Bromus berteroanus Colla (1836: 25

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 : 31-35

4. Bromus berteroanus Colla (1836: 25 , pl. 58). Figs. 11 , 12 .

Type :— CHILE. Cachapoal : Rancagua, 1830, Bertero 117 (holotype TO-8207!, isotypes P, SGO, US-1645128 ! with three fragment packets: one ex P, two ex MPU) .

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Trisetum litorale Philippi (1858: 92) . Trisetum trinii var. litorale (Phil.) Louis-Marie (1928: 243) . Type:— CHILE. cerca Coquimbo, en la piedras marítimas, 1837, Gay (holotype SGO-PHIL-063594! [Herb. Chil. No. 148, barcode SGO000000778]).

Avena paupercula Philippi (1858: 94) . Type :— CHILE. Concepción : prope Tomé unicum specimen invenit, November 1855, Germain s.n. (holotype SGO-PHIL-63589! [barcode SGO000000093 ], isotype BAA-00001524!).

Trisetum barbatum var. major Vasey (1893: 60) . Bromus barbatoides var. sulcatus Beal (1896: 615) , nom. illeg. superfl. Trisetum trinii var. majus (Vasey) Louis-Marie (1928: 243) . Type:— MÉXICO. 1888, E. Palmer 667 (holotype US- 1009576!, isotypes NY-1645070!, NY-16465076!, P-02630826). The US specimen and the protologue indicate only " México " as the locality, whereas the NY and P specimens identify the location as “Lower California, Guadalupe Island.”

Bromus barbatoides Beal (1896: 614) , replacement name. Replaced name: Trisetum barbatum Steudel (1854: 229) , nom. illeg. Blocking name: Trisetum barbatum Nees (1841: 345) . Type:— CHILE. Quillota, October 1829, Bertero 860 (holotype P- 00624386!, isotypes BAA-00004161!, MO-2114333!).

Bromus bicuspis Nees ex Steudel (1854: 322) . Danthonia pseudo-spicata Müller (1856: 348) , nom. illeg. superfl. Type:— CHILE. Valparaíso, Cuming 466 (holotype B-100250740!, isotypes CGE, K-308532!, K-308533!, US-103017! fragm. ex B, US-103016! fragm. ex CGE, W-0026507!).

Bromus trinii Desvaux (1853: 441) , replacement name. Replaced name: Trisetum hirtum Trinius (1836b: 300) . Blocking name: Bromus hirtus Lichtenstein ex Roemer & Schultes (1817: 654) . Trisetobromus hirtus (Trin.) Nevski (1934: 15) . Type:— CHILE: Chil. austr. Andes de Antuco, Poeppig, Pl. Chili III 33 (lectotype W-0026498!, designated here, isolectotypes W-18890242057!, W18890242058!, W-1889003750!, US-820872! fragm. ex P); Trinius cited two collections in the protologue of T. hirtum , the one chosen as the lectotype and “Chil. bor., in campis ad Concon, Poeppig s.n. (W-0026497!).”

Bromus trinii var. effusa Desvaux (1853: 442) . Trisetum trinii var. effusum (E. Desv.) Louis-Marie (1928: 243) . Type:— CHILE: Elqui: sobre las colinas maritimas en Coquimbo, por setiembre, Gay s.n. (lectotype P, designated by Pavlick et al. 2003: 187). Two syntypes were cited in the protologue, the lectotype and “Andes de Odessa, en noviembre, Gay s.n. (P- 00625385!)”. Pavlick et al. (2003) noted the collection from Coquimbo to be the holotype; this is treated as an error to be corrected to lectotype under Article 9.9 of the Code. We did not find this specimen among the images available online via the Pwebsite (accessed 19 July 2013).

Bromus trinii var. manicata Desvaux (1853: 441) . Trisetum trinii var. manicatum (E. Desv.) Louis-Marie (1928: 243) . Type:— CHILE. Biobío: Antuco, Poeppig s.n. (holotype B-10 0248068!, isotypes BAA-00001635!, P-00625388!).

Bromus trinii var. micranthera Desvaux (1853: 441) . Trisetum trinii var. micrantherum (E. Desv.) Louis-Marie (1928: 243) . Type:— CHILE. Valparaíso: Valparaíso, Anon. (holotype P-00625389!).

Bromus trinii var. pallidiflorus Desvaux (1853: 441) . Trisetum trinii var. pallidiflorum (E. Desv.) Louis-Marie (1928: 243) . Type:— CHILE. Santiago: Santiago, C. Gay s.n. (lectotype P-00625390!, designated here).

Bromus trinii var. stricta Desvaux (1853: 442) . Trisetum trinii var. strictum (E. Desv.) Louis-Marie (1928: 243) . Type:— CHILE. Elqui: Coquimbo, 1838, C. Gay s.n. (lectotype P-00625387!, designated here, isolectotypes BAA- 00001638!, US-865475!).

Bromus trinii var. excelsus Shear (1900: 25) . Bromus berteroanus var. excelsus (Shear) Pavlick (1995: 128) . Type:— UNITED STATES OF AMERICA. California: Inyo Co., Panamint Mountains, 1700 m, 31 March 1891, F.V. Coville & F. Funston 522 (holotype US-81596!).

Plants annual, often tufted, not rhizomatous. Culms (8–) 15–85 cm tall, 1–3(–4) mm wide at base, erect, usually glabrous below inflorescences, sometimes scabrous to pubescent; nodes 2–4, glabrous or pubescent. Leaf sheaths pubescent to pilose, hairs up to 1.5 mm long, or glabrous; auricles absent; ligules 1–3 mm long, glabrous or pubescent, erose-lacerate; blades up to 14 cm × 2–6 mm, flat, adaxial surfaces glabrous to pilose, hairs up to 2.5 mm long, abaxial surfaces glabrous to pubescent, hairs up to 0.5 mm long, sometimes restricted to margins, margins serrulate. Panicles (5–) 8–20 cm long, ± open, branches erect to ascending, glabrous or scabrous, 1–2(–4) spikelets per branch. Spikelets 1.5–2.1 cm long, 3–7(–8)-flowered, elliptic to lanceolate, terete to moderately laterally compressed; glumes glabrous, margins hyaline, midnerves glabrous, sometimes scabrous distally; lower glumes 6–12 mm long, lanceolate, 1(3)-nerved, green to purplish-green along the nerve(s), apex acute to attenuate; upper glumes 8–16 mm long, obovate-lanceolate, 3(5)-nerved, green to purplish-green along and between the nerve(s), apex mucronate, mucros up to 0.8 mm long; lemmas 6–15 mm long, lanceolate to linear-lanceolate, rounded over the backs, apices strongly bifid, teeth 2–3(–4) mm long, acuminate to awn-like, 7-nerved, light green to purplish-green along and between the nerves, variously pubescent to pilose, hairs up to 0.9 mm long; awns (7–) 10–20 mm long, arising 1.5 mm or more below the lemma apex, bent, twisted below middle; paleas narrower and shorter than the lemma, backs glabrous or scabrous, keels ciliate, cilia up to 0.3 mm long; anthers 1.5–2.5 mm long; caryopses 7−8.5 mm long. 2 n = 42 ( Knowles 1944, Bowden & Senn 1962, Klos et al. 2009).

Distribution: ―In México this species is known from Baja California and the Sierra Pinacate in Sonora ( Fig. 13 ). In the adjacent United States it is present in California, Oregon, Nevada, southern Utah and Arizona (Pavlick

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& Anderton 2007). It is native to South America, but its status in North America is unclear. Some authors have reported this species to be introduced in North America ( Hitchcock 1951, Villaseñor & Espinosa-García 2004, Pavlick & Anderton 2007), while others have considered it to be native ( Stebbins 1981, Peterson & Soreng 2007). Felger (2000) suggested that the Sierra Pinacate population may be a native Pleistocene relict.

Ecology: —Dry, open areas in sandy or gravelly, granitic or volcanic soils; associated with Fouquieria columnaris , Pachycereus pringlei ( Watson 1885: 368) Britton & Rose (1909: 422) and sclerophyllous shrubs. The species is a winter-spring ephemeral (annual) ( Felger 2000). Elevation: 10–1300 m.

Common Names: ―Chilean Chess (English).

Comments: ―The epithet berteroanus was given in honour of Italian botanist Carlo Luigi Giuseppe Bertero (1780–1831), who collected the type specimen. In the Sierra Pinacate in Sonora B. rubens may be displacing B. berteroanus ( Felger 2000) . This species is readily distinguished by its twisted and geniculate awns, character states that do not occur in any other Bromus species in North America. It is one of two species of sect. Neobromus, and the only one present in North America; the other, B. gunckelli Matthei (1986: 62) , is native to Chile.

Specimens Examined:― MÉXICO. Baja California: 2 mi Nof Parador Santa Ines , 27.0667°N, 111.9833°W, 670 m, 26 March 1979, J. R . Reeder & C. G . Reeder 7104 ( SD-116039 , US-2861045 ) ; 3 mi SE of Consuelo , 30.15°N, 115.7667°W, 75 m, 24 March 1970, R . Moran 17845 ( SD-76756 ) ; 3 mi WSW of El Socorro , 30.9583°N, 115.6875°W, 1050 m, 9 May 1978, R . Moran 25786 ( SD-100664 ) ; 5.3 km NE of Rancho Aguajito , 480 m, 19 April 1958, P. H . Raven, M . Mathias & J . Turner 12537 ( GH); Cedros Island , 28.22°N, 115.25°W, 18–20 March 1889, E . Palmer 659 ( GH, NY, 2 sheets); Cedros Island , copper mine canyon 1.5 mi from N end, [28.3667°N, 115.2167°W], 350 m, 27 March 1973, R . Moran 20332 ( SD-86929 ); Cedros Island , Nof Cañon Grande , top of Sern pine ridge, 28.1833°N, 115.2°W, 525 m, 30 March 1978, R . Moran 25417 ( SD-99680 ); Cedros Island , Punta Norte, 28.35°N, 115.2167°W, 510 m, 14 April 1983, T . Oberbauer, H . Wier & E . Wier 111 ( SD- 127654 ); Cedros Island , northern part of island in the vicinity of El Faro, 28.2833N, 115.25W 17 March 1994, J. P . Rebman, T . Nash, J . Delgadillo et al. 2417 (ASU-0010747), Cerro San Luis, 29.3167°N, 114.1167°W, 1300 m, 2 March 1963, R . Moran & J . Henrickson 10320 ( RSA-POM-165017 , SD-54520 ); E end of Sierra San Luis , 26–30

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mi Nof Punta Prieta , 29.36°N, 114.2833°W, 914–1097 m, 1 April 1950, H. S . Gentry & F . Cech 8912 (ASU- 0010746, DES-00009272, MICH-1119289 , US-2820899 ); Ensenada, [31.78°N, 116.6°W], 11 April 1882, M. E . Jones s.n. ( RSA-POM-112737 ); Guadaloupe Ranch, 32.5°N, 116.6°W, 6 April 1886, C. R . Orcutt s.n. (US); Guadalupe Island, [29.04°N, 118.28°W], 1875, E . Palmer 99 ( GH, NY, 3 sheets, one mixed with B. tectorum ); Guadalupe Island, [29.04°N, 118.28°W], 1889, E . Palmer 658 ( GH, MEXU-5964 , NY, 2 sheets) & 667 ( P- 002630826 ); Guadalupe Island, [29.04°N, 118.28°W], 29 March 1889, E . Palmer 898 (F-716074, GH, NY, P- 02632079 ); Guadalupe Island, along canyon leading Wfrom Lobster Camp , [29.04°N, 118.28°W], 29 April 1958, I.L. Wiggins & W. R . Ernst 205 ( GH); Guadalupe Island, canyonside Wof S Bluff , 28.8875°N, 118.275°W, 100 m, 27 April 1958, R . Moran 6700 ( SD-47590 ); Guadalupe Island, Melpomene drainage, [29.03°N, 118.275°W], 300 m, 29 April 1958, R . Moran 6729 ( SD-47591 ); Guadalupe Island, near mouth of Long Canyon, 29.0083°N, 118.2333°W, 100 m, 16 April 1970, R . Moran 17369 ( SD-74762 ); Guadalupe Island, S zone, [29.04°N, 118.28°W], 25 March 1982, A. E . Meling 37 ( SD-119488 ); Guadalupe Island, W Anchorage , 28.975°N, 118.2917°W, 20 m, 9 February 1957, R . Moran 5616 ( SD-46922 ); Guadalupe Island, [29.04°N, 118.28°W], 1 April 1889, E . Palmer 899 (F-716073, GH, NY, P-02632080 ); Hassler Cove, San Martín Island, 30.4833°N, 116.1083°W, 10 m, 10 April 1963, R . Moran 10498 ( SD-54331 , US); Nslope of Cerro San Juan de Dios , 30.1333°N, 115.1333°W, 1100 m, 1 May 1973, R. Moran 10687 ( RSA-POM-26201 , SD-87045 ); near San Juan Mine, 28.7167°N, 113.6333°W, 1200 m, 29 March 1960, R . Moran 8130 ( SD-60614 ); near Vallecito, 32.3833°N, 116.8833°W, 5 April 1886, C. R . Orcutt s.n. ( US-1009578 ); San Pedro near La Bocana, 28.4667°N, 113.4167°W, 250 m, 11 March 1966, R . Moran 12520 ( SD-63135 ); Santa María Plains 10–20 mi Sof Hamilton Ranch, 30.4°N, 115.8833°W, 6 April 1931, I. L . Wiggins 5199 ( GH, MICH-1119290 , NY, RSA-POM-26595 , US); Sierra San Borja, hillside near San Juan Mine, 28.7167°N, 113.6333°W, 1200 m, 29 March 1960, R . Moran 8130 ( RSA-POM- 172594 , US-2461509 ); ca. 0.5 km off of Hwy. 3 between Ensenada and H . de la Independencia, 0.2 km Nof km 87, 31.75°N, 115.8333°W, 1150 m, 12 August 1995, N. Snow & T . Prinzie 6421 ( MEXU); ca. 100 km Sof Ensenada on Rt. 1 near San Vicente , [31.11°N, 116.14°W], April 1967, Stephenson s.n. ( MSC-368365 ); Volcán Sureste, 12 km SW of Lázaro Cárdenas, 30.520742°N, 115.927466°W, 80 m, 21 February 2001, M. A . Baker & A . Pineda 13895 (ASU-0010745); San Quintín [Quentin] Bay, 30.560278°N, 115.9425°W, January 1889, E . Palmer 688 ( NY) & 686 ( P-02630823 ) . Sonora : Sierra Pinacate, summit of Pinacate Peak, [31.77°N, 113.49°W], 1290 m, 24 March 1970, R. S . Felger, T . Hansen, A . Woodin & J . Woodin 19449 ( RSA-POM-444025 , SD-190690 ); summit of Pinacate Peak, [31.77°N, 113.49°W], ca. 1250 m, 1 March 1987, R. S . Felger, C . Baker & G . Joseph 87-52 (RSA- POM-444057).

Figures

FIGURE 11. Bromus berteroanus. A. Glumes. B. Inflorescence. C. Lemma D. Spikelet. Illustration by C.T. Roché, reproduced from Barkworth et al. (2007) with permission.

FIGURE 12. Bromus berteroanus. Moran 6700 (SD-47590).

FIGURE 13. Geographical distribution of Bromus berteroanus in México.