmen , 9.5 mi NW of Pilares, 28.9318°N, 102.61°W, 1890 m, 7 September 2005, P . M . Peterson & J . Valdés-Reyna 18879 ( ANSM, CAN, MO, US); Mesa del Rosario , Santiago, 25.4333°N, 101.0167°W, 6 November 1983, I . Cagral C . 645 ( ANSM); Mpio. Arteaga , Sierra Rancho Nuevo, Cañón de San Juan, ca. 2 km al Nde San Juan, 25.2614°N, 100.3361°W, 2600 m, 31 August 1997, M . A . Carranza 2613, L . Zamora M . & D. Sánchez V. ( ANSM) ; Mpio. Arteaga , Ejido Agua Blanca, [25.44°N, 100.84°W], 2600 m, 22 July 1993, J . Garza 4 ( MEXU); Mpio. Arteaga , Ejido La Efigenia, [25.44°N, 100.84°W], 2460 m, 22 July 1993, P . Lobato 24 ( MEXU); Mpio. Arteaga , Sierra Catana, 40 km SW de Saltillo, [25.15°N, 101.18°W], J . A . Villarreal Q ., M .A. Carranza & J . Valdés R . 6576 ( MEXU); Mpio. Arteaga , Sierra de Arteaga, Cañón de Jamé, 3 mi al Ede Jamé, 25.35°N, 101.0336°W, 2180 m, 16 September 1989, P . S . Hoge 225, M . E . Barkworth, J . Valdés-Reyna & J . A . Villarreal Q. ( ANSM) ; Mpio. Arteaga , Sierra Madre Oriental, 3 km al Ede Los Lirios, 25.3669°N, 100.5833°W, 2604–2636 m, 22 October 2009, J . A . Villarreal 9473, L . Ramírez, M . Zárate & J . Ayala ( ANSM); Mpio. Ocampo , Sierra del Pino , Ejido Acebuches, Cañón La Vaca, 1850 m, 2 October 2000, M . A . Carranza 4072 & Ramírez ( ANSM); Mpio. Ramos Arizpe, Sierra de la Paila, Ejido el Cedral por el Cañón El Carmen, 1300–1600 m, J . Valdés-Reyna 2167 ( ANSM); Mpio. Saltillo , Sierra de Zapaliname, camino al Penitente, en vista del oso a orilla de camino Oeste de la Sierra, 25.3462°N, 101.95°W, 2953 m, 15 October 2005, E . Francisco García 155 & S . G . Gómez Pérez ( ANSM); Mpio. Saltillo , Cerro El Penitente, en exposición noroeste de la sierra, 25.3478°N, 100.9022°W, 2035 m, 16 October 2005, J . A . Encina 1796, F . J . Encina D., J . M . Guillermo E . & S .G. Gómez P. ( ANSM) ; Mpio. Saltillo , pies del Muerto, 25.355°N, 100.95°W, 2480–2560 m, 4 August 2000, J . A . Encina 550 & alumnos de la Escuela Técnica Forestal No. 3 ( ANSM); Mpio. Saltillo , along trail from camino El Cuatro to El Penitente, 25.3497°N, 100.908°W, 2840 m, 28 September 2007, P . M . Peterson, J . M . Saarela & S . G . Gómez Pérez 21123 ( CAN, MO, US) & 21125 ( CAN, MO, US); Mpio. Saltillo , along trail from camino El Cuatro to El Penitente, 25.3548°N, 100.93°W, 1628 m, 28 September 2007, P . M . Peterson, J . M . Saarela & S . G . Gómez Pérez 21121 ( CAN, MO, US); Mpio. Saltillo , along trail from El Cuatro to El Penitente, 25.3494°N, 100.908°W, 2925 m, 28 September 2007, P . M . Peterson, J . M . Saarela & S . G . Gómez Pérez 21130 ( CAN, MO, US); Mpio. Saltillo , Cañón de Santa Rosa - los Aguajes, 25.3294°N, 100.4325°W, 2375 m, 28 June 2001, J . A . Encina 719 con alumnos de la Escuela Técnica Forestal No. 3 ( ANSM); Mpio. Saltillo , above El Penitente on ridge top, 25.346°N, 100.9°W, 3086 m, 28 September 2007, P . M . Peterson, J . M . Saarela & S . G . Gómez Pérez 21135 ( CAN, MO, US); Mpio. Saltillo , above El Penitente, 25.345°N, 100.91°W, 3048 m, 28 September 2007, P . M . Peterson, J . M . Saarela & S . G . Gómez Pérez 21140 ( CAN, MO, US); Mpio. Saltillo , along camino "El Cuatro", Eof Saltillo, 25.3689°N, 100.91°W, 2356 m, 3 September 2005, P . M . Peterson & J . Valdés-Reyna 18814 ( ANSM, CAN, MO, US); Mpio. Saltillo , ca. 5 km Eof Saltillo (Las Palapas) up camino de Cuatro, then up trail towards cumbre, 25.3576°N, 100.9333°W, 2246 m, 20 September 2003, P . M . Peterson, J . Valdés-Reyna & R . H . Cárdenas 17865 ( ANSM, MO, US); Mpio. Saltillo , Eof Saltillo, 25.3468°N, 100.9016°W, 3070 m, 2 September 2005, P . M . Peterson & J . Valdés- Reyna 18799 ( ANSM, CAN, US); Mpio. Saltillo , Eof Saltillo, 25.3536°N, 100.92°W, 2400 m, 2 September 2005, P . M . Peterson & J . Valdés-Reyna 18782 ( ANSM, CAN, MO, US); Mpio. Saltillo , Eof Saltillo, 25.3556°N, 100.92°W, 2100 m, 2 September 2005, P . M . Peterson & J . Valdés-Reyna 18776 ( US) & 18779 ( ANSM, CAN, MO, US); Mpio. Saltillo , Cañon de San Lorenzo, 3 km Sof Saltillo, 23.3333°N, 100.9333°W, 2500 m, 29 August 1995, N . Snow, J . Valdéz Reyna & L . Arce González 6753 ( MEXU, MO); Mpios. Cuatro Ciénegas and Ocampo, Cañón del Agua, 27.0611°N, 102.4111°W, 1900 m, 26 July 1977, T . Wendt & J . Valdés-Reyna s.n. ( ANSM); Predio Santa Anita , en la Sierra de Rancho Nuevo al Nde San Juan de los Dolores, en exposición norte de la sierra, 25.4469°N, 100.6025°W, 2880 m, 10 September 2005, J . A . Encina 1436, F . J . Encina & J . López A. ( ANSM) ; Santa Rita, 2.5 mi Sof La Casita, [25.63°N, 100.34°W], 20 September 1948, Kenoyer & Crum 2915 ( GH); SE of San Antonio de Las Alazanas and SE of Saltillo along road up to Coahuilón, [25.26°N, 100.54°W], 2800 m, 17 October 1989, P . M . Peterson, J . Valdés-Reyna & J . A . Villarreal 08403 ( US); Sierra Catana, 40 km SW de Saltillo , 25.25°N, 101.2°W, J . A . Villarreal 6577, M . A . Carranza & H . Nieto ( ANSM); Sierra de la Concordia, 6 km al NE del Ejido La Casita , 25.2347°N, 100.3333°W, 2100–2300 m, 10 October 1996, J . A . Encina & J . Závala B . 282 ( ANSM) & 283 ( ANSM); Sierra de la Gloria , SE of Monclova, [26.81°N, 101.3°W], 4 March 1939, E . G . Marsh 1947 ( GH, F- 1254893); Sierra de Santa Rosa , [28.44°N, 102.06°W], 20 July 1938, E . G . Marsh 1399 ( GH, F-1254623); Sierra del Carmen , Ejido San Francisco, 28.9298°N, 102.47°W, 1490 m, 5 September 2005, P . M . Peterson, J . Valdés-

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Reyna & C. Sifuentes 18855 ( ANSM, CAN, MO-6192498 , US); Sierra El Astillero, 1.8 mi SW of Santa Rosa and 5.4 mi SW of San Pedro at trail head above water pump, 24.6349°N, 101.1132°W, 2550 m, 21 October 2007 , P. M. Peterson , J.M. Saarela & D. Stancik 21448 ( CAN, MO, US); Sierra El Jardín, 29.0878°N, 102.63°W, 2214 m, 3 September 2006 , P. M. Peterson & S. Lara-Contreras 19927 ( US); Sierra la Encantada, ca. 170 km de Múzquiz, cuesta de Malena, brecha a Boquillas de Carmen , 28.75°N, 102.3333°W, 8 September 1990 , M.A. Carranza C-830 et al. ( ANSM); Sierra Madre Oriental, road towards El Renacer de la Sierra and 36.5 mi Eof Arteaga , 25.2206°N, 100.3896°W, 2821 m, 27 September 2001 , P. M. Peterson & J. Valdés-Reyna 15950 ( ANSM, CAN, MO, US) & 15952 ( CAN, MO, US); 40 mi Sof Saltillo , [25.08°N, 100.36°W], July 1880 , E. Palmer 1372 ( GH, mixed sheet with B. densus ); Rancho Los Angeles, 48 km al sur de Saltillo, 25.4333°N, 100.6331°W, 1800 m, 16 July 1977 , F.A. Ibarro s.n. ( ANSM); La Siberia, Sierra de la Marta, cerca del Ejido Sta. Rita, ca. 20 km al SE de San Antonio de las Alzanas , 26.2333°N, 100.4167°W, 2400 m, 22 July 1983 , J. Valdés-Reyna et al. 159 [?] ( ANSM) ; Wslope of Sierra del Jardín above ( E of) Rancho El Caballo, 29.0083°N, 101.8944°W, 1400–2250 m, 16 September 1972 , F. Chiang , T. Wendt & M.C. Johnston 9293 ( NY); Sierra Madre Oriental, 5 mi Wof Chapultepec on cutoff road between Hwy. 54 & 57, 23 mi Sof Saltillo , 2410 m, 16 October 1995 , P. M. Peterson & M. B. Knowles 13271 ( US); Sierra El Pino, 33.5 km Wof Rancho El Cimarron , 1850 m, 14 September 1991 , P. M. Peterson & C. R. Annable 10651 ( US); Bosques de Montana, 5.7 mi Eof Highway 57 on road towards Los Lirios, 25.3878°N, 100.7056°W, 2144 m, 19 September 2012 , P. M. Peterson , K. Romaschenko & J. Valdés-Reyna 24574 ( US). Distrito Federal: 16 mi Eof Amecameca on road towards Puebla, in Parque Nacional Izta-Popo , 19.0918°N, 98.68°W, 3390 m, 13 October 2001 , P. M. Peterson & Y. Herrera-Arrieta 16167 ( CAN, MO, US); a las faldas del Cerro Teuhtli, Delegación Milpa Alta , [19.19°N, 99.02°W], 2470 m, 24 October 1985 , E. Manrique et al. 1239 ( MEXU- 1110814 ); Cerro Magdalena, Serranía de Ajusco , 19.2333°N, 99.1667°W, 23 October 1937 , E. Lyonnet & J. Elcoro 1881 ( MO, US-1746400 ); Delegación Tlalpan, Volcán El Pelado, [19.15°N, 99.22°W], 3200 m, July 1985 , A. Miranda et al. 28 ( MEXU); faldas del volcán Teuhtli, Delegación Milpa Alta, [19.19°N, 99.02°W], 2510 m, 13 November 1985 , A. Miranda & P. Guerrero 250 ( MEXU-1110803 ); San Angel, [19.34°N, 99.19°W], 12 November 1929 , M. St. Pierre 836 ( MICH-1119126 ); San Salvador Cuauhtenco, cerro San Bartolo Delegación Milpa Alta, [19.19°N, 99.09°W], 3070 m, 10 October 1985 , J.G. Serrano 268 ( MEXU); Topilejo, 19.2008°N, 99.1428°W, 5 December 1925 , G. Woronow & S. Juzepczuk 795 ( US-1274753 ); Vallede México , Topilejo, 19.2008°N, 99.1428°W, 2500 m, 3 August 1952 , E. Matuda 26201 ( MEXU, US-2119846 ); vertiente NE del Cerro Pelado, cerca de Parres, delegacíon de Tlalpan, [19.15°N, 99.22°W], 3400 m, 7 August 1983 , J. Rzedowski 38156 ( MEXU- 849547 , MO); Volcán Pelado , s.d., [19.15°N, 99.22°W] , J. López P. et al. PELE35- 2 ( MEXU-770226 ) ; Wof km 30 - Cuernavaca Hwy., [19.13°N, 99.18°W], 7 August 1940 , I.K. Langman 2612 ( US-2436903 ). Durango : 0.8 miS of Francisco I. Madero and 2.3 mi Nof Canoas , 22.6482°N, 104.2895°W, 2720 m, 2 October 2001 , P. M. Peterson & M.S. González-Elizondo 16030 ( CAN, MO, US); 18.4 mi Sof Mezquital on road towards Cuesta Blanca , 23.3475°N, 104.3331°W, 2382 m, 7 September 2006 , P. M. Peterson & F. Sánchez Alvarado 19975 ( CAN, US); 2 mi Sof Fancisco I. Madero and 1.2 mi Nof Canoas , 22.6322°N, 104.2926°W, 2700 m, 1 October 2001 , P. M. Peterson & M.S. González-Elizondo 16012a ( CAN, US); 2.7 mi Nof Fancisco I. Madero and 2.4 mi Sof Ciemega del Oro , 22.6891°N, 104.2777°W, 2800 m, 1 October 2001 , P. M. Peterson & M.S. González-Elizondo 16025 ( CAN, US) & 16026 ( CAN, US); 2.8 mi Nof Ciénega del Oro and 7.3 mi Nof Fancisco I. Madero, 22.7434°N, 104.2525°W, 2780 m, 1 October 2001 , P. M. Peterson & M.S. González-Elizondo 16028 ( CAN, MO, US); 20 miS of El Mezquital , 23.3154°N, 104.3434°W, 2522 m, 11 September 2003 , P. M. Peterson & F. Sánchez-Alvarado 17731 ( MO, NY, US); 3.4 mi NE of El Encinal on road towards Minas Promontoria , 25.1802°N, 105.1619°W, 2407 m, 2 October 2002 , P. M. Peterson , M. S. González-Elizondo & L.E. Brothers 16914 ( CAN, US); 4.7 mi NE of El Encinal on road towards Minas Promontoria , 25.1885°N, 105.1516°W, 2575 m, 2 October 2002 , P. M. Peterson , M. S. González-Elizondo & L.E. Brothers 16922 ( CAN, US); 40.3 mi Wof Huejuquilla towards Canoas , 22.6324°N, 104.2204°W, 2430 m, 30 September 2001 , P. M. Peterson & M.S. González-Elizondo 16004 ( CAN, MO, US); 5 mi Wof La Ventana on road towards La Guajolota, 22.9618°N, 104.4986°W, 2486 m, 12 September 2003 , P. M. Peterson & F. Sánchez-Alvarado 17763 ( US); 5 mi Wof Llano Grande along Durango-Mazatlan Hwy. , 23.8412°N, 105.271°W, 2499 m, 9 September 1967 , J. R. Reeder & C.G. Reeder 4904 ( MSC-231763 ; US- 3059166 ); 53 km Wof Durango on Hwy. 40 towards El Salto, 25.9161°N, 104.9904°W, 2460 m, 6 October 2002 , P. M. Peterson & L. E. Brothers 16942 ( US); ca. 1.5 km below Santa Bárbara along Arroyo Santa Bárbara , 23.6581°N, 105.4314°W, 2700 m, 16 September 2003 , P. M. Peterson , S. González-Elizondo & G.A. Teña-González 17801 ( CAN, MO, US); Coyotes Hacienda, 63 road mi W-SW of central Durango , 23.8167°N, 105.3333°W, 2400–2500 m, 1 September 1951 , J.H. Maysilles 7621 ( MICH-119155 , US-2119100 ); Mpio. Mezquital , 19.7 miS

ATAXONOMIC REVISION OF BROMUS (POACEAE)

Phytotaxa 185 (1) © 2014 Magnolia Press 17 of Mezquital on road towards Mesa La Gloria, 23.3172°N, 104.3381°W, 2500 m, 19 September 2005, P. M. Peterson & F. Sánchez Alvarado 19023 ( CAN, MO, US); Sierra Madre Occidental , 0.5 mi SE of Los Charcos, 23.016°N, 104.2897°W, 2690 m, 21 September 2005 , P. M. Peterson & F. Sánchez Alvarado 19048 ( US); Sierra Madre Occidental , 20 mi Sof El Mezquital, 23.3156°N, 104.3433°W, 2522 m, 11 September 2003 , P. M. Peterson & P. Catalán 17731 ( RSA-POM-725717 ); Sierra Madre Occidental, 3.2 mi SW of Las Bayas on road towards Ceballos , [23.53°N, 104.83°W], 2780 m, 10 October 2000 , P. M. Peterson , J. Cayouette & Y. Herrera-Arrieta 15419 ( US); Sierra Madre Occidental, 56 km Wof Durango along Hwy . 40 towards El Salto and 0.4 mi Eof entrance to Parque El Tecuán , 23.9109°N, 105.0252°W, 2536 m, 3 October 2007 , P. M. Peterson & J. M. Saarela 21251 ( CAN, US); Sierra Madre Occidental, 6.8 mi Eof Los Canoas on road towards Huejuquilla El Alto, 22.6349°N, 104.2163°W, 2460 m, 21 September 2005 , P. M. Peterson & F. Sánchez Alvarado 19062 ( US); Sierra Madre Occidental, 68 km Wof Durango on Hwy. 40 towards El Salto, 23.8827°N, 105.122°W, 2720 m, 8 October 2000 , P. M. Peterson & J. Cayouette 15401 ( CAN, MO, US); Sierra Madre Occidental, along Hwy . 40 from Durango City to El Salto, 30 mi Wof Durango, 2560 m, 24 September 1992 , K. W. Allred 5846 , T. Columbus & J. Valdés-Reyna ( ANSM); Sierra Madre Occidental, at 20 km mark up road towards Topia and 19.1 mi NW of Papasquiaro, 25.1158°N, 105.5838°W, 2580 m, 11 October 2000 , P. M. Peterson , J. Cayouette & M. S. González- Elizondo 15425 ( CAN, MO, US); 14.4 mi Sof Casas Blancas, and 2.5 mi Nof Unión y Progreso , 25.2567°N, 104.9731°W, 3140–3250 m, 29 September 2010 , P. M. Peterson , J. M. Saarela & K. Romaschenko 23687 ( US); Sierra Madre Occidental, 20 mi SE of Mezquital on road to Charcos , 2450 m, 7 November 1995 , P. M. Peterson , M. B. Knowles , C. Dietrich , S. Braxton & M. S. González-Elizondo 13635 - A ( US). Guanajuato : 2–4 km Eof Guanajuato, 21.0167°N, 101.22°W, 16 October 1952 , E. R. Sohns 291 ( US-2116761 ); ca. 12 mi from Guanajuato on road to Santa Rosa , 21.1°N, 101.13°W, 29 October 1946 , H. E. Moore, Jr. 1331 ( US-1963086 ); ca. 2 km NE of San Miguel, Cerro de la Presa 20.7167°N, 100.6833°W, 31 October 1952 , E. R. Sohns 498 ( US-2116916 ); ca. 2.5 km Sof Guanajuato, on the slopes of La Bufa, 20.99°N, 101.25°W, 18 October 1952 , E. R. Sohns 346 ( US- 2116796 ); ca. 6 km Eof Guanajuato, 21.0167°N, 101.19°W, 17 October 1952 , E. R. Sohns 299 ( MICH-1119151 , MO, US-2116766 ); Cerro Culiacán , cerca de la torre de microondas, Ede Victoria de Cortázar, [20.65°N, 101.33°W], 2800 m, 31 August 1981 , Rafael Guzmán 4465 ( MEXU); Cristo Rey en el Cerro del Cubilete , [21.01°N, 101.37°W], 2 September 1981 , A. A. Beetle 7376 ( MEXU-10153 ); Mpio. Cortázar, Cerro de Culiacan , [20.65°N, 101.33°W], 2800 m, 1 September 1981 , J. Sánchez Castellanos 97 - J ( MEXU); Mpio. Dolores Hidalgo, Ejido San Cristóbal, al sureste de la comunidad El Terreno, 20.9033°N, 101.0117°W, 2700 m, 2 July 1997 , J. J. Macías Cuéllar 843 - JJMC ( MEXU); Mpio. Guanajuato, camino Valencia-Cubilete , [21.02°N, 101.26°W], 2300 m, 2 September 1981 , J. Sánchez Catellanos 123 - J ( MEXU); Mpio. León , Vergel de la Sierra, 21.3197°N, 101.6278°W, 2627 m, 14 July 1997 , S. Rojas Villegas 494 ( MEXU); Mpio. Ocampo, Ejido "Laguna de Guadalupe ", [21.65°N, 101.48°W], 2210 m, 7 October 1992 , S. Rojas Villegas 11 ( MEXU-1089790 ); 0.5 km Eof Jofre, 21.5239°N, 100.4803°W, 2434 m, 16 September 2010 , P. M. Peterson , J. M. Saarela , K. Romaschenko , J. Herrera-Simoni & M. Herrera-Simoni 23423 ( US). Hidalgo : 13 mi Eof Pachuca, 20.1167°N, 98.5333°W, 2621 m, 11 September 1963 , J. R. Reeder & C. G. Reeder 3725 ( US-2473588 ); 20 km al NE de Zimapan, [20.74°N, 99.38°W], 2200 m, 28 October 1965 , L. González Quintero 3287 ( MICH-1119149 ); 4 mi Wof Tulancingo, 20.07°N, 98.43°W], 8 September 1962 , F. W. Gould 10164 ( MICH-1119137 ); by Río Panotlan, between Zacualtipán and Olotla on road to Metztitlan , 20.77°N, 98.6667°W, 1600–2000 m, 15 October 1949 , H. E. Moore, Jr. 5343 ( MEXU-143526 , MICH-1119130 , GH); ca. 2 mi Sof Barranca de los Marmoles along Pan-American Hwy ., 20.9667°N, 99.2°W, 2300 m, 4 August 1950 , J. R. Reeder , C. G. Reeder & L. N. Goodding 1642 ( RSA-POM- 253209 , US-2473575 ); ca. 46 mi Sof Tamazunchale on Hwy. 85, [21.06°N, 99.06°W], 28 September 1976 , F. W. Gould 14835 ( MEXU-957936 ); Jacala , [21.01°N, 99.19°W], 5000 ft, 8 July 1939 , V. H. Chase 7364 ( ANSM, MICH-1119281 , NY, GH, US-1763084 ); Mpio. Jacala , 21.0167°N, 99.1833°W, 1372 m, 26 June 1939 , V. H. Chase 7172 ( US-1763085 ); Mpio. Tepeapulco, Cerro del Tecolote , [19.7667°N, 101.8667°W], 2500 m, 19 August 1975 , A. Ventura A. 136 ( ANSM); Mpio. Tlanalapán , 3 km al NNE de Tepeapulco, and 1 km al Ede la carretera Tepeapulco-Sto. Tomás , 19.7833°N, 98.55°W, 2560 m, 16 September 1976 , S. D. Koch 76160 ( MEXU-270644 , MO, US-2824599 ); near Tula , 20.05°N, 99.35°W, 3–4 July 1905 , J. N. Rose , J. H. Painter & J. S. Rose 8357 ( US- 451850 ); Pachuca , 20.1167°N, 98.7333°W, 2438 m, 6–7 September 1910 , A. S. Hitchcock 6741 ( US-1009554 ); Real del monte Hidalgo , 20.1333°N, 98.6667°W, 1 October 1910 , I. W. Clokey 1911 ( US-726920 ). Jalisco: km 6 carretera La Manzanilla a Mazamitla , La Manzanilla, 19.97°N, 103.19°W, 1800 m, 21 July 1973 , C. Luis Diazhuna 4240 ( RSA-POM-302097 ) ; Nslopes of the Nevado de Colima , Wof summit of the N ridge, near junction of the old pack road to Zapotlán with Atenquique-Jazmín road, 19.65°N, 103.5333°W, 2100–2200 m, 15 October 1952 ,

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R. McVaugh 13509 ( MEXU-5983 , MICH-1119144 , US-2150832 ) ; NE slopes of the Nevado de Colima, below Canoa de Leoncito , 19.6167°N, 103.65°W, 2250–2550 m, 13 October 1952 , R. McVaugh 13501 ( MICH-1119141 , US-2118493 ); Sierra del Tigre, 3 mi Sof Mazamitla , 19.88°N, 103.02°W, 2100–2200 m, 16 September 1952 , R. McVaugh 12969 ( MEXU-5984 , ICH-1119147, US-2150768 ); Zapotlán , 19.75°N, 103.5°W, 1524 m, 22–25 September 1910 , A.S. Hitchcock 7241 ( US-1009555 ). Estado de México: 50 mi SE of San Juan del Río, Queretaro, on Hwy . 55, 19.76°N, 99.86°W, 2652 m, 6 September 1959 , T.R. Soderstrom 518 ( US-2378274 ); Mt. Popocatépetl , 19.0333°N, 98.6333°W, 2743 m, 5–6 August 1910 , A.S. Hitchcock 6011 ( US-1009553 ); a 3 km sobre la desviación a Rancho Viejo, carretera Toluca-Temascaltepec, 2610 m, 25 October 1983, Manrique, Jaramillo & Núñez 442 ( MEXU-1098033 ); carretera Tepotzotlán-Villa del Carbón , 1/ 2 km de la desviación Arcos del Sitio , 19.72°N, 99.46°W, 2300 m, 30 July 1981 , R. Guzmán M. 3938 ( MEXU-1110770 ); carretera Villa Victoria-El Oro , [19.64°N, 100.17°W], 2570 m, 9 August 1981 , R. Guzmán 4246 ( MEXU); Cerro La Mesa Ahumada, [19.8342°N, 99.2033°W], 2400 m, 6 September 1980, Romero & Rojas 410 ( MEXU-674482 ); Chapa de Mota, 19.9914°N, 99.5161°W, 2800 m, 1 November 2000, Adrian Vega 108 ( MEXU); Crucero Sultepec-camino al Nevado de Toluca, [19.14°N, 99.74°W], 7 September 1982 , R. Guzmán 6009 ( MEXU); entre Magdalena y Tapaxco , [19.7667°N, 100.0667°W], 9 August 1981 , R. Guzmán 4259 ( MEXU); Jilotepec , 19.97°N, 99.53°W, 2200 m, 21 September 1952 , E. Matuda 26709 ( MEXU); Mpio. Amecameca, km 20 carretera Amecameca-Tlamacas , [19.09°N, 98.69°W], 2 October 1992 , A. Miranda & G. Villegas 652 ( MEXU); Mpio. Donato Guerra, a 1.25 km al Nde Las Mesas de San Martín, 19.3478°N, 100.08°W, 2672 m, 29 October 1998 , J.A. García Ruíz 2 ( MEXU); Mpio. Huizquiluca, Rancho et Hielo, km 22 carretera Naucalpan-Toluca , [19.36°N, 99.58°W], 3050 m, 1 October 1992 , A. Miranda et al. 587 ( MEXU); Mpio. Jocotitlán, ladera SE del Cerro Jocotitlán , [19.74°N, 99.76°W], 3450 m, 17 October 1990 , M. González Ledesma & J. Gracía P. 291 ( MEXU-737772 ); 1 mi Wof Río Frio on road to Puebla from México City, 28 July 1950 , J. R. Reeder & C.G. Reeder 1495 ( RSA-POM-253392 ); 1.9 mi Wof Chiltepec on Hwy. 12, 18.9268°N, 99.8442°W, 2596 m, 10 October 2007 , P. M. Peterson , J. M. Saarela & M.J. Flores Villegas 21363 ( CAN, US); 12.1 mi NE of San Miguel, SW of Volcán Toluca , 19.0317°N, 99.9102°W, 2826 m, 10 October 2001 , P. M. Peterson & Y. Herrera-Arrieta 16140 ( CAN, US); 2 km al Wde San Miguel Tequixquiac , [19.91°N, 99.15°W], 23 June 1968, 2400 m , J. Rzedowski 25826 ( MICH-1119131 , MSC-223039 ); 2.8 mi Eof Ocuilán Arteaga , 18.9808°N, 99.3786°W, 2478 m, 11 October 2007 , P. M. Peterson , J. M. Saarela & M.J. Flores Villegas 21372 ( CAN, MO, US) & 21373 ( CAN, US); 20 mi Sof México City , [19.14°N, 99.02°W], 1 September 1946 , A. I. Ortonburger , J. B. Paxon & F. A. Barkley 16 M741 (F-1408939) ; 27 mi Sof San Juan del Río , [20.09°N, 99.7°W], 28 July 1960 , F.W. Gould 9205 ( MICH-1119122 ); 3 mi Wof Avila Camacho , 9000 ft, 9 August 1971 , L.H. Harvey 8791 ( MICH-1119292 ); 3.6 mi Sof Aculco on Hwy. 55, 20.0439°N, 99.87°W, 2602 m, 8 October 2007 , P. M. Peterson , J. M. Saarela & M.J. Flores Villegas 21332 ( CAN, MO, P-03216872 , US); 5.6 miNE of La Comunidad along Hwy. 134 towards Toluca , 19.1848°N, 99.87°W, 2980 m, 9 October 2001 , P. M. Peterson & Y. Herrera-Arrieta 16134 ( CAN, US) & 16135 ( CAN, MO, US); 55 km SE México City , 19.0674°N, 98.7858°W, 3200 m, 14 July 1942 , J.N. Weaver 791 ( US-2436904 ); ca. 1 mi W of Río Frio on road to Puebla from México City, 19.35°N, 98.6667°W, 28 July 1950 , J. R. Reeder & C.G. Reeder 1495 ( US-2473590 ); ca. 3 km (air) Wof Monte Río Frio Pass , (9 km Wof Río Frio), near top of Wslope Monte Río Frio, along Pan Am Hwy. , 19.35°N, 98.75°W, 2900 m, 18 August 1960 , H. H. Iltis , R. Koeppen & F.S. Iltis 1049 ( MICH-1119139 , US-2380090 ); ca. 5 km Wof San Andreas, Pedregal de San Angel , 19.37°N, 99.18°W, 13 October 1952 , E.R. Sohns 178 ( MICH-1119157 , US- 2116655 ); mile Sof Tlalnepantla , 19.55°N, 99.1833°W, 17 July 1940 , I.K. Langman 2457 ( US); Mpio. Amecameca, Zumpango, carretera al Paso de Cortés, 16 km al Edel entronque con la carretera Amecameca- Cuautla, Mor. 9., 19.06°N, 98.7°W, 3300 m, 20 November 1976 , S.D. Koch 76275 A ( US-2824582 ); Mpio. Texcoco, 2 km al SE de San Pablo Ixayoc, sobre el camino hacia el aguaje, 12.5 km al SE de Texcoco , 19.42°N, 98.83°W, 2670 m, 21 October 1976 , S. D. Koch 76213 ( MO, US-2824631 ); Mpio. Texcoco, Lado Sur de la Cañada de Aguas, 13.5 km al SE de Texcoco (11 km al ESE de Coatlinchón , por el camino que pasa., 19.41°N, 98.82°W, 2740 m, 29 July 1977 , S. D. Koch 77111 ( US-2832210 ); near La Cima, along old Cuernavaca Hwy. ( Rt. 95) between km 43 and 44, [19.58°N, 99.26°W], 1 September 1961 , E.K. Longpre 417 ( MSC-232149 ); near Toluca , 19.3°N, 99.65°W, 4 September 1903 , J. N. Rose & J.H. Painter 6796 ( NY, US-450355 ); Oro , 19.8°N, 100.1333°W, 2500 m, 28 September 1952 , E. Matuda et al. 27258 ( US-2119847 ); Paso de Cortés, [19.09°N, 98.64°W], 3680 m, 17 September 1958 , J.H. Beaman 2577 ( GH, MSC-162396 ); Tehuacán area, along Tehuacán-Orizaba highway just above Azumbilla , 18.7°N, 97.41°W, 1500–1800 m, 18 July 1961 , C. E. Smith Jr ., F. A. Peterson & N. Tejeda 3962 ( US-2382355 ); Temascaltepec District, Tequesquipan , 19.0578°N, 99.9506°W, 2480 m, 18 August 1932 , G.B. Hinton 133 ( GH, US-1612064 ); Toluca , 19.3°N, 99.65°W, 2682 m, 13 September 1910 , A.S. Hitchcock 6895 ( US-

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1009564); Volcán de Toluca , 19.1014°N, 99.7694°W, 3550 m, 18 October 1953, E. R . Sohns & E . Matuda 1011 ( US-2118917 ). Michoácan : 0.6 mi Nof Huajumbaro, 19.6906°N, 100.74°W, 2360 m, 8 October 2001, P. M . Peterson & Y . Herrera-Arrieta 16131 ( CAN, MO, US); 1 km al SE de Zíngiro, sobre el camino a Erongarícuaro , [19.56°N, 101.71°W], 2400 m, 2 November 1989, J . Rzedowski 49209 (F-2120708); 5.8 mi NE of Ario de Rosales off Hwy. 120, up road towards Microondas , 19.2115°N, 101.63°W, 2200 m, 8 October 2001, P. M . Peterson & O . Rosales 16121 ( CAN, MO, US); 9 mi Wof Zacapú , 19.82°N, 101.92°W, 2408 m, 18 October 1966, J. R . Reeder & C. G . Reeder 4747 ( US-2982194 ); above Lago de Pátzcuaro , 22 km (by road) NW of Quiroga on Hwy. 15, 19.75°N, 101.6667°W, 2050 m, 24 September 1965, K . Roe & E . Roe 2068 ( MICH-1119132 , US-3115393 ); ca. 1 km Sof El Tren , and km 10 NW from junction with MEX Hwy. 15 ( México-Morelia ), on road to and SSE of Zinapecuaro, 19.75°N, 100.75°W, 2300 m, 27 September 1978, H. H . Iltis, J . Doebley & A . Lasseigne 680 ( MEXU); ca. 12 mi W of Quiroga, 19.76°N, 101.69°W, 2134–2743 m, 3 October 1953, E. R . Sohns 801 ( US-2118723 ) & 802 ( US- 2118724 ); ca. 20 mi Wof Morelia, 19.63°N, 101.47°W, 1524–2134 m, 3 October 1953, E. R . Sohns 762 ( US- 2118691 ) & 765 ( US-2118694 ); cerca del Puerto de los Copales, 8 km al Ede Morelia , sobre el camino a Mil Cumbres, 2050 m, 8 October 1986, J . Rzedowski 40850 ( ANSM); Cerro Grande de Cujaruato, al SW de la Piedad , [19.99°N, 102°W], 2500 m, 16 November 1971, J . Rzedowski & R . McVaugh 552 ( MICH-1119146 ); Cima de Cuernavaca , September 1924, E . Lyonnet 47 ( GH, MO, NY); Mpio. de Tangancicuaro, Las Cañas, estribaciones inferiores noroccidentales del Cerro Patamban , [19.75°N, 102.34°W], 2500 m, 19 November 1971, J . Rzedowski & R . McVaugh 622 ( MICH-1119145 , NY); MEX 15, 56 mi Wof Morelia, 20.19°N, 102.39°W, 7000 ft, 22 August 1970, L. H . Harvey 8682 ( MICH-1119148 ); Morelia, Quinceo , 19.7167°N, 101.2333°W, 2800 m, 11 November 1909, G . Arséne 7286 ( US-1000494 ); Mpio. Ocampo, camino a Tintanueras , Ejido El Rosario, 19.59°N, 100.2656°W, 3400 m, 20 September 2001, Ma. Guadalupe Cornejo Tencrio & G . Ibarra Manríquez 314 ( MEXU- 1062900 , NY); Parte m s alta de Cerro Grande de Cujaruato , al SW de la Piedad , 20.1°N, 102.2°W, 2500 m, 16 November 1971, J . Rzedowski & McVaugh 552 ( US-3096843 , NY); 27 km to Zacapu , [19.82°N, 101.79°W], 27 August 1981, A. A . Beetle M-7088 ( MEXU-1088986 , MO); Mpio. Santa Clara del Cobre, Zirahuén , [19.44°N, 101.74°W], 2150 m, 3 September 1985, J. M . Escobedo 43 ( MEXU, RSA-POM-356516 ). Morelos: 14 km antes de llegar a Cuernavaca por la carretera federal, [18.99°N, 99.19°W], 25 July 1980, G . Andrade & C. H . Ramos s.n. ( MEXU-1089786 ); Cuernavaca-México City road, km 45[?], Nof Tres Cumbres, [19.18°N, 99.16°W], ca. 3100 m, 18 July 1954, G . B. van Schaack 3418 ( ANSM, MICH-1119129 ); Toro , 19.0939°N, 99.2414°W, 5 August 1924, G. L . Fisher 120 ( MO, NY, US-1207917 ); Lagunas Zempoala , 19.0494°N, 99.3175°W, 17 September 1938, E . Lyonnet 2495 ( US-1761291 ). Nuevo León: 10.6 mi NE of San Antonia de Peña Nevada on road towards La Siberia , 23.8238°N, 99.8757°W, 2653 m, 22 September 2002, P. M . Peterson, J . Valdés-Reyna & M . Sosa Morales 16783 ( ANSM, CAN, MO, US); 3 mi Nof La Siberia on road towards La Encantada, 23.8743°N, 99.8058°W, 2826 m, 21 September 2002, P. M . Peterson, J . Valdés-Reyna & M . Sosa Morales 16768 ( ANSM, CAN, US) & 16765 ( ANSM, CAN, US); 4 km W of Tepehuanes on road towards Zaragoza , 23.9349°N, 99.7312°W, 1980 m, 21 September 2001, P. M . Peterson & J . Valdés-Reyna 15863 ( CAN, MO, US) & 15872 ( CAN, US); 4.6 mi NE of Dulces Nombres , 23.9707°N, 99.54°W, 1625 m, 23 September 2001, P. M . Peterson & J . Valdés-Reyna 15930 ( ANSM, US); 54 km W of Linares on Hwy. 3, 24.5229°N, 99.9874°W, 2476 m, 22 September 2001, P. M . Peterson & J . Valdés-Reyna 15891 ( CAN, US); 7.1 mi Sof Zaragoza on road towards Ejido La Encantada , 23.9493°N, 99.7934°W, 1848 m, 21 September 2002, P. M . Peterson, J . Valdés-Reyna & M . Sosa Morales 16753 ( ANSM, CAN, MO, US); 8 km Sof La Cruata on Hwy. 3 towards Aramberri , 24.4421°N, 99.9553°W, 2313 m, 22 September 2001, P. M . Peterson & J . Valdés-Reyna 15889 ( ANSM, CAN, US); 9.8 mi Sof Zaragoza on road towards Ejido La Encantada , 23.9313°N, 99.7947°W, 2155 m, 21 September 2002, P. M . Peterson, J . Valdés-Reyna & M . Sosa Morales 16756 ( ANSM, CAN, MO, US); carretera San Roberto-Galeana, km 78, 25.3°N, 100.24°W, 1900 m, 2 September 1997, M . Castillo B. & P . Jauregui R . 285 ( MEXU); Cerro de la Silla, near Monterrey , 25.6333°N, 100.2333°W, 1575 m, 23 June 1939, L. H . Harvey 987 ( MICH-1119271 , US-1762719 ); Chipinque , [25.72°N, 100.38°W], 1 July 1947, F. A . Barkley, G. L . Webster & C . Rowell 7129 (F-1499765) & 7149 (F- 1500318); Hacienda Pablillo, Galeana , [24.8333°N, 100.38°W], 14 August 1936, M . Taylor s.n. (F-862670); Hacienda Vista Hermosa , 35 mi Sof Monterrey, [25.17°N, 100.31°W], 810 m, 25 June 1939, L. H . Harvey 1040 ( MICH-1119288 ); La Casita, [25.63°N, 100.34°W], 22 August 1948, Kenoyer & Crum 3051 ( GH); Mpio. Linares , El Pinal-Las Palmas, 1500 m, 10 October 1980, J . Ortiz E . s.n. ( ANSM); near Chipinque, on Sierra Anáhuac , Sof Monterrey, [25.72°N, 100.38°W], 1800 m, 8 June 1952, F. W . Gould 6324 ( MICH-1119133 , RSA-POM-108458 ); on Chipinque, Sierra Madre Mountains behind Monterrey , 25.9167°N, 100.3833°W, 7 September 1948, G. L . Webster & E . Aguirre 2941 ( US-2078874 ); Puerto del Aire , Santiago, 790 m, 22 May 1983, I . Cabral C . 305

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(ANSM); road to La Tinaja, 23.8873°N, 99.7956°W, 2500 m, 21 September 2002, P. M. Peterson, J. Valdés-Reyna & M. Sosa Morales 16763 ( ANSM, CAN, MO, US) ; Sof Zaragoza on road towards Ejido La Encantada, 23.9276°N, 99.7978°W, 2171 m, 21 September 2002, P. M. Peterson, J. Valdés-Reyna & M. Sosa Morales 16760 ( ANSM, CAN, MO, US) ; Sierra La Lagunita, 11.8 mi SE of Aramberri on road towards Agua Fría, 24.065°N, 99.737°W, 1817 m, 19 September 2002, P. M. Peterson, J. Valdés-Reyna & M. Sosa Morales 16703 ( ANSM, CAN, US) ; Sierra La Lagunita, 9.5 mi SE of Aramberri on road towards Agua Fría, 24.0605°N, 99.7599°W, 1450 m, 19 September 2002, P. M. Peterson, J. Valdés-Reyna & M. Sosa Morales 16699 ( ANSM, CAN, US) ; Sierra Las Cautivas, 1.0 mi Wof Dulces Nombres, 24.013°N, 99.5746°W, 1983 m, 26 September 2007, P. M. Peterson & J. M. Saarela 21083 ( CAN, US) ; Sierra Las Cautivas, 10.6 mi Eof Zaragoza on road towards Dulces Nombres, 23.992°N, 99.6846°W, 2270 m, 26 September 2007, P. M. Peterson & J. M. Saarela 21112 ( CAN, US) ; Sierra Las Cautivas, 12.0 mi Eof Zaragoza on road towards Dulces Nombres, 24.0001°N, 99.6747°W, 2088 m, 26 September 2007, P. M. Peterson & J. M. Saarela 21103 ( CAN, MO, US) ; Sierra Las Cautivas, 31.6 mi Wof Ejido Guayabas and 5.4 mi Wof Dulce Nombres, 23.9775°N, 99.5561°W, 1833 m, 25 September 2007, P. M. Peterson & J. M. Saarela 21073 ( CAN, US) ; Sierra Las Cautivas, 7.7 mi Eof Zaragoza on road towards Dulces Nombres, 23.9784°N, 99.7005°W, 2578 m, 27 September 2007, P. M. Peterson & J. M. Saarela 21114 ( CAN, MO, P- 03631105 , US) ; Sierra Las Cautivas, 8.7 mi SW of Dulces Nombres and 18.1 mi Eof Zaragoza, 23.9501°N, 99.6473°W, 2385 m, 26 September 2007, P. M. Peterson & J. M. Saarela 21100 ( CAN, US) ; Sierra Madre Mts., Monterrey, 25.52°N, 100.21°W, 15 July 1933, C. H. Mueller & M. T. Mueller 425 ( GH, US) ; Sierra Madre Mts., Monterrey, 25.52°N, 100.21°W, 23 July 1933, C. H. Mueller & M. T. Mueller 428 ( GH) ; Sierra Madre Oriental, 16 km NE of Sandia on road to La Ascension, 24.3071°N, 99.9921°W, 2000 m, 19 September 2001, P. M. Peterson & J. Valdés-Reyna 15824 ( CAN, US) ; Sierra Madre Oriental, ascent to Mesa de la Camisa, ca. 15 mi SW of Galeana, [24.61°N, 100.04°W], 22 July 1934, C. H. Mueller & M. T. Mueller 1168 (F-938865, GH, MICH-1119159 ) ; Sierra Madre Oriental, beyond Pablillo toward Santa Clara, ca. 15 m SW of Galeana, [24.62°N, 99.74°W], 18 July 1934, C. H. Mueller & M. T. Mueller 1078 ( GH) ; Sierra Madre Oriental, Dulces Nombres, Nuevo León, and just Eof border into Tamaulipas, 25.1583°N, 98.81°W, 1700 m, 14 July 1948, F. G. Meyer & D. J. Rogers 2778 ( US- 1962989 ) ; Sierra Madre, near Monterrey, 25.52°N, 100.21°W, 30 June 1888, C. G. Pringle 2052 ( US-1009570 ) ; W slopes of the Cerro, 23.8267°N, 99.8667°W, 2900 m, 22 September 2002, P. M. Peterson, J. Valdés-Reyna & M. Sosa Morales 16780 ( CAN, US) ; Wslopes of the Cerro, 23.8267°N, 99.87°W, 2900 m, 22 September 2002, P. M. Peterson, J. Valdés-Reyna & M. Sosa Morales 16778 ( CAN, US) ; ca. 11 mi Sof San Antonio in pine forest, [24.21°N, 99.63°W], 6 October 1976, F. W. Gould, K. W. Allred & C. Clark 14973 ( MEXU-842648 ) ; Sierra Madre Oriental, 2 mi Sof Los Mimbres and 2.6 mi Nof Ejido La Lagunita, 24.9509°N, 100.26°W, 2470 m, 10 September 2005, P. M. Peterson & J. Valdés-Reyna 18959 ( ANSM, CAN, US) ; Mpio. San Pedro Garza Garcia, Parque Ecológico Chipinque , vereda El Empalme, 25.6064°N, 100.96°W, 1266–1291 m, 6 October 2005, J. A. Encina 1539, F. J. Encina & I. Ramírez ( ANSM) ; Sierra Las Cautivas, 7.6 mi SW of Dulces Nombres and 19.2 mi Eof Zaragoza, 23.9459°N, 99.64°W, 2450 m, 26 September 2007, P. M. Peterson & J. M. Saarela 21098 ( CAN, US) ; Mpio. de Galeana, Arroyo Hondo, Hacienda San José de Raíces, 31 July 1935, C. H. Mueller 2290 (F-886780, GH, MICH-1119283 , NY, P-02630113 , US-1645342 ) ; Cerro Potosí, at NE summit of mountain, ca. 3650 m, 13 September 1960, J. H. Beaman 4467 ( GH, MSC-171507 , US-2381745 ) ; Mpio. Santa Catarina, edge of Parque Nacional Cumbres de Monterrey, slopes above Puerto del Conejo, 25.4931°N, 100.5839°W, 2650 m, 6 September 2010, P. M. Peterson , J. M. Saarela , K. Romaschenko , J. Valdés-Reyna & I. Cabral C. 23139 ( US) ; 7.7 mi Eof Hwy. 57 on MEX 31 towards Linares, 24.6828°N, 100.1725°W, 2850 m, 8 September 2010, P. M. Peterson, J. M. Saarela & K. Romaschenko 23199 ( US) ; Sierra Madre Oriental, 3 mi Wof Dieciocho de Marzo up road towards Cero Potosi, 2200 m, 18 October 1995, P. M. Peterson & M. B. Knowles 13326 ( US) . Oaxaca: 13.5 km Wof San Juan Mixtepec and 1.6 km Eof San Martín, [16.26°N, 96.44°W], 2480 m, 03 September 1990, P. M. Peterson, A. Campos-Villanueva 09766 ( US) ; 15 kms de San Miguel Peras, [16.94°N, 97.02°W], 21 July 1979, P. Guerrero 86 ( MEXU-1099493 ) ; después de Río Oro 2 km delante de Tamazulapa rumbo a México, 1850 m, 21 June 1980, A. A. Beetle M-4584 ( MEXU-1098040 ) ; Mpio. Tepelmeme al Nde El Rodeo ladera Wde Cerro Verde (Jaderoaria), 18.07°N, 97.31°W, 11 September 1990, J. Sánchez-Ken, P. Tenorio L. & A. Salinas T. 172 ( RSA-POM-544635 , F- 2128583) ; Sierra de San Felipe, 17.1°N, 96.85°W, 10,000 ft, 1894, C. L. Smith 942 (F-1524351, US-1009559 ) ; Sierra de San Felipe, 17.1°N, 96.85°W, 9500 ft, 18 September 1894, C. G. Pringle 4898 (CAN-159611, GH, MO, MSC-4592 , NY, P-02630110 , US, 3 sheets) . Puebla: 23.4 Eof Amecameca on road towards Puebla in Parque Nacional Izta-Popo, 19.0897°N, 98.6064°W, 3330 m, 13 October 2001, P. M. Peterson & Y. Herrera-Arrieta 16169 ( CAN, US) ; 3.2 mi Wof Xalitzintla and 27 mi Eof Amecameca, 19.0795°N, 98.5629°W, 2950 m, 13 October

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Phytotaxa 185 (1) © 2014 Magnolia Press 21 2001, P. M . Peterson & Y . Herrera-Arrieta 16176 ( CAN, MO, US); 8 km de Tehuacán rumbo a Orizaba , 18.46°N, 97.39°W, 2200 m, 14 October 1985, I . Núñez et al. 110 ( MEXU-1110773 ); 9 mi Eof paso de Cortés , 19.0801°N, 98.5592°W, 2904 m, 12 October 2007, P.M. Peterson, J. M . Saarela & M. J . Flores Villegas 21401 ( MO, US); along Tehuacán-Orizaba highway just above Azumbilla , [18.68°N, 97.47°W], 1500–1800 m, 18 July 1961, C. E . Smith, Jr., F. A . Peterson & N . Tejeda 3962 (F-1555797); ca. 16 mi Wof Texmelucan near the continental divide, 19.32°N, 98.5333°W, 2980 m, 31 August 1953, J. R . Reeder & C. G . Reeder 2206 ( RSA-POM-287293 , US-2473581 ); vicinity of San Luis Tulitlanapa, Cerro de Baxtle [Paxtle], 18.15°N, 97.4°W, July 1907, C. A . Purpus 2898 (F- 243754, GH, MO, NY, US-1009561 , US-840693 ); Chalchicomula , 18.9833°N, 97.45°W, 2700 m, 19 August 1910, A. S . Hitchcock 6277 ( US-1009560 ); Chalchicomula , 18.9833°N, 97.45°W, 2700 m, 19 August 1910, A. S . Hitchcock 88 2 ( GH, 2 sheets, F-715979, MO, NY); Esperanza 18.8667°N, 97.4°W, 2591 m, 28 August 1910, A. S . Hitchcock 6484 ( US-1009563 ); Esperanza , 18.8667°N, 97.4°W, 2660 m, 17 August 1905, H . Pittier 424 ( US- 570819 ); Hacienda Guadalupe, barranca de la Alseseca , vicinity of Puebla, 18.8333°N, 97.7333°W, 2120 m, 26 September 1907, G . Arséne 1505 ( MO, NY, US-1002657 ); Hacienda Santa Bárbara , vicinity of Puebla, 18.8333°N, 97.7333°W, 2150 m, 2 June 1907, G . Arséne 7193 ( US-1000495 ); Iztaccihuatl, Eside of mtn. above San Augustín , Wof Texmelucan, 19.17°N, 98.64°W, ca. 3570 m, 12 September 1958, J.H. Beaman 2540 ( GH, MSC-162395 ); Malinche , [19.074°N, 98.11°W], 3000 m, 9 November 1965, W . Boege 14 ( GH, MEXU-9432 ); Mpio. Cholula , 30 km al Ede Cholula, [17.03°N, 96.78°W], 3000 m, 30 June 1989, P . Dávila 396 ( MEXU-645154 ); Mpio. Nopalucan, 3 km antes de Huamantla por la carretera Amozoc Libres , 19.05°N, 98.22°W, 2420 m, 2 August 1981, L . Román M . 39 ( MEXU-1098037 ) & 45 ( MEXU-1098038 ); Mpio. Oriental, carretera Oriental-San Antonio Virreyes , faldas del cerro denominado "Tres Cerros", 19.4°N, 97.6347°W, 2350 m, 22 September 1993, Sergio Aguirre y C. S-129 ( MEXU); Mpio. S . Nicolás de los Ranchos, Buenavista , a 5 km al Ede Xalítzintla, 19.1°N, 98.55°W, 3300 m, 15 February 1988. P . Tenorio L . 15100 ( MEXU-548960 , US-3183529 ); Mpio. San Miguel Canoa, Xaxalpa , a 7 km al NE de San Miguel Canoa, 19.15°N, 98.22°W, 16 September 1988, P . Tenorio L . 15144 ( MEXU-551278 , RSA-POM-513693 ); Mpio. San Nicolás de los Ranchos, Paso de Cortés, 2 km Sof the rd to Amecameca, México, sobre la brecha Volcán Ixtaccihuatl , 19.12°N, 98.63°W, 3710 m, 1 November 1976, S. D. Koch 76246 ( MO, US-2824587 ); Mpio. Santiago Mihuatlán, 3 km de la desviación a Nicolás Bravo , 2060 m, 6 August 1981, S . Morales 64 ( MEXU-1110788 ); Mpio. Tetla, Ejido José María Morelos, 19.06°N, 98.2°W, 2600 m, 30 October 1985, A . Miranda 193 ( MEXU-1110783 ); Mt. Orizaba , 19.0167°N, 97.2667°W, 2438 m, 17–18 August 1910, A. S . Hitchcock 6264 ( US-1009566 ); Rancho Posadas , vicinity of Puebla, 19.0333°N, 98.1833°W, 2180 m, 6 August 1908, G . Arséne 1603 ( MO, US-1002637 , US-2306185 ) & 2288 ( US-1002632 , US-3466183 ); San Marcos , 18.4°N, 97.35°W, 2591 m, 29 August 1910, A. S . Hitchcock 6516 ( US-1009562 ); Sierra Negra, above Serdán , Cabecero , 19.05°N, 97.39°W, 2850 m, 25 September 1944, A. J . Sharp 441027 ( US-1939454 ); vicinity of Puebla, Rancho Posadas , 19.04°N, 98.21°W, 2180 m, 6 August 1908, G . Arséne 1603 ( GH) & 936 8 ( GH); Rancho Posadas, vicinity of Puebla, 19.0333°N, 98.1833°W, 67 m, 10 April 1909, Br. Nicolás & G . Arséne 270 ( NY, mixed sheet with B. carinatus var. marginatus ). Querétaro: Hwy. between Jalpan and Vizzarrón , at pass call Pinal de Amoles , 38 km SW of Jalpan, [21.14°N, 99.63°W], 8 October 1985, R . Spellenberg, J . Zimmerman, N . Zucker & G . Cunningham 8319 ( MEXU-560704 ); Mpio. Landa de Matamoros, Lobo , [20.7346°N, 100.202°W], 24 August 1982, R . Guzmán 5962 ( MEXU-1097975 ); Jardín Botánico Regional de Cadereyta "Ing. Manuel González de Cosío", [20.5875°N, 100.3928°W], 2150 m, 8 May 1992, R . Hernández M., J . Orozco H. & C . Orozco L . 9826 ( MEXU-631628 ); Mpio. Cadereyta, 4 km al sur de Vizarrón, carretera Cadereyta-Jalpan , 20.85°N, 99.7167°W], 2180 m, 17 September 1993, Jaramillo, Villegas & Miranda 788 ( MEXU-1110794 ); Mpio. Pinal de Amoles , Alrededores del poblado Pinal de Amoles , [21.135°N, 99.63°W], 2300 m, 29 September 1994, N . Rincón & L. G . Mora 204 ( MEXU); Mpio. Pinal de Amoles, Caseta Forestal Pinal de Amoles , 21.1333°N, 99.6267°W, 26 November 1981, A . Mora Benítez & Fco. J. Ramírez Rodriguez 468- AMB ( MEXU); just below Cerro El Zamorano , 20.9328°N, 100.1839°W, 3191–3250 m, 25 September 2012, P. M . Peterson, K . Romaschenko & S . Zamudio Ruíz 24677 ( US); near Cerro El Zamorano , 20.9331°N, 100.18°W, 3250–3373 m, 25 September 2012, P. M . Peterson, K . Romaschenko & S . Zamudio Ruíz 24694 ( US); 1.5 mi Nof Los Trigos, 20.9192°N, 100.1967°W, 2795 m, 25 September 2012, P. M . Peterson, K . Romaschenko & S . Zamudio Ruíz 24708 ( US); Sierra del Doctor, Cerro El Espolon , 20.7908°N, 99.565°W, 3140–3250 m, 26 September 2012, P. M . Peterson, K . Romaschenko & S . Zamudio Ruíz 24756 ( US); Sierra del Doctor , at 2 mi Wof El Doctor, 20.8583°N, 99.5992°W, 2867 m, 27 September 2012, P. M . Peterson, K . Romaschenko & S . Zamudio Ruíz 24769 ( US); just below Cerro El Zamorano , 20.9328°N, 100.1839°W, 3191–3250 m, 25 September 2012, P. M . Peterson, K . Romaschenko & S . Zamudio Ruíz 24678- B ( US) . San Luis Potosí : 1876, Schnaffner 1076 ( MEXU); 1876, J. G . Schaffner 129/1075 ( GH, NY, US-

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397109, US-821125 ); Álvarez, [22.0333°N, 100.6167°W], 13–23 July 1904, E. Palmer 171 ( GH); in mountains ca. 19 mi SW of San Luis Potosí on road to Aguascalientes, 21.94°N, 100.8°W, 19 July 1950 , J. R. Reeder , C. G. Reeder & L.N. Goodding 1398 ( US-2473579 ); between Puerta Huerta and Río Verde in the Sierra de Álvarez , 21.91°N, 100.1°W, 1150–2200 m, 11 September 1954 , E.R. Sohns 1196 ( US-2154452 ); Canyon del Lobo , Sierra de San Miguelito, 22.133°N, 100.983°W, 1900–2350 m, 10 September 1954 , E.R. Sohns 1163 ( US-2154425 ); Canyon del Muerto , ca. 3 km Wof Rodrigo, in the Sierra de San Miguelito, 21.8833°N, 100.98°W, 1800–2200 m, 18 September 1954 , E.R. Sohns 1326 ( US-2154564 ); Charcas , 23.1°N, 101.0333°W, July–August 1934 , A.F. Whiting 887 ( MICH-1119291 , US) & 980 ( MICH-1119128 , US); Charcas , Aguila Mt., 23.3833°N, 101.5333°W, July–August 1934 , A.F. Whiting 831 ( MICH-1119156 , US-1646157 ); exposición norte del Cerro del Lobo Villa de Arriaga , [21.91°N, 101.38°W], 2250 m, 16 November 1994 , J. L. Villalpando P. 204 ( MEXU); in mountains ca. 15 mi SW of San Luis Potosí on road to Aguascalientes, 22°N, 100.82°W, 18 July 1950 , J. R. Reeder , C. G. Reeder & L.N. Goodding 1380 ( US-2473578 ); near Puerta Huerta in the Sierra de Álvarez, 21.91°N, 100.1°W, 2200–2400 m, 4 September 1954 , E.R. Sohns 1032 ( US-2154311 ) & 1040 ( US-2154374 ); near the Minas de San Rafael in the Sierra de Guadalcázar, 22.2°N, 100.2833°W, 1900–2100 m, 3 October 1954 , E.R. Sohns 1512 ( US-2154705 ); near the village of San Francisco in the Sierra de San Miguelito ca. 25 km SW of San Luis Potosí, 22°N, 101.14°W, 2200–2400 m, 5 September 1954 , E.R. Sohns 1067 ( US-2154344 ) & 1069 ( US-2154346 ); Puerto Huerta , camino San Luis–Río Verde, 21.91°N, 100.1°W, 2200 m, 29 July 1954 , J. Rzedowski 161 ( US-2566609 ); Mpio. Guadalcázar , Ejido Minas de Plata, [22.62°N, 100.43°W], 1500 m, 18 November 1981 , H. Bravo 37 ( MEXU); Mpio. Villa de Zaragoza , Ejido San Francisco, 22.12°N, 100.74°W, 1800 m, 12 October 1981 , A. Bolaños 183 ( MEXU); San Luis Potosí , [22.16°N, 100.99°W], Schaffner 10361 ( MEXU-5768 ); 5.7 mi Eof Wadley, Sierra de Catorce, 23.59°N, 100.9031°W, 2131 m, 10 September 2010 , P. M. Peterson , J. M. Saarela & K. Romaschenko 23278 (US); Sierra del Jórdan, Sierra de Catorce , 23.5358°N, 100.8467°W, 2013 m, 11 September 2010 , P. M. Peterson , J. M. Saarela & K. Romaschenko 23312 ( US); Eof Wadley, Sierra de Catorce, 23.5664°N, 100.8661°W, 2300–2688 m, 11 September 2010 , P. M. Peterson , J. M. Saarela & K. Romaschenko 23307 ( US); Eof Wadley, Sierra de Catorce, 23.5722°N, 100.8842°W, 2300–2688 m, 11 September 2010 , P. M. Peterson , J. M. Saarela & K. Romaschenko 23294 ( US); 6.3 mi SE of Villa Zaragoza on road towards La Salitrera, 21.99°N, 100.6694°W, 1793 m, 14 September 2010 , P. M. Peterson , J. M. Saarela , K. Romaschenko , J. Herrera-Simoni & M. Herrera-Simoni 23368 ( US); Cerro de la Luz, Wof La Trinidad, 21.4078°N, 99.1022°W, 2795 m, 23 September 2010 , P. M. Peterson , J. M. Saarela & K. Romaschenko 23586 ( US); Cerro de la Luz, Wof La Trinidad, 21.4078°N, 99.1022°W, 3250–3373 m, 23 September 2010 , P. M. Peterson , J. M. Saarela & K. Romaschenko 23585 ( US) & 23586 ( US); Cerro El Pinon Blanco, 22.5206°N, 101.68°W, 2291–2744 m, 2 October 2012 , P. M. Peterson & K. Romaschenko 24881 ( US). Tamaulipas: 11 mi Sof Palmillas on the road to Tula, 23.2°N, 99.64°W, 18 September 1960 , J. Crutchfield & M. C. Johnston 5633 B ( MICH-1119150 , US-2462956 ); 14.6 mi NE of Dulces Nombres , 23.9983°N, 99.47°W, 1748 m, 23 September 2001 , P. M. Peterson & J. Valdés-Reyna 15918 ( CAN, US); carretera de Ciudad Victoria a Jaumave, 23.6283°N, 99.2044°W, 1060 m, 12 October 2007 , F.O. Zuloaga et al. 9706 ( ANSM); Filo y lado este de la Sierra Peña Nevada, [23.78°N, 99.66°W], 3500–3600 m, 5 July 1985, McDonald 1673 ( MEXU-1088981 ); Jaumave road ca. 13 mi SW of Ciudad Victoria , near summits of Sierra Madre, [23.58°N, 99.23°W], 1000 m, 13 May 1949 , R. McVaugh 10533 ( MICH-1119125 , NY); Mpio. Bustamante, El Capulín camino de terracería, 23.2167°N, 99.6833°W, 1610 m, 23 May 1994 , R. López 221 ( MEXU); Mpio. Miquihuana, Ej. El Aserradero, [23.56°N, 99.78°W], 2650 m, 3 September 1981 , R. Carranco 84 ( MEXU); Mpio. Tula. Cañon de la Lagartija, [22.99°N, 99.66°W], 1400 m, 21 October 1982 , G. Villegas 542 ( MEXU-1089016 ); Mpio. Victoria , Sierra Madre rumbo a Jaumave, [23.42°N, 99.39°W], 1200 m, 8 September 1981 , R. Carranco 91 ( MEXU- 1089013 ); Mpio. Victoria , Victoria-Atlas Cumbres, [23.73°N, 99.14°W], 1200 m, 21 July 1993 , J. Franco 142 (MEXU); on Eand Sslopes and summit of Peña Nevada, 23.7°N, 99.8°W, 19 July 1949 , L. R. Stanford , S. M. Lauber & L.A. Taylor 2575 ( RSA-POM-72561 , GH, MO, NY, US-2013061 ); Sierra Las Cautivas, 19.6 mi Wof Ejido Guayabas along road to Dulces Nombres , 23.9978°N, 99.47°W, 1755 m, 25 September 2007 , P. M. Peterson & J.M. Saarela 21057 ( CAN, US) & 21059 ( CAN, MO, US); 3 mi Nof Miquihuana, [23.57°N, 99.76°W], 15 July 1949 , L. R. Stanford , S. M. Lauber & L.A. Taylor 2480 ( GH, MO, RSA-POM-74021 ). Tlaxcala: 15 km Nof Apizaco , 19.5417°N, 98.15°W, 23 September 1953 , E.R. Sohns 617 ( US-2118588 ); between San Cristóbal and Calpulalpan , 19.5833°N, 98.61°W, 2134–2743 m, 22 September 1953 , E.R. Sohns 564 ( MEXU-171735 , P- 02630112 , US-2118545 ); ca. 3 mi NE of Tlaxco , 19.65°N, 98.08°W, 2743–3048 m, 23 September 1953 , E.R. Sohns 595 ( US-2118570 ); Cerro Zotoluca , al NE de Calpulalpan, 19.57°N, 98.56°W, 2600 m, 20 October 1985 , A. Miranda 186 ( MEXU-1089003 ); hills at Contadero Station , 19.4333°N, 98.5167°W, 2591 m, 30 August 1901 , C.G.

ATAXONOMIC REVISION OF BROMUS (POACEAE)

Phytotaxa 185 (1) © 2014 Magnolia Press 23

Pringle 8597 (CAN-159558, GH, MEXU, MO, MSC-4687, NY, RSA-POM-206109, US-396387, US-821087, US- 967967, US-1009565); Mpio. Calpulalpan, Aserradero, 19.59°N, 98.57°W, 6 September 1985, 2300 m, J . González Estevez s.n. ( MEXU-946222 ); near Huamantla, Eslope of La Malinche, 19.3°N, 98°W, 24 September 1953, E.R. Sohns 641 ( US-2118607 ); summit of Mt. Tlacuapanga , 19.2833°N, 98.15°W, 2774 m, 21 August 1944, A. J . Sharp & E . Hernández-Xolocotzi 44488 ( US-1939427 ). Veracruz: Mpio. Perote, 1.75 km al sur de el Conejo, [19.56°N, 97.25°W], 3570 m, 10 September 1992, B. V . Hernández 56 ( MEXU-1089766 ); Mpio. Perote, 5.5 km al ESE de Perote , 19.5372°N, 97.2°W, 2990 m, 10 September 1992, H. R . Sandoval 121 ( MEXU); Tlaxcala , ca. 14 mi SW of Mendoza, 18.72°N, 97.31°W, 2040 m, 17 August 1953, J. R . Reeder & C. G . Reeder 2011 ( RSA-POM-287185 , US). Zacatecas: Sierra Madre Occidental, 13 mi SW of Valparaíso and 13.8 mi NE of Huejuquilla , 22.6924°N, 103.71°W, 2180 m, 30 September 2001, P. M . Peterson & M. S . González-Elizondo 15968 ( NY, RSA-POM-732008 , US); Sierra Madre Occidental, 3.5 mi Wof Monte Escobedo at Uma Temoc A. C , 22.3152°N, 103.6°W, 2450 m, 3 October 2001, P. M . Peterson & M. S . González-Elizondo 16051 ( MO, RSA-POM-732040 , US); Sierra Madre Occidental, 8.6 mi NW of Monte Escobedo at Uma Temoc A. C , 22.3666°N, 103.61°W, 2480 m, 3 October 2001, P. M . Peterson & M. S . González-Elizondo 16044 ( MO, NY, RSA-POM-732042 , US); Cuajimalpa, La Venta, 19.3558°N, 99.3011°W, 2600 m, 19 September 1930, M . St. Pierre 916 ( MICH-1119127 , US-1537812 ); Mpio. Guadalupe, ladera norte del cerro de la Vírgen, 2490 m, August 1988, J . Balleza C . 1573 ( ANSM); Mpio. Miguel Auza , 2.5 km al Wde la Col. Emilio Carranza, por la brecha a la Sierra de Santa María , [22.1833°N, 101.8333°W], 2200 m, 2 September 1988, J.J. Balleza C . 1889 ( ANSM); Pedregal de San Angel , cerca de Coyoacán, 19.3°N, 99.18°W, 2 August 1952, J . Rzedowski 1391 ( US-2079374 ); San Angel, Briqueteri, 19.3733°N, 99.225°W, 2350 m, 12 September 1929, M . St. Pierre 836 ( US-1537811 ); Sierra de los Morones, near Plateado, 21.6667°N, 103.2°W, 1 September 1897, J. N . Rose 2727 ( US-301655 , US-821068 ); Sierra Los Cardos, Rancho El Robles, 17.8 mi NW of Jerez on road towards Monte de las Garcia , 22.6898°N, 103.2°W, 2820 m, 20 October 2007, P. M . Peterson, J. M . Saarela & D. Stancik 21436 ( CAN, MO, US); 9.5 mi Wof Sombrerete , [23.63°N, 103.65°W], 7900 ft, 18 August 1969, J . Taylor & C . Taylor 6256 ( NY); 4 km Eof Salaverna on the highway to Concepción del Oro , 24.1028°N, 100.9139°W, 2800 m, 1 July 1973, M. C . Johnston, T. L . Wendt & F . Chiang C . 11547 (NY).

Figures

FIGURE 1. Bromus anomalus. A. Habit. B. Inflorescence. C. Sheath and auricles. D. Spikelet. E. Glumes. Illustration by C.T. Roché, reproduced from Barkworth et al. (2007) with permission.

FIGURE 2. Bromus anomalus. Harvey 8501 (MICH-1119120).

FIGURE 3. Bromus anomalus inflorescence, Nuevo León, México (Peterson et al. 23139). Photo: J.M. Saarela.

FIGURE 4. Geographical distribution of Bromus anomalus in México and Central America.