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 : 88-91

11. Bromus frondosus (Shear) Wooton & Standley (1912: 144) . Figs. 38 , 39.

Basionym: Bromus porteri var. frondosus Shear (1900: 37) . Bromopsis frondosa (Shear) Holub (1973: 167) . Type:— UNITED STATES OF AMERICA. New México: Grant Co., Mangas Springs, 19 August 1897, J.G. Smith s.n. (holotype US- 81587!).

Plants perennial, not rhizomatous. Culms 54–84 cm tall, 2–4 mm wide at base, erect to spreading, glabrous below inflorescences, nodes 3–5, glabrous, occassionally with a few short hairs. Leaf sheaths sparsely to densely pilose, hairs up to 1.2 mm long; auricles absent; ligules 0.5–2 mm long, glabrous; blades up to 29 cm × 4–6 mm, flat, adaxial and abaxial surfaces usually glabrous, sometimes pubescent, hairs up to 1 mm long distally, margins serrulate. Panicles 9–13 cm × 3–6 cm, open, nodding, branches ascending to spreading, 0.5–5 cm long, shorter or longer than spikelets, scabrous, 1–4 spikelets per branch. Spikelets 1.6–3 cm long, 7–9-flowered, elliptic to lanceolate, terete to moderately laterally compressed; glumes glabrous, scabrous or minutely pubescent, hairs when present sometimes restricted to margins, margins hyaline, midnerves glabrous or scabrous; lower glumes 6–8.5 mm × 0.6–1.0 mm, lanceolate, 3-nerved, green along and between the nerves, apices acute; upper glumes 8–10 mm × 0.9–1.3 mm, oblong-ovate, 3-nerved, green along and between the nerves, apices acute or mucronate, mucros to 1 mm long; lemmas 9–12 mm × 1.5–2.5 mm, elliptic to lanceolate, rounded over the backs, apices truncate, 5–7- nerved, green along and between the nerves, backs glabrous or sparsely to densely pubescent, margins pubescent on lower third to half, hairs up to 1 mm long; awns 3.5–6 mm long, arising 0–0.5 mm below lemma apex, straight;

88 Phytotaxa 185 (1) © 2014 Magnolia Press

SAARELA ET AL.

paleas shorter than the lemmas, 9–10 mm long, backs glabrous or pubescent, keels ciliate, cilia up to 0.2 mm long; anthers 1.5–3.5 mm long; caryopses 6−8 mm long. 2 n = 14 ( Wagnon 1952).

Distribution: ―Native. In México B. frondosus is known from Chihuahua, Coahuila, Durango, Sinaloa and Sonora ( Fig. 40 ). It extends northwards to Colorado, New México and Arizona ( Pavlick & Anderton 2007).

Ecology: — Bromus frondosus is found on rocky slopes and rock outcrops, roadcuts and rocky meadows; associated with Pinus cembroides , Pinus spp. , Quercus , spp., Juniperus deppeana , Abies religiosa , Pseudotsuga menziesii , Cupressus arizonica , Rhamnus , Ceanothus , Arbutus , Arctostaphylos pungens , Cercocarpus Kunth in Humboldt (1824: 6), Salvia , Platanus wrightii Watson (1875: 349) , Senecio and Garrya flavescens Watson (1873: 301) . Elevation: 1295–2745 m.

Common Names:— Weeping brome (English); bromo frondoso (Spanish).

Comments: ― Bromus frondosus is distinguished by its 3-nerved lower glumes.

Specimens Examined:― MÉXICO. Chihuahua: Mpio. Madera, Laguna de Babícora , Arroyo Las Varas, [29.0556°N, 106.8611°W], 2300 m, 10 September 1994, G . Quintana & E. Estrada 3556 ( NY); "Rancho el peñasco" km 150 carretera Chihuahua-Cd. Juárez , [29.77°N, 107.37°W], 1720 m, 9 August 1979, M . Siqueiros 336 ( MEXU-1098029 ); 20.3 km NE of Ignacio Zaragoza on MEX 23 , [29.75°N, 107.64°W], 2400 m, 28 September 1989, P. M . Peterson & R. M . King 8154 ( CAN, US); 24.7 mi Nof San Juanito on road towards Cuauhtémoc , 28.2636°N, 107.4931°W, 2233 m, 5 September 2008, P. M . Peterson & J. M . Saarela 22028 ( CAN, US); 30.2 km N of San Juanito on road to Creel , [28.26°N, 107.61°W], 2235 m, 10 September 1989, P. M . Peterson, C. R . Annable & Y . Herrera-Arrieta 8002 ( CAN, MO, US); 34 mi Wof Balleza on road towards Guachochic , [27.12°N, 106.79°W], 2500 m, 29 October 1995, P. M . Peterson, M. B . Knowles, C. H . Dietrich & S. M . Braxton 13539 ( US); 8 mi Nof Santo Tomás , 28.81°N, 107.5667°W, 2010 m, 8 October 1953, J. R . Reeder & C. G . Reeder 2625 ( US- 2473583 ); ca. 23 (air) mi ENE of Villa Ahumada in NW canyon of Sierra de la Alcaparra NE of Rancho El Palmar , 30.6667°N, 106.1°W, 6000 ft [1828 m], 12 September 1973, J . Henrickson 12864 ( MEXU-1052580 , RSA-POM- 665758 ); Colonia Cumbres de Majalca , 33.8 km Wof MEX 45 , Nof Chihuahua City , [28.8°N, 106.46°W], 2190

ATAXONOMIC REVISION OF BROMUS (POACEAE)

Phytotaxa 185 (1) © 2014 Magnolia Press 89 m, 7 September 1989, P. M . Peterson, C. R . Annable & Y . Herrera-Arrieta 7969 ( CAN, US); km 11 entre San Bueneventura y El Carmen, carratera Nuevo Casas Grandes , 29.84°N, 107.47°W, 1600 m, 19 September 1955, E . Hernández X. & V . Mathus L . N-1941 ( GH); km 18 de San José Babícora a Santa Ana Babícora, 2200 m, 30 October 1954, E . Hernández X. & C . Tapia J . N-438 ( GH); Majalca (Pilares), [28.68°N, 106.11°W], 12 September 1939, L. H . Harvey 1485 ( GH, mixed sheet with B. carinatus var. carinatus ); Majalca , [28.73°N, 106.1°W], 18–20 September 1935, H . LeSueur 30 ( GH); Majalca , NW of Chihuahua, [28.73°N, 106.1°W], 2050–2100 m, 29 September 1934, F. W . Pennell 19299 ( GH); near Colonia Garcia , [29.97°N, 108.35°W], 7500 ft [2286 m], 14 September 1899, C. H. T . Townsend & C. M . Barber ( MSC-229571 ); Río Negro , 28 August 1937, H . LeSueur 0213 ( GH); San Ysidro, SW of Barranca, [28.5667°N, 107.45°W], 13 May 1929, Y . Mexia 2524 ( MICH-1119284 , NY, RSA-POM-364164 ); Sánchez , [31.77°N, 107.64°W], 8000 ft [2438 m], 12 October 1910, A. S . Hitchcock s.n. ( MEXU); Santa Clara Canyon , [29.72°N, 107.08°W], 18 August 1936, H . LeSueur 0123 ( GH); Sierra de los Órganos, 28.25°N, 104.75°W, 9 August 1937, H . LeSueur 211 ( GH, mixed sheet with B. carinatus var. marginatus ), US-1721654 ); Sierra Madre Mts. , Colonia Juárez, [30.3°N, 108.06°W], 6000 ft [1829 m], 12 September 1903, M. E . Jones s.n. ( RSA-POM-112405 ); Sierra Madre Mts. , Meadow Valley, [30.3°N, 108.06°W], 7000 ft [2134 m], 17 September 1903, M. E . Jones s.n. ( RSA-POM-112403 ); Sierra Madre Occidental, 12.1 mi W of San Juanito on road towards Baquiriachic , 27.9567°N, 107.76°W, 2530 m, 5 October 2000, P. M . Peterson & J . Cayouette 15358 ( CAN, US); Sierra Madre Occidental , 16.7 mi Wof Baquiriachic on Hwy. 16 towards Maycoba, 28.3543°N, 108.28°W, 2000 m, 4 October 2000, P. M . Peterson & J . Cayouette 15353 ( CAN, MO, US); 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 15368 ( CAN, US); Sierra Madres near Colonia Garcia , [29.97°N, 108.34°W], 7500 ft [2286 m], 14 September 1899, C. H. T . Townsend & C. M . Barber 327 (F-103052, MEXU-5764 , MICH- 1119280 , MSC-229571 , NY, 2 sheets); SW slope of Sierra de la Ranchería (on Rancho Candelaria ), 31.0167°N, 106.35°W, 1500–2180 m, 29 October 1972, T. L . Wendt, F . Chiang & M. C . Johnston 9949 ( MEXU-729813 , NY); 19.6 km Wof Balleza and 74.2 km Eof Guachochi, [26.9453°N, 106.4783°W], 2120 m, 18 September 1991, P. M . Peterson, C. R . Annable & J . Valdés-Reyna 10753 ( US); along dirt road to Cuesta Blanca in Sierra Brena SW of Colonia Juárez, 2256 m, 21 September 1992, K. W . Allred, T . Columbus & J . Valdés-Reyna 5727 ( ANSM); 19.6 km Wof Balleza and 74.2 km Eof Guachochi, 2120 m, 18 September 1991, P. M . Peterson, C. R . Annable & J . Valdés- Reyna 10753 ( US); Boycoyna, La Junta, Alrededores, 13 September 2003, R . Bye, M . Mendoza, G . Morales, J . Rodríguez & M. Hilerio 32463 ( US-3589121 ); Creel , Chih. 125 Creel km 90, Sitio 125 km 96.2, La Mesa de Yeguachi, 27°47.155’N, 107°38.611’W, 2248 m, 13 September 2003, R . Bye, M . Mendoza, G . Morales, J . Rodríguez & M. Hilerio 32541 ( US-3589119 ); Guachochi , Norogachi, Planicie, 16 September 2003, R . Bye, M . Mendoza, G . Morales, J . Rodríguez & M. Hilerio 32771 ( US-3589146 ). Coahuila: Mpio. Saltillo, area montañosa del Cañón " San Lorenzo", 25.3167°N, 101.95°W, 2000 m, M. G . Villaseñor s.n. ( ANSM); Sierra del Carmen , 28.9867°N, 102.9739°W, 2432 m, 30 August 1997, S . Wood, C . Crider & D. Doan-Crider s.n. ( ANSM); Maderas del Carmen , 2.0 mi from Campo Uno, up the road towards the summit, 29.0062°N, 102.6054°W, 2571 m, 22 September 2007, P. M . Peterson, J. M . Saarela, S . Lara Contreras & J . Reyna Álvarez 21029 ( CAN, MO, US); Sierra El Jardín, 29.0832°N, 102.6371°W, 2100 m, 3 September 2006, P. M . Peterson & S . Lara-Contreras 19941 ( CAN, US). Durango : Sierra Madre Occidental, 11 mi NE of Ojito de Camillones on road towards Papasquiaro, 25.0912°N, 106.136°W, 2580 m, 12 October 2000, P. M . Peterson, J . Cayouette & M. S . González-Elizondo 15436 ( US); Sierra Madre Occidental, Wof Ciudad Durango , 6 mi Wof El Salto, 23.7418°N, 105.452°W, 2745 m, 27 August 1958, J. R . Reeder & C. G . Reeder 3123 ( US-2473577 ); 30 km Wof Durango on Hwy. 40, just Eof Rio Chico crossing, 2200 m, 29 September 1988, P. M . Peterson & C. R . Annable 6007 ( US). Sinaloa: loop of the Río de Bavispe , NE Sonora, Cerro del Capulín , NW of Aribabi, 30.1°N, 109.083°W, 1875 m, 4 September 1939, L. H . Harvey 1693 ( US-1762717 ); Morelos, 30.8167°N, 109.2167°W, 15 September 1947, J . Vera-Santos 1939 ( US- 2079260 ); region of the Río de Bavispe , NE Sonora, El Bilito, NE of El Tigre , 30.61°N, 109.17°W, 1890 m, 12 October 1941, J . Vera-Santos 2128 ( US-1938696 ). Sonora: Arroyo Hondo, 11.5 km Eof El Kipor, 4 km Wof Chihuahua border on Son., 28.4417°N, 108.78°W, 1460 m, 11 September 1996, T. R . Van Devender, A.L. Reina G., G. Ferguson & L. Coyote 96-563 ( ARIZ-332596 , RSA-POM-610275 ); Cerro El Capulín, NW of Aribabi , [29.91°N, 109.99°W], 1875 m, 4 November 1939, L. H . Harvey 1693 ( MICH-1119279 ); El Bilito, NE of El Tigre , [30.59°N, 109.22°W], 6200 ft [1890 m], 12 October 1941, J . Vera Santos 2128 ( MICH-1119213 ) & 2136 ( NY); El Picacho del Pilar , 7400 ft [2255 m], 13 October 1941, J . Vera Santos 2167 ( MICH-1119124 ); El Puerto del Cumarito , Sierra de la Cabellera, [30.92°N, 109.11°W], 5400 ft [1646 m], 6 October 1941, J . Vera Santos 2079

90 Phytotaxa 185 (1) © 2014 Magnolia Press

SAARELA ET AL.

(MICH-1119282, NY); El Rancho de Robles, NE of El Tigre, [30.76°N, 109.18°W], 6000 ft [1829 m], 2–13 September 1941, J. Vera Santos 1939 (ARIZ-124777, MICH-1119278, MEXU-537473, NY, US-1938676); La Matancita, 1 mi Wof El Tigre, [30.76°N, 109.18°W], 4250 ft [1295 m], 30 August 1941, J. Vera Santos 1915 (ARIZ-124955, MICH-1119277, MEXU-537476, NY, US-1938674); Morelos, 15 September 1947, J. Vera Santos 1915 (MICH-119276); Nof Cananea Ejido José María Morelos, [30.98°N, 110.3°W], 25 September 1981, A.A. Beetle M-7869 (ARIZ-234251, MEXU, MO); region of the Río de Bavispe, NE Sonora, El Puerto del Cumarito, Sierra de la Cabellera, 30.0333°N, 109.3333°W, 1646 m, 6 October 1941, J. Vera-Santos 2079 (US-2463582, US- 1938690); Sierra Madre Occidental, 1.7 mi Wof Maycoba on Hwy. 16 towards Yécora, 28.4065°N, 108.68°W, 1720 m, 4 October 2000, P.M. Peterson & J. Cayouette 15351 ( US); Sierra Madre Occidental, 1.8 mi Wof Yécora on Hwy. 16 towards Hermosillo, 28.3583°N, 108.95°W, 1860 m, 3 October 2000, P.M. Peterson & J. Cayouette 15315, 15317 (CAN, MO, US); Sierra Madre Occidental, 12.3 mi Wof Yécora on Hwy. 16 towards Hermosillo, 28.3772°N, 109.05°W, 1810 m, 3 October 2000, P.M. Peterson & J. Cayouette 15305 ( US); Sierra Madre Occidental, 16 mi Wof Maycoba on Hwy. 16 towards Yécora, 28.4002°N, 108.8°W, 1520 m, 4 October 2000, P.M. Peterson & J. Cayouette 15346 ( US).

ATAXONOMIC REVISION OF BROMUS (POACEAE)

Phytotaxa 185 (1) © 2014 Magnolia Press 91

Figures

FIGURE 38. Bromus frondosus. A. Spikelet. B. Inflorescence. C. Glumes. D. Lemma. Illustration by C.T. Roché, reproduced from Barkworth et al. (2007) with permission.

FIGURE 40. Geographical distribution of Bromus frondosus in México.