FIGURE 52. Geographical distribution of Bromus lanatipes in México.
Cochise Co., Huachuca Mtns., September 1893, J.G. Lemmon s.n. (US, 2 sheets); Dos Cabezos, 31 October 1937, L.N. Goodding s.n. (US-1721928). New México: Gray’s Peak, 6500 ft, F.S. Earle 162 (US-1008675); Grant Co., about the S end of the Black Range, 6000 ft, 1904, O.B. Metcalfe s.n. (US-690339); Santa Fe Co., Glorieta, 2 July 1926, A.S. Hitchcock 22994 (US-1296512); Glorieta, 24 August 1910, E.O. Wooton s.n. (US-726304); Santa Fe, 3 July 1926, A.S. Hitchcock 23011 (US-1296516); Santa Fe, 1884, G. Vasey s.n. (US-1008802); Lincoln Co., Gray, 6000–6500 ft, July 1900, F.S. Earle & E.S. Earle 162 (US-382342); Ft. Stanton on El Paso road, 3 August 1937, L.N. Goodding 3514 (US-1723783); between Ft. Stanton and Ruidoso, 26 August 1937, L.N. Gooding & C.
Goodding 3519 (US-1723785); San Miguel Co., Las Vegas, 23 June 1891, L.H.D. s.n. (US-749847); vicinity of Las Vegas, 27 June 1928, G.E. Osterhout 7048 (US-1389144). Oklahoma: Cimarron Co., Mesa de Maya (Black Mesa), 3 mi N of Kenton, 9 July 1947, C.M. Rogers 4747 (US-2010965); N slopes of Black Mesa, 3 mi N of Kenton, 30 May 1952, U.T. Waterfall 10749 (US-2077766). Texas: Guadalupe Mts., 1881, [?] s.n. (US-132953);
Brewster Co., Chisos Mtns., Green Gulche, below Casa Grande, 26 August 1944, C.L. Lundell 13222 (US- 1913218); Jeff Davis Co., Sawtooth Mountain, Davis Mountains, 3 October 1926, E.J. Palmer 31833 (US- 1297432); Trans-Peco Texas, Davis Mtns., High Mt. Livermore, 22 July 1936, L.C. Hinckley 607a (US-1871685);
Uvalde Co., Black Mt., 13 August 1926, B.C. Tharp 4105 (US-1296999).
mm wide, midnerves glabrous proximally, scabrous distally, apices acute; lower glumes 6–11 mm long, narrowly lanceolate, 1-nerved, green to purple along nerve; upper glumes 10–17 mm long, lanceolate, 3-nerved, green to purple along and between nerves; lemmas 11–23 mm long, linear-lanceolate, rounded over the backs, apices bidentate, teeth 1–3 mm long, 5–7-nerved, green to purple along and between the nerves, glabrous or scabrous, margins hyaline, 0.2–0.4 mm wide; awns 12–30 mm long, inserted 1.5–4 mm below lemma apices, straight or arcuate, scabrous; paleas shorter and narrower than lemmas, backs glabrous or pubescent, keels ciliate, cilia 0.1–0.6 mm long; anthers 0.6–1.2 mm long; caryopses 8−11 mm long. 2n = 4x = 28 (Esnault 1984, Sánchez Anta et al. 1988, Sheidai & Fadaei 2005).
FIGURE 53. Bromus madritensis. A. Lemma. B. Spikelet. C. Inflorescence. Illustration by C.T. Roché, reproduced from Barkworth et al. (2007) with permission.
Distribution:―Introduced. In México B. madritensis is known only from Baja California (Fig. 55). In the United States its range includes California, southern Oregon and Arizona (Pavlick et al. 2007). Native to the Mediterranean region and Central and Atlantic Europe (Sales 1994).
Ecology:—Open, xeric, mostly disturbed sites. Elevation: 20–900 m.
Common Names:―Foxtail chess, madrid brome, Spanish brome, compact brome (English).
Comments:―Soderstrom & Beaman (1968) did not recognize B. madritensis in México, but it was treated by Gould & Moran (1981) for Baja California.
Bromus madritensis and B. rubens (sect. Genea) are part of the polymorphic B. madritensis complex, a group of morphologically similar taxa in which multiple taxa have been described (Scholz 1981, Sales 1993, Sales 1994).
The complex is distinguished from other taxa in sect. Genea by a combination of small lemmas and erect, contracted to somewhat contracted inflorescences during flowering (Sales 1994). They have mostly been recognized as distinct species (Gould & Moran 1981), although their recognition as species has been questioned on the basis of considerable morphological variation (Esnault 1984, Esnault & Huon 1985, Sales 1993). Sales (1994) conducted a multivariate analysis of the polymorphic B. madritensis complex and accepted one species and two subspecies (B. madritensis subsp. madritensis and B. madritensis subsp. rubens). Some recent treatments have
followed this circumscription (Jones et al. 1997, Saarela & Peterson 2012), whereas others have maintained the taxa as distinct species (Felger 2000, Aryavand 2002, Jessop et al. 2006, Pavlick & Anderton 2007).
FIGURE 54. Bromus madritensis. Moran 27265 (SD-102443).
FIGURE 55. Geographical distribution of Bromus madritensis in México.
Molecular studies have shed independent light on the evolutionary history of these taxa, informing their classification. Isozyme evidence suggests that the taxa have independent origins, in line with their recognition as species (Oja & Jaaska 1996, Oja 2002). A recent study of plastid and nuclear ribosomal loci, and the Waxy gene, demonstrated that these two taxa are allopolyploids that arose independently from different diploid parental taxa:
B. madritensis from a B. sterilis × B. fasciculatus Presl (1820: 39) cross, and B. rubens from a B. fasciculatus × B.
tectorum cross (Fortune et al. 2008). In both taxa the maternal genome is derived from the B. fasciculatus lineage (Fortune et al. 2008). Given their independent origins, we treat these taxa as species, a classification that reflects their evolutionary history. The morphological variation in the complex globally may reflect multiple origins of these taxa or hybridization among them.
The character states related to pubescence of the culms, glumes and lemmas given in the key may not always distinguish B. madritensis and B. rubens, as these characters apparently vary in the complex and B. madritensis can sometimes be pubescent (Sales 1994), but they distinguish the species in México based on the specimens examined here.
Specimens Examined:―MÉXICO. Baja California: 25 km SE of Tijuana, at bottom of Cañón la Presa, 32.3958°N, 116.8333°W, 190 m, 13 May 1982, R. Moran 30700 (SD-111163); at roadside 1.5 km NE of Las Delicias, ca. 17 km E of Ensenada, 31.9083°N, 116.425°W, 660 m, 20 May 1979, R. Moran 27265 (SD-102443);
between La Humarosa [Rumarosa] and Tecate, 32.53°N, 116.38°W, 27 April 1981, A.A. Beetle & R. Alcaraz M- 6745 (ARIZ-229626, MICH-1119160, MEXU); Guadalupe Island, S slope of cañon above NE Anchorage, 29.1542°N, 118.2833°W, 20 m, 14 February 1957, R. Moran 5688 (SD-47529); La Flor de Sol, 32.425°N, 116.95°W, 220 m, 22 June 1977, R. Moran 24273 (SD-97340); La Misión, between Ensenada and Tijuana, on steep slopes and in arroyo bottom along S side of river, 32.0936°N, 116.8694°W, 50 m, 18 April 1998, J. Rebman, P.
Flanagan & La Misión Community Group 5046 (RSA-POM-643071, SD-144705, SD-144706); Rancho ontiveros, Sern foothills of Otay Mountain just S of the US/MEX border between Tijuana and Tecate, along a side canyon of the Río Tecate just W of the MEX Hwy. 2 toll booth, 32.5461°N, 116.8544°W, 85 m, 26 April 2005, J. Rebman, J.
Delgadillo, M. White & K. Comer 11830 (SD-161420); San Carlos Canyon, S of Ensenada, [31.87°N, 116.59°W], 29 April 1981, A.A. Beetle & R. Alcaraz M 6600 (MEXU); San Isidoro, 30.7667°N, 115.5333°W, 900 m, 2 June 1975, R. Moran 22257 (SD-91369); Descando Valley, 32.1833°N, 116.8667°W, 15 m, 29 April 1972, R. Moran 19110 (MEXU, SD-83045).