Bromus diandrus Roth 1787

8. Bromus diandrus Roth (1787: 44). Figs. 29, 30.

Anisantha diandra (Roth) Tutin in Clapham et al. (1962: 1149). Anisantha diandra (Roth) Tutin ex Tzvelev (1963: 4). Type:— Gr. Bromoides, locustis maximus, lanuginosum, Italicum Hist. Nat.: 261. no. 444 (neotype OXF-Scheuchzer, designated by Sales 1993: 8). The holotype at B, collected by Roth, was destroyed during the war in 1943 (Sales 1993).

Bromus rigidus Roth (1790: 21). Genea rigida (Roth) Dumortier (1868: 67). Bromus madritensis var. rigidus (Roth) Babington in Sowerby ex Syme (1873: 161). Bromus villosus var. rigidus (Roth) Ascherson & Graebner (1901: 595). Anisantha rigida (Roth) Hylander (1945: 32). Bromus diandrus subsp. rigidus (Roth) Laínz (1967: 49). Anisantha diandra subsp. rigida (Roth) (Tzvelev 1976: 223). Bromus diandru s var. rigidus (Roth) Sales (1993: 9). Type:—“ Triandria Digynia. Bromus rigidus spiculis multifloris lato-lanceolatis rigidus, floribus diandris, panicula contracta. Roth, Catal. Bot. 1. P. 17. Habitat in Europa australi.” ([first-step] neotype B-Willd., designated by Sales 1993: 9, [second-step] neotype B-Willd. barcode no. B -W 02162 -02 0!, designated here, isoneotypes B-Willd. barcode nos. B -W 02162 -01 0!, B -W 02162 -03 0!, B -W 02162 -04 0!). Afolder in the Willdenow herbarium at Bincludes four sheets of the Roth collection that Sales (1993) designated as the neotype; there is a barcode on the folder cover (barcode no. B -W 02162 -00 0). Sales (1993) did not designate a particular specimen as neotype, thus a second-step neotype is selected here. The second-step neotype is a sheet with three flowering culms.

Plants annual. Culms 15–120 cm tall, 0.8–3 mm wide at base, erect or ascending, glabrous or weakly to moderately pubescent below inflorescences; nodes 2–3, blackish brown, glabrous. Leaf sheaths pubescent, hairs stiff and up to 0.6 mm long, or soft and up to 2.5 mm long; auricles absent; ligules 2.2–5 mm long, glabrous, erose-lacerate; blades 3–18 cm × 2–7 mm, flat, adaxial and abaxial surfaces pilose, hairs up to 1.5 mm long, margins smooth or pubescent. Panicles 6–30 cm × 2–25 cm, erect when young, sometimes spreading or nodding at maturity, branches erect to ascending, 0.3–5 cm long, usually shorter than spikelets, sometimes longer, scabrous to densely pubescent, 1–2 spikelets per branch. Spikelets 3.2–5.5 cm long (6.5–8 cm including awns), 5–8-flowered, linear-elliptic, moderately laterally compressed, widening at maturity with visible rachillas; glumes glabrous, margins hyaline, midnerves glabrous or scabrous, particularly distally, apices acute; lower glumes 12–26 mm, narrowly lanceolate to lanceolate, 1(–3)-nerved, green to purple along the nerves; upper glumes 18–35 mm long, lanceolate, 3(–5)-nerved, green to purple along nerves, lighter in colour and ± translucent between the nerves; lemmas 18–35 mm × 1–2 mm, linear-lanceolate, rounded over the backs, apices bifid, the cleft 2.2–5.2 mm deep, 7-nerved, green or sometimes

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purplish-green along and between nerves, backs sparsely to densely scabrous, sometimes with hairs up to 1 mm long at apex or along margins, occasionally with a dense tuft of hairs 0.1–0.2 mm long at base, margins hyaline (except at base), smooth or minutely serrulate, keels scabrous; awns 30–50 mm long, arising 3.5–6 mm below lemma apex, straight; paleas 13–16 mm long, shorter than lemmas, backs glabrous, keels ciliate, cilia up to 0.5 mm long; anthers 0.5–1.2 mm long; caryopses 10−12 mm long, terete in cross section to somewhat flattened dorsally. 2 n = 42, 56 (Oja & Laarmann 2002).

Distribution:―Introduced. In México B. diandrus is known from Baja California, where it has been present for over a century, and the Revillagigedo Islands (Colima), south of the Baja California peninsula (Fig. 31). Gould & Moran (1981: 30) reported B. diandrus to be "occasional to frequent" in Baja California. Bromus diandrus also occurs in central México in the states of México, Morelos, Puebla, Tlaxcala and Veracruz. It was not reported in central México by Soderstrom & Beaman (1968), but subsequent authors have reported it from Puebla (Beetle 1977, 1987, Dávila Aranda et al. 1990, Espejo-Serna et al. 2000), Estado de México (Beetle 1987, Espejo-Serna et al. 2000) and Distrito Federal (Espejo-Serna et al. 2000). Bromus diandrus was introduced to central México as early as the 1970s; a few collections were made in the 1970s and 1980s, and most collections were made more recently in the 1990s and 2000s. The species is likely spreading in this area. Also known from Guatemala (Cerro El Baul).

Bromus diandru s is native to Europe. It is widely introduced in western North America north of México, where it occurs from British Columbia south to California and Texas, with some scattered localities in the southeastern United States (Pavlick & Anderton 2007). In South America it occurs in Chile and Argentina (Planchuelo & Peterson 2000).

Ecology:— This species is widely distributed primarily in disturbed habitats along roadsides, rocky slopes, moist seeps and meadows; associated with Quercus spp., Pinus spp., Abies Miller (1754b: [unpaged]), Juniperus Linnaeus (1753: 1038), Arbutus Linnaeus (1753: 395), Rubus Linnaeus (1753: 492), Polygala Linnaeus (1753: 701), Solanum Linnaeus (1753: 184), Alnus Miller (1754b: [unpaged]), Baccharis, Sambucus Linnaeus (1753: 269), Prunus Linnaeus (1753: 473) and Penstemon Schmidel (1763). Elevation: 2400 m (Guatemala), 5–2200 m (México, Baja California), 1850–4267 m (México, except Baja California).

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Common Names: ―Ripgut brome, great brome (English); bromo frágil (Spanish).

Comments: ―Plants referred to B. diandrus in México and elsewhere in North America have previously been treated as B. rigidus Roth (e.g., Wiggins 1980). Both B. diandrus and B. rigidus have been reported from México (Espejo-Serna et al. 2000). Sales (1993) conducted a comprehensive survey of morphological, geographical and ecological variation in the B. diandrus / B. rigidus complex, and recognized these taxa as varieties: B. diandrus var. diandrus and B. diandrus var. rigidus. Sales (1993) noted the differences between the varieties are subtle and that it is often difficult to place specimens into one variety or the other. We recognize the taxon as B. diandrus, with no recognition of intraspecific taxa, as in other recent treatments in North America (Stubbendieck et al. 2003, Jessop et al. 2006, Pavlick & Anderton 2007, Saarela 2008, Saarela & Peterson 2012).

Bromus diandrus was not reported for Guatemala by Swallen & McClure (1955), but was reported for Guatemala by Soderstrom & Beaman (1968) and Pohl & Davidse (1994), based on the same collection (de Koninck 208) seen here for the country.

Specimens Examined:― GUATEMALA. Cerro El Baul, [14.83°N, 91.5°W], 2400 m, August 1954, M . de Koninck 208 (US-215372). MÉXICO. Baja California: 3.0 mi Eof Los Héroes de la Independencia, on road from Ensenada to San Felipe, 1040 m, 17 May 1979, J. R . Reeder & C. G. Reeder 7205 (SD-116088); Aguaje Vargas, Isla de Cedros, [28.19°N, 115.21°W], 600 m, 24 June 1977, L. M . Villareal de Puga 10641 (MEXU- 254480); Arroyo El Barbón, [32.05°N, 116.02°W], 6 May 1981, A . Preciado & M. Montoya 239 (MEXU- 1089759); arroyo mouth at El Descanso, 32.2042°N, 116.9125°W, 5 m, 27 March 1982, R . Moran 30105 (SD- 110757); Barranca de Salsipuedes, por la carreterra Tijuana-Ensenada, 32°N, 116.854°W, 28 April 1981, R . Guzmán M. 1258 (MEXU-1089755); ca. 1 mi Nof Camalú, 30.88°N, 116.0667°W, 120 m, 26 March 1979, J. R . Reeder & C. G. Reeder 7115 (SD-116041, US-2861043); camino entre el Sausal y San Antonio de las Minas, al N de Ensenada, 31.99°N, 116.58°W, 28 April 1981 R . Guzmán M. 1214 (MEXU); Cañon la Puerta, 6.0 km SW of Tecate, 32.5375°N, 116.6833°W, 500 m, 10 May 1980, R . Moran 28462 (MSC-276929, SD-105868); Cedros Island, Cerro de Cedros, 28.15°N, 115.2167°W, 1050 m, 16 April 1983, T . Oberbauer, H. Wier & E. Wier 46 (SD- 127652); Guadalupe Island, [29.03°N, 118.3°W], 19 April 1925, H. L . Mason 1543 (F-720871); Guadalupe Island, [29.03°N, 118.3°W], 24 April 1958, I. L . Wiggins & W. R. Ernst 77 (SD-48313); Guadalupe Island, [29.03°N, 118.3°W], 350 m, 30 April 1967, in bed of Arroyo Melpomene 2 mi from mouth, R . Moran 13773 (MSC-256754, RSA-POM-231537); Guadalupe Island, [29.03°N, 118.3°W], 3500 ft [1067 m], 15 June 1906, W. W . Brown Jr. 31 (GH); Guadalupe Island, Arroyo Melpomene, 28.9°N, 118.2833°W, 50 m, 8 June 2000, J. P . Rebman, T. Oberbauer & J. L. León de la Luz 6830 (SD-155061); Guadalupe Island, en route between NE Anchorage and springs, 29°N, 118.2667°W, 25 April 1958, I. L . Wiggins & W. R. Ernst 103 (US-2241595); Guadalupe Island, in arroyo bed, near mouth of Long Canyon, 29.0083°N, 118.2333°W, 40 m, 16 April 1970, R . Moran 17359 (SD-74870); Guadalupe Island, in bed of Arroyo Melpomene 2 mi from the mouth, 28.925°N, 118.2667°W, 350 m, 30 April 1967, R . Moran 13773 (SD-78963); Guadalupe Island, Nslope of island, slope below Hemizonia cliff, 29.1708°N, 118.3°W, 800 m, 24 May 1981, R. Moran 29595 (SD-108579); Guadalupe Island, NE Anchorage, 29.1625°N, 118.2833°W, 10 m, 30 April 1958, R . Moran 6758 (SD-47589); Guadalupe Island, near mouth of Long Canyon, [29.03°N, 118.3°W], 40 m, 16 April 1970, R . Moran 17359 (RSA-POM-364160); just Sof El Descanso, 32.1833°N, 116.8833°W, 10 m, 8 May 1977, R . Moran 24006 (SD-97268); La Rumorosa, km 65 carretera Mexicali-Tijuana, [32.6°N, 116.08°W], 1120 m, 27 April 1994, V . Jaramillo V., G. Villegas, A. Miranda 931 (MEXU-1089756); Los Coronados Islands, NE slope above Hotel Cove, S Island, 32.5792°N, 117.2458°W, 175 m, 7 May 1976, R . Moran 23126 (SD-95619); Mpio. Ensenada, "Cuesta del Lechero" km 35 carretera Ensenada-Ojos Negros, 31.9533°N, 116.3928°W, 700 m, 15 May 1997, L . Aragón M. 565 (MEXU-1089747); Mpio. Ensenada, a 22 km del Poblado de sitio Tomás 31.99°N, 116.58°W, 380 m, 8 April 1987, L . Elena López 127 (MEXU- 1110840); Mpio. Ensenada, Arroyo La Misión, Deleg. de Santa Rosa, [32.09°N, 116.85°W], 200 m, 12 March 1987, L . Elena López 7 (MEXU) & 9 (MEXU-1110796); Mpio. Ensenada, Cuenca del arroyo San Carlos, [29.61°N, 115.5°W], 30 April 1981, R . Guzmán 1315 (MEXU); Mpio. Ensenada, Delegación Francisco Zarco, Aguaje El Borreguero del rancho "El Dorado", 32.1369°N, 116.5331°W, 650 m, 14 May 1997, L . Aragón M. 551 (MEXU); Mpio. Ensenada, Ensenada, [31.87°N, 116.59°W], 10 m, 23 April 1979, M . Montoya 8 (MEXU- 1089758, MEXU-1017329); Mpio. Ensenada, Las Ánimas, 14 km al SE de Maneadero, 31.6333°N, 116.4833°W, 140 m, 7 May 1987, P . Tenorio L. & C. Romero de T. 1342 0 (MEXU-675500); Mpio. Ensenada, Coronel Esteban Cantu 14 km al SW de La Joya, 31.55°N, 116.6167°W, 100–570 m, 7 May 1987, P . Tenorio L. & C. Romero de T. 13362 (MEXU-801663, MEXU-837673); Mpio. Tecate, Santa Verónica, [32.46°N, 116.36°W], 930 m, 26 May

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Phytotaxa 185 (1) © 2014 Magnolia Press 75 1993, Jaramillo V ., Villegas G. & Domínguez O. 18 (MEXU-1089760); Rancho (solo) Sierra Blanca, Sierra Blanca, 32.075°N, 116.525°W, 675 m, 15 May 1976, R . Moran 23180 (SD-94914); Santa María Plains, 10–20 mi Sof Hamilton Ranch, 30.4°N, 115.8833°W, 8 April 1931, I. L . Wiggins 5200 (GH, MICH-1119285, NY, RSA-POM- 23502, US-1721755); Sierra Juárez, Wshore of Laguna Hanson, 32.0542°N, 115.075°W, 1610 m, 21 June 1980, R. Moran 28863 (SD-105481); Sierra San Pedro Mártir, Rancho Meling, [31.13°N, 115.19°W], 940 m, 27 May 1993, Jaramillo V ., Villegas G. & Domínguez O. 60 (MEXU); Sierra San Pedro Mártir, La Encantada, 30.9167°N, 115.4°W, 2200 m, 19 August 1967, R . Moran & R. F. Thorne 14326 (SD-76611); Sierra San Pedro Mártir, La Grulla, 30.8917°N, 115.4833°W, 2050 m, 9 June 1982, R . Moran 30938 (SD-111212); Mpio. Tecate, Juntas de Negi, 32.4922°N, 116.4606°W, 820 m, 12 May 1997, M. A . Vergara B. 85 (MEXU); between La Humarosa [Rumarosa] and Tecate, 32.53°N, 116.38°W, 27 April 1981, A. A . Beetle & R. Alcaraz M-6511 (ANSM, MICH- 1119217); Cañón de Agua Viva between Ensenada and Ojos Negros, [31.87°N, 116.62°W], 1 May 1981, A. A . Beetle & R. Alcaraz M. 6639 (MICH-1119218). Colima: Revillagigedo Islands, Guadalupe Island, 29°N, 118.2667°W, 19 April 1925, H. L . Mason 1543 (US-1319349). Estado de México: al oeste de Amecameca, por la carretera Amecameca-Cuautla, [19.13°N, 98.78°W], 2520 m, 8 February 1984, Manrique & Beetle 705- M (MEXU); Cuijingo (Delante de Juchitepec), [19.0839°N, 98.8517°W], 9 August 1994, B . Rodriguez C. s.n. (MEXU); Mpio. Amecameca, carretera Amecameca-Paso de Cortés, 19.0793°N, 98.719°W, 2785 m, 29 October 2000, H . Vibrans 6872 (MEXU); Mpio. Amecameca, San Pedro Nexapa, [19.09°N, 98.74°W], 2 October 1992, A . Miranda & G. Villegas 623 (MEXU); Mpio. Axapusco, Parque "Cerro Gordo" a 1.5 km al sur de la población de San Cristóbal Cuihuacan, 19.7606°N, 98.8256°W, 2740 m, 29 October 1999, R . Mendoza Domínguez 400 (MEXU); Mpio. Juchitepec, Cuijingo (adelante de Juchitepec), [19.0839°N, 98.8517°W], 9 August 1994, B . Rodriguez C. s.n. (ANSM); Mpio. Ocoyoacac, vía férrea al lado norte del pueblo, sobre cerro, cerca de la carretera Panorámica, 19.2756°N, 99.4599°W, 2692 m, 7 August 2001, H . Vibrans 7525 (MEXU-1029567); Mpio. Tenango de Arista, a 2.5 km al noroeste de San Fco. Tepexoxuca, [19.11°N, 99.6°W], 2865 m, 4 February 1993, A . Ramírez Abarca 488 (MEXU); Tenango del Aire 2 km saliendo, [19.17°N, 98.88°W], 1980 m, 27 September 1985, A . González & B. Rodríguez s.n. (MEXU-793444); 2.8 mi Eof Ocuilán Arteaga near cornfield, 18.9808°N, 99.3786°W, 2478 m, 11 October 2007, P. M . Peterson, J. M. Saarela & M. J. Flores Villegas 21371 (CAN, MO, US); 22.4 mi Eof junction of Hwys. 10 and 12 on Hwy. 12 towards Ixtapan de la Sal, 18.9301°N, 99.862°W, 2789 m, 10 October 2007, P.M. Peterson, J. M. Saarela & M. J. Flores Villegas 21357 (CAN, MO, US); 6.7 mi Eof Amecameca towards Paso de Cortés, 19.0706°N, 98.6923°W, 2488 m, 11 October 2007, P. M . Peterson, J. M. Saarela & M. J. Flores Villegas 21377 (CAN, MO, US); Mpio. Huitzilac, Rancho San Lorenzo km 53.5 de la carretera federal México-Acapulco (95), al SW del poblado Tres Marías, 19.65°N, 99.2333°W, 2660 m, 7 September 1988, I . Díaz V. 1158 (MEXU-508075). Morelos: km 24 carret. Ded. Cuernavaca-México, 19.06°N, 99.25°W, 1850 m, 29 March 1996, E . Oviedo 75 (MEXU); Mpio. Juchitepec, carretera Xochimilco-Oaxtepec 3 km desviación "Cicitec, [19.09°N, 99.02°W], 2000 m, 14 October 1993, J. A. Montaño 4 (MEXU); Mpio. Juchitepec, carretera Xochimilco-Oaxtepec 3 km desviación "Cicitec", [19.09°N, 99.02°W], 2000 m, 14 October 1993, J . Rosado 106 (MEXU). Oaxaca: 8.3 mi Nof San Cualimojoyas on road towards Santa María Yavesia, 17.1819°N, 96.4445°W, 2794 m, 20 September 2008, P. M . Peterson & J. M. Saarela 22318 (CAN, MO, US). Puebla: 1.2 mi W of Texmalaquila, just SE of Pico de Orizaba (Volcán Citlaltépetl), near Ebase of Sierra Negras, 18.9693°N, 97.3641°W, ± 1000 m, 3048–3353 m, 2 August 1977, E. Lehto, D. J. Pinkava, B. Parfitt & T. Reeves L-21976 (ASU-0010750); alpine zone of Pico de Orizaba (Volcán Citlaltépetl), 19.0331°N, 97.2728°W, ± 15000 m, 4267 m, 1 August 1977, T . Reeves, B. parfitt, E. Lehto & D.J. Pinkava R-5817 (ASU-0010749); 1 km al oeste de San Isidro Vaquerías, 3000 m, 27 June 1989, P . Dávila, P. Tenorio & J. Sánchez-Ken 358 (MEXU); 5.6 km SE of Ciudad Serdán on MEX Hwy. 140 to Tehuacán, [18.98°N, 97.44°W], 2540 m, 18 September 1990, P. M . Peterson & C. R. Annable 9956 (CAN, US); along the highway between Puebla and Córdoba near Orizaba near km 112 just Nof Puebla-Veracruz border, [18.83°N, 97.48°W], 2480 m, 24 February 1983, J. S . Miller & P. Tenorio L. 696 (MEXU- 776909); Ciudad Serdán, Guadalupe Sabinal, [18.98°N, 97.44°W], 19 February 2001, E . Gaspariano Martínez 115 (MEXU-1070140) & 116 (MEXU-1070141); Mpio. Atzizintla, El Mirador de las Cumbres de Maltrata, 29 km al O de Od. Mendoza, por la carretera a Puebla, [18.89°N, 97.32°W], 2500 m, 29 September 1984, S . D. Koch, J. Garciá, M. González & I. Hernández 8420 (F-2035538, MO). Morelos: a 10 km de Morelos Cañada rumbo a Ciudad Serdán y 3 km al Este Amazoquillo, 19.7606°N, 98.8256°W, 2360 m, 17 October 1995, A . Miranda, L. Aragón et al. 1084 (MEXU); Vertiente Sdel Pico de Orizaba, 1.0 km antes de Texmalaquilla por camino que viene de Atzitaintla, [18.98°N, 97.27°W], 14 September 1986, M. González-Kledesma & P. Vera C. 190 (SD-141347);

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Vertienta a Orizaba, supercarretera Puebla-Orizaba, 2300 m, 2 November 1972, W . Boege 2600 (GH). San Luis Potosí: San Luis Potosí, Virlet d'Aoust 1376 (P-02629888). Tlaxacala: Mpio. Terrenate, camino Zapata-Villarreal, [19.47°N, 97.921°W], 3000 m, 18 January 1990, L . Ordóñez 9 (MEXU) & 18 (MEXU). Veracruz: 0.8 mi Son the Orizaba / Puebla cuota (MEX 150) from its junction with the Veracruz / Puebla state line, [18.84°N, 97.29°W], 7700 ft [2346 m], 9 July 1990, S. & G. Jones 5323 (MICH-1119216); ca. 1 km Eof Puebla / Veracruz line, rte 150D, [18.84°N, 97.29°W], ca. 7000 ft [ca. 2133 m], 23 March 1970, R. W . Long & D. Burch 3081 (F-1952359); Mpio. Tehuipango, camino terracería Astacinga-Tehuipango, 18.525°N, 97.0556°W, 2400 m, 18 April 1997, H. R . Sandoval 467 (MEXU); supercarretera Puebla-Orizaba pendiente sur, [18.84°N, 97.29°W], 2500 m, 20 November 1972, L. W . Boege 2600 (MEXU) & 2609 (MEXU.