Revision of Muhlenbergia (Poaceae, Chloridoideae, Cynodonteae, Muhlenbergiinae) in Peru: classification,
phylogeny, and a new species, M. romaschenkoi
Paul M. Peterson1, Isidoro Sánchez Vega†2, Konstantin Romaschenko1, Diego Giraldo-Cañas3, Nancy F. Refulio Rodriguez4
1 Department of Botany MRC-166, National Museum of Natural History, Smithsonian Institution, Washington, DC 20013-7012, USA 2 Departamento de Biología, Herbario CPUN, Universidad Nacional de Cajamarca Apartado 55, Cajamarca, Perú 3 Herbario Nacional Colombiano, Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogotá DC, Colombia 4 ResearchGate, Invalidenstraße 115, 10115 Berlin, Germany Corresponding author: Paul M. Peterson ([email protected])
Academic editor:Muthama Muasya | Received 2 August 2018 | Accepted 1 December 2018 | Published 31 December 2018 Citation: Peterson PM, Sánchez Vega I, Romaschenko K, Giraldo-Cañas D, Refulio Rodriguez NF (2018) Revision of Muhlenbergia (Poaceae, Chloridoideae, Cynodonteae, Muhlenbergiinae) in Peru: classification, phylogeny, and a new species, M. romaschenkoi. PhytoKeys 114: 123–206. https://doi.org/10.3897/phytokeys.114.28799
Abstract
A taxonomic treatment, phylogeny based on analysis of six DNA sequence markers (ITS, ndhA intron, rpl32-trnL, rps3, rps16 intron and rps16-trnK) and classification of Muhlenbergia for Peru is given. Sev- enteen species and one presumed hybrid are recognised. Muhlenbergia romaschenkoi sp. nov. is newly de- scribed from the Río Huallaga Valley, northeast of Huánuco. The type of Podosemum angustatum [≡ Muh- lenbergia angustata] clearly aligns with what we had been referring to as the hybrid between this species and M. rigida. Therefore, we adopt the next available heterotypic name, Muhlenbergia coerulea, for what we had been calling M. angustata and change the hybrid designation to M. coerulea × M. rigida. Lectotypes are designated for Epicampes coerulea Griseb., Muhlenbergia affinis Trin., Muhlenbergia berlandieri Trin., Muhlenbergia beyrichiana Kunth, Muhlenbergia elegans var. atroviolacea Kuntze, Muhlenbergia elegans var.
subviridis Kuntze and Muhlenbergia phragmitoides Griseb.
† Deceased
http://phytokeys.pensoft.net
Copyright Paul M. Peterson et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Launched to accelerate biodiversity research
Resumen
Brindamos un tratamiento taxonómico, una filogenia basado en el análisis de seis marcadores de secuencia de ADN (ITS, intrón ndhA, rpl32-trnL, rps3, intrón rps16, rps16-trnK) y la clasificación de Muhlenbergia para Perú. Se reconocen diecisiete especies y un supuesto híbrido. Muhlenbergia romaschenkoi sp. nov. es descrita como especie nueva procedente del valle de Huallaga, al noreste de Huánuco. El tipo de Podosemum angustatum [≡ Muhlenbergia angustata] se alinea claramente con lo que nosotros habíamos referido como el híbrido entre esta especie y M. rigida. Por lo tanto, adoptamos el siguiente nombre heterotípico disponible, Muhlenbergia coerulea para lo que habíamos estado llamando M. angustata, y cambiamos la designación híbrida a M. coerulea × M. rigida. Los lectotipos son designados para Epicampes coerulea Griseb., Muhlen- bergia affinis Trin., Muhlenbergia berlandieri Trin., Muhlenbergia beyrichiana Kunth, Muhlenbergia elegans var. atroviolacea Kuntze, Muhlenbergia elegans var. subviridis Kuntze y Muhlenbergia phragmitoides Griseb.
Keywords
classification, ITS, lectotypification, Muhlenbergia, Peru, phylogeny, plastid DNA sequences, Poaceae, systematics, taxonomy
Introduction
Peru is located in the central and western part of South America between 0° and 18°
south latitude. The central Andes or backbone divides Peru into three great regions:
coast, mountain, and forest (Weberbauer 1936). The coastal region next to the Pacific Ocean is an extremely xeric, narrow, sandy plain that rises abruptly to the east. This region is occasionally transected by river valleys and the climate is very mild and warm.
The mountain region or ‘‘cordillera de Andes’’ consists of high peaks and extensive high plains (altiplano) that are sometimes dissected by deep, narrow valleys. The climate in this region can be very cold and it is not uncommon to have snow above 3800 m. The tropical forest region, east of the cordillera, extends into the Amazonian river basin where there are high levels of biodiversity associated with many different habitats. The climate here is very warm and humid with high levels of precipitation and there are large rivers including the Río Marañon and the Río Ucayali that feed into the Río Amazonas.
The Poaceae is a diverse family occupying a myriad of terrestrial habitats in Peru, with the exception of the permanent snow fields. The taxonomic knowledge of this family in Peru is incomplete and it is still possible to encounter novel species.
Hitchcock (1927), in his treatment of the grasses of Ecuador, Peru and Bolivia, recognised 12 species of Muhlenbergia Schreb., 10 of these being reported in Peru.
Standley (1936), in his Flora of Peru, recorded 10 species. More recently, Tovar (1993) produced an integral study of the Peruvian grasses, including 14 species of Muhlen- bergia. Brako and Zarucchi (1993) list 13 species as being present in Peru. All of these treatments are out of date and, subsequently, there have been new reports of grasses from Peru. As Muhlenbergia is a relatively large genus in North and Central America and is easily dispersed, a modern investigation was undertaken to more clearly under- stand its systematics and biogeography. In Peru, we report 17 species of Muhlenbergia (one of these is new) and one proposed hybrid.
The subtribe Muhlenbergiinae Pilg. (tribe Cynodonteae Dumort.) is a diverse as- semblage of 182 species represented by a single, monophyletic genus, Muhlenbergia (Peterson et al. 2010a, b; 2016; Soreng et al. 2017; Peterson et al. 2018). Species within Muhlenbergia are morphologically highly variable and are characterised in having mem- branous ligules (rarely a line of hairs); paniculate inflorescences that are rebranched or composed only of primary branches; spikelets that are usually solitary but sometimes in pairs or triads, with cleistogenes (self-pollinated flowers that do not open at matu- rity) occasionally present in the leaf sheaths; one floret (rarely more) per spikelet that is perfect, staminate or sterile; glumes that are awned or unawned; lemmas 3-nerved, apex awned or unawned; and a base chromosome number of x = 8−10 (Peterson et al.
1995, 1997, 2007a, b; Peterson 2000, 2003). Two subtypes of C4 photosynthesis based on nicotinamide adenine dinucleotide cofactor malic enzyme (NAD-ME) and phos- phoenolpyruvate carboxykinase (PCK) have been found in Muhlenbergia; subtypes in most species have been verified by anatomy and in a few species by biochemical assay (Gutierrez et al. 1974; Brown 1977; Hattersley and Watson 1992).
Based on seven molecular markers (nuclear ITS and plastid ndhA intron, ndhF, rps16-trnK, rps16 intron, rps3 and rpl32-trnL), Peterson et al. (2010b) provided a phy- logeny and classification for 124 species (68%) of the Muhlenbergiinae. They recog- nised five subgenera within Muhlenbergia: M. subg. Bealia (Scribn.) P.M. Peterson, M.
subg. Clomena (P. Beauv.) Hack., M. subg. Muhlenbergia, M. subg. Pseudosporobolus (Parodi) P.M. Peterson and M. subg. Trichochloa (P. Beauv.) A. Gray. Here we present an updated phylogeny of 146 species (80%) of Muhlenbergia with 13 new species based on six molecular markers (nuclear ITS and plastid ndhA intron, rpl32-trnL, rps3, rps16 intron and rps16-trnK), classification and a taxonomic revision of the genus for Peru.
Material and methods Phylogenetic analyses
Detailed methods for DNA extraction, amplification, sequencing and phylogenetic analysis are given in Peterson et al. (2010a, b, 2014a, 2015a, b, 2016). In brief, the phylogeny was estimated among members of Muhlenbergia based on the analysis of six molecular markers (nuclear ITS 1&2, and plastid ndhA intron, rpl32-trnL, rps3, rps16 intron, rps16-trnK). We sampled most species within subtribe Muhlenbergiinae and included outgroups: Distichlis scoparia (Nees ex Kunth) Arechav. (Monanthochloinae Pilg. ex Potztal), Willkommia sarmentosa Hack. (Traginae P.M. Peterson & Colum- bus) and Sporobolus indicus L. (Zoysieae Benth., Sporobolinae Benth.) [Peterson et al.
2010a, 2015a, 2014b, 2016; Soreng et al. 2017]. Voucher information and GenBank numbers for all samples (including the new ones) are given in Table 1.
The resulting plastid and ITS topologies were inspected for conflicting nodes with posterior probabilities (PP) ≥ 0.95. If no supported conflict was found, plastid and
ITS sequences were combined (Figure 1). When conflicting topologies were present, the datasets for inconsistently placed taxa were duplicated in the matrix. One set of the taxon was represented by only the corresponding plastid sequences, the other taxon set by only ITS sequences. The remaining positions for the duplicated datasets were then coded as missing data. We use this “taxon duplication” approach (Pirie et al.
2008; Pelser et al. 2010; Peterson et al. 2015b, 2016) to resolve our phylogenetic tree, minimising the diffusing effects of taxa with strongly supported incongruence between the plastid and ITS data and to represent their alternative placements in relation to the remaining phylogenetic groups, among which relationships are congruent.
Taxonomy
Specimens of Muhlenbergia from Peru were reviewed in the following herbaria: AAU, COL, CPUN, CUZ, F, HAO, K, LIL, LPB, MO, MOL, NY, US and USM. The first author (PMP) travelled and collected 3042 numbers (primarily grasses) while on seven expeditions to Peru between 1997 and 2008. Of these, 258 are collections of Muhlenber- gia and all are accounted for here. Synonymy is mainly limited to names of taxa described from South America. Additional synonyms accepted by us can be found in the Catalogue of New World Grasses, vol. II (Peterson et al. 2001) and on the Catalogue of New World Grasses web site (http://www.tropicos.org/Project/CNWG) that is continually updated within TROPICOS (http://www.tropicos.org). When counting culm nodes, it is best to start counting 1 cm above the base. Blade width is measured from margin to margin on a flat blade but not when the blade is tightly involute. Glabrous refers to without pu- bescence. Smooth indicates no prickle-hairs with broad bases and/or hooked or pointed apices (i.e. pubescence can occur on a smooth surface and a scabrous surface can be glabrous). Excluded species and an infrageneric classification of the accepted species of Muhlenbergia in Peru are presented at the end of the taxonomic treatment.
Results and discussion Phylogeny
A total of 226 new sequences from 146 species of Muhlenbergia are newly reported in GenBank (Table 1). The following 13 species are new in the phylogeny of Muhlenbergia:
M. capillipes (M.E. Jones) P.M. Peterson & Annable, M. ligularis (Hack.) Hitchc., M.
flabellata Mez, M. bushii R.W. Pohl, M. diversiglumis Trin., M. frondosa (Poir.) Fernald, M. glabrifloris Scribn., M. romaschenkoi P.M. Peterson, M. setaroides E. Fourn., M. syl- vatica (Torr.) Torr. ex A Gray, M. breviligula Hitchc., M. lehmanniana Henrard and M.
venezuelae Luces (Peterson et al. 2010b; 2018). Total aligned characters for individual regions and other parameters are noted in Table 2. The plastid−ITS sequences were com- bined in the analysis except where incongruence between datasets was detected (Fig. 1).
Figure 1. A, B Maximum-likelihood tree inferred from combined plastid (ndhA intron, rps16-trnK, rps16 intron, rps3, and rpl32-trnL) and ITS sequences. Thick branches indicate posterior probabilities of 0.95−1; species in red occur in South America; scale bar = 2%.
The Bayesian tree from the combined analysis of ITS and five plastid regions (ndhA intron, rpl32-trnL, rps3, rps16 intron and rps16-trnK) is well resolved with strong sup- port for the monophyly of Muhlenbergia, including M. ramulosa (Kunth) Swallen, sister to M. subg. Bealia−M. subg. Trichochloa, these all being in one clade that is sister to M. subg. Clomena−M. subg. Pseudosporobolus and sister to M. subg. Muhlenbergia.
(Fig. 1; posterior probability, PP = 0.95−1, shown with thick branches). Each of the five major clades, corresponding to the five subgeneric divisions that have been rec- ognised previously (Peterson et al. 2010b), include species that occur in Peru and/or South America (red) or as wide ranging species from the Americas. The species in each of these five subgenera share salient morphological characteristics or trends.
Species of M. subg. Bealia are strongly caespitose, never rhizomatous, annuals or perennials with pubescent margins or midnerves at least on the lower ½ of the lemma (only M. ligularis is without pubescence) and round, equal primary, secondary and tertiary vascular bundles without well-developed schlerenchyma (Peterson and Herrera Arrieta 2001; Peterson 2003; Peterson et al. 2010b). The Peruvian endemic, M. cax- amarcensis Lægaard & Sánchez Vega and the wide-ranging, M. ligularis (Central and South America), are members of this subgenus.
Figure 1. Continued.
Although the clade of species representing the M. subg. Trichochloa is strongly sup- ported in our analyses (Fig. 1), there is little resolution among members, suggesting very low levels of genetic divergence among the species in this subgenus. The low level of divergence may be a consequence of rapid speciation events. Within Muhlenbergia, this group is by far the most difficult to determine because there are very few morpho- logical differences among the taxa and discrete (nonplastic) characteristics are few. The species of M. subg. Trichochloa consist of robust perennials up to 3 metres tall with compressed-keeled or rounded basal sheaths, 1-veined glumes and unequal rectangular or obovate/elliptic secondary and tertiary vascular bundles with well-developed scle- renchyma girders, these usually with sclerosed phloem (Peterson and Herrera Arrieta 2001; Herrera Arrieta and Peterson 2007, 2017, 2018; Peterson et al. 2010b). In Peru, M. coerulea (Griseb.) Mez, M. coerulea×M. rigida, M. maxima Lægaard & Sánchez Vega (endemic) and M. rigida (Kunth) Kunth are placed in M. subg. Trichochloa.
Table 1. Taxon voucher (collector, number, and where the specimen is housed), country of origin, and GenBank accession for DNA sequences of rps3, rps16-trnK, rps16 intron, ndhA intron, rpl32-trnL, and ITS regions (bold indicates new accession); a dash (−) indicates missing data; an asterisk (*) indicates sequences not generated in our lab. NTaxonVoucherCountryrps3rps16-trnKrps16 intronndhA intronrpl32-trnLITS Outgroup 1Distichlis scoparia var. erinacea (Nees ex Kunth) Arechav.Peterson 17475, Soreng & Refulio-Rodriguez (US)
Argentina, NGU360034GU360501GU360477GU359480GU359803GU359334 euquen Mexico, 2Sporobolus indicus (L.) R. Br.Peterson 22025 & Saarela (US)GU360161GU360630GU360355GU359504GU359913GU359209 Chihuahua 3Willkommia sarmentosa Hack.Schweickerdt 2181 (US)South AfricaGU360252GU360645GU360343GU359545GU359924GU359194 Muhlenbergia Muhlenbergia ramulosa (Kunth) Mexico, 4Peterson 22447 & Saarela (US)GU360254GU360717GU360406GU359444GU359953GU359115 SwallenDurango Muhlenbergia ramulosa (Kunth) 5 Swallen
Peterson 24521 & Romaschenko (US)
Mexico, CoahuilaMK090816MK090876MK090848MK090728MK090773MK090686 6Muhlenbergia ramulosa (Kunth) SwallenPeterson 24697, Romaschenko & Zamudio Ruíz (US)Mexico, QueretaroMK090817MK090877MK090849MK090729MK090774MK090687 subg. Bealia 7Muhlenbergia annua (Vasey) SwallenPeterson 22022 & Saarela (US)Mexico, ChihuahuaHM143247HM143629HM143534HM143351HM143144HM143043 8Muhlenbergia arenicola BuckleyPeterson 19947 & Lara- Contreras (US)Mexico, CoahuilaGU360209GU360674GU360413GU359462GU359960GU359166 9Muhlenbergia argentea VaseyPeterson 22095 & Saarela (US)Mexico, ChihuahuaHM143248HM143630HM143535HM143352HM143145HM143044 10Muhlenbergia arizonica Scribn.Peterson 22173 & Saarela (US)Mexico, ChihuahuaHM143249HM143631HM143536HM143353HM143146HM143045 11Muhlenbergia biloba Hitchc.Peterson 7946, Annable & Herrera (US)Mexico, ChihuahuaGU360098GU360550GU360309–GU359859GU359258 12Muhlenbergia brevis C.O. Goodd.Peterson 22023 & Saarela (US)Mexico, ChihuahuaHM143253HM143635HM143540HM143357HM143150HM143049 13Muhlenbergia capillipes (M.E. Jones) P.M. Peterson & AnnablePeterson 20013 & Sánchez Alvarado (US)Mexico, DurangoMK090802MK090862MK090835MK090714MK090751MK090664 14Muhlenbergia caxamarcensis Lægaard & Sánchez VegaPeterson 21965, Soreng & Montoya Quino (US)Peru, La LibertadHM143256HM143639HM143543HM143360HM143153HM143052
NTaxonVoucherCountryrps3rps16-trnKrps16 intronndhA intronrpl32-trnLITS 15Muhlenbergia depauperata Scribn.Peterson 21293, Saarela & Flores Villegas (US)Mexico, ZacatecasHM143261HM143642HM143547HM143363HM143157HM143056 16Muhlenbergia eludens C. ReederPeterson 22188 & Saarela (US)Mexico, SinaloaHM143268HM143649HM143554HM143370HM143164HM143062 17Muhlenbergia filiformis (Thurb. ex S. Watson) Rydb.Peterson 10433, Annable & Weinpahl (US)USA, CaliforniaHM143272HM143653HM143558HM143374HM143168HM143066 18Muhlenbergia flavida VaseyPeterson 22237 & Saarela (US)Mexico, SinaloaHM143273HM143654HM143559HM143375HM143169HM143067 19Muhlenbergia fragilis SwallenPeterson 22194 & Saarela (US)Mexico, SinaloaHM143275HM143656HM143561HM143377HM143171HM143069 20Muhlenbergia ligularis (Hack.) Hitchc.
Peterson 12684, Annable, Lægaar
d & Soreng (US)Bolivia, La PazMK090808MK090868MK090842MK090720MK090760MK090674 21Muhlenbergia ligularis (Hack.) Hitchc.
Peterson 12792, Annable, Lægaar
d & Soreng (US)Bolivia, OruroMK090809MK090869MK090843MK090721MK090761MK090675 22Muhlenbergia ligularis (Hack.) Hitchc.Peterson 21513, Soreng, Torre & Rojas Fox (US)Peru, AncashMK090810MK090870MK090844MK090722MK090762MK090676 23Muhlenbergia ligularis (Hack.) Hitchc.Peterson 21912, Soreng & Montoya Quino (US)Peru, CajamarcaMK090811MK090871MK090845MK090723MK090763MK090677 24Muhlenbergia majalcensis P.M. PetersonPeterson 4519 & Annable (US)Mexico, ChihuahuaHM143295HM143675HM143577HM143394HM143190HM143088 25Muhlenbergia minutissima (Steud.) SwallenPeterson 22012 & Saarela (US)Mexico, ChihuahuaHM143300HM143680 − −HM143195HM143093 26Muhlenbergia minutissima (Steud.) Swallen
Peterson 24556 & Romaschenko (US)
Mexico, CoahuilaMK090813MK090873 − −MK090765MK090679 27Muhlenbergia shepherdii (Vasey) SwallenPeterson 22452 & Saarela (US)Mexico, DurangoGU360102GU360560GU360320GU359419GU359854GU359277 28Muhlenbergia sinuosa SwallenPeterson 7976, Annable & Herrera Arrieta (US)Mexico, ChihuahuaHM143323HM143704HM143603HM143419HM143219HM143117 29Muhlenbergia texana BuckleyPeterson 22016 & Saarela (US)Mexico, ChihuahuaHM143334HM143714HM143614 −HM143230HM143128 30Muhlenbergia torreyi (Kunth) Hitchc. ex BushPeterson 19429, Soreng, Salariato & Panizza (US)
Argentina, Catamar
caGU360266GU360720GU360267MK090744GU359992GU359118 31Muhlenbergia tricholepis (Torr.) ColumbusPeterson 22099 & Saarela (US)Mexico, ChihuahuaGU360103GU360559GU360305GU359418GU359853GU359278 32Muhlenbergia vaginata SwallenPeterson 22417 & Saarela (US)Mexico, Durango − HM143718HM143618MK090745HM143234MK090708
NTaxonVoucherCountryrps3rps16-trnKrps16 intronndhA intronrpl32-trnLITS subg. Clomena 33Muhlenbergia crispiseta Hitchc.Peterson 10768, Annable & Valdes Reyna (US)HM143258 − − − − − 34Muhlenbergia curvula SwallenPeterson 24830, Romaschenko, Rodriguez Avalos, Herrera- Simoni & Garcia Rodriguez (US)
Mexico, Aguascalientes − − − − MK090754MK090667 35Muhlenbergia flaviseta Scribn.Peterson 22409 & Saarela (US)Mexico, DurangoGU360250GU360685GU360410GU359448GU359957GU359127 36Muhlenbergia flaviseta Scribn.Peterson 24923, Romaschenko & González-Elizondo (US)Mexico, Durango − − MK090839 − MK090757MK090670 37Muhlenbergia filiculmis VaseyPeterson 11954 & Annable (US)USA, WyomingHM143271HM143652HM143557HM143373HM143167HM143065 38Muhlenbergia flabellata MezSenalobo 1985 (MO)Costa Rica, MK090805MK090865MK090838MK090717MK090756MK090669 39Muhlenbergia jonesii (Vasey) Hitchc.Peterson 4861 & Annable (US)USA, CaliforniaHM143289HM143669 − HM143390HM143185HM143083 40Muhlenbergia montana (Nutt.) Hitchc.Peterson 22046 & Saarela (US)Mexico, ChihuahuaHM143301HM143681HM143582HM143398HM143196HM143094 41Muhlenbergia peruviana (P. Beauv.) Steud.Peterson 22440 & Saarela (US)Mexico, DurangoGU360221GU360713GU360408GU359446GU359955GU359154 42Muhlenbergia peruviana (P. Beauv.) Steud.Peterson 24696, Romaschenko & Zamudio Ruíz (US)Mexico, Querétaro − − − − MK090767MK090680 43Muhlenbergia peruviana (P. Beauv.) Steud.Peterson 24721, Romaschenko & Zamudio Ruíz (US)Mexico, Querétaro − − − − MK090768MK090681 44Muhlenbergia peruviana (P. Beauv.) Steud.Peterson 24943, Romaschenko & Zamudio Ruíz (US)Mexico, Querétaro − − − − MK090769MK090682 45Muhlenbergia quadridentata (Kunth) Trin.Peterson 16103 & Rosales (US)Mexico, JaliscoHM143313HM143694 −HM143410HM143209HM143107 46Muhlenbergia straminea Hitchc.Peterson 15238 & Cayouette (US)USA, ArizonaHM143327HM143707HM143607MK090737HM143223HM143121 47Muhlenbergia virescens (Kunth) Trin.Peterson 22412 & Saarela (US)Mexico, DurangoHM143342HM143722HM143621HM143432HM143238HM143134
NTaxonVoucherCountryrps3rps16-trnKrps16 intronndhA intronrpl32-trnLITS subg. Muhlenbergia 48Muhlenbergia alamosae VaseyPeterson 22104 & Saarela (US)Mexico, ChihuahuaHM143243HM143626HM143530HM143347HM143140HM143039 49Muhlenbergia andina (Nutt.) Hitchc.Peterson 10432, Annable & Weinpahl (US)USA, CaliforniaHM143244 − HM143531HM143348HM143141HM143040 50Muhlenbergia appressa C.O. Goodd.Peterson 4183 & Annable (US)USA, ArizonaGU360211GU360676GU360415GU359443GU359962GU359164 51Muhlenbergia arsenei Hitchc.Peterson 15208 & Cayouette (US)Mexico, Baja California NorteHM143250HM143632HM143537HM143354HM143147HM143046 52Muhlenbergia beyrichiana KunthPeterson 20366, Soreng & Romaschenko (US)Peru, HuánucoGU360247GU360712GU360280GU359493GU359995GU359129 53Muhlenbergia brandegeei C. ReederPeterson 4760 & Annable (US)Mexico, Baja California SurGU360208GU360711GU360412GU359450GU359959GU359167 54Muhlenbergia breviseta Griseb. ex E. Fourn.McVaugh 22930 (US)Mexico − HM143636 − − − − 55Muhlenbergia bryophilus (Döll) P.M. PetersonColumbus 3565 (RSA)Peru, Cusco− − − − −GQ397862* 56Muhlenbergia bushii R.W. PohlBrant 4542 & O’Donnell (MO)USA, MissouriMK090801MK090861MK090834MK090713MK090750MK090663 57Muhlenbergia californica VaseyPeterson 5013 & Barron (US)USA, CaliforniaHM143254HM143637HM143541HM143358HM143151HM143050 58Muhlenbergia cenchroides (Humb. & Bonpl. ex Willd.) P.M. PetersonPeterson 22045 & Saarela (US)Mexico, ChihuahuaGU360143GU360578GU360274GU359403GU360011GU359259 59Muhlenbergia cenchroides (Humb. & Bonpl. ex Willd.) P.M. PetersonPeterson 24688, Romaschenko & Zamudio Ruíz (US)Mexico, Querétaro − − − − MK090752MK090665 60Muhlenbergia ciliata (Kunth) Trin.Peterson 22193 & Saarela (US)Mexico, SinaloaHM143257HM143640HM143544HM143361HM143154HM143053 61Muhlenbergia curtifolia Scribn.Peterson 5631 & Annable (US)USA, ArizonaHM143259HM143641HM143545HM143362HM143155HM143054 62Muhlenbergia diversiglumis Trin.Peterson 4611 & Annable (US)Mexico, ChihuahuaMK090804MK090864MK090837MK090716MK090755MK090668 63Muhlenbergia dumosa Scribn. ex VaseyPeterson 13438, Knowles, Dietrich & Braxton (US)Mexico, DurangoHM143265HM143646HM143551HM143367HM143161HM143059
NTaxonVoucherCountryrps3rps16-trnKrps16 intronndhA intronrpl32-trnLITS 64Muhlenbergia flexuosa Hitchc.Peterson 20373, Soreng & Romaschenko (US)Peru, HuánucoHM143274HM143655HM143560HM143376HM143170HM143068 65Muhlenbergia frondosa (Poir.) FernaldYatskievych 04-224, Smith, McKenzie & Nagel (MO)USA, MissouriMK090806MK090866MK090840MK090718MK090758MK090671 66Muhlenbergia glabrifloris Scribn.Brant 2659 (MO)USA, MissouriMK090807MK090867MK090841MK090719MK090759MK090672 67Muhlenbergia glauca (Nees) B.D. Jacks.Peterson 21180, Saarela, Gonzalez Elizondo, Rosen & Reid (US)Mexico, DurangoHM143277HM143659HM143564HM143380HM143174HM143071 68Muhlenbergia glomerata (Willd.) Trin.Peterson 20924, Saarela & Howard (US)USA, New YorkGU360253GU360716GU360407GU359445GU359954GU359114 69Muhlenbergia hakonensis (Hack. ex Matsum.) MakinoKim 2009858Korea (South) − − − − − HQ600507* 70Muhlenbergia himalayensis Hack. ex Hook. f.Soreng 5666, Peterson & Sun Hang (US)China, XizangHM143281HM143662HM143566HM143383HM143177HM143075 71Muhlenbergia huegelii Trin.Soreng 5344, Peterson & Sun Hang (US)China, SichuanHM143282HM143663HM143567HM143384HM143178HM143076 72Muhlenbergia japonica Steud.Soreng 5240, Peterson & Sun Hang (US)China, YunnanHM143287HM143667HM143571HM143388HM143183HM143081 73Muhlenbergia mexicana (L.) Trin.Ladd 13851 (US)USA, MissouriMK090812MK090872MK090846 − MK090764MK090678 74Muhlenbergia mexicana (L.) Trin.Peterson 5562 & Annable (US)USA, New MexicoHM143297HM143677HM143580HM143396HM143193HM143090 75Muhlenbergia mexicana var. filiformis (Torr.) Scribn.Peterson 20861 & Saarela (US)USA, New York.HM143298HM143678HM143579HM143395HM143192HM143091 76Muhlenbergia microsperma (DC.) KunthPeterson 21855 & Soreng (US)Peru, CajamarcaHM143299HM143679HM143581HM143397HM143194HM143092 77Muhlenbergia pauciflora BuckleyPeterson 22048 & Saarela (US)Mexico, Chihuahua − HM143686HM143587HM143403HM143201HM143099 78Muhlenbergia pectinata C.O. Goodd.Peterson 22108 & Saarela (US)Mexico, SonoraHM143306HM143687HM143588HM143404HM143202HM143100 79Muhlenbergia pereilema P.M. PetersonPeterson 22191 & Saarela (US)Mexico, SinaloaGU360245GU360710GU360282GU359519GU359993GU359131 80Muhlenbergia plumiseta ColumbusPeterson 20106, Hall, Alvarez Marvan & Alvarez Jimenez (US)Mexico, GuerreroGU360246GU360719GU360281GU359516GU359979GU359130
NTaxonVoucherCountryrps3rps16-trnKrps16 intronndhA intronrpl32-trnLITS 81Muhlenbergia polycaulis Scribn.Peterson 22092 & Saarela (US)Mexico, ChihuahuaHM143307HM143688HM143589MK090727HM143203HM143101 82Muhlenbergia porteri Scribn. ex BealPeterson 19846 & Lara Contreras (US)Mexico, CoahuilaHM143308HM143689HM143590HM143405HM143204HM143102 83Muhlenbergia ramosa (Hack. ex Matsum.) MakinoSoreng 5302, Peterson & Sun Hang (US)China, YunnanHM143314HM143695HM143595HM143411HM143210HM143108 84Muhlenbergia romaschenkoi P.M. PetersonPeterson 20331, Soreng & Romashchenko (US)Peru, Huánuco––−−MK090784MK096270 85Muhlenbergia schreberi J.F. Gmel.Peterson 19443, Soreng, Salariato & Panizza (US)
Argentina, TGU360214GU360679GU360404GU359456GU359950GU359161 ucuman Muhlenbergia setarioides 86Peterson 9897 & Annable (US)Mexico, OaxacaMK090823MK090883MK090853MK090736MK090785MK090697 E. Fourn. Muhlenbergia sobolifera (Muhl. ex 87Peterson 20834 & Saarela (US)USA, VirginiaHM143324MK090884HM143604HM143420HM143220HM143118 Willd.) Trin. 88Muhlenbergia spiciformis Trin.Peterson 22362 & Saarela (US)Mexico, OaxacaHM143326HM143706HM143606HM143422HM143222HM143120 Peterson 24580, Romaschenko Mexico, 89Muhlenbergia spiciformis Trin.−−−−MK090786MK090698 & Valdés-Reyna (US)Coahuila Muhlenbergia sylvatica (Torr.) 90Brant 4783 (MO)USA, MissouriMK090824MK090885MK090854MK090738MK090787MK090699 Torr. ex A. Gray Muhlenbergia tarahumaraMexico, 91Peterson 22053 & Saarela (US)HM143330HM143710HM143610HM143425HM143226HM143124 P.M. Peterson & ColumbusChihuahua Muhlenbergia tenella (Kunth) Mexico, 92Peterson 22141 & Saarela (US)HM143331HM143711HM143611 −HM143227HM143125 Trin.Chihuahua Muhlenbergia tenuiflora (Willd.) 93Peterson 15778 & Saarela (US)USA, Virginia.HM143332HM143712HM143612HM143426HM143228HM143126 Britton, Sterns & Poggenb. Muhlenbergia tenuifolia (Kunth) 94Peterson 22344 & Saarela (US)Mexico, OaxacaHM143333HM143713HM143613HM143427HM143229HM143127 Kunth Muhlenbergia tenuifolia (Kunth) Peterson 24462, Romaschenko Mexico, 95 − − − − MK090788MK090700 Kunth& Valdés-Reyna (US)Coahuila Muhlenbergia tenuifolia (Kunth) 96 Kunth
Peterson 24471 & Romaschenko (US)
Mexico, Coahuila − − − − MK090789MK090701 97Muhlenbergia tenuifolia (Kunth) Kunth
Peterson 24673 & Romaschenko (US)
Mexico, San Luis PotosíMK090825MK090886 − MK090739MK090790MK090702 98Muhlenbergia tenuifolia (Kunth) KunthPeterson 24730, Romaschenko & Zamudio Ruíz (US)Mexico, QueretaroMK090826MK090887 − MK090740MK090791MK090703
NTaxonVoucherCountryrps3rps16-trnKrps16 intronndhA intronrpl32-trnLITS 99Muhlenbergia tenuifolia (Kunth) Kunth
Peterson 24816, Romaschenko, Rodriguez Avalos, Herrera- Simoni & Garcia Rodriguez (US)
Mexico, AguascalientesMK090827MK090888 −MK090741MK090792MK090704 100Muhlenbergia tenuifolia (Kunth) Kunth
Peterson 24847 & Romaschenko (US)
Mexico, ZacatecasMK090828MK090889MK090855MK090742MK090793MK090705 101Muhlenbergia thurberi (Scribn.) Rydb.Peterson 5619 & Annable (US)USA, ArizonaHM143335HM143715HM143615HM143428HM143231HM143129 102Muhlenbergia uniseta (Lag.) ColumbusPeterson 22044 & Saarela (US)Mexico, ChihuahuaGU360128GU360577GU360278GU359392GU360012GU359260 103Muhlenbergia uniseta (Lag.) ColumbusPeterson 24712, Romaschenko & Zamudio Ruíz (US)Mexico, Queretaro − − − −MK090794MK090706 104Muhlenbergia uniseta (Lag.) ColumbusPeterson 24779, Romaschenko & Zamudio Ruíz (US)Mexico, Queretaro −− − −MK090795MK090707 subg. Pseudosporobolus 105Muhlenbergia alopecuroides (Griseb.) P.M. Peterson & Columbus
Peterson 20960, Saarela, Lara Contreras & Reyna Alvarez (US)Mexico, CoahuilaGU360224GU360688GU360426GU359425GU359976GU359152 106Muhlenbergia alopecuroides (Griseb.) P.M. Peterson & ColumbusPeterson 22008 & Saarela (US)Mexico, ChihuahuaGU360223GU360687GU360425GU359451GU359975GU359153 107Muhlenbergia alopecuroides (Griseb.) P.M. Peterson & Columbus
Peterson 24524 & Romaschenko (US)
Mexico, CoahuilaMK090798MK090858MK090831MK090711MK090747MK090660 108Muhlenbergia alopecuroides (Griseb.) P.M. Peterson & Columbus
Peterson 24885 & Romaschenko (US)
Mexico, San Luis PotosíMK090799MK090859MK090832MK090712MK090748MK090661 109Muhlenbergia arenacea (Buckley) Hitchc.Peterson 10624 & Annable (US)Mexico, CoahuilaGU360210GU360675GU360414GU359452GU359961GU359165 110Muhlenbergia asperifolia (Nees & Meyen ex Trin.) ParodiPeterson 15452, Soreng, Finot & Judziewicz (US)Chile, Region III (Atacama)HM143252HM143634HM143539HM143356HM143149HM143048 111Muhlenbergia atacamensis ParodiPeterson 19626, Soreng, Salariato, & Panizza, (US)Argentina, JujuyGU360115GU360595GU360489GU359382GU359879GU359344 112Muhlenbergia cuspidata (Torr. ex Hook.) Rydb.Hill 35331 (US)USAHM143260 −HM143546 −HM143156HM143055
