Bromus anomalus Ruprecht ex Fournier 1886

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) [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). 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. 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, consis

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