Rodríguez, 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 : 83-88

10. Bromus exaltatus Bernhardi (1841: 90) . Figs. 35 , 36 .

Bromus subalpinus Ruprecht in Galeotti (1842: 237), nom. nud. Bromus subalpinus Rupr. ex Fournier (1886: 128) , nom. inval., as synonym of B. exaltatus . Bromopsis exaltata (Bernh.) Holub (1973: 167) . Type:— MÉXICO. Anonymous s.n. (lectotype MO-2957788!, designated by Davidse & Pohl 1992: 100, isotypes LE-00000754!, LE-00000755!). Soderstrom & Beaman (1968) neotypified B. exaltatus based on their understanding that original material in the Bernhardi Herbarium (MO) was missing, as reported by Wagnon (1950) and Shear (1901). Their neotype followed a species concept used by Fournier (1886) and later Hitchcock (1913). However, two sheets of original material were later found at MO, one of which Davidse & Pohl (1992) selected as the lectotype of B. exaltatus , superseding the neotypification. One of the isotypes at LE (LE-00000754) is a mixed sheet with B. carinatus var. marginatus .

Plants perennial, not rhizomatous. Culms 70–120 cm tall, (1–) 2–3 mm wide at base, erect or ascending, glabrous or pubescent below inflorescences; nodes 2–4, weakly to densely pubescent. Leaf sheaths moderately to densely pilose, hairs up to 1.2 mm long; auricles absent or present; ligules 0.6–2.5 mm long, glabrous or pubescent, eroselacerate; blades up to 39 cm × 2.5–7 mm, flat, adaxial surfaces weakly to moderately pilose, hairs up to 0.8 mm long, abaxial surfaces glabrous, margins serrulate. Panicles 15–27 cm × 3–9 cm, open, nodding, branches ascending, 0.8–6 cm long, shorter or longer than spikelets, scabrous, 1–4 spikelets per branch. Spikelets 2.1–3.1(–3.4) cm long, 6–9-flowered, linear-elliptic, moderately laterally compressed; glumes glabrous or pubescent, hairs up to 0.8 mm long, margins narrowly hyaline, midnerves glabrous to scabrous distally or pubescent throughout, hairs up to 0.4 mm long; lower glumes (7–) 9–11 mm long, linear-lanceolate, 1- or 3-nerved, lateral nerves sometimes weak, green to purplish-green along and between nerves, apices attenuate to acute; upper glumes (9–) 11–14 mm long, lanceolate, 3-nerved, green to purplish-green along and between nerves, apices acute to mucronate, mucros up to 1.5 mm long; lemmas 12–16 mm long, linear-lanceolate, gradually narrowed to apices, apices acute to obtuse, 5–7-nerved, green to purplish-green along and between nerves, backs glabrous or weakly to

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Phytotaxa 185 (1) © 2014 Magnolia Press 83 moderately pubescent proximally, hairs appressed and up to 0.2(–0.3) mm long, usually glabrous distally, sometimes pubescent, hairs up to 0.2 mm long, margins pubescent with hairs up to 0.6 mm long, nerves scabrous; awns 2.5–6 mm long, straight, inserted up to 0.5 mm below lemma apex, straight; paleas shorter and narrower than the lemmas, backs puberulent, keels ciliate, cilia up to 0.2 mm long; anthers 1.7–2.5 mm long; caryopses 7.5−10 mm long, light brown. 2 n = 14 ( Pohl & Davidse 1971).

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Distribution: ―Native. Central México (Distrito Federal, Guerrero, Hidalgo, Jalisco, Estado de México, Michoácan, Oaxaca, Puebla, Querétaro, San Luis Potosí, Tlaxcala, Veracruz), Costa Rica (San José), Guatemala (Chimaltenango, Huehuetenango, Quezaltenango, Sacatepéquez, San Marcos, Totonicapan) and Panama (Chiriqui) ( Fig. 37 ). Bromus exaltatus is the only native Bromus species known from Panama.

Ecology:— This species occurs on steep rocky slopes, arroyos, wet meadows and roadsides; associated with Pinus spp. , P. hartwegii , P. rudis , Quercus spp. , Abies religiosa , Juniperus , Pseudotsuga menziesii , Cupressus , Arctostaphylos pungens , Lupinus , Arbutus , Alnus , Senecio Linnaeus (1753: 866) , Symphoricarpos , Coreopsis Linnaeus (1753: 907) and Buddleja . Elevation: 3250–3739 m ( Costa Rica), (2550–) 3000–4200 m ( Guatemala), 2400–3688 m ( México), (1700–) 2800–3170 m ( Panama). McVaugh (1983) noted B. exaltatus to be local or rare throughout its range.

Comments: ―Most specimens from Guatamela, Costa Rica and Panama have lemmas that are pubescent across the backs, while specimens from México have lemmas that are pubescent across the backs proximally and glabrous distally. Lemma margins are pubescent throughout the range of the species.

Bromus exaltatus is often confused with Festuca breviglumis Swallen (1950: 398) ( McVaugh 1983, J.M. Saarela & P.M. Peterson pers. obsv.), which is similarly distributed from central México to Costa Rica. McVaugh (1983) noted that F. breviglumis is more common than Bromus species in western México. Although superficially similar, the two species are readily distinguished by a number of characters. Bromus exaltatus differs from F. breviglumis in its closed sheaths [vs. open], anthers <3 mm long [vs. (3–)4–4.5(–5.3) mm long], lemmas that are pubescent marginally, proximally and sometimes distally [vs. glabrous or puberulent], lower glumes (5–) 9–11 mm long [vs. (3–)4–5(–8) mm long], upper glumes 11–14 mm long [vs. 5.5–7(–9) mm long], awns 2.5–6 mm long [vs. (6–)8–12(–20) mm long], palea keels ciliate [vs. scabridulous] and lemmas inserted <0.5 mm below the lemma apex [vs. 0.5–1.2 mm below the lemma apex] ( McVaugh 1983).

Specimens Examined:― COSTA RICA. Buenavista massif, road to Cerro Buvis, 3400m, 16 August 1969, A. E. Weston 5885 (CR-81595). San José: Cantón de Pérez Zeledón , Chirripó-Cuericí , 9.4661°N, 83.4944°W, 3400–3739 m, 20 July 1966, E. Alfaro 568 (INB-60022); Cordillera de Talamanca , Cerro de la Muerte , Asunción summit, [9.5°N, 83.67°W], 3250 m, 13 July 1966, R. W. Pohl & G. Davidse 10692 ( CR, F-1733108, US-3096597 ); Pérez Zeledón, P. N. Chirripó, Cuenca Térraba-Sierpe, 9.4512°N, 83.5046°W, 3350 m, 1 September 2000, A. Rodríguez 6542, V. Ramírez, G. Soto, L. Acosta & G. Sancho ( INB, MO); Valle de los Leones and in the lower part of the Valle de los Conejos along upper Río Talari , 9.45°N, 83.5167°W, 3250–3450m, 21–23 August 1971, W. C. Burger 8218 & L. Gómez P. (CR-54405, F-1824924, NY); Cerro de la Muerte, C. R. [9.57°N, 83.75°W], 3400 m, 30 August 1952, J. B. Carpenter 557 ( US-2117586 ); Chirripó, camino Sabana Leones, [9.32°N, 83.61°W], 5–13 November 1976, R. A. Ocampo 1494 (CR-63401); Chirripó, Vallede los Conejos, [9.32°N, 83.61°W], 5–13 November 1976, R. A. Ocampo 1463 & 1465 (CR-63336); Asunción summit, Cerro de la Muerte, [9.57°N, 83.75°W], 3335m, 22 July 1966, R. W. Pohl 10114 & C. Calderón (CR-140884) . GUATEMALA. Ixchiguan, San Marcos, [15.17°N, 91.93°W], 3200 m, 27 August 1977, D. N. Smith & B. Olson 765 (F-2069462); Totonicapán, on road between Huehuetenango and Sija , [14.91°N, 91.36°W], 3000–3450 m, 20 February 1939, P. C. Standley 65867 (F-987424). Huehuetenango: Sierra Cuchumatanes , 15.42°N, 91.34°W], 11100 ft [3383 m], 15 September 1934, A. F. Skutch 1242 ( GH, US-1637903 ); Sierra de los Cuchumatanes, between Tojiah and Chemal at km 319.5 on Ruta Nacional 9 N, [15.42°N, 91.34°W], ca. 3380 m, 30 July 1960, J. H. Beaman 3796 ( GH, MSC-172050 , US- 2381693 ); Aldea San Nicolás, Chinautla, [14.71°N, 90.5°W], 3160 m, 16 August 1976, D. N. Smith 317 (F- 2069942, F-2069943, F-1788847); between Tojquia and Caxin bluff, summit of Sierra de los Cuchumatanes, [15.54°N, 91.54°W], 3700 m, 6 August 1942, J. A. Steyermark 50151a ( US-1935058 , F-1201444); ladera sur del valle del Llano de la Ventura, Yac, Todos Santos, [15.77°N, 91.57°W], 3580 m, 7 August 1977, D. N. Smith 654 (F- 2069461); near Tuinimá, Sierra de los Cuchumatanes, [15.63°N, 91.52°W], 3300–3500 m, 6 July 1942, J. A. Steyermark 48273 (F-1201464). Quezaltenango: Volcán Santa Mariá, upper NE-facing slopes to summit of Volcáno, [14.76°N, 91.55°W], 3000–4200 m, 13 January 1949, J. A. Steyermark 34159 (F-1045771) .

Sacatepéquez: Volcán Agua, 5 December 1911, 14.465°N, 90.7431°W, 2000–3500 m, A. S. Hitchcock 9113 ( MICH-1119235 , S-1009548 ). San Marcos: Las Ventanas, San José Ojetenám, [14.77°N, 89.93°W], 3440 m, 11 October 1977, D. N. Smith 929 (F-1858162); upper slope of Volcán Tajumulco, between Las Canojas and top of ridge, 7 mi from San Sebastián, [15.03°N, 91.9°W], 3300–3900 m, 16 February 1940, J. A. Steyermark 35856 (F- 1052008). Totonicapan : on the Tecum Uman ridge at km 154 on Ruta Nacional no. 1, ca. 20 km Eof Totonicapan , [14.98°N, 91.41°W], ca. 3340 m, 13 August 1960, J. H. Beaman 4146 ( US-2381724 ); Tecum Uman Ridge at km

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154 on Ruta Nacional No. 1 ca 20 km Eof Totonicapan, [14.91°N, 91.36°W], ca. 3340 m, 13 August 1960, J. H. Beaman 4146 ( MSC-171998 ). MÉXICO. Chiapas: San Cristóbal Las Casas , 19 December 1972, 16.7100°N, 92.6200°W, 7200 ft., J. R. Reeder & C. G. Reeder 6068 ( MO); about 15 mi SE of Teopisca , 21 August 1953, 16.47°N, 92.29°W, J. R. Reeder & C. G. Reeder 2031 ( MO); Amatenango del Valle, 14 km SE of Teopisca along highway to Comitán , 16.5°N, 92.32°W, 2100 m , 15 November 1984, G. Davidse, M. Sousa S., O. Tellez V., E. Martínez L. & J. Davidese 29801 ( MO); NE edge of San Cristóbal Las Casas, 16.73°N, 92.63°W, 2250 m , D. E. Breedlove & G. Davidse 54749 ( MO); Zinacantán, near Paraje Nachij [Nachig], 16.72°N, 92.72°W , 28 October 1981, D. E. Breedlove & G. Davidse 53890 ( CAS, MO). Distrito Federal: 14 mi Eof Amecameca on road towards Puebla, in Parque Nacional Izta-Popo, 19.087°N, 98.68°W, 3330 m , 13 October 2001, P. M. Peterson & Y. Herrera- Arrieta 16166 ( CAN, MO, US); Llano Grande, cerca del Desierto de los Leones, Delegación Cuajimalpa, [19.32°N, 99.29°W], 3300 m , 18 January 1970, J. Rzedowski 26998 ( MICH-1119236 ); Valle de México, Cerro de Sta. Rosa, Contreras , 19.3°N, 99.2833°W, 2900 m , 5 November 1950, E. Matuda 18647 ( MEXU-230673 , US- 2040887 ). Guerrero: Cima del Cerro Teotepec , [17.45°N, 100.16°W], 3200 m , 24 August 1999, E. Domínguez 852 ( MEXU); Mpio. Chilpancingo, al Ede Omiltemi, rumbo a Cuevade la Viego , [17.56°N, 99.5°W] , 7 October 1985, G. Lozano Valdez 883 ( MEXU-486848 ); Mpio. General Heliodoro Castillo, Puerto Unión, ladera Cerro Teotepec , [17.45°N, 100.16°W], 3030 m , N. Diego, B. Ludlow & J. M. Davila 8205 ( MEXU); Mpio. Tlacotepec, Cerro Teotepec, [17.45°N, 100.16°W], 3350 m , 5 December 1963, J. Rzedowski 18153 ( ARIZ-237656 , MICH- 1119237 ). Hidalgo: Vallede México, Real del Monte , 20.1333°N, 98.6667°W, 2600 m , 12 August 1951, E. Matuda 21646 ( MEXU-91094 , US-2041604 ). Jalisco: 20.4 mi SW of Ciudad Guzmán, NE slopes of Nevado de Colima, 19.5971°N, 103.5869°W, 3245 m , 5 October 2001, P. M. Peterson & O. Rosales 16087 ( CAN, MO, US); 21.7 mi SW of Ciudad Guzmán , NE slopes of Nevado de Colima, 19.5962°N, 103.5896°W, 3282 m , 6 October 2001, P. M. Peterson & O. Rosales 16110 ( CAN, MO, US); 23.4 mi SW of Ciudad Guzmán , NE slopes of Nevado de Colima, 19.5935°N, 103.5923°W, 3415 m , 6 October 2001, P. M. Peterson & O. Rosales 16108 ( CAN, MO, P- 03216907 , US); Mpio. Tuxpan, 19 km al SO del Rancho de Las Milanes , brecha al Parque Nacional El Nevado, [19.56°N, 103.61°W], 3200 m , 20 Enero 1990, J. Villa C., S. D. Koch & J. Chavez L. 557 ( MICH-119231 ); NE slopes of the Nevado de Colima, above Canoa de Leoncito , 19.6167°N, 103.65°W, 3100–3300 m , 10 September 1952, R. McVaugh 12852 ( MEXU-772540 , MICH-1119232 , NY, US); road from Zapotlán to Mt. Nevada , 19.7°N, 103.5167°W, 1829–2743 m , 23–24 September 1910, A. S. Hitchcock 7152 ( US-1009479 ); Taludes de exposición N del Nevado de Colima, [19.56°N, 103.61°W] , 19 September 1980, A. A. Beetle & R. Guzmán M-5375 ( MEXU). Estado de México: circa urben México secus margines agrorum ubi Mays consita, January, W. Schaffner 42 ( P- 02630901 ); Mpio. Zinacantepec , a 4.93 km al SE de Ojo de Agua, 19.1786°N, 99.74°W, 3368 m , 11 October 1995, A. Ramírez Abarca 678 ( MEXU-1089789 ); 1.7 mi Eof Hwy. 10 along road to Nevado de Toluca , 19.1347°N, 99.8°W, 3688 m , 9 October 2007, P. M. Peterson, J. M. Saarela & M. J. Flores Villegas 21350 ( CAN, MO, P- 03216870 , US); junction of Hwy. 10 and road to Parque Nacional Nevado de Toluca , NW of the volcán, 19.1506°N, 99.8°W, 3572 m , 11 October 2001, P. M. Peterson & Y. Herrera-Arrieta 16155 ( CAN, MO, US); Mount Popocatépetl , 19.0222°N, 98.6278°W, 2700 m , 7 August 1910, A. S. Hitchcock 883 (F-715973, GH); Mpio. Amecameca, primera cañada al Nde la carretera al Paso de Cortés , sobre una brecha maderera que entra en el km 15, 19.06°N, 98.7°W, 3310 m , 10 December 1976, S. D. Koch 76323 ( US-2832267 ); Mt. Popocatépetl , 19.0333°N, 98.6333°W, 3048 m , 5–6 August 1910, A. S. Hitchcock 5985 ( NY, US-1009480 ) & 993 ( US-1009498 ). Oaxaca: Cañada El Derrumbadero, 7 km al NW de La Cofradía comunidad de San Pedro El Alto, [16.04°N, 96.47°W], 2800 m , 7 October 1998, A. G. Miranda M. & O. L. Hernández Martínez 188 ( MEXU-907865 ); Mpio. Zimatlán, Paraje Puesto Vigía, Comunidad de San Pedro El Alto , [16.04°N, 96.47°W], 3000 m , 30 November 1998, A. G. Miranda Moreno 798 ( MEXU); vicinity of Cerro Zempoaltepetl, 17.09°N, 95.88°W, 2900–3100 m , 10 August 1950, B. Hallberg 913 ( MEXU-58486 , MICH-1119228 , US). Puebla: 23.5 mi Ea of Amecaneca on road towards Puebla in Parque Nacional Izta-Popo , 19.089°N, 98.6°W, 3360 m , 13 October 2001, P. M. Peterson & Y. Herrera- Arrieta 16174 ( MO, US); Mpio. San Nicolás de los Ranchos, Declive Edel Paso de Cortés , 3.5 km (5 km por la brecha) al NE del entronque de la..., 19.0667°N, 98.4833°W, 3600 m , 6 December 1976, S. D. Koch 76315 ( US- 2832233 ). Queratero : 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 24681 ( US). San Luis Potosí: Virlet d'Aoust 1410 ( P- 02630902 ); 30 mi Eof San Luis Potosí, along Hwy. 86 to Rioverde , 22.06°N, 100.56°W, 2103 m , 13 July 1963, R. L. McGregor, L. J. Harms, A. J. Robinson, R. del Rosario & R. Segal 607 ( US). Sonora: along Arroyo El Kipor (Quipor) and on Cordon Las Taunas, from El Kipor E. to Tierra Panda (Las Taunas), 28.4°N, 108.5583°W, 1680 m ,

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T. R. Van Devender & A.L. Reina G. 95-936 ( ARIZ-322740 ). Tlaxacala: Parte alta de la Malinche, [19.38°N, 98.05°W], 26 June 1993 , E. Oviedo 39 ( MEXU-1089796 ); Volcán Malinche , NW slopes, approached from Huamantla , 19.3°N, 98°W, 3340 m, 3 September 1962 , D. Ugent, V. Ugent & R. Flores-C. 1436 ( US-2543137 ). Veracruz: Barranca El Caracol, Parque Nacional Cofre de Perote , 19.5°N, 97.1°W, 3200 m, 17 August 1983 , H. Narve F. 942 ( MEXU-726958 ); Mpio. Xico, Barranca de Morei, cerca de la rancheria Paso Panal , 4 km al Ede Tembladoras, 19.5°N, 97.15°W, 2900 m, 14 September 1983 , H. Narave F. 1008 ( MEXU-793415 ); Mt. Orizaba , 19.0167°N, 97.22°W, 25–26 July 1901 , J. N. Rose & R. Hay 5733 ( MEXU, US-395519 . PANAMA. Chiriqui: Volcán de Chiriqui, Potrero Muleteo to summit, [8.79°N, 82.52°W], 3500–4000 m, 13–15 July 1940 , R. E. Woodson, Jr . & R. W. Schery 408 ( GH, MO, US-1818226 ); Volcán de Chiriqui, Potrero Muleto to summit, Boquete District , [8.79°N, 82.52°W], 10,400 ft [3170 m], 18 July 1938 , M. E. Davidson 1047 (F-934768, US-1820854 ); Chiriqui Volcáno , [8.77°N, 82.5°W], 3000 m, 29–30 September 1911 , A. S. Hitchcock 8214 ( MICH-1119234 , US- 1009552 ) .

Figures

FIGURE 35. Bromus exaltatus. A. Spikelet. B. Lemma. C. Habit. D. Ligule. Based on Hitchcock 883 (GH). Illustration by Paulette Dennis © Paulette Dennis.

FIGURE 36. Bromus exaltatus. McVaugh 12852 (MICH-1119232).

FIGURE 37. Geographical distribution of Bromus exaltatus in México and Central America.