Bromus carinatus Hooker & Walker Arnott 1840

5. Bromus carinatus Hooker & Walker Arnott (1840: 403). Ceratochloa carinata (Hook. & Arn.) Tutin in Clapham et al. (1952: 1458). Type:— UNITED STATES OF AMERICA. California: Monterrey or San Francisco, at no great distance from the coast, 1833, D. Douglas s.n. (lectotype BM-000555843!, designated by Pavlick 1995: 40, isolectotypes E-00064818!, US-865522A fragm. ex BM, MO-2957183!, NY-346213!). Festuca pendulina Sprengel (1825: 356), replacement name. Replaced name: Bromus pendulinus Sessé ex Lagasca (1816: 4), nom. illeg. Blocking name: Bromus pendulinus Schrader (1810: 70). Bromus hookeri var. pendulinus (Spreng.) Fournier (1886: 128). Type:—Habitat in Nueva Hispania [México] unde semina advexit D. Sessé ann. 1804 [grown from seeds collected by Sessé in "Nova Hispania"] (holotype MA-14058, isotypes M, US-865450! fragm. ex M). Bromus luzonensis Presl (1830: 262). Triticum luzonense (Presl) Kunth (1833: 446). Type:— California or British Columbia: Haenke s.n. (holotype PR!). See Veldkamp (1990: 660) for typification and discussion of label errors. Also see comments below. Bromus hookerianus Thurber (1874: 493), replacement name. Replaced name: Ceratochloa grandiflora Hooker (1840: 253). Blocking name: Bromus grandiflorus Weigel (1772: 9), nom. illeg. superfl. Bromus carinatus var. hookerianus (Thurb.) Shear (1900: 60). Type:— UNITED STATES OF AMERICA. Oregon: plains of the Columbia River, 1826, D. Douglas s.n. (lectotype K, designated by Shear 1900: 61, isolectotype US-865712! fragm. ex K). Bromus oregonus Nuttall ex Hooker (1856: 18), nom. nud. Bromus oregonus Nuttall ex Shear (1900: 59), nom. inval., as syn. of Bromus carinatus. Type:— UNITED STATES OF AMERICA. Upper Missouri and Oregon territories, Geyer 244. Bromus virens Buckley (1862: 98). Type:— UNITED STATES OF AMERICA. Rocky Mountains and Columbia River, Nuttall s.n. (holotype PH-00008640!, isotypes BM-000578894!, US-865484 fragm.). Bromus hookeri var. schaffneri Fournier (1886: 127). Bromus schaffneri (E. Fourn.) Scribner & Merrill (1901: 30). Type:— MÉXICO: secus margines agrorum ubi Mays consit a, J.G. Schaffner 42 (holotype P-00647229!, isotype US-865501! fragm. exP). Bromus hookeri var. schlechtendalii Fournier (1886: 127). Bromus proximus var. schlechtendalii (E. Fourn.) Shear (1901: 245). MÉXICO. San Luis de Potosí, August 1891, Virletii 1420 (lectotype P-02632337!, designated here, isolectotype P-02632340!). The lectotype is chosen from among the five syntypes listed by Fournier (1886). Bromus virens var. minor Scribner ex Beal (1896: 614). Type:— UNITED STATES OF AMERICA. Oregon, 30 May 1884, T. Howell s.n. (holotype US-1007683!). Bromus carinatus var. californicus Shear (1900: 60). Type:— UNITED STATES OF AMERICA. California: San Diego, 24 May 1884, C.R. Orcutt 511a (lectotype US-1007777!, designated here). An annotation by P.M. Peterson in 2002 on the specimen indicates the lectotype is designated in "Contr. U.S. Natl. Herb. 2003 in ed.," but the lectotypification does not appear there. Bromus carinatus var. densus Shear (1900: 61). Type:— UNITED STATES OF AMERICA. California: Channel Islands, San Nicholas Island, about Opuntia, April 1897, B. Trask 12 (isotype US-340331!). Bromus carinatus var. linearis Shear (1900: 61). Type:— UNITED STATES OF AMERICA. California. 1875, Vasey (holotype US-81566!). Bromus laciniatus Beal (1896: 615). Ceratochloa laciniata (Beal) Holub (1973: 170). Type:— MÉXICO. Oaxaca: Sierra de San Felipe, alt. 9500 ft, 17.1°N, 96.85°W, 1894, C.G. Pringle 4897 (holotype MSC-4813!, isotypes E-00373824, GH, JE-00006481, K, MEXU-5884!, MO-2957787, NY, US-250847!). Bromus subvelutinus Shear (1900: 52). Type:— UNITED STATES OF AMERICA. Nevada: Reno, 1887, S.M. Tracy 249 (holotype US-81594!). Plants short-lived perennials, not rhizomatous. Culms up to 150 cm tall, 2–6 cm wide at base, erect, glabrous or pubescent below inflorescences; nodes 3–6, glabrous or pubescent. Leaf sheaths glabrous or pubescent to pilose, hairs up to 2 mm long, throats glabrous or pubescent; auricles absent; ligules (1–)2–3(–4) mm long, glabrous or pubescent, lacerate-erose; blades up to 33 cm × (2–)3–6(–11) mm, flat, abaxial and adaxial surfaces glabrous or pubescent, midveins narrowing beneath the collar, margins serrulate. Panicles 5–30 cm long, open, branches erect, ascending or spreading, 4–22 cm long including spikelets, pedicels longer or shorter than spikelets, glabrous, scabrous, or pubescent, 1–3(–9) spikelets per branch. Spikelets 2–4 cm long, 4–11-flowered, elliptic to lanceolate, strongly laterally compressed; glumes glabrous or pubescent, green to purple along and between the nerves, margins hyaline, midnerves glabrous to scabrous distally; lower glumes 6.5–12 mm long, lanceolate, 3–7-nerved, apices acute; upper glumes 9–15 mm long, shorter than the lowest lemma, obovate-lanceolate, 5–9-nerved, apices acute to acuminate; lemmas 11–20 mm long, lanceolate, laterally compressed, strongly keeled distally, apices acute to obtuse, 7–9-nerved, nerves usually not raised, glabrous, scabrous, or pubescent, hairs sometimes restricted to margins, hairs up to 0.5 mm long; awns 2–15 mm long, inserted 0–0.5 mm below the lemma apex, straight; paleas shorter than the lemmas, backs glabrous or pubescent, keels ciliate, cilia up to 0.2 mm long; anthers 0.5–4.5 mm long; caryopses 9−11 mm long. 2 n = 56 (Pohl & Davidse 1971). Distribution: ―Native. Bromus carinatus s.l. is widespread in México (Fig. 14). Ecology:— Meadows, pastures, barrancas and slopes; associated with Pinus arizonica, P. jeffreyi Balf. in Murray (1853: pl. s.n.), Abies religiosa (Kunth in Humboldt et al. 1817: 5) Schlechtendal (1830: 77), Pseudotsuga menziezii (Mirbel 1825: 63) Franco (1950: 74), Juniperus deppeana, Cupressus arizonica, Quercus spp., Q. rugosa Née (1801: 275), Arbutus xalapensis, Fallugia paradoxa (Don 1825: 576) Endl. ex Torr. in Emory (1848: 139), Arctostaphylos pungens, Salvia Linnaeus (1753: 23), Baccharis Linnaeus (1753: 860) and Ceanothus. Elevation 1500−3200 m. Common Names: ― California brome (English); basicuaáre, bromo de California, masiyague, pipillo (Spanish). Comments: ― Bromus carinatus is used as fodder in México (Saulés & Dávila Aranda 1992). Bromus carinatus seeds are used to make tejino or tesgüino, and when there is shortage of food they are apparently used as corn (Saulés & Dávila Aranda 1992). Tejino is "… a thick, milky, nutritious brew made from corn fermented with a local grass seed (basiáhuari)" (Kennedy 1963). Variation in the Bromus carinatus complex in North America— Bromus sect. Ceratochloa comprises the hexaploid (6x) B. catharticus complex native to South America, the octoploid (8x) B. carinatus complex native to North America, the North American duodecaploid (10x) B. arizonicus (Stebbins 1981), and recently discovered duodecaploid taxa from South America similar in morphology to B. ayacuchensis Saarela & P.M. Peterson in Saarela et al. (2006: 919), for which the chromosome number is unknown (Williams et al. 2011). There is extensive morphological variation within each of the hexaploid and octoploid complexes, which has led to multiple, often-conflicting taxonomic treatments for each. Recent genetical and morphological study of the B. catharticus complex supported recognition of two polymorphic species (Massa et al. 2001, 2004). Similar revisionary work is necessary to better characterize morphological and molecular variation in the B. carinatus complex throughout its natural range. The B. carinatus complex ranges throughout much of western North America, from Alaska to Central America. It has been introduced elsewhere, including Europe (Verloove 2012), China (Liu et al. 2006) and New Zealand (Stewart 1996). It comprises primarily cleistogamous plants and is extremely variable. Multiple taxa have been described in it on the basis of such characters as duration (i.e., annual, perennial, biennial), pubescence of the leaves and spikelets, blade width, panicle size and awn length (Hitchcock et al. 1969). In the early part of the twentieth century, multiple narrowly circumscribed taxa were generally recognized in western North America north of México (Piper 1906, Jepson 1912, Millspaugh & Nuttall 1923, Tidestrom 1925, Hitchcock 1951), whereas in more recent decades most authors have tended to recognize fewer taxa, often including much of the variation in a single species, B. carinatus, usually with infraspecific taxa (Hitchcock et al. 1969, Stebbins 1981, Jones et al. 1997, Pavlick & Anderton 2007). Pavlick (1995), however, recognized six taxa in North America north of México, distinguished primarily by differences in the vestiture of leaf sheaths and blades and panicle size: B. sitchensis Trin. in Bongard (1832: 173), B. aleutensis Trin. ex Griseb. in Ledebour (1852: 361), B. maritimus (Piper 1905: 148) Hitchc. in Jepson (1912: 177), B. carinatus (including vars. carinatus and marginatus), B. polyanthus Scribner ex Shear (1900: 56) and B. subvelutinus Shear (1900: 52). Pavlick & Anderton (2007) recognized the same taxa in the Flora of North America, except B. subvelutinus was included in B. carinatus var. carinatus. Numerous authors have commented on the extensive morphological variation in the complex, and have noted that there is considerable intergradation among taxa throughout the range of the complex (Shear 1900, Hitchcock 1951, Hitchcock et al. 1969, Pavlick 1995). Given the variation and difficulties in delimiting taxa in the B. carinatus complex, Stebbins (1981) suggested that all taxa be treated as a single species. No treatment so far has taken this approach. An AFLP study of B. carinatus s.l., based on 30 accessions collected throughout western North America, identified four major genetic lineages and five unique accessions, but the authors did not comment on morphological variation within the major lineages, or whether or not the lineages correspond to previouslyrecognized taxa in the complex (Massa & Larson 2005). Barkworth et al. (

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