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Phylogeny and taxonomic synopsis of Poa subgenus Pseudopoa (including Eremopoa and Lindbergella) (Poaceae, Poeae, Poinae)
expand article infoLynn J. Gillespie§, Robert John Soreng|, Evren Cabi, Neda Amiri§
‡ Canadian Museum of Nature, Ottawa, Canada
§ University of Ottawa, Ottawa, Canada
| Smithsonian Institution, Washington, United States of America
¶ Tekirdağ Namık Kemal University, Tekirdağ, Turkey
Open Access

Abstract

Eremopoa is a small genus of annual grasses distributed from Egypt to western China. Phylogenetic analyses of plastid and nuclear ribosomal DNA show that Eremopoa species, together with the monotypic genus Lindbergella and a single species of Poa (P. speluncarum), are nested within the genus Poa, in a clade that we accept as Poa subg. Pseudopoa. Here we accept seven species, four subspecies and four varieties in Poa subg. Pseudopoa. Five new combinations are made: Poa attalica, P. diaphora var. alpina, P. diaphora var. songarica, P. nephelochloides and P. persica subsp. multiradiata; P. millii is proposed as a replacement name for E. capillaris; and Poa sections Lindbergella and Speluncarae are proposed. We provide a diagnosis for Poa subg. Pseudopoa, synonymy for and a key to the taxa. Eight lectotypes are designated: Eragrostis barbeyi Post, Eremopoa nephelochloides Roshev., Glyceria taurica Steud., Nephelochloa tripolitana Boiss. & Blanche, Poa cilicensis Hance, Poa paradoxa Kar. & Kir., Poa persica var. alpina Boiss and Poa persica subsp. cypria Sam. Eremopoa medica is re-identified as a species of Puccinellia.

Keywords

Annuals, classification, DNA, Eremopoa, grasses, Lindbergella, phylogeny, Poa, Poaceae, taxonomy

Introduction

Eremopoa Roshev. is a small, primarily west and central Asian genus of annual grasses. Roshevitz (1934) named the genus Eremopoa (Greek: eremos = desert, poa = fodder / > bluegrass) and included six species of annuals for the former U.S.S.R. Up to that time, one or more of the taxa had been described or treated in Aira L. (Trinius 1835), Eragrostis Wolf (Post and Autran 1897), Festuca L. (Koch 1848), Glyceria R. Br. (Fischer and Meyer 1841, Steudel 1854), Nephelochloa Boiss. (Grisebach 1852, Boissier and Blanche 1859) and Poa L. (Trinius 1830, 1836, Steudel 1854, Boissier 1884, Hackel 1887, Stapf 1897, Ascherson and Graebner 1900). Poa persica Trin. is the type species of Eremopoa, Festuca sect. Pseudopoa K. Koch, Poa subgen. Pseudopoa (K. Koch) Stapf and P. sect. Pseudopoa (K. Koch) Hack. After Eremopoa was described, most authors accepted the genus (Grossheim 1939, Köie 1945, Bor 1960, 1968a, 1970, Pavlov and Gamajunova 1964, Tzvelev 1966, 1976, 1989, Scholz 1980, 1981, Tutin 1980, Czerepanov 1981, 1995, Cope 1982, Mill 1985, Clayton and Renvoize 1986, Watson and Dallwitz 1992, Soreng 2003, Valdés and Scholz 2006, Darbyshire 2007, Cabi and Doğan 2012, Nikiforova et al. 2012). Few taxonomists continued to refer the species to Poa (Samuelsson 1950, Kovalevskaja 1968). No revision of the genus as a whole exists.

Roshevitz (1934) differentiated the genus Eremopoa from Poa as: always annuals with long panicle branches arranged in half-whorls; glumes unequal, inferior 1-veined, superior 3-veined; lemmas with obscure keel and lateral veins, apex acuminate or briefly aristate; and callus without lanate hairs. Tzvelev (1976) added the following characteristics: lower glumes 2/7–2/3 the first lemma in length; lemmas somewhat keeled with 5 veins, apex gradually tapering, sometimes with a short cusp, somewhat scabrous due to very short spinules and often pilose in the lower part along the keel and marginal veins; callus obtuse, glabrous or almost glabrous; leaf sheaths closed only at the base and leaf blades flat or loosely folded. The genus is relatively easy to recognise as a set of annuals, whereas Poa has few annuals and those are distinct from species included in Eremopoa. However, none of the characters by themselves actually differentiates Eremopoa from Poa. In Poa, glumes can also be short, the lower one is commonly 1-veined, the upper one normally 3-veined. Lemmas in Poa are usually distinctly keeled, with soft hairs at least on the keel and with an obtuse, acute or acuminate apex. They are rarely weakly keeled (e.g. in sect. Secundae), sometimes glabrous (ca. 15% of spp.) and rarely produce a minute cusp (a cusp occurs more often than acknowledged in the literature, but is usually irregularly expressed). In Poa, a dorsal tuft of hairs on the callus is present in 2/3 of the species. In the other species, the callus is sometimes glabrous or has a minute or more developed crown of hairs around the base of the lemma. In addition, Poa leaf sheaths are only infrequently closed at the base, most being closed more than 1/10 the length, and leaf blade form runs the gamut from flat and thin to tough and involute. Panicle branches in Poa are infrequently whorled with 6 or up to 9 branches per lower node, the normal range is 1 to 5. Although panicle branches are commonly numerous (ranging up to 27) in Eremopoa, with most taxa usually having over 5, E. altaica (Trin.) Roshev. has 1–5(–7) and E. songarica (Schrenk ex Fisch. & C.A. Mey.) Roshev. varies widely with (1–)3–8(–12). Eremopoa species are annual with some extreme features usually not found in Poa, but, other than abundantly branching panicles, those characteristics are broached in all cases. No one has doubted that Eremopoa was closely related to Poa.

The taxa placed in Eremopoa range from Egypt (Sinai and north coast) across the northern Middle East (Israel, Lebanon, Syria, Iraq, Turkey [Anatolia], Iran), to Afghanistan, Pakistan, northwest India (Himachal Pradesh, Kashmir), western China (Tibet and Xinjiang), north through Transcaucasia into the Caucasus mountains of Russia and across central Asia in Turkmenistan, Uzbekistan, Tajikistan, Kyrgyz Republic and Kazakhstan. Two taxa have been observed elsewhere as waifs: E. persica in western Europe (France, Norway) and E. altaica (Trin.) Roshev. in Canada (see references in Taxonomy section). The geographic region with the most diversity of Eremopoa taxa is clearly Asia Minor; nearly all of the accepted species occur in Turkey.

There have been many differences of opinion on the species and infraspecific ranks to accept in Eremopoa (Table 1). Roshevitz (1934) treated six species in his new genus in the former U.S.S.R (E. altaica, E. bellula (Regel) Roshev., E. oxyglumis (Boiss.) Roshev., E. multiradiata (Trautv.) Roshev., E. persica and E. songarica). Tzvelev (1976) reduced these six species to two species, E. persica and E. altaica, with two and three subspecies, respectively, all of which were accepted as species by Czerepanov (1981, 1995). Scholz (1980, 1981) described two new species, E. attalica H. Scholz from Turkey and E. medica H. Scholz from Azerbaijan. The type of E. medica (holotype at W, isotype at B) was determined to be a species of Puccinellia Parl. (Soreng pers. obs. 2015). Mill (1985) treated six species in Turkey, including two new species, E. capillaris R.R. Mill and E. mardinensis R.R. Mill. Rahmanian et al. (2014) accepted four species in Iran, including E. medica and E. persica with three varieties.

Classification history of Eremopoa and other taxa here accepted in Poa subg. Pseudopoa. Species and infraspecific taxa accepted by Roshevitz (1934) and authors of major floras and the region covered by their treatments are given. The last column provides the corresponding names in Poa accepted here.

Roshevits (1934) Roshevits (in Köie 1945) Bor (1970) Tzvelev (1976, 1983) Cope (1982) Mill (1985) Czerepanov (1995) Zhu et al. (2006) Gabrieljan and Oganesian (2010) Rhamanian et al. (2014) Euro+Med (on-line) Here
USSR SW Iran Iran, Afghanistan, w. Pakistan, n.w. Iraq, s. Turkmenistan, s.e. Azerbaijan USSR Pakistan Turkey USSR China (Xinjiang, Xizang) Armenia Iran Europe, Transcaucasia, Turkey, Levant, North Africa whole range
E. persica E. persica E. persica E. persica E. persica E. persica E. persica E. persica E. persica E. persica Poa persica
var. persica subsp. persica subsp. persica var. persica subsp. persica
var. major
E. multiradiata (= var. songarica) subsp. multiradiata subsp. multiradiata E. multiradiata E. multiradiata E. multiradiata (= persica var. persica) E. multiradiata subsp. multiradiata
E. altaica E. altaica E. altaica E. altaica P. diaphora E. altaica P. diaphora
subsp. altaica subsp. altaica subsp. diaphora subsp. altaica subsp. diaphora
var. diaphora
E. songarica var. songarica subsp. songarica subsp. songarica E. songarica E. songarica E. songarica var. songarica subsp. songarica var. songarica
E. bellula E. bellula (pp. = altaica, pp = songarica) (= altaica s.l.) (indirectly referenced, not accepted) (pp. = altaica, pp = songarica) E. bellula (= var. alpina?)
P. persica var. alpina (under oxyglumis) (indirectly referenced, not accepted) var. alpina
E. oxyglumis E. oxyglumis (= var. songarica) subsp. oxyglumis subsp. oxyglumis (=E. songarica) E. oxyglumis subsp. oxyglumis E. oxyglumis E. persica var. oxyglumis subsp. oxyglumis subsp. oxyglumis
E. attalica E. attalica P. attalica
E. capillaris E. capillaris P. millii
E. mardinensis E. mardinensis (= P. persica subsp. multiradiata)
E. nephelochloides E. nephelochloides (Iran) E. nephelochloides P. nephelochloides
E. medica (= Puccinellia sp.)
P. sintenisii
P. speluncarum

Bor’s genus Lindbergella (Bor 1968b, 1969) comprises a single annual species that is morphologically similar to Eremopoa. It differs from Eremopoa only in having firmer lemmas that are 3-veined and obscurely apiculate and panicles with 1–5 branches that are smooth. Lindbergella sintenisii (H. Lindb.) Bor was originally published as Poa sintenisii by Lindberg (1942) and also as P. persica var. cypria by Samuelsson (1950), the type of which is a syntype of P. persica var. alpina Boissier (1884). The species is endemic to Cyprus.

The first molecular data on Eremopoa, generated by our lab in 2004/2005, indicated that E. songarica was nested within Poa. That data was first published by Gillespie et al. (2007) using chloroplast DNA sequences from the trnT-trnL-trnF region. Based on this same data, inclusion of Eremopoa in Poa was already applied in the Flora of China account (Zhu et al. 2006, as P. subg. Pseudopoa (K. Koch) Stapf) and was continued in Gillespie et al. (2008, 2010), Soreng (2004+) and Soreng et al. (2010, 2015a, 2017a). Although nested within Poa, Eremopoa was positioned on a long branch separate from other Poa clades, justifying its recognition as a distinct subgenus, P. subg. Pseudopoa (Gillespie et al. 2007).

We published our initial DNA results for only one species of Eremopoa (E. songarica) based on trnT-trnL-trnF and, subsequently, nuclear ribosomal (nrDNA) ITS and ETS sequence data (Gillespie et al. 2007, 2008, 2010, Soreng et al. 2010). We subsequently sequenced two additional plastid regions (matK and rpoB-trnC) and added data for Eremopoa persica (Cabi et al. 2017, as Poa persica). A DNA analysis of ITS sequence data by Hoffmann et al. (2013) showed Lindbergella sintenisii was also nested within Poa near Eremopoa. Since then, we have accumulated nrDNA and plastid sequence data for most of the Eremopoa taxa and L. sintenisii and sampled many more species of Poa from Turkey and around the world. Analysis of our accumulated phylogenetic data on Eremopoa is presented here. All Eremopoa taxa were nested well within Poa, and P. speluncarum J.R. Edm. and L. sintenisii were found to be nested within or sister to the set of Eremopoa species. Here we place these taxa in Poa subg. Pseudopoa and present a taxonomic synopsis of all the species and infraspecies, as well as a key to the taxa we currently accept. Further study is needed before a comprehensive revision of the subgenus can be produced.

Methods

Collections of Eremopoa at E and G (except those not available for loan), several from P and two type specimens from BM and B were loaned to RJS at US. Other material was examined by RJS at B, K, LE, P, US and herbaria in Turkey (ANK, ISTE, NKU). Fieldwork in which 38 specimens of Eremopoa were collected by us was conducted in Kyrgyz Republic (RJS 2006) and Turkey (RJS and associates 1994, 2013, 2014, 2015; LJG & RJS and associates 2011; EC was a co-collector on the 2011 to 2015 expeditions). Additional material was obtained from R. Hand (Lindbergella sintenisii) and M. Assadi and M. Amini-Rad (Iranian Eremopoa).

The molecular phylogenetic analysis included 77 samples: 15 Eremopoa, 56 Poa, 1 Lindbergella and 5 outgroup samples (Appendix 1). A diverse set of Poa species was chosen to represent the majority of sections, including all sections in southwest Asia. Outgroup taxa were chosen to include representatives of the two taxa (Phleum L. and Milium L.) and one clade considered most closely related to Poa (Gillespie et al. 2010, Soreng et al. 2015b). Sequences of Lindbergella and the majority of Eremopoa samples, plus many matK and rpoB sequences, are new to this study (Appendix 1). For simplicity, due to the confusing taxonomy and nomenclature, we refer to Eremopoa taxa using names at the species level in the Results, trees and Appendix 1 (see Table 1 for their corresponding names in Poa). The collection TARI 135082 was previously identified as E. medica (Rahmanian et al. 2014), but was re-determined by RJS as P. persica subsp. persica.

DNA was extracted from silica gel dried or herbarium leaf material as described in Gillespie et al. (2008). Three plastid markers (matK, rpoB-trnC and trnT-trnL-trnF [TLF]) and two nuclear ribosomal DNA (nrDNA) markers (internal transcribed spacer [ITS] and external transcribed spacer [ETS]) were sequenced. Amplification and sequencing protocols, including primers used, were described in our previous studies, as follows: ITS and TLF (Gillespie et al. 2008); ETS (Gillespie et al. 2009, 2010); matK and rpoB-trnC (Soreng et al. 2015b). Sequences were assembled, edited, aligned and concatenated using Geneious ver. 6.1.5 (http://www.geneious.com). The MAFFT ver. 7.017 plugin (Katoh and Standley 2013) was used for alignment, followed by manual adjustment. All samples are complete for all markers, except for several samples with missing ends. The molecular study was conducted at the Canadian Museum of Nature; sequencing was mostly performed by NA, analyses by LJG.

Maximum parsimony (MP) analyses were performed in PAUP* 4.0b10 (Swofford 2002) using the heuristic search command with default settings, including tree bisection-reconnection (TBR) swapping, saving all multiple shortest trees (Multrees) with a maximum number set to 100,000. Branch support was assessed using MP bootstrap analyses performed in PAUP* with heuristic search strategy, 10,000 bootstrap replicates, each with ten random addition sequence replicates, saving ten trees per replicate.

Bayesian Markov chain Monte Carlo analyses were conducted in MrBayes (Ronquist et al. 2011). Optimal models of molecular evolution for individual markers were first determined using the Akaike information criterion (AIC; Akaike 1974) conducted through likelihood searches in jModeltest with default settings (Darriba et al. 2012). Models were set at GTR + Γ for ITS, ETS and rpoB-trnC partitions and GTR + I + Γ for matK and TLF partitions based on the AIC scores and the models allowed in MrBayes. Two independent runs of four chained searches were performed for either two or three million generations (analyses were stopped when split frequency of 0.005 was reached or closely approached), sampling every 500 generations, with default parameters. A 25% burn-in was implemented prior to summarising a 50% majority rule consensus tree and calculating Bayesian posterior probabilities (pp).

MP heuristic searches and bootstrap analyses were performed initially on the separate marker alignments. Strict consensus trees were examined for conflicting topologies with incongruence identified by branch conflicts with ≥75% bootstrap support (BS). No supported incongruence was found between ITS and ETS trees, nor amongst the three plastid trees. Further MP and Bayesian analyses were performed on the separate concatenated nrDNA (77 samples, 1251 aligned characters) and plastid (77 samples, 4465 characters) alignments. Since supported incongruence was detected between the nrDNA and plastid strict consensus trees, species and clades determined to be incongruent were removed prior to performing analyses on the concatenated combined nrDNA and plastid alignment (68 samples, 5599 aligned characters). Trees were viewed in FigTree v1.4.0 (Rambaut 2006+). Clade designations follow Soreng et al. (2010) with modifications as in Cabi et al. (2017) and Soreng et al. (2017b), wherein well-supported major clades are assigned letters.

Results

Plastid and nrDNA Bayesian trees are given in Fig. 1 with summary statistics in Suppl. material 1. There are 100 new sequences reported in GenBank and these are given in Appendix 1. MP trees (bootstrap values shown below branches in Fig. 1) were very similar to the Bayesian trees with a few minor unsupported differences. Major clades (shown by letter and colour in Fig. 1) are identical in both nrDNA and plastid trees, with two exceptions: Poa arctica R. Br. and P. sect. Secundae members (P. curtifolia Scribn., P. secunda J. Presl and P. stenantha Trin.), each belonging to different major clades in the two trees. The position of three major clades differs significantly between the nrDNA and plastid trees: J clade (sect. Jubatae: P. jubata A. Kern.), S clade (sects. Stenopoa and Abbreviatae) and V clade (sect. Pandemos: P. trivialis L.). Poa major clades have been described elsewhere (Gillespie et al. 2007, 2008, 2009, Soreng et al. 2010, 2017b, Cabi et al. 2017); here we focus on the position of Eremopoa.

Figure 1. 

Poa nrDNA and plastid Baysian analyses showing placement of Eremopoa and Lindbergella. Bayesian 50% majority rule consensus trees of nrDNA ITS and ETS (left) and plastid data (trnT-trnL-trnF, matK and rpoB-trnC) (right). Bayesian posterior probabilities are shown above branches, MP bootstrap values below branches. Outgroups are not shown. Major clades are indicated by colour and capital letter. Taxa shown in black belong to different major clades in plastid and nrDNA trees.

Eremopoa species, together with Lindbergella sintenisii and Poa speluncarum, form a clade (E clade) in both nrDNA and plastid trees, but are strongly supported only in the plastid analysis (pp = 1, BS = 99%). All E. multiradiata, E. oxyglumis, E. persica and E. songarica accessions form a strongly supported clade (core Eremopoa clade) in both trees (pp = 1, BS = 100%). In the plastid analysis E. attalica, L. sintenisii and P. speluncarum form a strongly supported clade (pp = 1, BS = 100%), with L. sintenisii sister to E. attalica (pp = 1, BS = 96%). In the nrDNA tree, E. attalica and P. speluncarum are sister taxa (pp = 0.99, BS = 77%) and Lindbergella is weakly supported as sister to this clade plus the core Eremopoa clade (pp = 0.97, BS = 59%). Within the core Eremopoa clade, all E. oxyglumis and E. songarica samples form a strongly supported clade in the nrDNA analysis (pp = 1, BS = 100%), whereas in the plastid analysis, these samples are divided between two strongly supported clades corresponding to E. oxyglumis plus one E. songarica sample (IRAN 20357, identification needs confirmation) (pp = 1, BS = 89%) and all remaining samples of E. songarica (pp = 1, BS = 100%). Eremopoa multiradiata and E. persica samples do not form a clade in either analysis, although all except one (E. persica, Soreng 9215) are strongly supported as a clade (pp = 1, BS = 95%) in the plastid tree.

The combined nrDNA and plastid Bayesian tree with proportional branch lengths is shown in Fig. 2. Prior to analysis, species and clades with positions incongruent (branch conflicts with ≥ 75% BS) between the nrDNA and plastid trees were removed, including Lindbergella sintenisii, P. arctica, P. sect. Secundae species and the J, S, and V clades. The E clade is strongly supported, as are its two subclades, E. attalica-P. speluncarum and the core Eremopoa clade (all pp = 1, BS = 100%). Both subclades are on long branches and separated by considerable genetic distance. The core Eremopoa clade is subdivided into two strongly supported clades: E. multiradiata-E. persica (pp = 0.99, BS = 96%) and E. oxyglumis-E. songarica (pp = 1, BS = 94%). Eremopoa oxyglumis and three of four accessions of E. songarica each form moderately or strongly supported clades (pp = 1, BS = 86%; pp = 1, BS = 100%, respectively).

Figure 2. 

Poa combined nrDNA and plastid Baysian analysis showing placement of Eremopoa. Bayesian 50% majority rule consensus tree of combined nrDNA (ITS and ETS) and plastid data (trnT-trnL-trnF, matK and rpoB-trnC). Bayesian posterior probabilities are shown above branches, MP bootstrap values below branches. Major clades are indicated by colour and capital letter; outgroups are shown in black.

In the combined nrDNA and plastid tree (Fig. 2), the E clade is strongly supported as sister (pp = 1, BS = 100%) to a clade comprising Poa supersects. Homalopoa (H clade) and Poa (P clade) and the N clade (P. sect. Nanopoa plus unassigned species). In the nrDNA analysis, the E clade is strongly supported as sister to clades P+H (not differentiated), N, and X (represented here by P. arctica) (Fig. 1). In the plastid analysis, the E clade is sister to a larger clade comprising clades H, N, and P, plus J, S and V (Fig. 1).

Discussion

Our molecular analyses of plastid and nuclear ribosomal DNA strongly support the position of Eremopoa and Lindbergella within the genus Poa. Eremopoa and Lindbergella were united in a clade along with Poa speluncarum with strong support in the plastid and combined trees (weak support in the nuclear tree). We call this set the E clade (Soreng et al. 2010, Cabi et al. 2017) and accept it as Poa subg. Pseudopoa. In its recent usage, this subgenus was initially considered to include only Eremopoa (Zhu et al. 2006, Gillespie et al. 2007); here it is expanded to include Lindbergella and P. speluncarum.

Within the E clade, three taxa of southwest Turkey and Cyprus, E. attalica, P. speluncarum and Lindbergella sintenisii, are phylogenetically isolated from all the other species of Eremopoa sampled (the core Eremopoa clade). All three taxa formed a strongly supported clade in the plastid tree, while in the nuclear tree only the first two species form a clade and L. sintenisii is sister to this clade plus the core Eremopoa clade. The position of L. sintenisii is moderately supported as incongruent between the nuclear and plastid trees suggesting that the genus may be of hybrid origin; however, further studies are needed to confirm incongruence over lack of support.

All Eremopoa taxa sampled, excluding E. attalica, form a strongly supported clade in all trees, called here the core Eremopoa clade. This clade includes two strongly supported subclades in the combined nuclear-plastid tree, corresponding to E. persica s.l. and E. altaica s.l. In the first subclade, E. multiradiata is nested amongst E. persica samples, as is the sample originally determined as E. medica (TARI 35082). The E. multiradiata sample (Soreng 9240) comes from the type locality of E. mardinensis in SW Turkey and is a good match for that species, but we believe that E. mardinensis should be treated as a synonym of E. multiradiata. The E. altaica s.l. subclade in the combined tree includes a strongly supported and divergent clade of three E. songarica samples and a clade of E. oxyglumis plus one sample of E. songarica (identification needs confirmation). The position of E. songarica (tetraploid) with E. oxyglumis (diploid and hexaploid) is strongly supported in the combined and nuclear trees, but is weakly supported with E. persica (diploid) in the plastid tree. This, together with ploidy level, is suggestive of a possible hybrid origin for E. songarica, but this hypothesis needs to be further explored.

As noted in the introduction and Table 1, there has been no consensus on the taxonomy of Eremopoa species. Bor (1970, p. 49) wrote “As far as the genus Eremopoa Roshev. is concerned I am prepared to accept two species only: Eremopoa persica (Trin.) Roshev. and E. bellula (Regel) Roshev.” He considered E. songarica, multiradiata and oxyglumis “only worthy of varietal rank” as the single taxon, E. persica var. songarica. Tzvelev (1976), Cope (1982) and Mill (1985) dismissed the E. bellula form as indistinct, yet it was maintained as a species by Bor (1970) and Rahmanian et al. (2014). As such, the array of taxa has been treated as a series of species, subspecies or varieties. The taxonomy proposed by Tzvelev (1976) seems the most useful for treating E. persica s.l. and E. altaica s.l.; each is treated as a separate species with subspecies. His classification, supported by molecular data, is adopted here with some minor modifications.

Here, we present a synopsis of P. subg. Pseudopoa based on our current understanding. Further herbarium and molecular study is needed before a more comprehensive revision of the subgenus can be produced. We treat all Eremopoa species, Lindbergella sintenisii and P. speluncarum in P. subg. Pseudopoa. We merge all Eremopoa taxa and L. sintenisii into Poa and treat the Eremopoa taxa as five species. Poa diaphora Trin. is the correct name for E. altaica within Poa. Two subspecies, subsp. diaphora and oxyglumis (Boiss.) Soreng & G.H. Zhu, are recognised in P. diaphora based in part on their mostly clear separation in the plastid analyses and morphological distinctions. Subspecies diaphora includes three difficult to distinguish varieties: var. diaphora (formerly E. altaica), var. alpina and var. songarica (formerly E. songarica). Poa persica includes two subspecies and is clearly separated from both P. diaphora subspecies in the analyses. Most Eremopoa taxa already have names in Poa or the epithets used in Eremopoa are available in Poa (with one exception).

Taxonomy

Poa subg. Pseudopoa (K. Koch) Stapf in J. D. Hooker, Fl. Brit. India 7(22): 337. 1897 [1896].

Festuca [unranked] Pseudopoa K. Koch, Linnaea 21(1[4]): 409. 1848. Poa sect. Pseudopoa (K. Koch) Hack., Nat. Pflanzenfam. 2(2): 73. 1887. Eremopoa Roshev., Fl. URSS 2: 429, 756. 1934. Type. Poa persica Trin. ≡ Festuca persica (Trin.) K. Koch.

Lindbergia Bor, Svensk Bot. Tidskr. 62: 467, 1968 (nom. illeg. hom., non Kindb., 1897). Lindbergella Bor, Svensk Bot. Tidskr. 63: 368. 1969. Type. Poa sintenisii H. Lindb. ≡ Lindbergella sintenisii (H. Lindb.) Bor.

Emended diagnosis

Like species of other Poa subgenera, but annual (P. speluncarum a weak stooling perennial) and differing from other annual species of Poa by combination of sheath margins fused only near the base (basal sheaths fused to 16%, top sheath 4–12% [to 50% in P. speluncarum]), panicle branches scabrous along angles (P. sintenisii smooth), arranged in whorl-like groups of 5 to 27 per node (sometimes fewer in P. diaphora and P. sintenisii), sometimes the lower whorls of branches naked or with only a few sterile spikelets, flowers bisexual, glumes short (lower glume 2/7–2/3 (–3/4) the first lemma in length), 1-veined (3-veined in P. sintenisii), apex sharply pointed, sometimes apiculate, rachilla internodes exposed, scaberulous, callus glabrous (or with a short crown of hairs in P. sintenisii), lemmas membranous to subchartaceous (P. sintenisii chartaceous), 3–5 veined, the intermediate veins faint when present, laterally compressed, but the keel not pronounced, glabrous or keel and marginal veins short sericeous (also sericeous between the veins in P. sintenisii), but keel scabrous distal to the hairs.

Distribution

Southwest Asia from Israel, Lebanon, Cyprus and Turkey eastwards through Transcaucasia, Iran, central Asia to western China and northwest India. Sporadic elsewhere, possibly adventive on Egypt’s North African coast but native east of the Red Sea, adventive in Europe and Canada.

Notes

A subgenus of seven species with several infraspecies, distributed mainly in semi-arid midlands to uplands (usually 300 m plus) to alpine, with winter spring / summer drought precipitation pattern, often along trails and roads, cultivated fields and pastures, around puddles, shallow springs, swales and vernal pools, snow beds, in pine/oak forests to open grasslands and deserts, also in shallow caves, in shallow sandy or stony soils or screes of igneous or metamorphic rocks of igneous or sedimentary origin, including pumice, lava, serpentine, shale, sandstone, limestone and marble.

Key to Poa subgen. Pseudopoa taxa and other annual species of Poa in the coincident geographic region

Plants annual (infrequently perennial or perenniating); anthers mostly 0.2–1 mm (to 1.7 mm in the weak stemmed, stooling perennial P. speluncarum, to 2.8 mm in the annual species Poa persica).

1 Palea keels soft hairy, never scabrous; callus glabrous (Poa sect. Micrantherae) 2
Palea keels scabrous at least in part (if hairy in part, then distally scabrous); callus glabrous or hairy 3
2 Anthers 0.2–0.5 mm long; panicle branches ascending, spikelets congested along the branches; plants light green Poa infirma Kunth
Anthers 0.5–1 mm long; panicle branches spreading to ascending, spikelets moderately congested along the branches; plants darker green Poa annua L.
3 Spikelets ovate; lemma keels densely villous medially, many hairs over 0.5 mm long; callus with a plicate web; anthers 0.4–0.8 mm long; panicles short (to 5 cm long), branches terete, smooth or sparsely scabrid, with 1–2 branches per node; upper culm sheath margins fused 25–35(–50)% their length; plants of vernal swales, Albania, Croatia, Greece, Bulgaria and European part of Turkey (Poa sect. Jubatae) Poa jubata
Spikelets generally lanceolate; lemma keels glabrous or sericeous, hairs less than 0.3(–0.5) mm long; callus glabrous or with a short crown of hairs; anthers 0.2–2.8 mm long; panicles short or long, branches angled, smooth or scabrous, mostly with 2 to 27 branches per node, commonly appearing whorled; upper culm sheath margins fused 4–12% their length (40–50% in P. speluncarum); plants of Cyprus, Anatolian Turkey, southwards and eastwards across Asia into China (Poa subg. Pseudopoa, incl. Eremopoa) 4
4 Uppermost culm sheath margins fused 40–50% their length; spikelets mostly 1-flowered; lemmas glabrous; callus glabrous; anthers 1.1–1.7 long; plants feeble, stooling perennials of caves and shady cool moist places in the Taurus Mts. of Turkey (rare) (Poa sect. Speluncarae) Poa speluncarum
Uppermost culm sheath margins fused 4–12% their length; spikelets (1–)2 to 10-flowered; lemmas glabrous or pubescent; callus glabrous or with a minute crown of hairs; anthers 0.2–2.8 mm long; plants slender tufted annuals 5
5 Lemmas 3-veined, apex slightly apiculate, lemmas and paleas subcoriaceous, sericeous along the keel(s) and marginal veins and between the veins; panicle branches smooth, mostly 1–5 at lower nodes; callus glabrous or with a short crown of hairs; plants endemic to Cyprus (usually on serpentine substrates) (Poa sect. Lindbergella) Poa sintenisii
Lemmas 5-veined (veins commonly faint), apex infrequently apiculate, lemmas and paleas subchartaceous to subcoriaceous, glabrous between the veins or throughout; panicle branches scabrous, (1–)5–27 at lower nodes; callus glabrous; plants widespread, but not in Cyprus (Poa sect. Pseudopoa) 6
6 Panicles with 1 to 3 lower whorls of 7 or more sterile/naked or mostly sterile branches; panicles 7–20 cm long, effusely branched; lemmas 2–2.5 mm long, sericeous along the keel and marginal veins; spikelets 1–4(–6)-flowered 7
Panicles not or infrequently with some sterile lower branches; panicles 2–21 cm long, effusely to sparsely branched; lemmas 1.8–4.5 mm long, glabrous or sericeous along the keel and marginal veins; spikelets 1–12-flowered 8
7 Anthers 1.1–1.5 mm long; ligules 1.5–2.5 mm long; branches 7–20 per lower whorl; spikelets 1–4(–6)-flowered; plants of Zagros Mts., Iran P. nephelochloides
Anthers 0.8–1 mm long; ligules 1–1.5 mm long; branches 7–15 per lower whorl; spikelets 1–3-flowered; plants of Taurus Mts., Turkey P. attalica
8 Anthers (1.2–)1.4–2.8 mm long; lemma apex blunt or obtuse to acutely pointed, with a broad membranous margin (P. persica s.l.) 9
Anthers 0.2–1.3 mm long; lemma apex acute or narrowly acute to acuminately pointed, with a narrow membranous margin (blunt or slightly pointed in P. millii but then with 13–27 branches at lower panicle nodes) 10
9 Lemmas all glabrous or rarely with a few hairs near the base of the keel or marginal veins; spikelets (4–)5–10(–12)-flowered; panicles usually ¼–½ the plant height; anthers 1.5–2.8 mm long P. persica subsp. multiradiata
Lemmas (at least of the lowest flower in a spikelet) minutely sericeous along the keel and marginal veins for ¼–⅔ the length; spikelets (2–)3–7(–9)-flowered; panicles usually ⅖–⅔ the plant height; anthers (1.2–)1.4–1.8 mm long P. persica subsp. persica
10 Anthers mostly 0.2–0.6 mm long; lemmas 1.8–4.5 mm long, apex sharply pointed, usually glabrous, infrequently sparsely puberulent along the keel with one or a few soft hairs scattered near the base; spikelets (1–)2–3(–5)-flowered; plants 2–40 cm tall 11
Anthers 0.6–1.3 mm long; lemmas 2.3–3 mm long, apex acute and sharply pointed to obtuse and blunt, at least the lowest lemma in a spikelet evenly sericeous (hairs ca. 0.1–0.3(–3.5) mm long, stiff, appressed) along the keel in the proximal ¼–½ and along the marginal veins near the base; spikelets 3–5(–9)-flowered; plants mostly 15–60 cm tall 13
11 Lemmas 3.5–4.5 mm long; panicles (2–)3–8(–9) cm long, branches 1–5(–7) at lower nodes, divaricately rebranched and relatively stout, spikelets usually sparse and few; plants mostly 5–25(–30) cm tall P. diaphora subsp. diaphora var. diaphora
Lemmas 1.8–3.5(–3.8 in large specimens with many spikelets) mm long; panicles 2–15(–20) cm long, branches (1–)3–8(–12) at lower nodes, divaricately rebranched or not, capillary to somewhat stout, spikelets sparse to crowded, few to many; plants 2–40 cm tall 12
12 Plants low growing, with dense fascicles of rebranching culms; culms 2–6 cm tall, with lateral inflorescences from lower culm leaves; panicles contracted to open, 1.5–4 cm long, included in tuft of basal leaves or slightly exerted; lemmas 3–3.5 mm long; plants alpine P. diaphora subsp. diaphora var. alpina
Plants low growing or taller, without fascicles of rebranching culms; culms solitary to several, mostly 10–40 cm tall, without lateral inflorescences; panicles effuse, usually more than 5 cm long, usually exerted; lemmas 1.8–3.5 mm long; plants of various habitats P. diaphora subsp. diaphora var. songarica
13 Spikelet pedicels mostly 2–5 mm long; panicle branches 5–18 at lower nodes, stiffly spreading, lower whorls never naked or with rudimentary spikelets; lemma apices acutely pointed; anthers 0.6–1.0(1.1) mm long P. diaphora subsp. oxyglumis
Spikelet pedicels mostly 5–10 mm long: panicle branches (9–)13–27 at lower nodes, slender, slightly flexuous, lower whorls sometimes with a few branches that are naked or with some rudimentary spikelets in addition to normal spikelets; lemma apices obtuse to acute, blunt or slightly pointed; anthers 0.8–1.3 mm long P. millii

Poa subg. Pseudopoa sect. Pseudopoa (K. Koch) Hack., Nat. Pflanzenfam. 2(2): 73. 1887.

Emended description

Tufted annuals. Leaf sheaths keeled, margins fused for 4–12% their length; blades flat to convolute, surfaces scabrous. Panicles open, with (1–)3–27 branches at lower nodes, lower whorls sometimes sterile; branches ascending to widely spreading, scabrous angled, with pedicels mostly equalling or up to 4× longer than their spikelets. Spikelets 1–10-flowered; glumes unequal, 1st glume 1-veined, 2nd glume 3-veined, usually reaching to less than ⅔ the adjacent lemma; rachilla internodes terete, scabrous; callus smooth, glabrous, with a round disarticulation scar; lemmas laterally compressed, weakly keeled, glabrous or short sericeous in lower half of the keel and also along the marginal veins, between veins smooth or scabrous, glabrous (rarely sericeous), 5-veined, intermediate veins obscure to distinct, margins narrowly to broadly scarious, apex obtuse to acuminate, sometimes briefly muticus. Flowers perfect, ovaries glabrous, anthers 0.2–2.8 mm long; caryopsis 1.5–2.5 mm long, narrowly elliptical, laterally compressed, fused to the palea, solid, hilum ⅛–⅙ the grain in length.

Poa attalica (H. Scholz) Soreng, Cabi & L.J. Gillespie, comb. nov.

Eremopoa attalica H. Scholz, Willdenowia 10(1): 33, f. 1. 1980.

Type

Turkey. Antalya, “nordwestl. Antalya bei Termessos, ausgetrockneter Gebirgsbach”, 300 m, 23 Jul 1979, Kehl s.n. (holotype: B! [B-100272775])

Distribution

Turkey (western Taurus Mts.).

Notes

We provisionally retain this species in sect. Pseudopoa, despite its divergent phylogenetic placement. The species is morphologically similar to other members of the section. As noted by Mill (1985), it is most like Poa nephelochloides Roshev., but the anthers are smaller. Some populations of P. millii approach P. attalica and are problematical to separate (see under P. millii). Further molecular study is needed to determine if the three species are closely related and if a new section is warranted.

Poa diaphora Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math., Seconde Pt. Sci. Nat. 4,2(1): 69–70. 1836.

Aira altaica Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg Divers Savans 2: 526. 1835. Nephelochloa altaica (Trin.) Griseb., Fl. Ross. 4(13): 367. 1852. Poa diaphana Boiss., Fl. Orient. 5: 611. 1884, nom. inval. Eremopoa altaica (Trin.) Roshev., Fl. URSS 2: 431. 1934.

Type

“Sterilissimus salsuginosis deserti editi Tschujae”, [1800–3000 m], July 1832, A. Bunge (lectotype, designated by Tzvelev 1976, pg. 480, and marked in herbarium: LE! [Trinius herbarium microform image 424-A4! p.p. Bunge 1832]; isolectotypes: LE [3 specimens: TRIN-2620.01! with original description (Trinius herbarium microform 312-A1), Trinius herbarium microform images 424-A3!, 424-A5!], K [K000789849 image!; specimen labelled “Aira altaica Trin. Altai”, “Acad. St. Petrop, mis. 8br 1835” is a good match for LE type material]). See Soreng et al. (1995) for explanation of Trinius herbarium citations.

Distribution

Egypt (Sinai Peninsula) to China (Xinjiang, Xizang).

Notes

Separating the four forms of Poa diaphora s.l. treated here is often difficult. Here we choose to recognise two subspecies as divided in the molecular plastid analysis. Subspecies diaphora and oxyglumis are most easily separated by the minute anthers (0.2–0.6 mm) combined with glabrous or nearly glabrous lemmas in the former and slightly longer anthers (0.6–1.1 mm) combined with hairy lemma keels and marginal veins in the latter. The other forms, diaphora s.s., songarica and alpina are essentially intergrading and are here treated as varieties in subsp. diaphora.

The specimen K000789848 (image!) (“Al. Bunge” ex hrbr. Alexandri Lehmann, Reliquiae botanicae, original det “Poa diaphora Tr.”) might be original material of Aira altaica, but RJS doubts it as it is not a good match for LE types; it is a taller plant more like K00789847 (also Reliquiae Lehmannianae), which is Bunge material collected 20 May 1842, in Karakum desert.

Poa diaphora subsp. diaphora var. diaphora

Fig. 3A

Poa persica var. diaphora (Trin.) Asch. & Graebn., Syn. Mitteleur. Fl. 2: 437. 1900. Eremopoa altaica (Trin.) Roshev. subsp. altaica.

Distribution

China (Xinjiang, Xizang), Kazakhstan, Kyrgyz Republic, Pakistan, Russia (Altai Mts.), Tajikistan, Turkey.

Notes

A single specimen recorded from Turkey (Kars Prov., Litvinov 4790 US ex K) evidently belongs to this variety and was also cited by Mill (1985) under E. songarica.

Figure 3. 

Poa subgenus Pseudopoa sect. Pseudopoa. A P. diaphora subsp. diaphora var. diaphora, Chu, Kyrgyz Republic (Soreng et al. 7537) B, C P. persica subsp. persica, Adiyaman, Turkey (Soreng et al. 9215) B habit C closeup of base of plant showing keeled leaf sheaths and caniculate blades D, E P. persica subsp. multiradiata, Mardin, Turkey (Soreng et al. 9240) D habit E spikelet showing glabrous lemmas. Photos by R.J. Soreng.

Poa diaphora subsp. diaphora var. alpina (Boiss.) Soreng, Cabi & L.J. Gillespie, comb. nov.

Poa persica var. alpina Boiss., Fl. Orient. 5: 610. 1884.

Type

Turkey. Plantae Lyciae, ad fonts reginis alpinae montis Elmalu, 25 Jun 1860, E. Bourgeau 271 (lectotype, here designated: G [G00330280 image!]; isolectotypes: G [G00380172 image!, p.p. central and right top two samples], G [G0038173 image!], K [K-000789856 image!]).

Distribution

Armenia, Azerbaijan, Afghanistan, Georgia, Iran, Kyrgyz Republic, Pakistan, Turkey and Turkmenistan(?).

Notes

This taxon, accepted as Eremopoa bellula by several authors (see Names of Uncertain Application below), was first recognised infraspecifically by Boissier (1884) as Poa persica var. alpina. The variety is common in the highest elevations at which the genus occurs, in the alpine of Turkey, Iran and Afghanistan to the Pamir mountains, reaching 4000 m. Further study is needed to clarify the distinction of var. alpina from var. diaphora and these from Eremopoa bellula, as the material placed here appears heterogenous.

Of the six syntypes of var. alpina cited by Boissier (Bourgeau 271, hab. in alpinis, montes supra Elmali Lyciae [G00380172, G0038173, G00330280, K000789856]; Kotschy 12, Tarus Cilicicus, 5–6000’; Prairies humides de la region alpine du Taurus, au Boulgarmden [as 12d: G00330281, K000789851 image!]; Balansa s.n., Jul-Aug 1855 [K000789857, P02358251 p.p. bottom right]; Blanche s.n., Libani cacuminal; Kotschy 477, mons Kuh Delu Persiae australis, 10 Jun 1842 [BM000959359 image!, E!, G00308632 image!, P02358251! p.p. “fo. pygmaea” bottom left]), we select Bourgeau 271 as the lectotype as it is typical of the form. As noted by Samuelsson (1950), the Sintenis syntype (mons Troodos, Cypri) represents a separate form that is treated here as Poa sintenisii. Poa persica var. “minor” Boiss. (cited by Mill, in Fl. Turkey 9: 492. 1985) is a nomen nudum since it is a herbarium name on Bourgeau 271, syntype of var. alpina Boiss.; this name is also inscribed on Kotschy 12d (p.p. G00308174), but the latter is original material, not a syntype, mentioned by Boissier (1884).

Poa diaphora subsp. diaphora var. songarica (Schrenk ex Fisch. & C.A. Mey.) Soreng, Cabi & L.J. Gillespie, comb. nov.

Glyceria songarica Schrenk ex Fisch. & C.A. Mey., Enum. Pl. Nov. 1: 1–2. 1841. Nephelochloa songarica (Schrenk ex Fisch. & C.A. Mey.) Griseb., Fl. Ross. 4(13): 367. 1852. Nephelochloa persica var. songarica (Schrenk ex Fisch. & C.A. Mey.) Regel, Trudy Imp. S.-Peterburgsk. Bot. Sada 7: 603. 1881. Poa songarica (Schrenk ex Fisch. & C.A. Mey.) Boiss., Fl. Orient. 5: 611. 1884. Poa persica var. songarica(Schrenk ex Fisch. & C.A. Mey.) Stapf, Fl. Brit. India 7(22): 337. 1897 [1896]. Eremopoa songarica (Schrenk ex Fisch. & C.A. Mey.) Roshev., Fl. URSS 2: 431, pl. 32, f. 11. 1934. Eremopoa persica var. songarica (Schrenk ex Fisch. & C.A. Mey.) Bor, Grass. Burma, Ceylon, India & Pakistan 532. 1960. Eremopoa altaica subsp. songarica (Schrenk ex Fisch. & C.A. Mey.) Tzvelev, Bot. Zhurn. (Moscow & Leningrad) 51(8): 1104. 1966. Poa diaphora subsp. songarica (Schrenk ex Fisch. & C.A. Mey.) Soreng & G.H. Zhu, Fl. China vol. 22: 266. 2006. Poa songarica var. argaea Hausskn. & Bornm. ex R.R. Mill, Fl. Turkey & E. Aegean Isl. 9: 491. 1985, nom. inval., as syn. of Eremopoa songarica.

Poa paradoxa Kar. & Kir., Bull. Soc. Imp. Naturalistes Moscou 864. 1841, nom. illeg. hom. Poa subtilis Kar. & Kir., Bull. Soc. Imp. Naturalistes Moscou 15(3): 532. 1842. nom. nov. (cited Poa paradaxa Kar. & Kir., 1941 [entry no.] 926). Type protologue. Hab. in herbosis ad rivulum Ai deserti Soongoro-Kirghisici, Jun, Karelin & Kiriloff. Type: Hab. in herbosis ad rivulum Ai deserti Soongoro-Kirghisici, Jun 1840, Karelin & Kiriloff (Herb. Fischer no. 504) (lectotype, here designated: LE!; isotypes: P [P02663383!], W [W0028251 image!]).

Type

Ad fl. Karatal versus montes Karatau, 13 June 1840, H. Schrenk s.n. (holotype: LE; isotype: LE).

Distribution

Afghanistan, Armenia, Azerbaijan, China (Xizang), Georgia, Iran, Israel, Kazakhstan, Kyrgyz Republic, Tajikistan, Turkey, Turkmenistan and Uzbekistan.

Notes

Poa diaphora var. songarica was recently recorded (as Eremopoa songarica; determination verified here) from one locality in northernmost Israel (Danin and Fragman-Sapir 2016+). It was collected as a waif in Canada (Manitoba) in the 1950s (Stevenson 1965, as E. persica; Darbyshire 2007, as E. altaica: re-identified here), but is apparently not persistent (Darbyshire 2007, B.A. Ford, pers. comm, 2018).

Tzvelev (1976, pg. 480) cited “In herbidis Songaria ad rivulum Tschulak [Jun 1841], Karelin & Kiriloff 2123” (LE!) as type of P. subtilis (duplicates at BM000959360 image!, K000789846 image!, BR0000006600860 image!, P02663388!, P02663405!), but the type is the one [1840] collection cited by Karelin and Kiriloff (1841) distributed as Herb. Fischer no. 504.

Poa diaphora subsp. oxyglumis (Boiss.) Soreng & G.H. Zhu, Fl. China 22: 266. 2006.

Poa persica var. oxyglumis Boiss., Fl. Orient. 5: 610. 1884. Eremopoa oxyglumis (Boiss.) Roshev., Fl. URSS 2: 430, 756, pl. 32, f. 9–10. 1934. Eremopoa persica var. oxyglumis (Boiss.) Grossh., Fl. Kavkaza (ed. 2) 1: 268. 1939. Eremopoa altaica subsp. oxyglumis (Boiss.) Tzvelev, Bot. Zhurn. (Moscow & Leningrad) 51(8): 1104. 1966. Eremopoa persica var. oxyglumis (Boiss.) Rahmanian, Iran. J. Bot. 21(11): 214. 2014. nom. inval. isonym.

Type

Turkey. In collibus prope Baibout, 17 Jul 1963, E. Bourgeau (lectotype, designated by Tzvelev 1976, pg. 479: LE! [LE00009676]; isolectotypes: LE [LE00009678 image!], P [P02358146! pp a, P03142400!]).

Distribution

Armenia, Azerbaijan, Georgia, China (Xizang), Kyrgyz Republic, Pakistan, Tajikistan, Turkey, Turkmenistan and Uzbekistan.

Notes

Most accounts have recognised this taxon at one rank or another, except Mill (1985) who treated it as a synonym of E. songarica. Several collections were cited in the original protologue: Tchihatcheff, Hab. in Ponto; Balansa, Ponto Lazico ad Djimil [Balansa 1549 G00308631, E, LE!, P02014318 (= subsp. oxyglumis), P02014317 (= P. persica subsp. multiradiata), US!]; Huet, Erzurum [G00330279, G00308633]; E. Bourgeau, Armenia, in collibus et agris in cultis Armeniae Turcicae ad Gumuchkhane.

Poa millii Soreng, Cabi & L.J. Gillespie, nom. nov.

Eremopoa capillaris R.R. Mill, Fl. Turkey & E. Aegean Isl. 9: 624, 490. 1985 (non Poa capillaris L. 1753). Eremopoa persica var. ramosissima Azn. ex R.R. Mill, Fl. Turkey & E. Aegean Isl. 9: 490. 1985, nom. inval.

Type

Turkey. Adana, distr. Feke, Sencan Dere nr Gurumze, 1300 m, 30 May 1952, P.H. Davis, Dodds & Cetic 19681 (holotype: E! [E00196495]; isotypes: BM! [BM000959355], K! [K000789852]).

Distribution

Turkey (central and eastern Taurus Mts. and adjacent ranges).

Notes

Morphologically Poa millii is intermediate between P. persica subsp. persica and P. attalica. However, we are not sure which of these it is actually related to or if it is a hybrid between them. The type approaches P. persica in having anthers 1.2–1.3 mm long and P. attalica in having abundant branching and sometimes having some sterile branches amongst the lower branch whorls. Much of the material of P. millii from further west than the type location from the Taurus Mts. has smaller anthers and is problematical to separate from P. attalica.

Poa nephelochloides (Roshev.) Soreng, Cabi & L.J. Gillespie, comb. nov.

Eremopoa nephelochloides Roshev., in Köie, M., Beitr. Fl. Sudwest Iran I. Danish Sci. Invest. Iran In K. Jessen & R. Sparck. (Eds) Danish Sci. Invest. Iran, pt. 4: 50. 1945. Eremopoa persica var. nephelochloides Roshev., nom. inval. as syn. of E. nephelochloides.

Type

Iran. 60 km north of Dizful, 3 May 1937, M. Köie 475 (lectotype, here designated: C [C10016935 image!]; isolectotype: LE).

Distribution

Iran (Zagros Mts.).

Notes

Due to its sterile whorls of branches, this species seems very close to Poa millii and P. attalica, but may be a derivative of P. persica since it has longer anthers than the previous taxa. Roshevits cited two gatherings of Köie: “Kechwar, 700 m (3 May 1937; no. 475). Chah-Bazan, 500 m” (Kechvar is about 60 km north of Dizful). The specimen at C has the same date and collection number as Roshevits cited and was annotated by Roshevits as this taxon; we select it as the lectotype. The anthers are ca. 1.1–1.2 mm as measured from the C photo and other characters seem to match P. attalica. The anther length is given as 1.5 mm in Roshevits’ diagnosis. The specimen clearly has the hyaline lemma apices of P. persica s.l. (in contrast to P. diaphora). However, these features are also present in the type of E. capillaris (=P. millii). Poa attalica has shorter anthers, ca. 0.8 to 1 mm, on the type (anthers not described by Scholz 1980 or Mill 1985). Poa nephelochloides and P. attalica may represent the same species, diagnosed as different from P. persica by sterile branches and from Nephelochloa orientalis Boiss. by glabrous lemmas (P. nephelochloides has pubescent lemmas). However, Poa nephelochloides and P. attalica are geographically isolated by over 1500 km and have different anther lengths.

Poa persica Trin., Mém. Acad. Imp. Sci. St.-Pétersbourg, Sér. 6, Sci. Math. 1(4): 373. 1830.

Festuca persica (Trin.) K. Koch, Linnaea 21(1[4]): 410. 1848. Nephelochloa persica (Trin.) Griseb., Fl. Ross. 4(13): 366. 1852. Poa pamphylica Boiss., Diagn. Pl. Orient., ser. 1, 13: 58. 1854[1853?], nom. inval. as syn. of Poa persica. Eremopoa persica (Trin.) Roshev., Fl. URSS 2: 430, pl. 32, f. 8. 1934.

Type

Iran: in collibus ad Akar-Tschai prob. Karabagh, 1400–1900 m, 27 May 1829, Szowits 246 (lectotype, designated by Tzvelev 1976, pg. 479: LE! [photo E000327964!, TRIN-microform 434-B4!]; isolectotypes: LE [TRIN-2666.02!, TRIN-microform 434A8!, 434-B1!, 434-B2!, 434-B3!]).

Notes

Other original material includes: Iran, Prov. Aderbeidschan. distr. Khoi, ad Seidchadzi, 18 May 1828, Szovits 258 (LE!, LE0009678 [image!], LE0009679, LE0009680 [image!], LE0009681 [image!], W0028250 [image!]; In apricis prov. Aderbeidschan e Karabahg, Fischer [herb. Fischer] (K000789867 [image!]). Poa persica has two major variations: subsp. persica with pubescent lemmas and relatively narrower panicle length to plant height ratio; and subsp. multiflora with glabrous lemmas and relatively greater panicle length to plant height ratio, and often more flowers per spikelet.

Poa persica subsp. persica

Fig. 3B, C

Eremopoa persica var. typica Grossh., Trudy. Bot. Inst. Azerbaidzh. Fil. Akad. Nauk. S.S.S.R. 8: 268. 1939, nom. inval. Eremopoa persica var. persica. 1960.

Poa cilicensis Hance, Ann. Sci. Nat., Bot., sér. 4, 18: 234. 1862. Type protologue. In Tauro cilicio, Kotschy 529. Type. In monte Tauro, aestate 1836, Kotschy 529, this from hb. H.F. Hance [via Reed 1887] no. 7498 (lectotype, here designated: BM! [BM000551484, right hand plant (2 left hand specimens are Poa diaphora var. songarica and are clearly excluded from Hance’s description written on the sheet)]; isolectotype: P! [P02642319]).

Glyceria taurica Steud., Syn. Pl. Glumac. 1: 286. 1854 (non Poa taurica E. Pojarkova, 1965, Poa × taurica H.N. Pojark., 1963). Type protologue. In monte Tauro, 1836, Kotschy (Kotschy hrbr.). Type. In monte Tauro, Aestate,1836, Kotschy 529 (lectotype, here designated: P! [P02642319]; isolectotype: BM [BM000551484 image!]).

Distribution

Armenia, Azerbaijan, Georgia, Egypt (north coast, possibly adventive), Iran, Iraq, Lebanon, Pakistan, Syria, Turkey; waif in France (introduced in wool, Marseille, H. Roux, P06768417!, P03370109!; RJS determination, 2015) and Norway (Greuter et al. 1984+).

Notes

Although Kotschy’s herbarium is mainly at W, a search of the W herbarium website did not turn up Kotschy 529 except as the genus Arenaria from Tauro cilicio or a Scrophularia from Persia. Kotschy 528 at W is a Poa of the P. bulbosa complex from “In monte Tauro” in 1836. Presumably the earlier 1836 set was broken up and 529 ended up at BM and P. The anthers in the G. taurica lectotype are 1.8 mm long and the lemmas are pubescent along the keel and marginal veins.

Poa persica subsp. multiradiata (Trautv.) Soreng, Cabi & L.J. Gillespie, comb. nov.

Fig. 3D, E

Poa palustris var. multiradiata Trautv., Trudy Imp. S.-Peterburgsk. Bot. Sada 4: 406. 1876. Poa multiradiata (Trautv.) Regel, Trudy Imp. S.-Peterburgsk. Bot. Sada 7: 620. 1880. Eremopoa multiradiata (Trautv.) Roshev., Fl. URSS 2: 430, t. 32. 1934. Eremopoa persica subsp. multiradiata (Trautv.) Tzvelev, Zlaki SSSR 479. 1976.

Nephelochloa tripolitana Boiss. & Blanche, Diagn. Pl. Orient., ser. 2, 4: 133–134. 1859. Poa persica var. major Boiss., Fl. Orient. 5: 610–611. 1884. Type protologue. Hab. ad margines semitarum inter hortos ad Tripolium Syriae (Blanche), circa Byrouth in Libano (Gaillardot). Type. Lebanon. S. Tripoli, dans les bords des chemins, 16 May 1854, Blanche 1267 (lectotype, here designated: JE [JE00005064 ex herb. Gaillardot, image!]). Note. Two of the original specimens turned up in our search, Blanche 1267 (JE00005064 ex herb. Gaillardot) and Gaillardot s.n. (JE00005065 ex herb Gaillardot no. 2323 [image!]). Blanche in 1869 (P02530724) might also be original material, with a distribution date rather than a collection date.

Eragrostis barbeyi Post, Bull. Herb. Boissier 5: 760–761. 1897. Type protologue. Habitat in collibus prope Midyat (Mardin), no. 38. Type. Turkey. Midyat, Hillsides, May 1895, 38 Barbey (lectotype, here designatedby Nada Sinno Saoud & RJS: BEI! (image seen by RJS!)). Note. The BEI sheet has “No. 55 38 Barbey, 1895” (55 was originally written as 54 but the 4 written over by 5).

Eremopoa mardinensis R.R. Mill, Fl. Turkey & E. Aegean Isl. 9: 624, 488. 1985.Type. Turkey. Mardin, Mardin to Nusaybin, 8 km from Mardin, 850 m alt., shallow limestone gully, 22 May 1957, P. H. Davis & D. Hedge 28491 (holotype: E! [E00196494]).

Type

Armenia rossica, prope monasterium Kiptschach, 1875, G. Raddi. Type: Armenia rossica: prope monasterium Kiptschach in monte Alagos, Jun 1875, G. Radde 124 (holotype: LE! [photo E00326521!]; isotypes: LE, LE, W [W19160014191 image!]).

Distribution

Armenia, Georgia, Iran, Lebanon, Pakistan, Syria and Turkey.

Notes

The presence of hairs on the lemmas in material treated as “multiradiat” is confused in the literature. Mill (1985) indicates that E. multiradiata and E. persica s.s. have lemma keels hairy in the lower ⅓–½. We concur with Tzvelev (1976), who keyed E. persica subsp. persica as lemmas short pilose along the base of keel and marginal veins and subsp. multiradiata as lemmas glabrous or with a few solitary hairs.

Mill (1985) distinguished his new species Eremopoa mardinensis from E. multiradiata based on its glabrous lemmas, 8–12-flowered spikelets and florets strongly divergent from the rachilla. However, subsp. multiradiata also has glabrous lemmas (as noted above) and divergent florets (when spikelets are in flower) and its (4)5–9(10)-flowered spikelets overlap in number; therefore, we treat E. mardinensis as a synonym of E. multiradiata. The type material of Eragrostis barbeyi is from the same place as E. mardinensis and is clearly the same form (spikelets many-flowered); Nephelochloa tripolitana, with ca. 12–14-flowered spikelets, also appears to belong to this form. If E. mardinensis were accepted as a species, the basionym names Eragrostis barbeyi or Nephelochloa tripolitana would have priority.

Poa subg. Pseudopoa sect. Speluncarae Soreng, Cabi & L.J. Gillespie, sect. nov.

Type

Poa speluncarum J.R. Edm.

Diagnosis

Differing from Poa sect. Pseudopoa in being perennial and stooling, with top culm sheath margins fused 40–50% their length and from almost all Poa in proximal spikelets being 1-flowered.

Poa speluncarum J.R. Edm., Fl. Turkey & E. Aegean Isl. 9: 623. 473. 1985.

Type

Turkey. C4, Konya, distr. Ermenek, Kamis Dere between Ermenek and Oyuklu Dag., floor of caverns, 1400–1500 m, 14 Aug 1949, P. H. Davis 16180 (holotype: K! [K000641325]; isotype: E! [E00367874]).

Distribution

Turkey (central Taurus Mts.).

Notes

Poa speluncarum was described by Edmondson (1985) as an annual species of Poa sect. Ochlopoa Asch. & Graebn (≡ Poa sect. Micrantherae Stapf. Type: Poa annua). Our investigation found it to be a feeble, stooling perennial with sparsely scabrous panicle branches, uppermost sheaths closed up to half their length, spikelets sparsely scaberulous, mostly 1-flowered, the distal-most ones frequently 2(–3) flowered, anthers 1.1–1.7 mm, caryopsis 1.7–1.8 mm long, hilum 0.3 mm long and grain adherent to the palea. DNA data have clearly placed it in the Poa clade that includes Eremopoa species (E clade), either as sister to P. attalica (nuclear data) or as sister to P. attalica + P. sintenisii (plastid data). The species is odd in subgenus Pseudopoa for its perennial habit (albeit weak) and more closed sheaths, and in Poa generally by its mostly uniflorous spikelets. It is a very rare species that lives in the backs of shallow, moist, cool caves in the Taurus Mts., along with other cave endemics.

Poa subg. Pseudopoa sect. Lindbergella (Bor) Soreng, Cabi & L.J. Gillespie, sect. nov.

Lindbergia Bor, Svensk Bot. Tidskr. 62: 467, 1968 (nom. illeg. hom., non Kindb., 1897).

Lindbergella Bor, Svensk Bot. Tidskr. 63: 368. 1969.

Type

Poa sintenisii H. Lindb. ≡ Lindbergella sintenisii (H. Lindb.) Bor.

Diagnosis

Differing from Poa sect. Pseudopoa in: panicle branches smooth; lower glume 3-veined, up to 3/4 as long as the lower lemma; lemmas 3-veined, relatively firm, sericeous on keel marginal veins and sides; callus with short crown of hairs, the hairs 0.2 mm long; and palea keels sericeous in part.

Poa sintenisii H. Lindb., Årsbok-Vuosik. Soc. Sci. Fenn. 20 B (7): 5. 1942 (emend. Lindberg 1946).

Lindbergia sintenisii (H. Lindb.) Bor, Svensk Bot. Tidskr. 62: 467. 1968. Lindbergella sintenisii (H. Lindb.) Bor, Fl. Cyprus 63: 368. 1969.

Poa persica subsp. cypria Sam., Ark. Bot., n.s. 1(9): 417. 1950 [1951].Type. Cyprus. auf dem Troodos, 20 Jun 1880, P. Sintenis 881 (lectotype, here designated: S; isolectotypes: B [B 10 0365891!], LD [LD1808162 image!, LD1808226 image!], G?, K [K000789835 image!, K000789836 image!, K000789837 image!], W [W0012225 image!, W0033518 image!, W00096518 image!, W0019026 image!]).

Type protologue

Cyprus. In pineto (P. pallasiana) in m. Troodos lecta est. 1939. Type. Cyprus. Troodos in pineto juxta via huad procul ab “Olympus Camp Hotel”, 22 Jun 1939, H. Lindberg s.n. (holotype: S [S-11-34137 image!]; isotypes: S [S-G-4941 image!], K [K000789839 image!], LD [LD1807330 image!], W [image!]).

Distribution

Cyprus (Mt. Troodos, endemic to serpentine rocks).

Names of uncertain application within Poa subgen. Pseudopoa

Festuca bellula Regel, Trudy Imp. S.-Peterburgsk. Bot. Sada 7: 594. 1881. Eremopoa bellula (Regel) Roshev., Fl. URSS 2: 431, pl. 32, f. 12. 1934.

Type protologue

Ad fontes calidos Araschan Bulak in Turkestania occidentali, Krause s.n. Type: Taschkenter Alatau, Araschan Bulak, 11 Jun 1871, (Hieronymous) Krause s.n. (holotype: LE [only one collection cited]).

Notes

Eremopoa bellula was applied by several authors to small densely tufted alpine annual plants of south-central and southwest Asia, which we recognise as P. diaphora var. alpina (based on Poa persica var. alpina Boissier [1884]). Tzvelev (1976, pg. 480) noted that the holotype collection of E. bellula appeared to be a mix of altaica (diaphora) and songarica forms (“p.p. max” = E. altaica subsp. songarica , somewhat intermediate between this subsp. and subsp. altaica, and “p.p. minor” = E. altaica subsp. altaica); he considered E. bellula to be a synonym of E. altaica subsp. songarica. Further study is needed to clarifiy the placement of Eremopoa bellula and determine if it is synonymous with P. diaphora var. alpina.

Eremopoa glareosa Gamajun., Bot. Mater. Gerb. Inst. Bot. Akad. Nauk Kazahsk. SSR 2: 2. 1964.

Type protologue

Usbekistanica, Tian Schan Occid., Bostandyk, fonts Aksar-sai, 28 Jul 1949, N. V. Pavlov s.n. (holotype: AA).

Notes

Tzvelev (1976, pg. 480) included E. glareosa as a synonym under E. altaica subsp. songarica, but noted that it is somewhat intermediate between this taxon and E. altaica subsp. altaica. As the protologue indicates the plants are 10–28 cm tall, with 3 to 4 florests per spikelet, spikelets 4–7 mm long and anthers 2.5 mm long, this is more likely to be Poa persica, perhaps subsp. multiradiata, since no pubescence is indicated.

Festuca heptantha K. Koch, Linnaea 21(3): 410. 1848. Poa heptantha (K. Koch) Steud., Syn. Pl. Glumac. 1: 255. 1854.

Type protologue

Im Hochgebirge, auf sumpfigen Wiesen, auf Urgestein, 5500 ft, C. Koch s.n. (holotype: B, probably destroyed).

Note

There is no location in the species protologue beyond the article title “Beitrage zu einer Flora des Orients”. Tzvelev (1976) indicated this name and the next, Festuca polygama, probably apply to Eremopoa persica and that the types of these were in Berlin (B). Clayton et al. 2002+ (GrassBase) reflect the same information. RJS was unable to locate type material of either of these two names at B, P or via internet searches.

Festuca polygama K. Koch, Linnaea 21: 409. 1848. Poa polygama (K. Koch) Steud., Syn. Pl. Glumac. 1: 255. 1854.

Type protologue

“Aus dem Wilhelm’schen Herbr als Poa persica.” Type: Wilhelms (holotype: B, probably destroyed).

Notes

Tzvelev (1976) indicates “Caucasus?”, but there is no location in the species protologue beyond the article title “Beitrage zu einer Flora des Orients”.

Excluded names

Eremopoa medica H. Scholz, Willdenowia 11(1): 96. 1981.

Type

Persia, Prov. Azerbaijan occid.: In pratis paludosis SE Shahpur versus lacum Rezaiyeh (Urmia), 1300 m; 12 Jun 1971, Rechinger 41820 (holotype: W [W1972-0000975 image!; isotypes: B! [B 10_0272774], GZU [GZU000201751 image!], WU [WU0033125 image!]).

Notes

The type collection of Eremopoa medica is clearly a perennial species of Puccinellia (possibly P. gigantea (Grossh.) Grossh.) with lemmas rounded on the back, a distinct short crown of callus hairs and papillae common on vegetative structures (pedicels and leaves). Material cited as E. medica in Rahmanian et al. (2014, fig. 5) appears to us to be Poa persica subsp. persica; their description and illustration indicate an annual habit, pubescent lemmas and panicles with 10 or more branches per whorl. The single specimen (TARI 35082) cited was included in our molecular analysis and formed a clade with other P. persica accessions in all trees.

Invalid names, not vouchered

Festuca amherstiana Nees, Ill. Bot. Himal. Mts. 417. 1839, nom. nud., name in list, no voucher.

Notes. Kew GrassBase (Clayton et al. 2002+) indicates it is equal to E. persica. The specimen K00078950 (ex P) (image!), Voyage V. Jacquemont aux Indes orient. no. 1902, has this name on the label. The specimen is certainly P. diaphora, not P. persica.

Acknowledgements

We thank the curators and staff at the following herbaria for loans and/or specimen images: E, G, BEI, BM, B, K, LE, P, US, ANK, ISTE, NKU and W. Ralf Hand kindly sent us leaf material and a duplicate of Lindbergella sintenisii from Cyprus; Mostafa Assadi and Mohammad Amini-Rad kindly sent leaf material from Iran from the TARI and IRAN herbaria. Nada Sinno Saoud kindly provided an image of Eragrostis barbeyi from the Post herbarium (BEI). We thank Roger Bull for assistance with the molecular research and Paul Peterson, Stephen Wagstaff and Clifford Morden for their helpful reviews. Part of the molecular study was performed by NA as part of her Masters thesis at the University of Ottawa and Canadian Museum of Nature. LJG acknowledges the support of the Canadian Museum of Nature and the Natural Sciences and Engineering Research Council of Canada (NSERC).

References

  • Ascherson PFA, Graebner ORPP (1900) Synopsis der Mitteleuropaïschen Flora, Vol. 2, part 1. W. Engelmann, Leipzig.
  • Boissier PE (1884) Ordo CXLI. Gramineae. In: Boissier PE (Ed.) Flora Orientalis, vol. 5. H. Georg, Basilae, 432–692.
  • Boissier PE (1859) Gramineae. In: Boissier PE (Ed.) Diagnoses Plantarum Orientalium Novarum, ser 2. (4). B. Hermann, Lipsiae, 125–146.
  • Bor NL (1960) Grasses of Burma, Ceylon, India & Pakistan. Pergamon Press, Oxford, 767 pp.
  • Bor NL (1968a) Gramineae. In: Townsend CC, Guest ER, Al-Rawi A (Eds) Flora of Iraq, vol. 9. Ministry of Agriculture of the Republic of Iraq, Baghdad, 1–588.
  • Bor NL (1968b) Lindbergia Bor: A new genus of grasses from Cyprus. Svensk Botanisk Tidskrift 62: 467–470.
  • Bor NL (1969) A correction. Svensk Botanisk Tidskrift 63: 368–369.
  • Bor NL (1970) Gramineae. In: Rechinger KH (Ed.) Flora Iranica, vol. 70. Akademische Druck und Verlagsanstalt, Graz, Austria, 1–573.
  • Cabi E, Doğan M (2012) Poaceae. In: Güner A, Aslan S, Ekim T, Vural M, Babaç MT (Eds) Türkiye Bitkileri Listesi. Nezahat Gökyiğit Botanik Bahçesi ve Flora Araştırmaları Derneği Yayını, Istanbul, 690–756.
  • Cabi E, Soreng RJ, Gillespie LJ (2017) Taxonomy of Poa jubata and a new section of the genus (Poaceae). Turkish Journal of Botany 41: 404–415. https://doi.org/10.3906/bot-1611-28
  • Clayton WD, Renvoize SA (1986) Genera Graminum. Kew Bulletin, Additional Series 13: 1–389.
  • Cope TA (1982) Poaceae, In: Nasir E, Ali SI (Eds) Flora of Pakistan, Vol. 143. Univ. of Karachi, Karachi, 1–678.
  • Czerepanov SK (1981) Sosudistye rasteniya SSSR [Vascular Plants of the USSR]. Nauka, Leningradskoe Otd-nie, Leningrad, 1–509.
  • Czerepanov SK (1995) Vascular plants of Russia and Adjacent States (the former USSR). Cambridge University Press, Cambridge, 1–532.
  • Darbyshire SJ (2007) Eremopoa Roshev. In: Flora of North America Editorial Committee (Eds) Flora of North America North of Mexico, Vol. 24. Oxford University Press, New York, 618–619.
  • Darriba D, Taboada GL, Doallo R, Posada D (2012) jModelTest2: More models, new heuristics and parallel computing. Nature Methods 9(8): 772. https://doi.org/10.1038/nmeth.2109
  • Edmondson JR (1985) Poa. In: Davis PH (Ed.) Flora of Turkey and East Aegean Islands, Vol. 9. Edinburgh University Press, Edinburgh, 470–486.
  • Fischer FEL, Meyer CA (1841) Enumerato Plantarum Novarum, Vol. 1. G. Fisheri, Petropoli, 1–77.
  • Gabrieljan ET, Oganesian ME (2010) Poaceae. In: Takhtajan AL (Ed.) Flora Armenii 11. Gantner ARG, Ruggell, 1–547.
  • Gillespie LJ, Archambault A, Soreng RJ (2007) Phylogeny of Poa (Poaceae) based on trnT-trnF sequence data: Major clades and basal relationships. Aliso 23(1): 420–434. https://doi.org/10.5642/aliso.20072301.33
  • Gillespie LJ, Soreng RJ, Bull RD, Jacobs SWL, Refulio Rodríguez N (2008) Phylogenetic relationships in subtribe Poinae (Poaceae, Poeae) based on nuclear ITS and plastid trnT-trnL-trnF sequences. Botany 86(8): 938–967. https://doi.org/10.1139/B08-076
  • Gillespie LJ, Soreng RJ, Jacobs SWL (2009) Phylogenetic relationships of Australian Poa (Poaceae: Poinae), including molecular evidence for two new genera, Saxipoa and Sylvipoa. Australian Systematic Botany 22(6): 413–436. https://doi.org/10.1071/SB09016
  • Gillespie LJ, Soreng RJ, Paradis M, Bull RD (2010) Phylogeny and reticulation in subtribe Poinae and related subtribes (Poaceae) based on nrITS, ETS, and trnTLF data. In: Seberg O, Petersen G, Barfod AS, Davis JI (Eds) Diversity, Phylogeny, and Evolution in the Monocotyledons. Aarhus University Press, Aarhus, 589–617.
  • Grisebach AHR (1852) Ordo CXLII Gramineae, In: Ledebour CF von (Ed.) Flora Rossica, Vol. 4. Sumtibus Librariae E. Schweizerbart, Stuttgartiae, 324–484.
  • Grossheim AA (1939) Flora Kavkaza (ed. 2), Vol. 1. Akademiia Nauk S.S.S.R. Azerbaidzhanskii filial, Baku, Azerbaijan, 1–402.
  • Hackel E (1887) Gramineae. In: Engler HGA, Prantl K (Eds) Die Natürlichen Pflanzenfamilien, Vol. 2 (2). Engelmann, Leipzig, 1–97.
  • Karelin GS, Kiriloff IP (1841) Enumeratio plantarum in desertis Songariae orientalis. Moscow, 233 pp.
  • Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution 30(4): 772–780. https://doi.org/10.1093/molbev/mst010
  • Koch K (1848) Beitrage zu einer Flora des Orients. Linnaea 21(1[4]): 289–743.
  • Köie ME (1945) Beitrag zur Flora Sudwest-Irans I. Danish Science Investigations in Iran 4: 9–56.
  • Kovalevskaja SS (1968) Poa. Opredelitel’. Rastenii Srednei Azii 1: 127–138.
  • Lindberg H (1942) En botanisk resa till Cypern 1939: foredrag vid tilltradendet av ordforande-skapet i Finska Vetensaps-Societeten den 29 April 1941. Årsbok-Vousikirja Societas Scientiarum Fennica 20 B(7): 1–12.
  • Lindberg H (1946) Iter cyprium contributio ad cognitionem florae insulae Cypri. Acta Societatis Scientiarum Fennicae, Series B. Opera Biologica 7: 3–37.
  • Mill RR (1985) Eremopoa, Nephelochloa, Catabrosella, Colpodium, Hyalopoa, Catabrosa, Paracolpodium. In: Davis PH (Ed.) Flora of Turkey and East Aegean Islands, Vol. 9. Edinburgh University Press, Edinburgh, 486–501.
  • Nikiforova OD, Ovchinnikova SV, Lipin AS, Blasova N, Olonova MV (2012) Poaceae. In: Baikov KS (Ed.) Conspectus Florae Rossiae Asiaticae: Plantae Vasculares, Academia Scintiarum Rossica, Divisio Sibirca, Novosibirsk, 518–576.
  • Pavlov NV, Gamajunova AP (1964) Addenda ad “Flora Kasachstanica” I-IV. Botanicheskie Materialy Gerbariia Instituta Botaniki Akademii Nauk Kazakhskoi SSR 2: 10–13.
  • Post GE, Autran EJB (1897) Plantae Postianae, quas enumerant G. -E. Post and E. Autran, Novas species descripsit G. -E. Post, Fasciulus VIII. Bulletin de l’Herbier Boissier 5: 754–761.
  • Rahmanian S, Saeidi H, Assadi M, Rahiminejad MR (2014) A taxonomic revision of the genus Eremopoa Roschv. (Poaceae, Poeae) in Iran. Iranian Journal of Botany 20(1): 8–15.
  • Rambaut A (2006+) FigTree, Tree Figure Drawing Tool Version 1.4.3. Institute of Evolutionary Biology, University of Edinburgh. http://tree.bio.ed.ac.uk/ [Accessed January 2018].
  • Roshevitz RU (1934) Eremopoa. In: Komarov VLL (Ed.) Flora URSS, vol. 2 Gramineae Juss. Akademii Nauk SSSR, Leningrad, 1–778.
  • Samuelsson G (1950 [1951]) In: Rechinger KH (Ed.) Zur Flora von Cypren (Reliquiae Samuelssonianae II.). Arkiv för Botanik, n.s. 1(9): 417.
  • Scholz H (1980) Eine neue Eremopoa-Art (Gramineae) aus der Turkei. Willdenowia 10(1): 33–36.
  • Scholz H (1981) Drei neue Gramineen aus Iran und Libyen. Willdenowia 11(1): 95–100.
  • Soreng RJ (2003) Eremopoa Roshev. In: Soreng RJ, Peterson PM, Davidse G, Judziewicz EJ, Zuloaga FO, Filgueiras TS, Morrone (Eds) Catalogue of New World Grasses (Poaceae): IV. Subfamily Pooideae, Contributions from the United States National Herbarium 48. Smithsonian Institution, Washington, D.C., 310–311.
  • Soreng RJ, Gillespie LJ, Bull RD (2010) Phylogeny and reticulation in Poa based on plastid trnTLF and nrITS sequences with attention to diploids. In: Seberg O, Petersen G, Barfod AS, Davis JI (Eds) Diversity, Phylogeny and Evolution of Monocotyledons, Aarhus University Press, Aarhus, 619–644.
  • Soreng RJ, Gillespie LJ, Consaul LL (2017b) Taxonomy of the Poa laxa group, including two new taxa from Arctic Canada and Greenland, and Oregon, and a re-examination of P. sect. Oreinos. Nordic Journal of Botany 35: 513–538. https://doi.org/10.1111/njb.01507
  • Soreng RJ, Gillespie LJ, Koba H, Boudko E, Bull RD (2015b) Molecular and morphological evidence for a new grass genus, Dupontiopsis (Poaceae tribe Poeae subtribe Poinae s.l.), endemic to alpine Japan, and implications for the reticulate origin of Dupontia and Arctophila within Poinae s.l. Journal of Systematics and Evolution 53(2): 138–162. https://doi.org/10.1111/jse.12146
  • Soreng RJ, Peterson PM, Annable CR (1995) Trinius Herbarium, Komarov Botanical Institute, St. Petersberg (Guide to the microform collection). IDC BT– 16: 1–66.
  • Soreng RJ, Peterson PM, Romaschenko K, Davidse G, Teisher JK, Clark LG, Barberá P, Gillespie LJ, Zuloaga FO (2017a) A worldwide phylogenetic classification of the Poaceae (Gramineae) II: An update and a comparison of two 2015 classifications. Journal of Systematics and Evolution 55(4): 259–290. https://doi.org/10.1111/jse.12262
  • Soreng RJ, Peterson PM, Romaschenko K, Davidse G, Zuloaga FO, Judziewicz EJ, Filgueiras TS, Davis JI, Morrone O (2015a) A worldwide phylogenetic classification of the Poaceae (Gramineae). Journal of Systematics and Evolution 53(2): 117–137. https://doi.org/10.1111/jse.12150
  • Stapf O (1897) Poa. In: Hooker JD (Ed.) The Flora of British India 7(22). L. Reeve & Co., London, 337–346.
  • Stevenson GA (1965) Notes on the more recently adventive flora of the Brandon area, Manitoba. Canadian Field Naturalist 79: 174–177.
  • Steudel EG (1854) Synopsis Plantarum Glumacearum, Vol. 1. J. B. Metzler, Stuttgart, 1–475.
  • Swofford DL (2002) PAUP*: Phylogenetic analysis using parsimony (and other methods), version 4.0b10. Sinauer Associates, Sunderland.
  • Trinius CB (1830) Graminum genera, quedam speciesque complures definitionibus novis. Mémoires de l’Académie Impérial des Sciences Saint-Pétersbourg, Série 6, Sciences, mathématiques, physiques et naturelles 1(4): 353–416.
  • Trinius CB (1835) [Poaceae] In: Bunge AA. ) Verzeichniss der Im Jahre 1832 im Östichen Theile Des Altai-Gebirges Gesammelten Pflanzen. Ein Supplement zur Flora Altaica. Mémoires presentés a l’Académie Impérial des Sciences Saint-Pétersbourg pars des divers savans 2: 523–608 [526–529].
  • Trinius CB (1836) Graminum in hisce actis a se editorum generibus ac speciebus supplementa addit Mémoires de l’Académie Impérial des Sciences Saint-Pétersbourg, sér. 6, pt. 2 Sciences Naturelles 4, 2(1): 69–70.
  • Tutin TG (1980) Gramineae. In: Tutin TG, Heywood VH, Burges NA, Valentine DH, Walters SM, Webb DA (Eds) Flora Europaea, Vol. 5. Cambridge University Press, Cambridge, 118–267.
  • Tzvelev NN (1966) Ob ecology-geograicheskikh rasakh v semeistve zlakov Gramineae I ikh taxksonomischeskom range. Botaničeski Žurnal (Moscow & Leningrad) 51(8): 1104.
  • Tzvelev NN (1976) Zlaki SSSR. Nauka Publishers, Leningrad Section, Leningrad, 1–788 [English translation: 1983. Grasses of the Soviet Union, 2 vols. Amerind Publishing Co. New Delhi.]
  • Watson L, Dallwitz MJ (1992) Grass Genera of the World. CAB International, Cambridge, 1038 pp.
  • Zhu G, Liu L, Soreng RJ, Olonova MV (2006) Poa. In: Wu C-Y, Raven PH, Hong D-Y (Eds) Flora of China, Vol. 22. Science Press, Beijing; Missouri Botanical Garden Press, St. Louis, 266–309.

Appendix 1

Eremopoa, Lindbergella, Poa and outgroup samples used in the phylogenetic analyses. Ingroup samples are arranged by plastid clade (pl), nuclear clade (nr) and section. Voucher information (herbarium indicated in parentheses) and country of origin are provided; where there is no collector or collector number, the herbarium specimen number is given. GenBank Accession numbers are provided for ITS, ETS, trnT-trnL-trnF, matK and rpoB-trnC sequences for each sample; those in BOLD are new to this study.

pl nr Section Taxon Voucher Country ITS ETS TLF matK rpoB-trnC
A A Alpinae Poa alpina L. Gillespie 6299 (CAN) USA, Colorado GQ324483 GQ324287 DQ353985.2 KM523888 KM524001
A A Alpinae Poa badensis Haenke ex Willd. Hajkova et al. 2004-12 (US) Bulgaria GQ324490 GQ324295 GQ324402 KY378861 KY378827
A A Alpinae Poa ligulata Boiss. (JACA 166095) Spain GQ324522 GQ324346 GQ324432.2 KY378876 KY378842
A A Alpinae Poa thessala Boiss. & Orph. Gillespie et al. 10400 (CAN) Turkey KM523802 KM523729 KM524088 KM523901 KM524014
A A Arenariae Poa bactriana subsp. glabriflora (Roshev.) Tzvelev Gauba (IRAN 21237) Iran KX118734 KX118716 KX118751 MH921344 MH921369
A A Arenariae Poa bulbosa L. Catalan 13-2000 (UZ) Spain EU792388 GQ324297.2 AH015557.3 KY378863 KY378829
A A Arenariae Poa bulbosa subsp. vivipara (Koeler) Arcang. Soreng & Soreng 5814 (US) USA, Nevada (introd.) GQ324492 GQ324298 GQ324404 MH921345 MH921370
A A Arenariae Poa sinaica Steud. subsp. sinaica Soreng & Cabi 9249 (US) Turkey KX118748 KX118731 KX118766 KY378886 KY378852
A A Arenariae Poa timoleontis Heldr. ex Boiss. Soreng et al. 7509-1 (US) Greece KX118750 KX118732 KX118768 MH921354 MH921379
E E Lindbergella Lindbergella sintenisii (H. Lindb.) Bor Hand 6102 (US) Cyprus MH921326 MH921310 MH921393 MH921342 MK060117
E E Pseudopoa Eremopoa attalica H. Scholz Gillespie et al. 10612 (CAN) Turkey MH921313 MH921297 MH921380 MH921329 MH921355
E E Pseudopoa Eremopoa multiradiata (Trautv.) Roshev. Soreng & Cabi 9240 (US) Turkey MH921314 MH921298 MH921381 MH921330 MH921356
E E Pseudopoa Eremopoa oxyglumis (Boiss.) Roshev. Gillespie & Levin 10313 (CAN) Turkey MH921316 MH921300 MH921383 MH921332 MH921358
E E Pseudopoa Eremopoa oxyglumis Gillespie et al. 10578 (CAN) Turkey MH921317 MH921301 MH921384 MH921333 MH921359
E E Pseudopoa Eremopoa oxyglumis Gillespie et al. 10584 (CAN) Turkey MH921318 MH921302 MH921385 MH921334 MH921360
E E Pseudopoa Eremopoa oxyglumis Soreng & Cabi 8855 (US) Turkey MH921315 MH921299 MH921382 MH921331 MH921357
E E Pseudopoa Eremopoa persica (Trin.) Roshev. Assadi & Vosoughi (TARI 24939) Iran MH921321 MH921305 MH921388 MH921337 MH921363
E E Pseudopoa Eremopoa persica Mozaffarian (TARI 53671) Iran MH921320 MH921304 MH921387 MH921336 MH921362
E E Pseudopoa Eremopoa persica Soreng & Cabi 9215 (US) Turkey KY378812 KY378823 KY378816 KY378879 KY378845
E E Pseudopoa Eremopoa persica Yazdanfard (IRAN 51968) Iran MH921319 MH921303 MH921386 MH921335 MH921361
E E Pseudopoa Eremopoa persica Mozaffarian & Nowrozi (TARI 35082) Iran MH921322 MH921306 MH921389 MH921338 MH921364
E E Pseudopoa Eremopoa songarica (Schrenk ex Fisch. & C.A. Mey.) Roshev. Assadi & Mozaffarian (TARI 36867) Iran MH921324 MH921308 MH921391 MH921340 MH921366
E E Pseudopoa Eremopoa songarica Iranshahr (IRAN 20357) Iran MH921323 MH921307 MH921390 MH921339 MH921365
E E Pseudopoa Eremopoa songarica Soreng & Güney 4165 (US) Turkey EU792400 GQ324311 DQ353988.2 KY378868 KY378834
E E Pseudopoa Eremopoa songarica Soreng & Cabi 9320 (US) Turkey MH921325 MH921309 MH921392 MH921341 MH921367
E E Speluncarae Poa speluncarum J.R. Edm. Soreng et al. 8202 (US) Turkey MH921328 MH921312 MH921395 MH921353 MH921378
H P+H unclassified Poa pseudobulbosa Bor Soreng et al. 8246 (US) Turkey KX118747 KX118729 KX118765 MH921352 MH921377
H P+H Acutifoliae Poa planifolia Kuntze Peterson et al. 19233 (US) Argentina KM523800 KM523727 KM524087 KM523896 KM524009
H P+H Brizoides Poa poiformis (Labill.) Druce Gillespie et al. 7381 (CAN) Australia GQ324534 GQ324361 GQ324445 KM523897 KM524010
H P+H Homalopoa s.l. Poa reflexa Vasey & Scribn. Soreng 7422 (US) USA Colorado GQ324543 KX118730 GQ324450 KY378882 KY378848
H P+H Homalopoa s.s. Poa asiae-minoris H. Scholz & Byfield Soreng et al. 8100 (US) Turkey MH921327 MH921311 MH921394 MH921343 MH921368
H P+H Homalopoa s.s. Poa chaixii Vill. Soreng 4677 (US) Russia EU792404 GQ324299 EU854590 KM523890 KM524003
H P+H Homalopoa s.s. Poa chaixii Soreng 7524 (US) Germany GQ324493 GQ324300 GQ324405 MH921346 MH921371
H P+H Homalopoa s.s. Poa masendarana Freyn & Sint. Assadi (TARI 73254) Iran KX118743 KX118725 KX118761 MH921351 MH921376
H P+H Homalopoa s.s. Poa occidentalis Vasey Peterson & Valdes Rena 18918 (US) Mexico KU756540 KU763436 KU763514 KY378877 KY378843
H P+H Homalopoa s.s. Poa remota Forselles Soreng et al. 7540 (US) Kyrgyz Republic GQ324545 GQ324372 GQ324452 KY378883 KY378849
H P+H Madropoa Poa fendleriana (Steud.) Vasey Gillespie 6292 (CAN) USA, Colorado EU792403 GQ324319 DQ354027 KY378869 KY378835
H P+H unclassified (supersect. Homalopoa) Poa calycina (J. Presl) Kunth Peterson et al. 17923 (US) Peru EU792425 KU763395 EU792467 KY378864 KY378830
H P+H unclassified (supersect. Homalopoa) Poa marshallii Tovar Peterson et al. 21546 (US) Peru KM523799 KM523726 KM524086 KM523895 KM524008
J J Jubatae Poa jubata A. Kern. Soreng et al. 9029-2 (US) Turkey KY378810 KY378820 KY378814 KY378873 KY378839
J J Jubatae Poa jubata Soreng et al. 9266 (US) Turkey KY378811 KY378821 KY378815 KY378874 KY378840
M M Micrantherae Poa infirma Kunth Catalan 3-2000 (UZ) Spain GQ324516 GQ324334 GQ324427 KY378871 KY378837
M M Micrantherae Poa supina Schrad. Soreng & Cayouette 5950-2 (US) USA, cult. (from Europe) EU792387 GQ324383 DQ353984 KY378888 KY378854
N N Nanopoa Poa trichophylla Heldr. & Sart. ex Boiss. Soreng et al. 7508 (US) Greece GQ324554 GQ324386 GQ324461 KY378889 KY378855
N N unclassified Poa dolosa Boiss. & Heldr. Soreng et al. 7495-1 (US) Greece GQ324502 GQ324312 GQ324414 KM523891 KM524004.2
N N unclassified Poa iconia var. pelasgis (H. Scholz) Soreng Gillespie et al. 10492 (CAN) Turkey KX118744 KX118726 KX118762 MH898827 MH898844
N N unclassified Poa ursina Velen. Stoneberg SH17 (US) Bulgaria GQ324527 GQ324352 GQ324437 KY378892 KY378858
N S Secundae Poa curtifolia Scribn. Soreng & Soreng 6347c-1 (US) USA, Washington EU792394 KY378819 DQ353994.2 KY378867 KY378833
N S Secundae Poa secunda J. Presl. subsp. secunda Soreng & Soreng 5812 (US) USA, Nevada EU792393 KU763450 DQ353991 KY378884 KY378850
N S Secundae Poa stenantha Trin. Soreng & Soreng 6068-1 (US) USA, Alaska KU756554 KU763455 DQ354057.2 KY378887 KY378853
P P+H Macropoa Poa densa Troitsky Soreng & Cabi 9306 (US) Turkey KX118738 KX118720 KX118755 MH921347 MH921372
P P+H Macropoa Poa bucharica Roshev. Soreng et al. 7662 (US) Kyrgyz Republic KX118735 KX118717 KX118752 KY378862 KY378828
P P+H Macropoa Poa diversifolia (Boiss. & Balansa) Hack. ex Boiss. Gillespie et al. 10529 (CAN) Turkey KX118739 KX118721 KX118756 MH921348 MH921373
P P+H Macropoa Poa iberica Fisch. & C.A. Mey. Soreng et al. 7977 (US) Russia, Cabardino-Balkaria KX118741 KX118723 KX118758 MH921349 MH921374
P P+H Macropoa Poa longifolia Trin. subsp. longifolia Soreng et al. 7945 (US) Russia, Cabardino-Balkaria KX118742 KX118724 KX118760 MH921350 MH921375
P P+H Macropoa Poa sibirica Roshev. subsp. sibirica Olonova 2003-45 (CAN) Russia, Khakasia GQ324547 KY378824 GQ324455 KY378885 KY378851
P P+H Poa Poa irkutica Roshev. Kasanovskiy 2002-7 (CAN) Russia, Irkutsk EU792402 GQ324335 DQ354007.2 KY378872 KY378838
P P+H Poa Poa pratensis L. subsp. pratensis Gillespie et al. 10592 (CAN) Turkey KX118746 KX118726 KX118764 KY378880 KY378846
P X Malacanthae Poa arctica R. Br. subsp. arctica Gillespie & Aiken 5701 (CAN) Canada, Nunavut GQ324487 GQ324291 DQ354009 KY378860 KY378826
R R Parodiochloa Poa cookii (Hook.f.) Hook.f. Hennion Gen1 (P) Subantarctic Islands, Crozet I. EU792383 GQ324306 EU792454 KY378866 KY378832
R R Parodiochloa Poa flabellata (Lam.) Raspail Wright 9NSG (not vouchered) South Georgia Islands EU792381 GQ324321 EU792453 KM523892 KM524005
S S Abbreviatae Poa flexuosa Sm. subsp. flexuosa Brochmann 2000-3-1 (CAN) Norway GQ324520 GQ324342 GQ324418 KY378875 KY378841
S S Abbreviatae Poa pseudoabbreviata Roshev. Soreng & Soreng 6032-1 (US) USA, Alaska EU792398 GQ324370 DQ353997 KY378881 KY378847
S S Stenopoa Poa biebersteinii H.N. Pojark. (cf) Gillespie & Cabi 10327 (CAN) Turkey KY944706 KY944668 KY987089 KY944622 KY987044
S S Stenopoa Poa glauca Vahl Gillespie 5804 (CAN) Canada, Nunavut AY237839 GQ324324 GQ324421 KY378870 KY378836
S S Stenopoa Poa palustris L. Gillespie 6461 (CAN) Canada, Ontario EU792396 KY378822 DQ354000 KY378878 KY378844
S S Tichopoa Poa compressa L. Gillespie 6457 (CAN) Canada, Quebec EU792395 KY378818 DQ354003 KY378865 KY378831
V V Pandemos Poa trivialis L. subsp. trivialis Soreng 4681-1 (US) USA, Maryland (introd.) GQ324555 GQ324387 GQ324462 KY378891 KY378857
V V Pandemos Poa trivialis subsp. sylvicola (Guss.) H. Lindb. Gillespie et al. 10368 (CAN) Turkey KY378813 KY378825 KY378817 KY378890 KY378856
Y Y Sylvestres Poa autumnalis Elliott Soreng 4680 (US) USA, Maryland EU792379 GQ324294 DQ353979 KM523889 KM524002
Y Y Sylvestres Poa saltuensis Fernald & Wiegand Gillespie 7043 (CAN) Canada, Ontario EU792378 GQ324374 EU792451 KM523899 KM524012
Y Y Sylvestres Poa wolfii Scribn. Soreng & Soreng 5800 (US) USA, Missouri EU792377 GQ324389.2 AH015556.2 KY378893 KY378859
outgroup Arctagrostis latifolia (R. Br.) Griseb. Gillespie et al. 6586 (CAN) Canada, Nunavut EU792351 GQ324245 DQ353969 KM523924 KM523954
outgroup Milium effusum L. Soreng 7771 (US) Sweden KM523785 KM523711 KM524072 KM523870 KM523983
outgroup Nicoraepoa andina (Trin.) Soreng & L.J. Gillespie Soreng & Soreng 7182 (US) Chile EU792354 GQ324275 DQ353971 KM523874 KM523987
outgroup Phleum montanum K. Koch Gillespie et al. 10614-2 (CAN) Turkey KM523793 KM523720 KM524081 KM523883 KM523996
outgroup Phleum pratense L. Soreng 7943 (US) Russia, Stavropol KM523796 KM523723 KM524084 KM523886 KM523999