Revision of Muhlenbergia (Poaceae, Chloridoideae, Cynodonteae, Muhlenbergiinae) in Peru: classification, phylogeny, and a new species, M. romaschenkoi Author Peterson, Paul M. Author Vega, Isidoro Sanchez Author Romaschenko, Konstantin Author Giraldo-Canas, Diego Author Rodriguez, Nancy F. Refulio text PhytoKeys 2018 114 123 206 http://dx.doi.org/10.3897/phytokeys.114.28799 journal article http://dx.doi.org/10.3897/phytokeys.114.28799 1314-2003-114-123 CB70FFE82A46DE6DFFEBFFF8F43A6230 2533527 4 . Muhlenbergia cenchroides (Humb. & Bonpl. ex Willd.) P.M. Peterson, Caldasia 31(2): 280, f. 2 C-D . 2009. Fig. 3C, D Aegopogon cenchroides Humb. & Bonpl. ex Willd., Sp. Pl. 4(2):899. 1806. Type: Venezuela, Sucre, Cumana , F.W.H.A. Humboldt & A.J.A. Bonpland s.n. (holotype: B-W-01637-020 [image!]; isotypes: P!, US-75957 fragm. ex P!). Aegopogon geminiflorus Kunth, Nov. Gen. Sp. (quarto ed.) 1:133, t. 43. 1815 (1816). Aegopogon cenchroides var. geminiflorus (Kunth) Griseb., Abh. Koenigl . Ges. Wiss. Goettingen 24:301. 1879. Type: Venezuela, Amazonas, inter Cerro Duida et Rio Tamatama, juxta Esmeralda, May, F.W.H.A. Humboldt & A.J.A. Bonpland s.n. (holotype: P; isotype: US-75956 fragm. ex P!). Hymenothecium quinquesetum Lag., Gen. Sp. Pl. 4. 1816. Aegopogon quinquesetus (Lag.) Roem. & Schult., Syst. Veg. 1:805. 1817. Type: Mexico, Mexico Iperio, Ludovicus Nee (holotype: MA; isotype: BAA-00002156 [image!]). Hymenothecium trisetum Lag., Gen. Sp. Pl. 4. 1816. Aegopogon trisetus (Lag.) Roem. & Schult., Syst. Veg. 2:805. 1817. Aegopogon cenchroides var. trisetus (Lag.) E. Fourn., Mexic. Pl. 2:72. 1886. Type: Mexico, Mexico Imperio (holotype: MA; isotype: BAA-00002158 [image!]). Aegopogon setifer Nees, Linnaea 19(6):691. 1847. Type: Mexico, A. Aschenborn 132 (holotype: B; isotypes: FR-0036375 [image!], FR-0036376 [image!], US-75953 fragm. ex B!). Aegopogon cenchroides var. multisetus E. Fourn., Mexic. Pl. 2: 72. 1886. Type: Mexico, Moran. in rupibus, 1840, H. Galeotti 5808 (lectotoype: BR, designated by P.M. Peterson, Contr. U.S. Natl. Herb. 41: 10. 2001; isolectotypes: P!, US-75958 fragm. ex P!). Description. Caespitose perennials often sprawling, occasionally with stolons. Culms (10-) 25-55 cm long, glabrous below the nodes; internodes glabrous. Leaf sheaths mostly 0.8-8 cm long, shorter than the internodes, glabrous; ligules 1-2 mm long, apex acute, lacerate; blades 1.5-6 cm long, 0.5-2 mm wide, flat, scaberulous above, smooth beneath. Panicles 2-8 cm long, 0.5-1.2 cm wide, open, loosely-flowered with recemosely arranged branches; primary branches 2-4 mm long, excluding the awns, one per node, often purplish. Spikelet fasciles of three with one sessile or subsessile perfect spikelet and two short-pedicelled lateral spikelets staminate or sterile, the pedicels less than 0.2-0.5 mm long and the other two spikelets short-pedicelled, the pedicels about 0.7-1.2 mm long; glumes (1-)1.5-2.8 mm long, oblong and wider distally, 1-veined, apex deeply notched, awned, the awns 2-4 mm long, lobes triangular, acute; lemmas 2.5-3 mm long, fusiform, 3-awned, the central awns 5-13 mm long, lateral awns 2-3 mm long; paleas 2.5-3 mm long, puberulent, apex awned, the awns 1-2 mm long; anthers 1.6-1.8 mm long, yellowish to purplish. Caryopses 1-1.4 mm long, fusiform. 2 n = 40, 60, 80. Figure 5. A-D Muhlenbergia coerulea (Griseb.) Mez A habit B culm and panicle C ligule D spikelet E-K Muhlenbergia coerulea (Griseb.) Mez x Muhlenbergia rigida (Kunth) Kunth E culm and panicle F spikelet G floret H glumes I lemma J paleas, dorsal and ventral view K lodicules, stamens, and pistil. A-D drawn from P.M. Peterson, R.J. Soreng & J. Montoya Quino 21957 (US) E-K drawn from P.M. Peterson & N. Refulio Rodriguez 14860 (US). Distribution. Muhlenbergia cenchroides ranges from throughout Mexico , Central America to South America in Bolivia, Brazil, Colombia, Ecuador, Peru, Guyana and Venezuela. Ecology . Muhlenbergia cenchroides occurs on rocky slopes, canyons, cliffs, roadcuts, arroyos, seeps and meadows associated Baccharis spp., Festuca myuros , Alnus , Polylepis , Fucaria , Salvia spp., Urochloa , Salvia spp., Calceolaria spp., Eupatorium , Festuca , Puya , Rubus , Schizachyrium , Cenchrus clandestinus , Calamagrostis spp., Muhlenbergia spp., Hyptis , Oxalis , Aristida , Begonia , Adiantum, Bidens , Lepechinia , Oreocallis grandiflora (Lam.) R. Br., Desmodium , Cortaderia bifida Pilg., Cortaderia jubata (Lemoine ex Carriere) Stapf, Sporobolus , Monnina , Carex , Cheilanthes , Eragrostis , Lupinus , Lycopodium , Jarava , Nassella , Werneria , Agave , Eucalyptus , Thalictrum and Chusquea ; 1430-3850 m. Flowering December through September. Comments. Muhlenbergia cenchroides can be separated from M. bryophilus in having anthers 1.6-1.8 mm long, the perennial habit with (10-) 25-55 cm tall culms and sessile or inconspicuously pedicelled perfect spikelets usually associated with two staminate or sterile pedicelled spikelets ( Tovar 1993 ; Giraldo-Canas and Peterson 2009 ). Muhlenbergia cenchroides lies in a clade with M. tarahumara , M. uniseta and M. bryophilus in M. subg. Muhlenbergia (Fig. 1B ; Peterson et al. 2010b ). The multiple sampled individuals of M. cenchroides and M. uniseta in the current analyses do not resolve into reciprocally monophyletic lineages, but one sublineage, including accessions of each species, is strongly supported. This strongly suggests gene flow among individuals of these two species. Specimens examined. Peru. Ancash : Prov. Bolognesi, Paramarca, cerros al E de Chiquian , 3680 m, 8 Apr 1952, E. Cerrate 1396 (MOL, USM); Prov. Carhuaz, Cordillera Negra, 22 km SW of Shupluy and Rio Santa, 9°17'19.7"S , 77°44'39.3"W , 3770 m, 13 Mar 2008, P.M. Peterson 21652, R.J. Soreng, M.I. La Torre & J.V. Rojas Fox (US, USM); Prov. Corongo, Cordillera Blanca, 3 km E of Yanac, 8°35'55.6"S , 77°50'4.8"W , 2849 m, 18 Mar 2008, P.M. Peterson & R.J. Soreng 21786 (US, USM); Prov. Corongo, Cordillera Blanca, E of Yungay, 3400 m, 5 Apr 1988, S.A. Renvoize & S. Laegaard 5055 (CPUN, K); Prov. Huaraz, W side of Cordillera Blanca, S of Quebrada Ishinca, 9°24'55.8"S , 77°32'45.8"W , 2885 m, 12 Mar 2008, P.M. Peterson 21614, R.J. Soreng, M.I. La Torre & J.V. Rojas Fox (US, USM); Prov. Huaraz-Recuay border, W side of Rio Santa Canyon, 19 km S of Huaraz, 9°40'58.1"S , 77°28'29.7"W , 3202 m, 11 Mar 2008, P.M. Peterson 21600, R.J. Soreng, M.I. La Torre & J.V. Rojas Fox (US, USM); Prov Huaylas, Cordillera Blanca, encima de Caraz, 3200 m, A Weberbauer 3234 (MOL); Prov. Huaylas, Huata, M.I. La Torre 2793 (USM); Prov. Pallasca, 17 km N of Huandoval and 6 km S of Pallasca, 3360 m, 27 Mar 1997, P.M. Peterson & N. Refulio Rodriguez 13927 (US, USM); 3.4 km N of Huandoval towards Huascaschuque, 8°18'59.5"S , 77°58'9.5"W , 2949 m, 21 Mar 2008, P.M. Peterson & R.J. Soreng 21817 (US, USM); Prov. Huari, near Laguna de Puruhay, M.I. La Torre 3692 (USM); San Marcos, A. Cano 13505 (USM); Huascaran National Park, Quebrada Rurichinchay, D. Smith 12659, A. Gonzales & D. Maldonado (USM); Prov. Recuay, Cordillera Blanca, 16 km E of Catac on road towards Chavin de Huantar, 3630 m, 21 Mar 1997, P.M. Peterson & N. Refulio Rodriguez 13833 (US, USM); Prov. Yungay, Cordillera Blanca, 20 km ENE of Yungay, 9°6'26.2"S , 77°41'8.2"W 3247 m, 15 Mar 2008, P.M. Peterson 21712, R.J. Soreng, M.I. La Torre & J.V. Rojas Fox (US, USM); Quebrada Llauganuco, 24 km ENE of Yungay, 9°5'46.3"S , 77°40'19.3"W , 3309 m, 16 Mar 2008, P.M. Peterson 21739, R.J. Soreng, M.I. La Torre & J.V. Rojas Fox (US, USM); Prov. Yungay, Huascaran National Park, Llanganuco sector, Maria Josefa trail between Chinancocha and Pucayacu, 77°39'W , 9°05'S , 3700-3850 m, 7 May 1985, D.N. Smith 10512 (CPUN, MO). Ayacucho : Ayna, L. Aucasime 258 (USM); Seqseqa, L. Aucasime 81 (USM); Prov. Huanca Sancos, 27 km NW of Putajasa and 3 km S of Sacsamarca, 13°57'51.1"S , 74°18'41.5"W , 3650 m, 25 Feb 2002, P.M. Peterson 16278, A. Cano, M. I. La Torre, A. Ramirez & D. Susanibar Cruz (US, USM). Cajamarca : near Cajamarca, 2700 m, 11 Mar 1986, B, Becker & F.M. Terrones H. 601 (LPB); Cajamarca to Celendin , 95 km from Cajamarca, above Quillimbas, 3000 m, 22 Mar 1988, S.A. Renvoize & S. Laegaard 4868 (CPUN, K); Prov. Cajamarca, 16 km W of Central Plaza of Cajamarca up road towards Cumbemayo on Avenida Peru, 3440 m, 31 Mar 1997, P.M. Peterson & N. Refulio Rodriguez 14012 (US, USM); Prov. Cajamarca, Dist. Banos del Inca, Tres Tingos, margen Oeste del rio Chonta, 2900 m, 2 Mar 2002, I. Sanchez Vega 11261 (CPUN); Prov. Cajamarca, Cerro Huairapongo-Banos del Inca, 19 Nov 1965, I. Sanchez Vega 126 & Julio Mercado (CPUN); Prov. Cajamarca, Fundo Aylambo, a 5 km antes de la ciudad, 8 May 1969, I. Sanchez Vega 385 & W. Ruiz Vigo (CPUN); Prov. Cajamarca, cerca al Paso El Gavilan , 3100 m, 19 Apr 1975, I. Sanchez Vega 1407, P. Brandelard & J. Sanabria (CPUN); Prov. Cajamarca, Laguna de Moyococha, 2550 m, 29 Aug 1965, I. Sanchez Vega 92 (CPUN); Prov. Cajamarca, Dist. San Juan, lugar Huacraruco, 2350 m, 19 Jun 1993, J. Cabanillas S. & J. Guevara B. 615 (CPUN); Choten , en el Arboretum de CICAFOR, a 4 km de la carretera Pacasmayo-Cajamarca , 2990 m, 23 May 1981, I. Sanchez Vega 2562, V. Torrel & E. Medina (CPUN); Prov. Cajamarca, Dist. Cajamarca. Paso El Gavilan , 3100 m, 26 Jun 1966, I. Sanchez Vega 239 (CPUN); Prov. Cajamarca, ladera que converge al Valle Cajamarca, Lla Colpa, 2900 m, 22 Apr 1975, I. Sanchez Vega 1416, P. Brandelard, J, Sanabria & W. Ruiz Vigo (CPUN); Prov. Cajamarca, Sais Atahualpa de Porcon , en el Arboretum de CICAFOR, 3400 m, 16 May 1981, I. Sanchez Vega 2512, V. Torrel P. & E. Medina (CPUN); Prov. Cajamarca, Dist. Namora, Hacienda Polloquito, 3000 m, 6 May 1967, I. Sanchez Vega 321 (CPUN);12 km N of Cajamarca on road towards Bambamarca, 3020 m, 14 Mar 2000, P.M. Peterson & N. Refulio Rodriguez 14855 (CPUN, US, USM); S of Paso El Gavelan, 10 air km S of Cajamarca, 7°15'37.2"S , 78°30'34.5"W , 2544 m, 26 Mar 2008, P.M. Peterson 21875, R.J. Soreng & I. Sanchez Vega (US, USM); cumbre El Gavilan , R. Ferreyra 3255 (USM); Prov. Celendin, 4 air km W of Celendin, 6°54'13.5"S , 78°10'8.4"W , 2687 m, 29 Mar 2008, P.M. Peterson 21925, R. J. Soreng & J. Montoya Quino (US, USM); Prov. San Ignacio, 31 km E of Sondor on road towards Tabaconas, 2490 m, 31 Mar 2000, P.M. Peterson & N. Refulio Rodriguez 15128 (CPUN, US, USM); Prov. San Miguel, Niepos, 2200 m, 30 Oct 1985, S. Llatas 1503 (CPUN); Prov. San Pablo, Dist. San Pablo, lugar El Molino, 2200 m, 12 Jun 1993, J. Sanchez Vega 663 (CPUN). Cusco : Prov. Calca, Manto a Amparaes, 2650 m, 18 Apr 1966, C. Vargas C. 17393 (LPB); Prov. Cusco, Socorro-K'inka , C. Vargas C. 13161 (USM); Ruinas de Sacsahuaman , 3500 m, 6 May 1983, S.G. Beck 8365 (LPB); Mandor, abajo de Machupicchu, 2000 m, 13 Apr 1957, H. Ellenberg 988 (LPB); Cusco-Huancaro hwy, P. Gutte & G. Muller 9425a (USM); Machupicchu, camino entre la ciudad inca de Machupicchu y el Puente del Inca, 2000 m, 18 Sep 2004, D. Giraldo-Canas 3759 (COL). Huancavelica : Prov. Huancavelica, Huando, O. Tovar 1254 (USM). Huanuco : Prov. Huanuco , Carpish, cumbre entre Huanuco y Tingo Maria , 2700 m, 1 Oct 1950, R. Ferreyra 8070 (MOL); Prov. Huanuco , Carpish, cumbre entre Huanuco y Tingo Maria , R. Ferreyra 10022 (USM); 48.6 km NE of Huanuco towards Tingo Maria , T. Plowman & P.M. Rury s.n. (USM); Prov. Pachitea, 21 air km ENE of Huanuco , 9°51'53.1"S , 76°2'55.1"W , 2498 m, 6 Mar 2007, P.M. Peterson 20341, R.J. Soreng & K. Romaschenko (US, USM); Canyon of the Rio Grande, 44 air km E of Huanuco , P.M. Peterson 20363, R.J. Soreng & K. Romaschenko (US, USM). Junin : Prov. Huancayo, near Paian , O. Tovar 9311 (USM); Cerro al E de Huancayo, 3900 m, 2 May 1954, O. Tovar 2173 (US, USM); Prov. Tarma, entre Huancapistana, 2200 m, A. Weberbauer 2042 (MOL). La Libertad : Prov. Bolivar, entre el desvio a Uchumarca y Santa Luisa, 7°04'S , 77°49'W , 3700 m, 11 Nov 2001, I. Sanchez Vega 11194, M. Dillon & G. Iberico (CPUN); Prov. Bolivar, E side of Cerro Salumpuy, 8 air km NW of Bolivar, 7°7'39.2"S , 77°46'9.9"W , 3390 m, 30 Mar 2008, P.M. Peterson 21942, R. J. Soreng & J. Montoya Quino (US, USM); 3 air km ESE of Longotea on road to Bolivar, 7°3'27.9"S , 77°51'2.2"W , 3202 m, 31 Mar 2008, P.M. Peterson 21961, R. J. Soreng & J. Montoya Quino (US, USM). Lambayeque : Prov. Ferrenafe , Inkawasi, S. Llatas Quiroz 4122 (USM). Pasco : Prov. Oxapampa, Rio Boqueria, 26 km from Oxapampa via Rio Yamaquizu, D. Smith 1807, A. Pretel & L. Acosta (USM). Piura : Prov. Huancabamba, Porculla, 2200 m, 10 May 1992, S. Llatas Quiroz & H. De la Cruz S. 3098 (CPUN). San Martin : Prov. Mariscal Caceres , puerto de Norte de Rio Abiseo, A. Cano 7466, B. Young & J. Roque (USM).