A 313 plastome phylogenomic analysis of Pooideae: Exploring relationships among the largest subfamily of grasses.

In this study, we analyzed 313 plastid genomes (plastomes) of Poaceae with a focus on expanding our current knowledge of relationships among the subfamily Pooideae, which represented over half the dataset (164 representatives). In total, 47 plastomes were sequenced and assembled for this study. This is the largest study of its kind to include plastome-level data, to not only increase sampling at both the taxonomic and molecular levels with the aim of resolving complex and reticulate relationships, but also to analyze the effects of alignment gaps in large-scale analyses, as well as explore divergences in the subfamily with an expanded set of 14 accepted grass fossils for more accurate calibrations and dating. Incorporating broad systematic assessments of Pooideae taxa conducted by authors within the last five years, we produced a robust phylogenomic reconstruction for the subfamily, which included all but two supergeneric taxa (Calothecinae and Duthieeae). We further explored how including alignment gaps in plastome analyses oftentimes can produce incorrect or misinterpretations of complex or reticulate relationships among taxa of Pooideae. This presented itself as consistently changing relationships at specific nodes for different stripping thresholds (percentage-based removal of gaps per alignment column). Our summary recommendation for large-scale genomic plastome datasets is to strip alignment columns of all gaps to increase pairwise identity and reduce errant signal from poly A/T bias. To do this we used the "mask alignment" tool in Geneious software. Finally, we determined an overall divergence age for Pooideae of roughly 84.8 Mya, which is in line with, but slightly older than most recent estimates.

Miocene-Pliocene speciation, introgression, and migration of Patis and Ptilagrostis (Poaceae: Stipeae).

Genetic interchange between American and Eurasian species is fundamental to our understanding of the biogeographical patterns, and we make a first attempt to reconstruct the evolutionary events in East Asia that lead to the origin and dispersal of two genera, Patis and Ptilagrostis. We conducted a molecular phylogenetic study of 78 species in the tribe Stipeae using four plastid DNA sequences (ndhF, rpl32-trnL, rps16-trnK, and rps16 intron) and two nuclear DNA sequences (ITS and At103). We use single copy nDNA gene At103 for the first time in the grasses to elucidate the evolutionary history among members of the Stipeae. Ampelodesmos, Hesperostipa, Oryzopsis, Pappostipa, Patis, and Stipa are found to be of multiple origins. Our phylograms reveal conflicting positions for Ptilagrostis alpina and Pt. porteri that form a clade with Patis coreana, P. obtusa, and P. racemosa in the combined plastid tree but are aligned with other members of Ptilagrostis in the ITS tree. We hypothesize that Ptilagrostis still retains the nucleotype of an extinct genus which transited the Bering land bridge from American origins in the late Miocene (minimum 7.35-6.37 mya) followed by hybridization and two plastid capture events with a Trikeraia-like taxon (7.96 mya) and para-Patis (between 5.32 and 3.76 mya). Ptilagrostis porteri and Patis racemosa then migrated to continental North America 1.7-2.9 mya and 4.3-5.3 mya, respectively.

Taxonomic revision of Muhlenbergia (Poaceae, Chloridoideae, Cynodonteae, Muhlenbergiinae) in Central America: phylogeny and classification.

A taxonomic treatment of 38 species of Muhlenbergia, a phylogeny based on analysis of six DNA sequence markers, and classification of Muhlenbergia for Central America (Belize, Costa Rica, El Salvador, Guatemala, Honduras, Nicaragua, Panama; and Campeche, Chiapas, Quintana Roo, Tabasco, and Yucatán, México) is given. With the support from a molecular phylogeny we describe Muhlenbergiasubg.Ramulosaesubgen. nov. In our treatment we place M.gigantea (younger name) as a synonym of M.mutica. Lectotypes are designated for the names Agrostismicrosperma Lag., Epicampesgigantea E. Fourn., Lamarckiatenella DC., Muhlenbergiaadspersa Trin., M.diversiglumis Trin., M.exilis E. Fourn., M.flabellata Mez, M.setarioides E. Fourn., Pereilemaciliatum E. Fourn., P.crinitumvar.cirratum E. Fourn., Podosemumciliatum Kunth, P.tenuissimum J. Presl, and Schellingiatenera Steud.

A molecular phylogeny and classification of Leptochloa (Poaceae: Chloridoideae: Chlorideae) sensu lato and related genera.

BACKGROUND AND AIMS: Leptochloa (including Diplachne) sensu lato (s.l.) comprises a diverse assemblage of C(4) (NAD-ME and PCK) grasses with approx. 32 annual or perennial species. Evolutionary relationships and a modern classification of Leptochloa spp. based on the study of molecular characters have only been superficially investigated in four species. The goals of this study were to reconstruct the evolutionary history of Leptochloa s.l. with molecular data and broad taxon sampling. METHODS: A phylogenetic analysis was conducted of 130 species (mostly Chloridoideae), of which 22 are placed in Leptochloa, using five plastid (rpL32-trn-L, ndhA intron, rps16 intron, rps16-trnK and ccsA) and the nuclear ITS 1 and 2 (ribosomal internal transcribed spacer regions) to infer evolutionary relationships and revise the classification. KEY RESULTS: Leptochloa s.l. is polyphyletic and strong support was found for five lineages. Embedded within the Leptochloa sensu stricto (s.s.) clade are two Trichloris spp. and embedded in Dinebra are Drake-brockmania and 19 Leptochloa spp. CONCLUSIONS: The molecular results support the dissolution of Leptochloa s.l. into the following five genera: Dinebra with 23 species, Diplachne with two species, Disakisperma with three species, Leptochloa s.s. with five species and a new genus, Trigonochloa, with two species.

A phylogeny of the Triraphideae including Habrochloa and Nematopoa (Poaceae, Chloridoideae).

To investigate the evolutionary relationships among species of the tribe Triraphideae (including two monotypic genera, Habrochloa and Nematopoa), we generated a phylogeny based on DNA sequences from nuclear ribosomal (ITS) and four plastid markers (rps16-trnK, rps16 intron, rpl32-trnL, and ndhA intron). Habrochloa and Nematopoa form a clade that is sister to Neyraudia and Triraphis. Member of the Triraphideae have paniculate inflorescences, 3-veined, marginally ciliate lemmas, usually with hairy lateral veins, that are apically bifid and awned from between a sinus. A description of the Triraphideae and key to the genera is provided, and the biogeography is discussed, likely originating in Africa.

Allotetraploid origin and divergence in Eleusine (Chloridoideae, Poaceae): evidence from low-copy nuclear gene phylogenies and a plastid gene chronogram.

BACKGROUND AND AIMS: Eleusine (Poaceae) is a small genus of the subfamily Chloridoideae exhibiting considerable morphological and ecological diversity in East Africa and the Americas. The interspecific phylogenetic relationships of Eleusine are investigated in order to identify its allotetraploid origin, and a chronogram is estimated to infer temporal relationships between palaeoenvironment changes and divergence of Eleusine in East Africa. METHODS: Two low-copy nuclear (LCN) markers, Pepc4 and EF-1α, were analysed using parsimony, likelihood and Bayesian approaches. A chronogram of Eleusine was inferred from a combined data set of six plastid DNA markers (ndhA intron, ndhF, rps16-trnK, rps16 intron, rps3, and rpl32-trnL) using the Bayesian dating method. KEY RESULTS: The monophyly of Eleusine is strongly supported by sequence data from two LCN markers. In the cpDNA phylogeny, three tetraploid species (E. africana, E. coracana and E. kigeziensis) share a common ancestor with the E. indica-E. tristachya clade, which is considered a source of maternal parents for allotetraploids. Two homoeologous loci are isolated from three tetraploid species in the Pepc4 phylogeny, and the maternal parents receive further support. The A-type EF-1α sequences possess three characters, i.e. a large number of variations of intron 2; clade E-A distantly diverged from clade E-B and other diploid species; and seven deletions in intron 2, implying a possible derivation through a gene duplication event. The crown age of Eleusine and the allotetraploid lineage are 3·89 million years ago (mya) and 1·40 mya, respectively. CONCLUSIONS: The molecular data support independent allotetraploid origins for E. kigeziensis and the E. africana-E. coracana clade. Both events may have involved diploids E. indica and E. tristachya as the maternal parents, but the paternal parents remain unidentified. The habitat-specific hypothesis is proposed to explain the divergence of Eleusine and its allotetraploid lineage.

A classification of the Chloridoideae (Poaceae) based on multi-gene phylogenetic trees.

We conducted a molecular phylogenetic study of the subfamily Chloridoideae using six plastid DNA sequences (ndhA intron, ndhF, rps16-trnK, rps16 intron, rps3, and rpl32-trnL) and a single nuclear ITS DNA sequence. Our large original data set includes 246 species (17.3%) representing 95 genera (66%) of the grasses currently placed in the Chloridoideae. The maximum likelihood and Bayesian analysis of DNA sequences provides strong support for the monophyly of the Chloridoideae; followed by, in order of divergence: a Triraphideae clade with Neyraudia sister to Triraphis; an Eragrostideae clade with the Cotteinae (includes Cottea and Enneapogon) sister to the Uniolinae (includes Entoplocamia, Tetrachne, and Uniola), and a terminal Eragrostidinae clade of Ectrosia, Harpachne, and Psammagrostis embedded in a polyphyletic Eragrostis; a Zoysieae clade with Urochondra sister to a Zoysiinae (Zoysia) clade, and a terminal Sporobolinae clade that includes Spartina, Calamovilfa, Pogoneura, and Crypsis embedded in a polyphyletic Sporobolus; and a very large terminal Cynodonteae clade that includes 13 monophyletic subtribes. The Cynodonteae includes, in alphabetical order: Aeluropodinae (Aeluropus); Boutelouinae (Bouteloua); Eleusininae (includes Apochiton, Astrebla with Schoenefeldia embedded, Austrochloris, Brachyachne, Chloris, Cynodon with Brachyachne embedded in part, Eleusine, Enteropogon with Eustachys embedded in part, Eustachys, Chrysochloa, Coelachyrum, Leptochloa with Dinebra embedded, Lepturus, Lintonia, Microchloa, Saugetia, Schoenefeldia, Sclerodactylon, Tetrapogon, and Trichloris); Hilariinae (Hilaria); Monanthochloinae (includes Distichlis, Monanthochloe, and Reederochloa); Muhlenbergiinae (Muhlenbergia with Aegopogon, Bealia, Blepharoneuron, Chaboissaea, Lycurus, Pereilema, Redfieldia, Schaffnerella, and Schedonnardus all embedded); Orcuttiinae (includes Orcuttia and Tuctoria); Pappophorinae (includes Neesiochloa and Pappophorum); Scleropogoninae (includes Blepharidachne, Dasyochloa, Erioneuron, Munroa, Scleropogon, and Swallenia); Traginae (Tragus with Monelytrum, Polevansia, and Willkommia all embedded); Tridentinae (includes Gouinia, Tridens, Triplasis, and Vaseyochloa); Triodiinae (Triodia); and the Tripogoninae (Melanocenchris and Tripogon with Eragrostiella embedded). In our study the Cynodonteae still include 19 genera and the Zoysieae include a single genus that are not yet placed in a subtribe. The tribe Triraphideae and the subtribe Aeluropodinae are newly treated at that rank. We propose a new tribal and subtribal classification for all known genera in the Chloridoideae. The subfamily might have originated in Africa and/or Asia since the basal lineage, the Triraphideae, includes species with African and Asian distribution.

Eriocoma valdesii, a new species from México (Poaceae, Stipeae).

Eriocoma valdesii sp. nov., is described and illustrated. The new species was found growing on calcareous rocky slopes and hillsides between 1700-2721 m in Coahuila, Nuevo León, San Luis Potosí, and Tamaulipas. The new species is morphologically similar to Eriocoma lobata but differs in having ligules (2-) 4.5-8.5 mm long with acute to narrowly acute and lacerate apices and florets with a sharp-pointed callus. In addition, we include a key to the species of Eriocoma in northeastern México.

ResumenSe describe e ilustra una nueva especie, Eriocoma valdesii sp. nov. La nueva especie se encontró creciendo en laderas calcáreas rocosas y laderas entre 1700­2721 m en Coahuila, Nuevo León, San Luis Potosí y Tamaulipas. La nueva especie es morfológicamente similar a Eriocoma lobata, pero difiere en tener lígulas de (2­) 4.5­8.5 mm de largo con ápices y flósculos agudos a estrechamente agudos y lacerados con un callo puntiagudo. Además, incluimos una clave para las especies de Eriocoma en el noreste de México.

A phylogeny of species near Agrostis supporting the recognition of two new genera, Agrostula and Alpagrostis (Poaceae, Pooideae, Agrostidinae) from Europe.

Based on a molecular DNA phylogeny of three plastid (rpl32-trnK, rps16 intron, and rps16-trnK) and nuclear ITS regions investigating 32 species of Agrostidinae, we describe two new genera, Agrostula gen. nov. with a single species and Alpagrostis gen. nov. with four species; provide support for five species in a monophyletic Podagrostis; and include a small sample of 12 species of a monophyletic Agrostis s.s. (including the type and most species of Neoschischkinia), that separates into two clades corresponding to A. subg. Agrostis and A. subg. Vilfa. Agrostula differs from Agrostis in having leaf blades with pillars of sclerenchyma which are continuous between the adaxial and abaxial surface of the blades, dorsally rounded glumes with blunt to truncate and erose to denticulate apices, florets ½ the length of the glumes, lemmas equally wide as long, widest at (or near) apex, apices broadly truncate, irregularly 5 to 7 denticulate to erose, awnless, anthers longer than the lemmas, and rugose-papillose caryopses. Alpagrostis differs from Agrostis in having geniculate basally inserted awns and truncate lemma apices with lateral veins prolonged from the apex in (2)4 setae. The following eight new combinations are made: Agrostula truncatula, Agrostula truncatula subsp. durieui, Alpagrostis alpina, Alpagrostis alpina var. flavescens, Alpagrostis barceloi, Alpagrostis setacea, Alpagrostis setacea var. flava, and Alpagrostis schleicheri. In addition, we provide a key separating Agrostula and Alpagrostis from Agrostis s.s. and other genera previously considered as synonyms of Agrostis; lectotypify Agrostis alpina Scop., A. schleicheri Jord. & Verl., A. truncatula Parl., and A. truncatula var. durieui Henriq.; and neotypify A. setacea Curtis.

ResumenSobre la base de una filogenia molecular de ADN de tres regiones plastidiales (rpl32-trnK, rps16 intrón y rps16-trnK) e ITS nuclear de 32 especies de Agrostidinae, describimos dos nuevos géneros, Agrostula gen. nov. con una sola especie, y Alpagrostis gen. nov. con cuatro especies; mostramos el apoyo para las cinco especies dentro de Podagrostis monofilético; e incluimos una pequeña muestra de 12 especies de Agrostis s.s (que incluye el tipo y la mayoría de las especies de Neoschischkinia), este último dividido en dos subclados que corresponden a A. subg. Agrostis y A. subg. Vilfa. Agrostula se diferencia de otras especies de Agrostis por tener láminas foliares con haces de esclerénquima continuos entre las superficies adaxial y abaxial de los limbos, glumas de dorso redondeado y ápice embotado a truncado y eroso a denticulado, antecios de ½ de la longitud de las glumas, lemas tan anchas como largas, lo más ancho en o cerca del ápice, ápices anchamente truncados, irregularmente 5 a 7 denticulados o erosos, sin arista, anteras más largas que los lemas y cariopsis rugosa-papilosa. Alpagrostis se diferencia de otras especies de Agrostis por tener aristas geniculadas insertas basalmente y ápices de lema truncados con venas laterales que se prolongan en (2)4 arístulas apicales. Presentamos las siguientes ocho nuevas combinaciones: Agrostula truncatula, Agrostula truncatula subsp. durieui, Alpagrostis alpina, Alpagrostis alpina var. flavescens, Alpagrostis barceloi, Alpagrostis setacea, Alpagrostis setacea var. flava y Alpagrostis schleicheri. Además, proporcionamos una clave que separa Agrostula y Alpagrostis de Agrostis s.s. y otros géneros previamente considerados como sinónimos de Agrostis, lectotipificamos Agrostis alpina Scop., A. schleicheri Jord. & Verl., A. truncatula Parl. y A. truncatula var. durieui Henriq. y neotipificamos A. setacea Curtis.

New combinations and updated descriptions in Podagrostis (Agrostidinae, Poaceae) from the Neotropics and Mexico.

Based on morphological study and corroborated by unpublished molecular phylogenetic analyses, five grass species of high-mountain grasslands in Mexico, Central and South America, Agrostis bacillata, A. exserta, A. liebmannii, A. rosei, and A. trichodes, are transferred to Podagrostis and bring the number of species of this genus recognized in the New World to ten. The name Apera liebmannii is lectotypified and epitypified. We provide an updated genus description for Podagrostis, and updated species descriptions, images, and notes on the new combinations. The diagnostic characteristics differentiating Podagrostis from Agrostis are: a) palea that reaches from (2/3) ¾ to almost the apex of the lemma; b) florets that usually almost equal the length of the glumes or are at least ¾ the length of the glumes; c) rachilla extension present and emerging from under the base of the palea as a slender short stub (rudimentary or up to 1.4 mm long, sometimes obscure in most florets in P. rosei), smooth or scaberulous, glabrous or distally pilulose (hairs < 0.3 mm long); d) lemmas usually awnless, sometimes with a short straight awn 0.2-0.6 mm long, inserted medially or in the upper 1/3 of the lemma, not surpassing the glumes (awn well-developed, straight or geniculate and inserted in lower 1/3 of lemma, not or briefly surpassing glumes in P. rosei). We include a generic key to distinguish the species of Podagrostis from other similar genera in Latin America and a key to distinguish the species of Podagrostis now accepted as occurring in these areas.

ResumenBasado en estudios morfológicos y corroborado por datos no publicados de análisis filogenéticos, las cinco gramíneas de alta montaña de los pastizales en México, Centroamérica y Sur América Agrostis bacillata, A. exserta, A. liebmannii, A. rosei, y A. trichodes, son transferidos a Podagrostis incrementando a diez el número de especies en el nuevo mundo de este reconocido género. El nombre de Apera liebmannii también es lectotipificado y epitipicado. Proporcionamos una descripción actualizada del género Podagrostis, y descripciones actualizadas de las especies, imágenes y comentarios sobre las nuevas combinaciones. Las características diagnósticas que diferencian a Podagrostis de Agrostis son: a) pálea de 2/3-¾ hasta casi el ápice de la lemma; b) espiguillas, generalmente casi iguales en longitud a las glumas o de al menos ¾ la longitud de las glumas; c) extensión de la raquilla presente y emergiendo desde debajo de la base de la pálea como un trozo corto y delgado (desde rudimentario hasta de 1.4 mm de largo, a veces oscuro en la mayoría de los flósculos en P. rosei), liso o escabérulo, glabro o piloso distalmente (pelos < 0.3 mm de largo); d) lemmas generalmente sin arista, o a veces con arista corta y recta de 0.2­0.6 mm de largo, insertada medialmente o en el tercio superior de la lemma, sin sobrepasar las glumas (arista bien desarrollada, recta o geniculada e insertada en la parte inferior 1/3 de lemma, no o superando brevemente las glumas en P. rosei). Incluimos una clave para distinguir Podagrostis de otros géneros similares presentes en América Latina y una clave para distinguir las especies de Podagrostis ahora aceptadas por estar presentes en estas áreas.

A key to the North American genera of Stipeae (Poaceae, Pooideae) with descriptions and taxonomic names for species of Eriocoma, Neotrinia, Oloptum, and five new genera: Barkworthia, ×Eriosella, Pseudoeriocoma, Ptilagrostiella, and Thorneochloa.

Based on earlier molecular DNA studies we recognize 14 native Stipeae genera and one intergeneric hybrid in North America. We provide descriptions, new combinations, and 10 illustrations for species of Barkworthia gen. nov., Eriocoma, Neotrinia, Oloptum, Pseudoeriocoma gen. nov., Ptilagrostiella gen. nov., Thorneochloa gen. nov., and ×Eriosella nothogen. nov. The following 40 new combinations are made: Barkworthiastillmanii, Eriocomaalta, E.arida, E.arnowiae, E.bloomeri, E.bracteata, E.contracta, E.coronata, E.curvifolia, E.hendersonii, E.latiglumis, E.lemmonii, E.lemmoniissp.pubescens, E.lettermanii, E.lobata, E.nelsonii, E.nelsoniissp.dorei, E.nevadensis, E.occidentalis, E.occidentalisssp.californica, E.occidentalisssp.pubescens, E.parishii, E.parishiissp.depaupertata, E.perplexa, E.pinetorum, E.richardsonii, E.robusta, E.scribneri, E.swallenii, E.thurberiana, E.wallowaensis, ×Eriosellacaduca, Pseudoeriocomaacuta, P.constricta, P.editorum, P.eminens, P.hirticulmis, P.multinodis, Ptilagrostiellakingii, and Thorneochloadiegoensis. A key to the native and introduced genera of North American Stipeae, and an overview of the tribe in North America and worldwide are given. Lectotypes are designated for Eriocomacuspidata Nutt., Fendleriarhynchelytroides Steud., Stipabloomeri Bol., Stipacoronata Thurb., Stipamembranacea Pursh, Stipamormonum Mez, Stiparichardsonii Link, and Stipawilliamsii Scribn. Achnatherum s.s. and Piptatherum s.s. are now restricted to Eurasia and the Mediterranean/Asia, respectively.

ResumenBasados en estudios anteriores de ADN molecular, reconocemos 14 géneros nativos de Stipeae y un híbrido intergenérico en América del Norte. Se presentan descripciones, nuevas combinaciones, y 10 ilustraciones para las especies de Barkworthia gen. nov., Eriocoma, Neotrinia, Oloptum, Pseudoeriocoma gen. nov., Ptilagrostiella gen. nov., Thorneochloa gen. nov. y ×Eriosella nothogen. nov. Se realizan las siguientes 40 nuevas combinaciones: Barkworthiastillmanii, Eriocomaalta, E.arida, E.arnowiae, E.bloomeri, E.bracteata, E.contracta, E.coronata, E.curvifolia, E.hendersonii, E.latiglumis, E.lemmonii, E.lemmoniissp.pubescens, E.lettermanii, E.lobata, E.nelsonii, E.nelsoniissp.dorei, E.nevadensis, E.occidentalis, E.occidentalisssp.californica, E.occidentalisssp.pubescens, E.parishii, E.parishiissp.depaupertata, E.perplexa, E.pinetorum, E.richardsonii, E.robusta, E.scribneri, E.swallenii, E.thurberiana, E.wallowaensis, ×Eriosellacaduca, Pseudoeriocomaacuta, P.constricta, P.editorum, P.eminens, P.hirticulmis, P.multinodis, Ptilagrostiellakingii y Thorneochloadiegoensis. Se presenta una clave para los géneros nativos e introducidos de las especies norteamericanas, y una visión general de la tribu en América del Norte y en todo el mundo. Se designan lectotipos para Eriocomacuspidata Nutt., Fendleriarhynchelytroides Steud., Stipabloomeri Bol., Stipacoronata Thurb., Stipamembranacea Pursh, Stipamormonum Mez, Stiparichardsonii Link y Stipawilliamsii Scribn. Achnatherum s.s. y Piptatherum s.s. ahora están con distribución restringida- a Eurasia y el Mediterráneo/Asia, respectivamente.

Páramo Calamagrostis s.l. (Poaceae): An updated list and key to the species known or likely to occur in páramos of NW South America and southern Central America including two new species, one new variety and five new records for Colombia.

Calamagrostis (syn. Deyeuxia), as traditionally circumscribed, is one of the most speciose genera from páramo grasslands of northwest South America and southern Central America and often dominates these high-elevation habitats. However, it remains difficult for researchers to accurately identify the species due to a lack of floristic treatments for most of the countries containing páramo, with the distribution of many species still very poorly known. In an effort to ameliorate this, we present an updated list and identification keys in English and Spanish (as electronic appendix) to the species of Calamagrostis s.l. known or likely to occur in the páramos of Peru, Ecuador, Colombia, Venezuela, Costa Rica and Panama. Fifty-four species are accepted, constituting 47 species currently circumscribed in Calamagrostis and seven species recently transferred to Deschampsia. Included within this are two new species, Calamagrostiscrispifolius and Deschampsiasantamartensis, which are described and illustrated. Both new species are found in páramos of the Sierra Nevada de Santa Marta (departamento Magdalena), on the northernmost tip of Colombia, with C.crispifolius also found in the Serrania de Perija on the border with Venezuela. Calamagrostiscrispifolius differs from all other species of Calamagrostis s.l. by the presence of strongly curled, readily deciduous leaf blades, amongst numerous other characteristics including open inflorescences with generally patent branches, small spikelets, (3.5-)4-5.5 mm long, with sessile florets and a rachilla prolongation reaching from 2/3 to almost the apex of the lemma, with short hairs (< 1 mm long). Deschampsiasantamartensis is similar to Deschampsiahackelii (=Calamagrostishackelii) from austral South America but differs by its broad, rigid and erect, strongly conduplicate blades, 1.5-2.5 mm wide when folded, ligules of innovations 0.5-1 mm long, truncate or obtuse, ligules of upper flowering culms 3-4 mm long, broadly shouldered with an attenuate central point, ellipsoid spike-like panicle, 3-5.5 long × 1.5-2.5 cm wide, lemma surfaces moderately to lightly scabrous between the veins, lemma apex acute to muticous, entire, rachilla extension often absent and inside of the floret often with hyaline shiny sinuous trichomes to 1 mm long, emerging from the base of the ovary. We also present a broader circumscription of the common species Deschampsiapodophora (=Calamagrostispodophora), with the new variety D.podophoravar.mutica described and illustrated. Deschampsiapodophoravar.mutica principally differs from var. podophora by florets lacking awns and larger habit i.e. multiple taller culms with longer and wider leaf blades forming tussocks, with inflorescences often held within sheaths. Nomenclatural changes are presented, with Deyeuxiamacrostachya newly synonymised under C.macrophylla and C.pittieri, C.pubescens and Deyeuxiapubescens newly synonimised under C.planifolia. Lectotypes are designated for Agrostisantoniana, Calamagrostispisinna, Deyeuxiamacrostachya and Deyeuxiasodiroana. We also document and give notes on five new records of Calamagrostis for Colombia: C.carchiensis, C.guamanensis, C.heterophylla, C.pisinna and C.rigida.

ResumenCalamagrostis (syn. Deyeuxia), como traditionalmente está delimitado, es uno de los géneros con mayor número de especies registradas para las zonas paramunas de Sudamérica y el sur de Centroamérica, en donde a menudo es un elemento dominante en hábitats de alta montaña tropical. Sin embargo, es aun difícil para investigadores identificar con precisión las especies de este género, principalmente por la falta de tratamientos florísticos para la mayoría de países que contienen páramo, con la distribución de muchas especies todavía poca conocida. Con el fin de realizar un aporte al conocimiento de las gramíneas tropicales, se presenta una lista actualizada de nombres y claves de identificación taxonómica en inglés y español (como appendice electrónico) de las especies de Calamagrostis s.l. conocidas o que probablemente se encuentran en los páramos de Perú, Ecuador, Colombia, Venezuela, Costa Rica y Panamá. Como resultado, 54 especies son aceptadas, de las cuales 47 especies están circunscritas en Calamagrostis y siete especies están recientemente transferidas a Deschampsia. Como novedades taxonómicas, se presentan dos nuevas especies, Calamagrostiscrispifolius y Deschampsiasantamartensis, las cuales están descritas e ilustradas. Estas nuevas especies crecen en páramos de la Sierra Nevada de Santa Marta (departamento de Magdalena), en la punta más al norte de Colombia, con C.crispifolius también presente en la Serranía de Perijá en la frontera con Venezuela. Calamagrostiscrispifolius difiere de las otras especies de Calamagrostis s.l. por la presencia de láminas foliares fuertemente crispadas y fácilmente caducas, entre otras características incluyendo inflorescencias abiertas con ramas generalmente patentes, espiguillas pequeñas, (3.5­)4­5.5 mm long., con antecios sésiles y una prolongación de la raquilla llegando desde 2/3 hasta casi el ápice de la lema, con tricomas cortos (< 1 mm long.). Deschampsiasantamartensis es parecida a Deschampsiahackelii (=Calamagrostishackelii) de Sudamérica austral, pero difiere por sus láminas foliares siendo anchas, rígidas y rectas, y fuertemente conduplicadas, 1.5­2.5 mm de ancho cuando plegada, lígulas de las innovaciones 0.5­1 mm long., truncadas u obtusas, lígulas de la parte superior de las cañas floríferas 3­4 mm long., anchas con un punto central atenuado, panojas elipsoides y espiciforme, 3­5.5 long. × 1.5­2.5 cm de ancho, superficies de las lemas moderada a levemente escabroso entre las venas, ápices de las lemas agudas a múticas, enteras, prolongación de la raquilla a menudo ausente, y la presencia de tricomas hialinos, brillantes, y sinuosos de hasta 1 mm long., los cuales salen de la base del ovario. Por último, se presenta una circunscripción más amplia de la especie común Deschampsiapodophora (=Calamagrostispodophora), con la nueva variedad D.podophoravar.mutica, la cual es descrita e ilustrada. Deschampsiapodophoravar.mutica difiere principalmente de la var. podophora porque en sus antecios las aristas están ausentes, y su hábito es más grande i.e. con múltiples cañas largas con láminas foliares más largas y anchas, con inflorescencias a menudo escondidas dentro de las vainas. Los cambios nomenclaturales que se presentan son Deyeuxiamacrostachya sinonimizada bajo C.macrophylla, y C.pittieri, C.pubescens y Deyeuxiapubescens sinonimizada bajo C.planifolia. Designamos los lectotipos para Agrostisantoniana, Calamagrostispisinna, Deyeuxiamacrostachya y Deyeuxiasodiroana. También documentamos y damos notas de cinco nuevos registros de Calamagrostis para la flora de Colombia: C.carchiensis, C.guamanensis, C.heterophylla, C.pisinna, y C.rigida.

Poa laegaardiana, a new species from Ecuador (Poaceae, Pooideae, Poeae, Poinae).

Poa laegaardianasp. nov., is described and illustrated. The new species was found growing on sandy, volcanic soil in Festuca-Calamagrostis dominated grasslands southwest of Ambato and 2.2 km from Fecundo Vela in Provincia de Bolivar. The new species is morphologically similar to Poa gigantea but differs in having glumes 3/4 to 7/8 as long as the adjacent lemmas, a callus with a sparse, short, dorsal tuft of woolly hairs, culms 50-72 cm tall and spikelets 4.1-4.8 mm long. In addition, we include a key to the narrow-spikelike panicled species of Poa in Ecuador.

ResumenSe describe e ilustra una nueva especie, Poa laegaardianasp. nov. La especie se encontró creciendo en suelo arenoso y volcánico en pastizales dominados por Festuca-Calamagrostis al suroeste de Ambato y a 2.2 km de Fecundo Vela en la Provincia de Bolívar. La nueva especie es morfológicamente similar a Poa gigantea, pero se distinque por glumas ¾ a 7/8 tan larga como los lemmas adyacentes, un callo con un mechón escaso, corto, dorsal de pelos lanosos, culmos de 50­72 cm de altura, y espiguillas 4.1­4.8 mm de largo. Además, incluimos una clave para las especies de Poa con panículas espiciformes de Ecuador.

Phylogeny of Muhlenbergia subg. Pseudosporobolus, including M. spatha (Poaceae, Chloridoideae, Cynodonteae, Muhlenbergiinae) now found in Zacatecas, Mexico.

Muhlenbergia spatha, previously known only from near the type locality in San Luis Potosí, is reported from two localities in Zacatecas, Mexico. Historically, botanists have overlooked this diminutive annual. To clarify affinities of M. spatha, we present a molecular phylogeny emphasising species in M. subg. Pseudosporobolus using sequence data from two plastid markers (rpl32-trnL and rps16 intron) and nrDNA ITS. In addition, we include an updated description, illustration and discussion of the habitat of M. spatha.

Resumen Muhlenbergia spatha, anteriormente conocida solo cerca de la localidad tipo en San Luis Potosí, se reporta en dos localidades en Zacatecas, México. Históricamente, los botánicos han pasado por alto esta diminuta anual. Para aclarar las afinidades de M. spatha, presentamos una filogenia molecular que enfatiza especies en M. subg. Pseudosporobolus usando datos de secuencia de dos marcadores plástidos (rpl32-trnL e rps16 intron) y nrADN EIT. Además, incluimos una descripción actualizada, ilustración y discusión del hábitat de M. spatha.

Monograph of Diplachne (Poaceae, Chloridoideae, Cynodonteae).

Diplachne P. Beauv. comprises two species with C4 (NAD-ME) photosynthesis. Diplachne fusca has a nearly pantropical-pantemperate distribution with four subspecies: D. fusca subsp. fusca is Paleotropical with native distributions in Africa, southern Asia and Australia; the widespread Australian endemic D. f. subsp. muelleri; and D. f. subsp. fascicularis and D. f. subsp. uninervia occurring in the New World. Diplachne gigantea is known from a few widely scattered, older collections in east-central and southern Africa, and although Data Deficient clearly is of conservation concern. A discussion of previous taxonomic treatments is provided, including molecular data supporting Diplachne in its newer, restricted sense. Many populations of Diplachne fusca are highly tolerant of saline substrates and most prefer seasonally moist to saturated soils, often in disturbed areas. Some populations of Diplachne fusca in southern Asia combine nitrogen-fixation, high salinity tolerance and palatibilty to livestock, which should be pursued with further research for purposes of soil reclamation. Diplachne fusca subsp. uninervia is the most invasive of the subspecies and is becoming weedy in some non-native areas, including in the Old World. This monograph provides detailed descriptions of all taxa, a key to the species and subspecies, geographic distributions and information on the anatomy of leaves, stems, lemmatal micromorphology and discussions of the chromosome numbers. Lectotypes are designated for: Atropis carinata Grisb.; Diplachne acuminata Nash; Diplachne capensis (Nees) Nees var. concinna Nees; Diplachne capensis (Nees) Nees var. obscura Nees, Diplachne capensis (Nees) Nees var. prolifera subvar. minor Nees, Diplachne halei Nash, Diplachne maritima E.P. Bicknel, Diplachne muelleri Benth., Diplachne reverchonii Vasey, Diplachne tectoneticola Backer, Leptochloa imbricata Thurb., Leptochloa neuroglossa Peter, Leptochloa uninervia var. typica fo. abbreviata Parodi, Triodia ambigua R. Br. and Triodia parviflora R. Br.

Revision of Muhlenbergia (Poaceae, Chloridoideae, Cynodonteae, Muhlenbergiinae) in Peru: classification, phylogeny, and a new species, M.romaschenkoi.

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. Seventeen species and one presumed hybrid are recognised. Muhlenbergiaromaschenkoi sp. nov. is newly described from the Río Huallaga Valley, northeast of Huánuco. The type of Podosemumangustatum [≡ Muhlenbergiaangustata] 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, Muhlenbergiacoerulea, for what we had been calling M.angustata and change the hybrid designation to M.coerulea × M.rigida. Lectotypes are designated for Epicampescoerulea Griseb., Muhlenbergiaaffinis Trin., Muhlenbergiaberlandieri Trin., Muhlenbergiabeyrichiana Kunth, Muhlenbergiaelegansvar.atroviolacea Kuntze, Muhlenbergiaelegansvar.subviridis Kuntze and Muhlenbergiaphragmitoides Griseb.

ResumenBrindamos 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. Muhlenbergiaromaschenkoi sp. nov. es descrita como especie nueva procedente del valle de Huallaga, al noreste de Huánuco. El tipo de Podosemumangustatum [≡ Muhlenbergiaangustata] 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, Muhlenbergiacoerulea para lo que habíamos estado llamando M.angustata, y cambiamos la designación híbrida a M.coerulea × M.rigida. Los lectotipos son designados para Epicampescoerulea Griseb., Muhlenbergiaaffinis Trin., Muhlenbergiaberlandieri Trin., Muhlenbergiabeyrichiana Kunth, Muhlenbergiaelegansvar.atroviolacea Kuntze, Muhlenbergiaelegansvar.subviridis Kuntze y Muhlenbergiaphragmitoides Griseb.

A 250 plastome phylogeny of the grass family (Poaceae): topological support under different data partitions.

The systematics of grasses has advanced through applications of plastome phylogenomics, although studies have been largely limited to subfamilies or other subgroups of Poaceae. Here we present a plastome phylogenomic analysis of 250 complete plastomes (179 genera) sampled from 44 of the 52 tribes of Poaceae. Plastome sequences were determined from high throughput sequencing libraries and the assemblies represent over 28.7 Mbases of sequence data. Phylogenetic signal was characterized in 14 partitions, including (1) complete plastomes; (2) protein coding regions; (3) noncoding regions; and (4) three loci commonly used in single and multi-gene studies of grasses. Each of the four main partitions was further refined, alternatively including or excluding positively selected codons and also the gaps introduced by the alignment. All 76 protein coding plastome loci were found to be predominantly under purifying selection, but specific codons were found to be under positive selection in 65 loci. The loci that have been widely used in multi-gene phylogenetic studies had among the highest proportions of positively selected codons, suggesting caution in the interpretation of these earlier results. Plastome phylogenomic analyses confirmed the backbone topology for Poaceae with maximum bootstrap support (BP). Among the 14 analyses, 82 clades out of 309 resolved were maximally supported in all trees. Analyses of newly sequenced plastomes were in agreement with current classifications. Five of seven partitions in which alignment gaps were removed retrieved Panicoideae as sister to the remaining PACMAD subfamilies. Alternative topologies were recovered in trees from partitions that included alignment gaps. This suggests that ambiguities in aligning these uncertain regions might introduce a false signal. Resolution of these and other critical branch points in the phylogeny of Poaceae will help to better understand the selective forces that drove the radiation of the BOP and PACMAD clades comprising more than 99.9% of grass diversity.

Unraveling the evolutionary dynamics of ancient and recent polyploidization events in Avena (Poaceae).

Understanding the diversification of polyploid crops in the circum-Mediterranean region is a challenging issue in evolutionary biology. Sequence data of three nuclear genes and three plastid DNA fragments from 109 accessions of Avena L. (Poaceae) and the outgroups were used for maximum likelihood and Bayesian analyses. The evolution of cultivated oat (Avena sativa L.) and its close relatives was inferred to have involved ancient allotetraploidy and subsequent recent allohexaploidy events. The crown ages of two infrageneric lineages (Avena sect. Ventricosa Baum ex Romero-Zarco and Avena sect. Avena) were estimated to be in the early to middle Miocene, and the A. sativa lineages were dated to the late Miocene to Pliocene. These periods coincided with the mild seasonal climatic contrasts and the Mediterranean climate established in the Mediterranean Basin. Our results suggest that polyploidy, lineage divergence, and complex reticulate evolution have occurred in Avena, exemplifying the long-term persistence of tetraploids and the multiple origins of hexaploids related to paleoclimatic oscillations during the Miocene-Pliocene interval in the circum-Mediterranean region. This newly-resolved infrageneric phylogenetic framework represents a major step forward in understanding the origin of the cultivated oat.

A revision of Poa subsection Aphanelytrum (Poaceae, Pooideae, Poaeae, Poinae); and a new species, Poa auriculata.

In this study the peculiar Andean grass genus Aphanelytrum, with two species, is reduced to Poa subsect. Aphanelytrum comb. & stat. nov. A third species, Festuca reclinata, is assigned to the subsection, which shows states transitional between a more typical Poa and Aphanelytrum. Poa subgen. Poa supersect. Homalopoa sect. Dioicopoa subsect. Aphanelytrum comb. & stat. nov. is characterized in having stooling perennials with decumbent to spreading culm bases that continuously branch and often root at low to mid-culm nodes, glabrous spikelets with long rachillas 1.2-4.2 mm long, short glumes less than ½ the length of the florets, and lemmas with bifid apexes that are mucronate to short-awned. We provide for the three species taxonomic discussions, morphological and anatomical descriptions, keys, illustrations, and a list of specimens. Also, we provide two new names, Poa hitchcockiana nom. nov. and Poa sanchez-vegae nom. nov., and one new combination, Poa reclinata comb. nov. A new species, Poa auriculata sp. nov. from Peru, not thought to be a member of Poa subsect. Aphanelytrum, is presented. It is the first in the genus with prominent auricles. In addition, we place Poa apiculata in Poa subgen. Poa supersect. Homalopoa sect. Dioicopoa subsect. Tovarochloa comb. & stat. nov.

ResumenEn este estudio el género peculiar de gramíneas andinas Aphanelytrum, representado por dos especias, ha sido reducido a Poa subsecc. Aphanelytrum comb. & stat. nov. Una tercera especie es incluida en la subsección, Festuca reclinata, esta presenta características transicionales entre las especies más típicas de Poa y Aphanelytrum. Poa subgen. Poa supersecc. Homalopoa secc. Dioicopoa subsecc. Aphanelytrum comb. & stat. nov. se caracteriza por incluir plantas perennes, stoloniferas, con los culmos decumbentes o extensos, continuamente ramificados, a menudo enraizados en los nudos basales hasta los intermedios, las espiguillas son glabras con rachillas largas de 1.2­4.2 mm de longitud, las glumas más cortas son de la ½ de longitud de los flósculos y las lemas con los ápices bífidos son mucronadas hasta breve-aristadas. Nosotros proveemos para las tres especias discusiones taxonómicas, descripciones morfológicas y anatómicas, claves, ilustraciones y la lista de las muestras usadas. Tambien, proveemos dos nombres nuevos - Poa hitchcockiananom. nov. y Poa sanchez-vegaenom. nov., y una combinación -Poa reclinatacomb. nov. Una nueva especie - Poa auriculatasp. nov. de Perú es presentada. Esta es la especie primera del género con prominentes aurículas. En adición, transferimos Poa apiculata a Poa subgen. Poa supersecc. Homalopoa secc. Dioicopoa subsecc. Tovarochloa comb. & stat. nov.

An updated checklist and key to the open-panicled species of Poa L. (Poaceae) in Peru including three new species, Poa ramoniana, Poa tayacajaensis, and Poa urubambensis.

We provide an updated checklist and key to the 30 Poa species with open panicles from Peru which includes previously circumscribed Dissanthelium and Aphanelytrum species, new taxon records, and three undescribed species. Poa compressa, Poa grisebachii, and Poa leioclada are recorded from Peru for the first time. A number of species are placed in synonymy: Poa carazensis, Poa ferreyrae and Poa tovarii are synonymized under the name Poa fibrifera; Poa adusta (tentatively) and Poa pilgeri are synonymized under Poa candamoana; Poa superata is synonymized under Poa grisebachii; and Poa paramoensis is synonymized under Poa huancavelicae. Included within this treatment are three new species, Poa ramoniana, Poa tayacajaensis and Poa urubambensis, which are described and illustrated. Poa ramoniana, found growing near lakes in high elevation Puna grasslands of Junín, is similar to a small form of Poa glaberrima, but differs in having rhizomes and growing to only 5 cm tall. Poa tayacajaensis, found from shrublands on Andean slopes of Huancavelica and Huánuco, bears similarities to Poa aequatoriensis but differs in having shorter lemmas which are pubescent between the veins, densely scabrous sheaths with smooth, glabrous throats, and shorter ligules. Poa urubambensis, a common element of the undisturbed Polylepis forest understory of the Cordillera Urubamba, Cusco, is distinct from all other members of open-panicled Poa's by having glabrous lemmas with a smooth and glabrous callus, and notably small anthers. The type material for the name Poa adusta is discussed and a lectotype is selected.

ResúmenAportamos una lista actualizada y una clave para las 30 especies de Poa con panículas abiertas de Perú que incluye las especies de Dissanthelium y Aphanelytrum anteriormente circunscritas, nuevos registros de taxones y tres especies no descritas. Poa compressa, Poa grisebachii, y Poa leioclada se registran para Perú por primera vez. Un número de especies son sinonimizadas: Poa carazensis, Poa ferreyrae y Poa tovarii son sinonimizadas bajo el nombre de Poa fibrifera; Poa adusta (tentativamente) y Poa pilgeri son sinonimizadas bajo el nombre de Poa candamoana; Poa superata es sinonimizada bajo Poa grisebachii; y Poa paramoensis es sinonimizada bajo Poa huancavelicae. Dentro de este tratamiento se incluyen dos especies nuevas, Poa ramoniana, Poa tayacajaensis y Poa urubambensis que a continuación se describen e ilustran. Poa ramoniana, que se encuentra creciendo en pastizales de alta elevación cercanos a lagos en la Puna de Junín, es similar a la forma pequeña de Poa glaberrima, pero se diferencia por tener rizomas y crecer hasta sólo 5 cm de altura. Poa tayacajaensis, que se encuentra en matorrales de las laderas andinas de Huancavelica y Huánuco, tiene un parecido a Poa aequatoriensis, pero se diferencia por tener lemas cortas que son pubescentes entre las venas, vainas densamente escabrosas con suaves gargantas glabras, y lígulas cortas. Poa urubambensis, un elemento común de sotobosque no perturbado de los bosques de Polylepis de la Cordillera Urubamba, Cusco, se distingue de todos los otros miembros de Poa con panícula-abierta por tener lemas glabras con un callo suave y glabro, y sus notables anteras pequeñas. El material tipo del nombre Poa adusta es discutido y un lectotipo es seleccionado.