Research Article |
Corresponding author: Lynn J. Gillespie ( lgillespie@mus-nature.ca ) Academic editor: Clifford Morden
© 2018 Lynn J. Gillespie, Robert John Soreng, Evren Cabi, Neda Amiri.
This is an open access article distributed under the terms of the CC0 Public Domain Dedication.
Citation:
Gillespie LJ, Soreng RJ, Cabi E, Amiri N (2018) Phylogeny and taxonomic synopsis of Poa subgenus Pseudopoa (including Eremopoa and Lindbergella) (Poaceae, Poeae, Poinae). PhytoKeys 111: 69-101. https://doi.org/10.3897/phytokeys.111.28081
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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.
Annuals, classification, DNA, Eremopoa , grasses, Lindbergella , phylogeny, Poa , Poaceae , taxonomy
Eremopoa Roshev. is a small, primarily west and central Asian genus of annual grasses.
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
Classification history of Eremopoa and other taxa here accepted in Poa subg. Pseudopoa. Species and infraspecific taxa accepted by
Roshevits (1934) | Roshevits (in |
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Rhamanian et al. (2014) | Euro+Med (on-line) | Here |
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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 (
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
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 (
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
DNA was extracted from silica gel dried or herbarium leaf material as described in
Maximum parsimony (MP) analyses were performed in PAUP* 4.0b10 (
Bayesian Markov chain Monte Carlo analyses were conducted in MrBayes (
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
Plastid and nrDNA Bayesian trees are given in Fig.
PoanrDNA and plastid Baysian analyses showing placement of Eremopoa and Lindbergella. Bayesian 50% majority rule consensus trees of nrDNAITS 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.
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.
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 (
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
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).
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.
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.
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.
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.
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 |
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.
Eremopoa attalica H. Scholz, Willdenowia 10(1): 33, f. 1. 1980.
Turkey. Antalya, “nordwestl. Antalya bei Termessos, ausgetrockneter Gebirgsbach”, 300 m, 23 Jul 1979, Kehl s.n. (holotype: B! [B-100272775])
Turkey (western Taurus Mts.).
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
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.
“Sterilissimus salsuginosis deserti editi Tschujae”, [1800–3000 m], July 1832, A. Bunge (lectotype, designated by
Egypt (Sinai Peninsula) to China (Xinjiang, Xizang).
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 persica var. diaphora (Trin.) Asch. & Graebn., Syn. Mitteleur. Fl. 2: 437. 1900. Eremopoa altaica subsp. altaica.
China (Xinjiang, Xizang), Kazakhstan, Kyrgyz Republic, Pakistan, Russia (Altai Mts.), Tajikistan, Turkey.
A single specimen recorded from Turkey (Kars Prov., Litvinov 4790 US ex K) evidently belongs to this variety and was also cited by
Poa subgenus Pseudopoa sect. Pseudopoa. AP. diaphora subsp. diaphora var. diaphora, Chu, Kyrgyz Republic (Soreng et al. 7537) B, CP. 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, EP. persica subsp. multiradiata, Mardin, Turkey (Soreng et al. 9240) D habit E spikelet showing glabrous lemmas. Photos by R.J. Soreng.
Poa persica var. alpina Boiss., Fl. Orient. 5: 610. 1884.
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!]).
Armenia, Azerbaijan, Afghanistan, Georgia, Iran, Kyrgyz Republic, Pakistan, Turkey and Turkmenistan(?).
This taxon, accepted as Eremopoa bellula by several authors (see Names of Uncertain Application below), was first recognised infraspecifically by
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
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!]).
Ad fl. Karatal versus montes Karatau, 13 June 1840, H. Schrenk s.n. (holotype: LE; isotype: LE).
Afghanistan, Armenia, Azerbaijan, China (Xizang), Georgia, Iran, Israel, Kazakhstan, Kyrgyz Republic, Tajikistan, Turkey, Turkmenistan and Uzbekistan.
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 (
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.
Turkey. In collibus prope Baibout, 17 Jul 1963, E. Bourgeau (lectotype, designated by
Armenia, Azerbaijan, Georgia, China (Xizang), Kyrgyz Republic, Pakistan, Tajikistan, Turkey, Turkmenistan and Uzbekistan.
Most accounts have recognised this taxon at one rank or another, except
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.
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]).
Turkey (central and eastern Taurus Mts. and adjacent ranges).
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.
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.
Iran. 60 km north of Dizful, 3 May 1937, M. Köie 475 (lectotype, here designated: C [C10016935 image!]; isolectotype: LE).
Iran (Zagros Mts.).
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
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.
Iran: in collibus ad Akar-Tschai prob. Karabagh, 1400–1900 m, 27 May 1829, Szowits 246 (lectotype, designated by
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.
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!]).
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+).
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 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]).
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!]).
Armenia, Georgia, Iran, Lebanon, Pakistan, Syria and Turkey.
The presence of hairs on the lemmas in material treated as “multiradiat” is confused in the literature.
Poa speluncarum J.R. Edm.
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.
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]).
Turkey (central Taurus Mts.).
Poa speluncarum was described by
Lindbergia Bor, Svensk Bot. Tidskr. 62: 467, 1968 (nom. illeg. hom., non Kindb., 1897).
Lindbergella Bor, Svensk Bot. Tidskr. 63: 368. 1969.
Poa sintenisii H. Lindb. ≡ Lindbergella sintenisii (H. Lindb.) Bor.
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.
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!]).
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!]).
Cyprus (Mt. Troodos, endemic to serpentine rocks).
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]).
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
Usbekistanica, Tian Schan Occid., Bostandyk, fonts Aksar-sai, 28 Jul 1949, N. V. Pavlov s.n. (holotype: AA).
Im Hochgebirge, auf sumpfigen Wiesen, auf Urgestein, 5500 ft, C. Koch s.n. (holotype: B, probably destroyed).
There is no location in the species protologue beyond the article title “Beitrage zu einer Flora des Orients”.
“Aus dem Wilhelm’schen Herbr als Poa persica.” Type: Wilhelms (holotype: B, probably destroyed).
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!]).
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
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.
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).
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 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 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 | Poaiberica Fisch. & C.A. Mey. | Soreng et al. 7977 (US) | Russia, Cabardino-Balkaria | KX118741 | KX118723 | KX118758 | MH921349 | MH921374 |
P | P+H | Macropoa | Poa longifolia subsp. longifolia | Soreng et al. 7945 (US) | Russia, Cabardino-Balkaria | KX118742 | KX118724 | KX118760 | MH921350 | MH921375 |
P | P+H | Macropoa | Poa sibirica 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 subsp. pratensis | Gillespie et al. 10592 (CAN) | Turkey | KX118746 | KX118726 | KX118764 | KY378880 | KY378846 |
P | X | Malacanthae | Poa arctica 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 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 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 |
Table S1. Characteristics of the DNA alignments and data partitions and parameters and summary statistics of the PAUP and Bayesian analyses
Data type: (measurement/occurence/multimedia/etc.)
Explanation note: Five DNA sequence alignments for Poa were analysed: ETS, ITS, matK, rpoB-trnC and trnT-trnL-trnF (TLF). For each data partition (five individual markers, plastid, nuclear and combined), the number of samples and the total number of aligned characters are given. For the PAUP analyses, the following statistics are given: the number of parsimony informative (PI) characters, percentage of characters that are parsimony informative, maximum parsimony (MP) tree length (L), number of most parsimonious trees, consistency index excluding uninformative characters (CI) and retention index (RI). Parameters used and statistics of the Bayesian analyses, as determined by the Akaike Information Criterion (AIC) implemented in jModeltest, are given as follows: likelihood score (-InL), number of substitution schemes, substitution rates (rAC, rAG, rAT, rCG, rCT, rGT), character state frequencies (fA, fC, fG, fT), substitution model, proportion of invariable sites and gamma shape parameter.