TAXON 64 (3) • June 2015: 445–467 Peterson & al. • Phylogeny and classification of the Eleusininae

A molecular phylogeny and classification of the Eleusininae with a new genus, Micrachne (Poaceae: Chloridoideae: Cynodonteae)

Paul M. Peterson,1 Konstantin Romaschenko1,2 & Yolanda Herrera Arrieta3

1 Smithsonian Institution, Department of Botany, National Museum of Natural History, Washington D.C. 20013-7012, U.S.A. 2 M.G. Kholodny Institute of Botany, National Academy of Sciences, Kiev 01601, Ukraine 3 Instituto Politécnico Nacional, CIIDIR Unidad Durango-COFAA, Durango, C.P. 34220, Mexico Author for correspondence: Paul M. Peterson, [email protected] ORCID: PMP, http://orcid.org/0000-0001-9405-5528; KR, http://orcid.org/0000-0002-7248-4193

DOI http://dx.doi.org/10.12705/643.5

Abstract The subtribe Eleusininae (Poaceae: Chloridoideae: Cynodonteae) is a diverse group containing about 212 species in 31 genera found primarily in low latitudes in Africa, Asia, Australia, and the Americas, and the classification among these genera and species is poorly understood. Therefore, we investigated the following 28 Eleusininae genera: Acrachne, Afrotrichloris, Apochiton, Astrebla, Austrochloris, Brachyachne, Chloris, Chrysochloa, Coelachyrum, Cynodon, Daknopholis, Dinebra, Diplachne, Disakisperma, Eleusine, Enteropogon, Eustachys, Harpochloa, Leptochloa, Lepturus, Lintonia, Microchloa, Ochthochloa, Oxychloris, Saugetia, Schoenefeldia, Stapfochloa, and Tetrapogon. The goals of our study were to reconstruct the evolutionary history of the subtribe Eleusininae using molecular data with increased species sampling compared to earlier studies. A phylogenetic analysis was conducted on 402 samples, of which 148 species (342 individuals) were in the Eleusininae, using four plastid (rpl32-trnL spacer, ndhA intron, rps16-trnK spacer, rps16 intron) and nuclear ITS 1 & 2 (ribosomal internal transcribed spacer) sequences to infer evolutionary relationships and revise the classification. We found strong support for the following Eleusininae lineages: Acrachne, Apochiton, Astrebla, Austrochloris, Chloris, Chrysochloa, Cynodon, Daknopholis, Dinebra, Diplachne, Disakisperma, Eleusine, Enteropogon, Eustachys, Leptochloa, Lepturus, Micrachne gen. nov., Stapfo­ chloa, and Tetrapogon, and moderate support for Harpochloa and Microchloa. Four species of Brachyachne, including the type, are imbedded within Cynodon; Oxychloris scariosa (monotypic) is sister to Harpochloa-Microchloa; Coelachyrum is polyphyletic since C. lagopoides, Apochiton burttii, and C. poiflorum form a grade, with the latter species sister to Eleusine; Schoenefeldia appears paraphyletic since Afrotrichloris martinii and Schoenefeldia transiens together are sister to Schoenefeldia gracilis; Saugetia fasciculata, Enteropogon brandegeei, and E. chlorideus are embedded within Tetrapogon; Lintonia nutans and Ochthochloa compressa are embedded in Chloris; and Enteropogon mollis and Chloris exilis are embedded in Leptochloa. Our plastid and ITS analyses show rearrangement of lineages within Acrachne and Chloris, indicating possible hybridiza­ tion events or evidence for multiple origins. The molecular results support recognition of a new genus, Micrachne with five species and emendation of Stapfochloa with six species. We provide 22 new combinations, Chloris flagellifera, Ch. nutans, Cynodon ambiguus, C. prostratus, Diplachne divaricatissima, Leptochloa anisopoda, L. exilis, Micrachne fulva, M. obtusi­ flora, M. patentiflora, M. pilosa, M. simonii, Stapfochloa berroi, S. canterae, S. ciliata, S. elata, S. parvispicula, Tetrapogon brandegeei, T. chlorideus, T. fasciculatus, T. pleistachyus, and T. roxburghiana; a new name, Cynodon simonii; and lectotypify Eleusine flagillifera.

Keywords Brachyachne; Chloris; Cynodon; Diplachne; Enteropogon; ITS; Leptochloa; Micrachne, plastid DNA sequences; Stapfochloa; systematics; Tetrapogon

Supplementary Material Electronic Supplement (Figs. S1–S2) and alignment are available in the Supplementary Data section of the online version of this article at http://www.ingentaconnect.com/content/iapt/tax

INTRODUCTION (Apochiton C.E.Hubb., Austrochloris Lazarides, Daknopholis Clayton, Harpochloa Kunth, Ochthochloa Edgew., Oxychloris The subtribe Eleusininae Dumort. (Poaceae: Chlorid­ Lazarides, Pommereulla L.f., Rheochloa Filg. & al., Sclero­ oideae: Cynodonteae) is a morphologically diverse group, dactylon Stapf, Stapfochloa H.Scholz), 16 genera contain 10 characterized by having racemose inflorescences (with a or fewer species: Acrachne Wight & Arn. ex Chiov. (3 spp.), few exceptions) and contains about 212 species in 31 genera Afrotrichloris Chiov. (2 spp.), Astrebla F.Muell. (4 spp.), found primarily in low latitudes in Africa, Asia, Australia, Brachyachne (Benth.) Stapf (9 spp.), Chrysochloa Swallen and the Americas. Ten of the genera are currently monotypic (4 spp.), Coelachyrum Hochst. & Nees (5 spp.), Cynodon Rich.

Received: 7 Nov 2014 | returned for (first) revision: 24 Jan 2015 | (last) revision received: 18 Mar 2015 | accepted: 20 Mar 2015 || publication date(s): online fast track, n/a; in print and online issues, 25 Jun 2015 || © International Association for Plant Taxonomy (IAPT) 2015

Version of Record 445 Peterson & al. • P hylogeny and classification of the Eleusininae TAXON 64 (3) • June 2015: 445–467

(9 spp.), Diplachne P.Beauv. (2 spp.), Disakisperma Steud. MATERIALS AND METHODS (4 spp.), Eleusine Gaertn. (10 spp.), Leptochloa P.Beauv. (5 spp.), Lintonia Stapf (2 spp.), Microchloa R.Br. (6 spp.), Saugetia Taxon sampling. — Our sampling consists of 402 samples, Hitchc. & Chase (2 spp.), Schoenefeldia Kunth (2 spp.), and representing 224 species of grasses, of which 148 species are Tetrapogon Desf. (5 spp.), and 5 genera contain 13 or more included in the Eleusininae and 70 species are included in five species: Chloris Sw. (59 spp.), Dinebra Jacq. (23+ spp.), Entero­ other tribes and subtribes: Centropodieae with a single species, pogon Nees (18 spp.), Eustachys Desv. (13 spp.), and Lepturus Triraphideae with two species, Eragrostideae with ten species, R.Br. (15 spp.) (Clayton & al., 1974, 2006; Peterson & al., 2010a, Zoysieae with seven species; and within the Cynodonteae we 2012, 2014a; Soreng & al., 2014, 2015). include the Tripogoninae with six species, Pappophorinae with In a large phylogenetic study of the Chloridoideae Kunth three species, Traginae with three species, Hilariinae with two ex Beilschm. based on seven DNA sequence markers (ITS, species, Muhlenbergiinae with two species, Scleropogoninae ndhA intron, ndhF, rps16-trnK, rps16 intron, rps3, rpl32-trnL) with two species, Boutelouinae with a single species, Mon­ Eleusininae was monophyletic in the ITS, plastid, and com­ anthochloinae with three species, Orcuttiinae with two species, bined plastid/ITS phylograms (Peterson & al., 2010a). In the Gouiniinae with three species, Cteniinae with a single species, combined plastid/ITS tree, Cynodon appeared paraphyletic Trichoneurinae with a single species, Gymnopogoninae with six with at least two species of Brachyachne embedded within species, Farragininae with two species, Perotidinae with four it; an additional strongly supported clade of three species of species, Triodiinae with a single species, and Aeluropo­dinae Brachyachne was found outside of Cynodon; Chloris, Leptu­ with three species (see Appendix 1). Our ingroup sampling rus, Microchloa, and Tetrapogon were monophyletic, although was primarily focused on 28 morphologically similar genera Chloris appeared polyphyletic in the ITS tree; Astrebla was (Acrachne, Afrotrichloris, Apochiton, Astrebla, Austrochloris, paraphyletic with Schoenefeldia gracilis Kunth embedded Brachyachne, Chloris, Chrysochloa, Coelachyrum, Cynodon, within (but not in the plastid analysis); and Enteropogon, Daknopholis, Dinebra, Diplachne, Disakisperma, Eleusine, Eustachys, Leptochloa, and Tetrapogon were polyphyletic Enteropogon, Eustachys, Harpochloa, Leptochloa, Lepturus, (Peterson & al., 2010a). Based on five DNA sequence mark­ Lintonia, Microchloa, Ochthochloa, Oxychloris, Saugetia, ers (rpl32-trnL, ndhA intron, rps16 intron, ccsA, ITS) the spe­ Schoenefeldia, Stapfochloa, Tetrapogon), most of which have cies of Leptochloa were realigned into four genera (Dinebra, been previously included in the Eleusininae (Peterson & al., Diplachne, Disakisperma, Leptochloa) in the Eleusininae 2010a). The dataset for these 28 genera includes 71% (148/209) and a new genus, Trigonochloa P.M.Peterson & N.Snow, was of the species currently placed in these genera by Clayton & al. erected to accommodate two species in the subtribe Peroti­ (2006–). In addition, we include two species of Neobouteloua dinae P.M.Peterson & al. (Peterson & al., 2012, 2014a; Snow Gould and three species of Dactyloctenium Willd. that are & Peterson, 2012a). Emphasizing possession of an ovoid cary­ currently in the Cynodonteae, incertae sedis. Outside of the opsis that is broadly concave on the hilar (dorsal) side, Scholz Chloridoideae, three species of Danthonioideae (Capeochloa & Müller (2004) placed Chloris lamproparia Stapf in a mono­ cincta subsp. sericea (N.P.Barker) N.P.Barker & H.P.Linder, typic genus, Stapfochloa. Other members of Anderson’s (1974) Danthonia compressa Austin, Rytidosperma pallidum (R.Br.) “Chloris ciliata Sw. complex”, of which C. lamproparia is a A.M.Humphreys & H.P.Linder), one species of Aristidoideae member, have trigonous caryopses without a concave surface. (Aristida gypsophila Beetle), and one species of Panicoideae Stapfochloa and members of the Chloris ciliata complex have (Chasmanthium latifolium (Michx.) H.O.Yates, phylogenetic never been sampled in a molecular study. root) were chosen as the outgroup. A complete list of taxa, The aim of our paper is to discern the evolutionary rela­ voucher information, and GenBank numbers can be found in tionships, test the monophyly, and present a new classification Appendix 1. All collections gathered by PMP after 1998 were of genera in the Eleusininae, excluding only Pommereulla, collected in silica but the majority of samples used in this study Rheochloa and Sclerodactylon of which we have no good were taken from herbarium specimens. material. To accomplish the immediate goal of understand­ To compare ITS and combined plastid incongruence ing the evolutionary history of the Eleusininae we are using among species of the Chloris clade we provide a tanglegram a large dataset of species of Chloridoideae. Here, we present using Enteropogon macrostachyus (Hochst. ex A.Rich.) a new phylogenetic analysis for 148 of the 212 species that Munro ex Benth. and Leptochloa digitata (R.Br.) Domin. are part of the Eleusininae based on analysis of ITS and four as outgroups. Since there is hard incongruence involving plastid regions (rpl32-trnL, ndhA, rps16, rps16-trnK). The the placement of Acrachne outside of the Eleusininae in the number of species sampled within many of these 28 Eleusini­ ITS tree and within the Eleusininae in the plastid tree, we nae genera is expanded compared to earlier studies and for the removed all ITS sequences from this genus in our combined first time we are including species of Afrotrichloris, Dakno­ plastid + ITS tree. pholis, Harpochloa, Ochthochloa, and Oxychloris (Peterson DNA extraction, amplification, and sequencing. — All & al., 2007, 2010a, 2012, 2014a). We discuss morphological procedures were performed in the Laboratory of Analytical and anatomical characters supporting relationships and pro­ Biology (LAB) at the Smithsonian Institution. DNA isolation, pose changes to the classification. In addition, we include amplification, and sequencing of rpl32-trnL spacer and ndhA a description of a new genus, Micrachne, and emend the intron (small single-copy region), rps16-trnK spacer and rps16 description of Stapfochloa. intron (large single-copy region), and ITS was accomplished

446 Version of Record TAXON 64 (3) • June 2015: 445–467 Peterson & al. • Phylogeny and classification of the Eleusininae

following procedures outlined in Peterson & al. (2010a, b). We generations, sampling once per 100 generations. The analy­ specifically targeted four of the plastid regions that proved to be sis was run until the value of the standard deviation of split most informative in our previous studies on chloridoid grasses sequences dropped below 0.01 and the potential scale reduction (Peterson & al., 2010a, b, 2011, 2012, 2014a, b). factor was close to or equal to 1.0. The fraction of the sampled Phylogenetic analyses. — We used Geneious v.5.3.4 values discarded as burn-in was set at 0.25. Posterior prob­ (Drummond & al., 2011) for contig assembly of bidirectional abilities of 0.95–1.00 were considered to be strong support. sequences of rpl32-trnL, ndhA intron, rps16 intron, rps16-trnK, and ITS regions, and MUSCLE v.3.5 (Edgar, 2004) to align consensus sequences and adjust the final alignment. We identi­ RESULTS fied models of molecular evolution for the cpDNA and nrDNA regions using jModelTest (Posada, 2008) and applied maximum Phylogenetic analyses. — Fifty-four percent (893/1639) of likelihood (ML) and Bayesian searches to infer overall phylog­ the sequences used in our study are newly reported here, 46% eny. The combined datasets were partitioned in accordance (746/1639) are from previous studies, and 18.5% (371/2010) are with the number of the markers used. Nucleotide substitution missing (Appendix 1). Total aligned characters for individual models selected by Akaike’s information criterion, as imple­ regions are noted in Table 1. Plastid rpl32-trnL and ITS had the mented in jModelTest v.0.1.1, were specified for each partition highest sequencing success of 93.8% and 92.6%, respectively, (Table 1). The ML analysis was conducted with GARLI v.0.951 whereas recovery of other regions across the entire dataset (Zwickl, 2006). The ML bootstrap analysis was performed ranged from 66.3% to 82.6%. with 1000 replicates, with 10 random addition sequences per Incongruences between the ITS and combined plas- replicate. The output file containing trees of ML found for each tid phylograms. — Summary plastid, ITS, and combined bootstrap dataset was then read into PAUP where the majority- plastid + ITS abbreviated trees are compared (Fig. 1). Sepa­ rule consensus tree was constructed. Bootstrap (BS) values of rate expanded trees for ITS and combined plastid markers 90%–100% were interpreted as strong support, 70%–89% as are given in Figs. S1–S2 (Electr. Suppl.). All three trees pro­ moderate, and 50%–69% as weak. vide strong support (BS 93–96/PP 1.00) for the monophyly Bayesian posterior probabilities (PP) were estimated using of the Eleusininae (in part). Within Eleusininae, the order of a parallel version of MrBayes v.3.1.2 (Huelsenbeck & Ronquist, divergence is: Diplachne, Acrachne, Dinebra, Coel­achyrum 2001; Ronquist & Huelsenbeck, 2003) where the run of eight lagopoides (Burm.f.) Senaratna–Apochiton–C. poiflorum Markov chain Monte Carlo iterations was split between an Chiov.–Eleusine, and remaining genera (plastid tree); Apo­ equal number of processors. Bayesian analysis was initiated chiton–Coelachyrum poiflorum–Eleusine, Diplachne, and with random starting trees and was initially run for 4 million remaining genera (ITS tree); and Acrachne, Dinebra, Coel­

Table 1. Characteristics of rpl32-trnL, ndhA intron, rps16 intron, rps16-trnK and ITS, and parameters used in Bayesian analyses indicated by Akaike information criterion. ndhA rps16 Combined rpl32-trnL intron intron rps16-trnK plastid data ITS Overall Total aligned characters 1117 1373 1110 1146 4746 857 5603 Sequencing success (%) 93.8 66.3 70.7 82.6 78.3 92.6 81.2 Number of new sequences 219 (58%) 135 (51%) 137 (48%) 181 (54%) 672 (53%) 221 (59%) 893 (54%) Likelihood score (–lnL) 8149.6 7692.53 5965.44 7383.91 20188.86 Number of substitution types 6 6 6 6 – 6 – Model for among-site rate variation gamma gamma gamma gamma – gamma – Substitution rates 0.7009 1.4804 1.1730 1.2472 – 1.2221 – 1.6018 2.9329 1.2242 2.6893 3.0333 0.3286 0.6893 0.3295 0.4504 1.6482 1.0772 2.4887 1.3903 1.9104 0.9412 1.0554 3.2724 1.9526 2.6182 4.9300 1.0000 1.0000 1.0000 1.0000 1.0000 Character state frequencies 0.3809 0.3652 0.4022 0.3094 – 0.2403 – 0.1400 0.1336 0.0972 0.1364 0.1908 0.1136 0.1448 0.1630 0.1399 0.2416 0.3654 0.3563 0.3374 0.4142 0.3272 Proportion of invariable sites 0.1336 0.3039 0.2059 0.1640 – 0.2380 – Substitution model GTR + I + G TVM + G TIM3 + I + G TIM3 + G – GTR + I + G – Gamma shape parameter (α) 0.7828 1.0849 0.9792 1.5740 – 1.0171 –

Version of Record 447 Peterson & al. • Phylogeny and classification of the Eleusininae TAXON 64 (3) • June 2015: 445­ –467 Chloris Astrebla Dinebra Lepturus Cynodon Eleusine Micrchloa Acrachne Eustachys Diplachne Leptochloa Tetrapogon Chrysochloa Stapfochloa Enteropogon Microachne Disakisperma Neobouteloua Dactyloctenium Apochiton burttii Afrotrichloris martinii Austrochloris dichanthioides Coelachyrum lagopoides Coelachyrum poiflorum Schoenefeldia gracilis Schoenfeldia transiens Saugetia fasciculata Daknopholis boivinii Oxychloris scariosa Harpochloa flax 85 100 100 100 100 100 100 99 100 100 100 100 100 100 100 100 100 93 62 98 93 100 100 100 100 100 100 100 100 100 100 100 96 94 100 58 COMBINED 88 65 68 78 86 96 82 Aeluropodinae, Triodinae, Aeluropodinae, Triodinae, Cteninae, Farragininae, Gouniinae, Gymnopogoninae Orcuttinae, Perotidinae, Trichoneuriinae 94 Eleusininae 62 68 75 80 Triraphideae Eragrostideae Zoysieae Boutelouinae, Hilariinae, Monantochloinae, Muhlenbergiinae Triodinae Pappophorinae, Traginae,Tripogoninae, OUTGROUP Centropodieae Chloris Astrebla Dinebra Lepturus Cynodon Eleusine Acrachne Eustachys Diplachne Microchloa Leptochloa Micrachne Tetrapogon Chrysochloa Stapfochloa Enteropogon Disakisperma Neobouteloua Dactyloctenium Aeluropodinae Apochiton burttii Austrochloris dichanthioides Afrotrichloris martinii Harpochloa flax Coelachyrum poiflorum Schoenefeldia gracilis Schoenefeldia transiens Saugetia fasciculata Daknopholis boivinii Oxychloris scariosa 82 99 95 85 100 100 100 100 100 100 100 100 100 100 100 100 100 96 98 100 59 92 55 99 98 100 100 100 100 100 95 97 55 100 78 likelihoodtrees. Numbers branchesabove are bootstrap values;thick branches represent statistically supported relation - 100 ITS 82 90 maximum 80 62 Triodinae, Cteninae, Farragininae, Gouniinae, Gymnopogoninae Triodinae, Orcuttinae, Perotidinae, Trichoneuriinae 89 71 93 57 Eleusininae plastid + ITS) 70 Boutelouinae, Hilariinae, Monantochloinae, Muhlenbergiinae Pappophorinae, Traginae,Tripogoninae Triraphideae OUTGROUP Centropodieae Eragrostideae Zoysieae and combined ( , Chloris Dinebra Astrebla Eleusine Lepturus Diplachne Cynodon Acrachne Leptochloa Eustachys Microchloa Tetrapogon Micrachne Enteropogon Chrysochloa Stapfochloa Disakisperma Neobouteloua ITS Dactyloctenium , Austrochloris dichanthioides Afrotrichloris martinii Apochiton burttii Coelachyrum lagopoides Harpochloa flax Coelachyrum poiflorum Schoenefeldia gracilis Schoenefeldia transiens Saugetia fasciculata Daknopholis boivinii Oxychloris scariosa 99 92 98 97 96 94 98 100 100 100 100 93 81 81 81 94 98 77 85 93 83 91 97 61 89 60 62 97 100 100 –100 and/or posterior probability of 0.95–1.00); thin branches indicate unsupported relationships; green branches are derived from the plastid analysis; blue branches are derived from from derived are branches blue analysis; plastid the from derived are branches green relationships; unsupported indicate branches thin 0.95–1.00); of probability posterior and/or –100 76 89 67 PLASTID 69 70 Orcuttinae, Perotidinae, Trichoneuriinae Cteninae, Farragininae, Gouniinae, Gymnopogoninae Asummary comparison ofthe plastid 67

63 96 Fig. 1. Fig. ships (high bootstrap of the ITS analysis; red branches indicate support not found in either the ITS or the plastid phylogeny; black branches are found in all three analyses; arrow indicates the Eleusininae clade. Triraphideae Centropodieae Eragrostideae Zoysieae OUTGROUP 448 Aeluropodinae, Boutelouinae, Hilariinae, Monantochloinae, Triodinae, Muhlenbergiinae, Pappophorinae, Traginae,Tripogoninae, Eleusininae TAXON 64 (3) • June 2015: 445–467 Peterson & al. • Phylogeny and classification of the Eleusininae

achyrum lagopoides–Apochiton–C. poiflorum–Eleusine, Within the Chloris s.str. clade there was hard incongru­ Diplachne, and remaining genera (plastid & ITS tree). In the ence between the combined plastid and ITS datasets; there­ ITS tree Dinebra is sister to the Aeluropodinae (–/0.58) and fore, we did not combine them but provide a detailed tang­ Acrachne is sister to Dactyloctenium (100/1.00), both outside legram (Fig. 3). Both the ITS and combined plastid trees are the Eleusininae. well resolved and the monophyly of Chloris is strongly sup­ Two clades, Acrachne and Diplachne, showed “hard” ported (100/1.00) but their interrelationships differ in each data incongruence and were excluded from the datasets. We refer partition. The four strongly supported clades (100/1.00) in the to hard incongruence as clades with strong bootstrap and pos­ ITS tree labeled A–D nearly correspond to the clades with terior probability, while soft incongruence are clades with weak the same letters in the combined plastid tree. In the combined or no bootstrap and posterior probability less than 95%. We plastid tree clades A and C are each strongly supported (91, used only plastid sequences to represent the Acrachne clade 100/1.00), clade D is moderately supported (87/1.00), and clade in the combined analysis (Fig. 2). However, we retained the B is poorly supported (–/0.69). In the ITS tree, species in A-B full set of sequences for Dinebra and Diplachne where the ITS and C-D form strongly supported clades (95/1.00) whereas in signal completely overrides plastid signal. Due to differences the combined plastid tree, species in A-C form a strongly sup­ in rates of evolution in plastid and nuclear DNA as well as a ported clade (99/1.00) and species in B-D form a moderately notoriously high rate of homoplasy in ITS data, our separate supported clade (71/0.99). Lintonia nutans Stapf and Ochtho­ plastid and ITS phylogenetic trees have different patterns of chloa compressa (Forssk.) Hilu are included within Chloris in resolution (Fig. 1). Combination of the ITS and plastid data the ITS tree and are sister to Chloris s.str. in the plastid tree. In improves branch support significantly within the Eleusininae the combined plastid tree Ochthochloa is sister to the remain­ tree by providing resolution for 54 major nodes with moderate ing members of Chloris and Lintonia is sister to clade A-C to strong support compared to 39 with only plastid data and (99/1.00). In the ITS tree Ochthochloa is sister to Lintonia and 38 with only ITS data. Seven of those nodes gained support in two samples of Chloris mossambicensis K.Schum. (66/0.75), the combined analysis. and all of these are sister to clade A-B (95/1.00). In the ITS and Phylogenetic trees of Eleusininae. — The ML tree from combined plastid tree clade A (C. barbata Sw. and C. prieurii the combined plastid (rpl32-trnL, ndhA, rps16-trnK, rps16) Kunth) and B (C. amethystea Hochst., C. andropogonoides and ITS regions (Fig. 2) is well resolved with strong support E.Fourn., C. cucullata Bisch., C. latisquamea Nash, C. mos­ (99–100/1.00) for clades of the following genera: Acrachne, sambicensis, C. × subdolichostachya Müll.Hal., C. verticillata Dinebra, Apochiton­ , Eleusine, Diplachne (including Dinebra Nutt.) are each composed of the same species, although their divaricatissima (S.T.Blake) P.M.Peterson & N.Snow), Lepto­ interrelationships differ. In the ITS tree clade C contains three chloa (including Chloris exilis Renvoize and Enteropogon species (C. gayana Kunth, C. pilosa Schumach., C. virgata mollis (Nees) Clayton), Astrebla sister to Austrochloris, Sw.) whereas in the combined plastid tree C. orthonoton Döll Disakisperma, Enteropogon, Chloris, Tetrapogon (including and C. rufescens Lag. are additional members of clade C; these Chloris roxburghiana Schult., Enteropogon brandegeei (Vasey) two species are sister to clade C in the ITS tree, but support Clayton, E. chlorideus (J.Presl) Clayton, and Saugetia fascicu­ for this topology is weak. In the combined plastid tree clade D lata Hitchc. & Chase), Lepturus sister to Daknopholis boivinii contains nine species (C. cruciata (L.) Sw., C. cubensis Hitchc. (A.Camus) Clayton, four species of Brachyachne that do not & Ekman, C. divaricata R.Br., C. ekmanii Hitchc., C. halophila include the type (= Micrachne, gen. nov.), Eustachys, Chryso­ Parodi, C. pectinata Benth., C. pycnothrix Trin, C. sagraeana chloa, three species of Chloris (C. canterae Arechav., C. cili­ A.Rich., C. submutica Kunth) whereas in the ITS tree clade ata, C. elata Desv.) with Stapfochloa lamproparia (Stapf) D contains these nine species plus three additional species H.Scholz, and Cynodon (including four species of Brachy­ (C. orthonoton, C. rufescens, C. texana (Vasey) Nash). Chloris achne, of which one is the type). Three species of Microchloa castilloniana Lillo & Parodi is sister to the B-D clade in the are in a moderately supported (85/1.00) clade that is sister to combined plastid tree (71/0.99) and is sister to clade C in the Harpochloa falx (L.f.) Kuntze, and together these are sister ITS tree (100/1.00). to Oxychloris scariosa (F.Muell.) Lazarides. Coelachyrum appears polyphyletic since C. lagopoides and C. poiflorum do not form a clade. The latter species, C. poiflorum, is sister DISCUSSION to Eleusine (100/1.00) whereas C. lagopoides is basal to the Apochiton–C. poiflorum–Eleusine clade (98/1.00). Schoene­ Within the Eleusininae we found strong support for the feldia appears paraphyletic since Afrotrichloris martinii Chiov. following lineages recognized as genera: Acrachne, Apochi­ (type) and Schoenefeldia transiens (Pilg.) Chiov. form a strongly ton, Astrebla, Austrochloris, Chloris, Chrysochloa, Cynodon, supported clade (100/1.00) that is sister to Schoenefeldia graci­ Daknopholis, Dinebra, Diplachne, Disakisperma, Eleusine, lis (type). Together these three species form an unsupported Enteropogon, Eustachys, Leptochloa, Lepturus, Micrachne, clade (–/0.72). Deeper clades with maximum support (100/1.00) Stapfochloa, and Tetrapogon; and moderate support for Harpo­ include: Chloris-Tetrapogon-Lepturus-Microchloa-​Micrachne- chloa and Microchloa. Four species of Brachyachne, including Eustachys-Chrysochloa-Stapfochloa-Cynodon, Microchloa- the type, are embedded within Cynodon; Oxychloris scariosa Micrachne-Eustachys-Chrysochloa-​Stapfochloa-Cynodon, (monotypic) is sister to Harpochloa-Microchloa; Coelachyrum Chrysochloa-Stapfochloa-Cynodon, and Stapfochloa-Cynodon. appears polyphyletic since C. lagopoides, Apochiton burttii

Version of Record 449 Peterson & al. • P hylogeny and classification of the Eleusininae TAXON 64 (3) • June 2015: 445–467

O U T G R O U P Fig. 2A–C. Maximum likelihood tree inferred from combined plastid A (rpl32-trnL, ndhA, rps16, rps16-trnK) and ITS sequences. Numbers Centropodieae above branches are bootstrap values; numbers below branches are 82 posterior probabilities; color indicates native distribution (see legend); 1.00 Triraphideae 100 vertical bars indicate our classification; ● indicates the Eleusininae 1.00 Eragrostideae clade; scale bar = 3%.

100 Zoysieae 1.00 TRIPOGONINAE 56 100 0.61 84 PAPPOPHORINAE 1.00 1.00 TRAGINAE 93 1.00 MUHLENBERGIINAE 89 1.00 SCLEROPOGONINAE 0.75 HILARIINAE 0.73 75 100 BOUTELOUINAE 1.00 1.00 MONANTHOCHLOINAE 100 Neobouteloua paucirracemosa 98 1.00 Neobouteloua lophostachya 1.00 100 Dactyloctenium robecchii 1.00 80 Dactyloctenium geminatum 1.00 Dactyloctenium scindicum

68 ORCUTTIINAE 1.00 80 GOUINIINAE 88 1.00 100 CTENIINAE 1.00 0.97 100 TRICHONEURINAE 1.00 68 GYMNOPOGONINAE 0.94 65 FARRAGININAE 0.80 75 0.99 PEROTIDINAE TRIODIINAE 0.81 AELUROPODINAE Acrachne racemosa Zavattari 1196 Acrachne racemosa Jackson 1408 100 Acrachne racemosa Strohbach 1308 Acrachne racemosa Zavattari 1195 1.00 Acrachne racemosa Pappi 5990 Acrachne 62 Acrachne racemosa Rattray 1303 0.97 Dinebra chinensis 88 100 Dinebra squarrosa 1.00 1.00 Dinebra retroflexa Snow 6950 100 Dinebra retroflexa Ndegwa 610 1.00 100 Dinebra xerophila 65 1.00 Dinebra marquisensis 0.96 98 Dinebra panicea var. mucronata 100 1.00 97 Dinebra panicea var. brachiata Peterson 20086 24H02 1.00 Dinebra panicea var. brachiata Peterson 22185 17D06 1.00 98 Dinebra panicoides 72 98 100 1.00 Dinebra viscida Snow 6528a 1.00 Dinebra viscida Peterson 22184 1.00 1.00 65 Dinebra caudata 0.66 Dinebra aquatica ELEUSININAE 98 Dinebra coerulescens Dinebra 1.00 100 Dinebra scabra 1.00 Dinebra nealleyi 63 100 Dinebra haareri 1.00 1.00 Dinebra somalensis 55 Dinebra neesii 0.61 98 100 Dinebra ligulata 1.00 1.00 Dinebra southwoodii 95 Dinebra decipiens var. peacockii 1.00 100 78 Dinebra decipiens var. decipiens Waterhouse 5948 91 1.00 1.00 Dinebra decipiens var. decipiens Snow 7328 100 Dinebra decipiens var. asthenes Snow 7327 1.00 1.00 Dinebra decipiens var. asthenes Snow 7355 Coelachyrum lagopoides 98 100 Apochiton burttii Greenway 11513 1.00 1.00 Apochiton burttii Peterson 24163 100 Coelachyrum poiflorum 1.00 100 Eleusine multiflora Qing Liu 100 100 1.00 Eleusine multiflora Peterson 24272 1.00 75 Eleusine multiflora Davidse 30264 Eleusine jaegeri Hilu 203 North America 0.98 100 Eleusine jaegeri Kindeketa 762 96 1.00 100 Eleusine jaegeri Peterson 24299 1.00 Eleusine floccifolia Hilu 2285 South America 1.00 Eleusine africana Qing Liu 090 100 Eleusine africana Peterson 24072 100 1.00 Eleusine africana Kayombo 424 1.00 Eleusine africana Seydel 2798 Africa Eleusine africana Seydel 3889 Eleusine tristachya Beck 3227 Eleusine 100 Eleusine tristachya Qing Liu 107 1.00 Eleusine tristachya Hilu 2464 Australia + Pacific Eleusine tristachya Zardini 35791 Eleusine indica Peterson 21362 93 Eleusine indica Peterson 24429 Eleusine indica Peterson 23802 SE Asia + Japan 1.00 Eleusine indica Chase 9922 Eleusine indica Brown 628 Eleusine coracana Qing Liu 093 Eleusine coracana Qing Liu 103 Fig. 2B Eleusine coracana Peterson 24112 450 0.03 Eleusine kigeziensis Purseglove 3384 TAXON 64 (3) • June 2015: 445–467 Peterson & al. • Phylogeny and classification of the Eleusininae

B Fig. 2A Dinebra divaricatissima 100 Diplachne fusca subsp. uninervia Peterson 21305 1.00 Diplachne fusca subsp. muelleri Diplachne fusca subsp. uninervia Peterson 20786 Diplachne fusca subsp. fusca Peterson 24322 Diplachne Diplachne fusca subsp. fusca Walsh 6558 Leptochloa digitata 100 100 Enteropogon mollis Proosdij 813 1.00 Enteropogon mollis Steyermark 29192 1.00 94 100 Chloris exilis Anderson 37049 1.00 1.00 Chloris exilis Anderson 37190 91 94 100 Leptochloa pluriflora Swallen 10100 1.00 1.00 1.00 Leptochloa pluriflora Sohns 1258 100 Leptochloa pluriflora Peterson 15048 Leptochloa crinita Leptochloa 1.00 Leptochloa virgata Peterson 15088 100 Leptochloa virgata Vargas 2710 Leptochloa virgata Rimachi 8359 1.00 Leptochloa chloridiformis Myndel 2662 Leptochloa chloridiformis Myndel 3741 78 100 Austrochloris dichanthioides 0.95 Astrebla lappacea 1.00 100 1.00 64 Astrebla pectinata Astrebla 0.78 Astrebla elymoides 100 Disakisperma yemenicum Peterson 24254 1.00 Disakisperma yemenicum Verdcourt 3275 100 100 Disakisperma eleusine Oakes 1242 1.00 100 1.00 Disakisperma eleusine Snow 6982 1.00 Disakisperma obtusiflorum Belsky 527 100 77 63 Disakisperma obtusiflorum Peterson 24211 1.00 0.99 94 68 0.66 Disakisperma obtusiflorum Peterson 24198 1.00 Disakisperma obtusiflorum Peterson 24243 Disakisperma 1.00 98 Disakisperma dubium Peterson 8105 1.00 Disakisperma dubium Peterson 24554 100 Disakisperma dubium Peterson 24566 1.00 Disakisperma dubium Peterson 24665 99 Disakisperma dubium Peterson 24845 0.99 Disakisperma dubium Peterson 22334 65 Disakisperma dubium Peterson 24905 0.97 Disakisperma dubium Peterson 24453 Schoenefeldia gracilis 100 65 Afrotrichloris martinii Herlocker 459 0.72 100 1.00 Afrotrichloris martinii Hemming 3407 1.00 1.00 100 Schoenefeldia transiens Spjut 3920 1.00 Schoenefeldia transiens Greenway 9781 Schoenefeldia transiens Peterson 24216 100 Enteropogon ramosus Peterson 14367 1.00 100 Enteropogon ramosus Saarela 1663 1.00 Enteropogon acicularis Enteropogon sechellensis Fosberg 49814 88 Enteropogon sechellensis Peterson 23815 100 0.97 Enteropogon sechellensis Peterson 23833 1.00 Enteropogon macrostachyus Smook 9026 86 Enteropogon macrostachyus Oakes 1250 88 1.00 Enteropogon macrostachyus Laegaard 15902 1.00 87 Enteropogon monostachyus Barber 2607 92 1.00 Enteropogon monostachyus Clayton 5954 1.00 71 Enteropogon barbatus Kabuye 575 Enteropogon 0.99 64 Enteropogon macrostachyus Peterson 24197 0.97 Enteropogon macrostachyus Peterson 24228 87 84 Enteropogon dolichostachyus Blake 23488 1.00 1.00 56 Enteropogon dolichostachyus Lazarides 7416 61 0.67 Enteropogon dolichostachyus Gould 13680 0.90 Enteropogon rupestris Vesey 4973 90 Enteropogon rupestris Gillet 13082 1.00 58 Enteropogon rupestris Bogdan 4026 1.00 Enteropogon rupestris McKinnon 124

100 1.00 Chloris (see Fig. 3) Saugetia fasciculata 100 71 Enteropogon brandegeei Enteropogon chlorideus Gould 12679 1.00 0.97 100 98 Enteropogon chlorideus Meyer s.n. 1.00 Enteropogon chlorideus Sohns 384 1.00 100 Enteropogon chlorideus Leavenworth 1900 0.66 100 Tetrapogon tenellus Peterson 23794 North America 1.00 62 1.00 Tetrapogon tenellus Peterson 24244 0.95 95 Tetrapogon bidentatus Bogdan 3903 1.00 Tetrapogon bidentatus Greenway 12778 85 Tetrapogon 100 Chloris roxburghiana Peterson 24194 South America 1.00 1.00 64 Chloris roxburghiana Peterson 23812 53 Chloris roxburghiana Peterson 24225 63 0.98 Tetrapogon villosus Ash 2561 94 0.79 Tetrapogon villosus Johannes s.n. Africa 100 Tetrapogon cenchriformis BN 7701 1.00 1.00 70 Tetrapogon cenchriformis Peterson 24240 0.64 Tetrapogon cenchriformis Greenway 8844 Australia + Pacific Tetrapogon cenchriformis Peterson 24192 Daknopholis boivinii 100 75 Lepturus gasparricensis 1.00 100 1.00 Lepturus geminatus SE Asia + Japan 1.00 99 Lepturus repens Peterson 23835 96 1.00 52 Lepturus repens Whistler 9853 Lepturus Lepturus radicans 1.00 0.52

0.03 Fig. 2C

Version of Record 451 Peterson & al. • P hylogeny and classification of the Eleusininae TAXON 64 (3) • June 2015: 445–467

C Fig. 2B

Oxychloris scariosa Harpochloa falx 76 Microchloa altera Brand 315 1.00 85 100 Microchloa altera Robinson 6362 1.00 100 1.00 Microchloa altera Malaisse 16572 Microchloa caffra Smook 10441 1.00 100 Microchloa caffra Peterson 24288 Microchloa 1.00 100 100 Microchloa kunthii Peterson 24048 1.00 100 1.00 Microchloa kunthii Hagerup 455 1.00 100 Microchloa kunthii Peterson 24255 1.00 100 Microchloa kunthii Peterson 24716 1.00 Microchloa kunthii Peterson 22152 100 1.00 100 Brachyachne fulva 100 Brachyachne patentiflora 1.00 1.00 96 Brachyachne obtusiflora Schweinfurth 2150 Micrachne 1.00 Brachyachne obtusiflora Laegaard 16295 Eustachys paspaloides 100 Eustachys tenera 62 1.00 82 Eustachys glauca 0.92 80 0.99 99 Eustachys petraea Taylor 12539 0.94 Eustachys petraea Strong 3124 1.00 100 Eustachys neglecta 1.00 Eustachys floridana Eustachys distichophylla Eustachys 73 Eustachys retusa 93 0.94 99 Eustachys glabrescens Pedersen 6121 1.00 0.61 53 1.00 Eustachys glabrescens Pedersen 6307 0.68 99 Eustachys calvescens Chase 10971 1.00 Eustachys calvescens Rojas 6633 Eustachys caribaea 100 Chrysochloa orientalis 1.00 100 Chrysochloa hindsii 1.00 Chrysochloa annua Chrysochloa Chloris canterae Schinini 5321 Chloris canterae Pedersen 9597 100 Chloris canterae Smith 01 1.00 Chloris canterae var. grandiflora 100 Chloris elata Webster 10789 1.00 Chloris elata Swallen 14461 570.80 96 Chloris elata Spiaggi 40 Stapfochloa 0.68 1.00 Chloris elata Beck 4454 Chloris elata Acosta Solis 12667 Chloris ciliata Lundell 14872 100 0.59 86 60 Chloris ciliata Peterson 24632 1.00 0.99 0.99 Chloris ciliata Peterson 24622 Stapfochloa lamproparia Cynodon maritimus var. vaginiflorus Cynodon maritimus 89 Brachyachne convergens 1.00 Brachyachne ambigua 0.68 Brachyachne tenella 100 96 Brachyachne ciliaris Mullen s.n. 0.60 Brachyachne ciliaris Latz 14224 1.00 1.00 97 Cynodon hirsutus Cynodon incompletus Oakes 663 1.00 90 1.00 Cynodon incompletus Russell 2408 0.69 Cynodon incompletus Smook 1052 90 Cynodon radiatus Laegaard 21611 1.00 Cynodon radiatus Clayton 5836 85 Cynodon aethiopicus Vesey-Fitzgerald 6475 0.92 0.97 Cynodon aethiopicus Peterson 24268 76 0.87 Cynodon aethiopicus Peterson 24318 1.00 Cynodon plectostachyus Cynodon nlemfuensis Peterson 23791 0.79 52 Cynodon nlemfuensis Peterson 24113 0.94 0.84 68 Cynodon nlemfuensis Peterson 24058 1.00 Cynodon nlemfuensis Peterson 24207 Cynodon nlemfuensis Molina 25803 66 Cynodon aethiopicus Smook 4140 1.00 Cynodon nlemfuensis Oakes 1602 Cynodon nlemfluensis Laegaard 19845 Cynodon nlemfuensis Laegaard 19876 Cynodon nlemfuensis Hansen 13002 Cynodon Cynodon nlemfuensis Lobo 2099 62 Cynodon nitidus North America 0.97 Cynodon erectus Asplund 10905 Cynodon erectus Martin 1814 Cynodon affinis Booth 1251 0.55 Cynodon affinis Collins 63 South America 57 Cynodon affinis Cabrera 5149 0.71 Cynodon aristiglumis Cynodon hirsutissimus Cynodon mucronatus Fosberg 273 Africa Cynodon mucronatus Hermann 8502 0.81 97 Cynodon transvaalensis Smook6710 1.00 Cynodon transvaalensis Laegaard 20560 Australia + Pacific Cynodon aristulatus Fassett 26000 Cynodon aristulatus Killip 31600 Cynodon aristulatus Egler 42-314 0.68 Cynodon pascuus Moran 12518 SE Asia + Japan Cynodon dactylon Peterson 22000 57 Cynodon dactylon var. pilosus Peterson 24626 0.84 Cynodon dactylon var. pilosus Saarela1606 Cynodon dactylon var. pilosus Marsh 141 82 Cynodon dactylon var. longiglumis Peterson 19701 Cynodon dactylon var. longiglumis Dunkle 8366 0.03 1.00 Cynodon dactylon var. longiglumis Dunkle 8652

452 Version of Record TAXON 64 (3) • June 2015: 445–467 Peterson & al. • Phylogeny and classification of the Eleusininae

C.E.Hubb, and C. poiflorum appear in a grade, with the lat­ combined plastid tree (Fig. 2A) indicates that the E. cora­ ter species sister to Eleusine; Schoenefeldia appears paraphy­ cana–E. indica–E. kigeziensis S.M.Phillips–E. tristachya letic since Afrotrichloris martinii and Schoenefeldia transiens (Lam.) Lam. clade is sister to five accessions of E. africana. together are sister to Schoenefeldia gracilis; Saugetia fascicu­ In the plastid tree of Liu & al. (2011) E. africana, E. kigeziensis, lata (monotypic), Enteropogon brandegeei, and E. chlorideus and E. coracana share a recent common ancestor and these are are embedded within Tetrapogon; and Lintonia nutans (mono­ sister to E. indica–E. tristachya. The ITS sequence marker in typic) and Ochthochloa compressa (monotypic) are embedded our study seems to be the primary marker responsible for this in Chloris. incongruence since E. africana, E. coracana, and E. indica Acrachne, Apochiton, Coelachyrum, Dactyloctenium, form a clade based solely on plastid sequences (see Electr. Dinebra, and Eleusine. — In order to detect different plastid Suppl.; Fig. S2). However, after realignment of the rpl32-trnL haplotypes and to test the consistency of phylogenetic signal sequences for accessions of Liu 090 (E. africana) and Liu provided by plastid DNA markers, we included six samples 093, 103 (E. coracana) used here and in Liu & al. (2011), we of Acrachne racemosa from five countries in east and south­ found that they still aligned with our E. africana and E. cora­ ern Africa, and these form a highly supported clade with no cana–E. indica–E. kigeziensis–​E. tristachya clades, respec­ genetic variability (Fig. 2A). Based on possessing ornamented tively. Our results support the recognition of E. africana at fruits with free pericarps, Phillips (1995) considered Eleusine, the specific level, rather than as a subspecies of E. coracana Acrachne, and Dactyloctenium to be closely related. Addition­ or E. indica (Phillips, 1995; Cope, 1999). ally, the species in these three genera share inflorescences with Sister to Eleusine is Coelachyrum poiflorum, a relationship digitately to subdigitately arranged racemes usually with bi- that has been found in previous studies (Peterson & al. 2010a, seriate and secund spikelets (Watson & Dallwitz, 1992; Clayton 2012, 2014a; Liu & al. 2011; Snow & al., 2013). Coel­achyrum & al, 2006–). Our data support this assumption, at least in part, lagopoides and C. poiflorum do not form a clade but are in a since Acrachne is strongly supported sister of Dactyloctenium grade separated by two samples of Apochiton burttii Phillips in the ITS tree outside of the Eleusininae, but in the combined (1995) indicated, “Coelachyrum is a rather heterogeneous plastid tree Acrachne is the second basal split in the Eleusininae assemblage … the species vary considerably in inflorescence (Fig. 1). Phillips (1972) noted that species of Acrachne can some­ structure and hairiness of the spikelets.” Emilio Chiovenda, times be mistaken for Eleusine multiflora Hochst. ex A.Rich. who first described C. poiflorum, later transferred the spe­ and that the latter species is somewhat intermediate between cies into Eleusine (Chiovenda, 1898, 1912). There are quite a the two genera by sharing acropetal floret maturation (from the few morphological differences between C. poiflorum and spe­ base upwards), mucronate lemmas, and a delicate pericarp that cies of Eleusine, such as dorsiventrally compressed caryopses ruptures shedding the grain from the floret. However, Eleusine (versus ellipsoid to trigonous) and lemmas with long shaggy does not share a recent common ancestor with Acrachne in hairs (versus glabrous; Cope, 2007). We still lack samples for any of our trees and E. multiflora is clearly part of Eleusine three species of Coelachyrum (C. brevifolium Hochst. & Nees suggesting that these characters are homoplasious. Further [type], C. longiglume Napper, C. piercei (Benth.) Bor) so we investigation of Acrachne and Dactyloctenium using low-copy hesitate to make any taxonomic changes at the moment. The het­ nuclear genes is needed to test for hybridization and subsequent erogeneous nature of Coelachyrum was observed in Snow & al. genomic exchange or introgression, given their discordant posi­ (2013), in which C. yemenicum (Schweinf.) S.M.Phillips was tions in the plastid and ITS trees (Romaschenko & al., 2014). resolved within the Disakisperma clade, prompting the com­ In our ITS tree, Dinebra is unresolved outside of the bination Disakisperma yemenicum (Schweinf.) P.M.Peterson Eleusininae, a result also found in Peterson & al. (2012). We & N.Snow. verify the placement of two additional species of Dinebra (see Afrotrichloris, Astrebla, Austrochloris, Daknopholis, Fig. 2A) for the first time using molecular data: D. aquatica Diplachne­ , Disakisperma, Enteropogon, Leptochloa, Lepturus, (Scribn. & Merr.) P.M.Peterson & N.Snow and D. haareri Saugetia, Schoenefeldia, and Tetrapogon. — Within Disaki­ (Stapf & C.E.Hubb.) P.M.Peterson & N.Snow, where the sperma, we include a larger sample of D. dubium (Kunth) former is an unsupported sister to D. coerulescens (Steud.) P.M.Peterson & N.Snow (eight versus two individuals, see P.M.Peterson & N.Snow in a larger strongly supported Snow & al., 2013) in order to test intraspecific variation in the clade that includes D. caudata (K.Schum.) P.M.​Peterson & ​ species (Fig. 2B). There appear to be no gross morphological N.Snow, D. panicoides (J.Presl) P.M.Peterson & N.Snow, features or geographic components separating Peterson 8105 and D. viscida (Scribn.) P.M.Peterson & N.Snow; and the latter and Peterson 24554 from the other six accessions even though is a strongly supported sister to D. somalensis (Stapf) P.M.Peter​ ­ they form a strongly supported clade (phylogenetic signal in son &​ N.Snow, a clade whose deep relationships are unclear. ITS) sister to the other six accessions. Three species, D. eleusine Based on plastid markers, previous studies of Eleusine (Nees) P.M.Peterson & N.Snow, D. obtusiflorum (Hochst.) indicated that E. africana Kenn.-O’Byrne and E. coracana P.M.Peterson & N.Snow, and D. yemenicum, which form a (L.) Gaertn. (finger millet) share a recent common ances­ strongly supported clade, have clavicorniculate hairs on the tor (Liu & al., 2011, 2013). We here offer one new insight lemmas whereas D. dubium, the sister species to this clade, has within Eleusine because our results differ from these. Based acute or rounded lemmatal hairs (Snow & al., 2103). on five samples of E. africana, five samples of E. indica Within the Diplachne clade we include two samples (L.) Gaertn., and three samples of E. coracana our ITS/ of D. fusca (L.) P.Beauv. ex Roem. & Schult. subsp. fusca and

Version of Record 453 Peterson & al. • Phylogeny and classification of the Eleusininae TAXON 64 (3) • June 2015: 445–467 Chloris 100 1.00 62 0.89 62 0.98 0.003 71 0.99 0.69 Leptochloa digitata 0.77 99 B 53 1.00 0.99 87 1.00 D 94 1.00 52 53 0.80 68 0.78 0.98 91 1.00 0.78 82 95 52 0.80 1.00 1.00 A 88 63 1.00 1.00 63 1.00 100 1.00 C 98 1.00 Enteropogon macrostachyus 100 1.00 100 1.00 100 Lintonia nutans 1.00 Peterson 23974 Peterson 23949 Peterson 24262 Peterson 24070 Chloris texana 96 70 70 Bidgood 3170 Chloris cubensis 1.00 1.00 0.86 Peterson 24184 Peterson 24305 Chloris pectinata Chloris divaricata Pedersen 6378 Chloris amethystea 95 Chloris sagraeana 1.00 66 0.97 154 Chloris verticillata Chloris castilloniana 1 68 0.96 Chloris submutica Reeder 4845 . Chloris x subdolichostachya Chloris submutica McGregor 706 Chloris halophila Peterson 13970 PLASTID Chloris halophila Laegaard 71450 Chloris cucullata Isely 10647 Chloris submutica Peterson 22393 Chloris submutica Peterson 24842 Chloris submutica Peterson 24939 Chloris submutica Peterson 24560 Chloris ekmanii Liogier 12416 Chloris pycnothrix Harley 17042 Chloris cucullata Crutchfield 5475 Chloris ekmanii Liogier 15884 Chloris barbata King 4038 Chloris prieurii Laegaard 17061 Chloris pycnothrix Peterson 24172 Chloris latisquamea Stanford s.n. Chloris pycnothrix Chloris pycnothrix Chloris pycnothrix Chloris pycnothrix Chloris prieurii Laegaard 17863 Chloris pycnothrix Peterson 22278 Chloris barbata Ernst 1507 Chloris cruciata Correll 50431 Chloris barbata Bentley 102 Allen s. n Chloris cruciata Correll 48667 Chloris pycnothrix Chloris pycnothrix Chloris mossambicensis Oakes 1211 Chloris pycnothrix Chloris barbata Saarela 1830 Chloris barbata Saarela 1740 Chloris latisquamea Fleetwood s.n. Chloris pycnothrix Chloris andropogonoides Swallen 10091 Chloris mossambicensis Peterson 24219 Chloris andropogonoides Swallen 10232 Chloris mossambicensis Peterson 23803 Chloris andropogonoides Peterson 18839 Chloris barbata Peterson 23795 Chloris barbata Peterson 22255 Chloris andropogonoides Cory 52467 Chloris divaricata var. cynodontoides Chloris rufescens Chloris pilosa Chloris virgata Peterson 24286 PRI Chloris pilosa Brenan 9601 Chloris gayana Batty 125 BAR Chloris gayana Balleza 1788 Chloris orthonoton Harley 16629 Chloris gayana Peterson 24253 Ochthochloa compressa USD A Chloris gayana Peterson 23972 Chloris gayana Peterson 23997 Chloris orthonoton Pedersen 5817 Ochthochloa compressa Gloves 485 Chloris virgata Peterson 249 1 Chloris virgata Peterson 24789 Chloris virgata Peterson 24659 Chloris pilosa Peterson 23919 EKM PYC SUB HAL MOS LAT AND CRU Chloris virgata Peterson 24954 Ochthochloa compressa Rechinger 29410 Chloris virgata Peterson 24854 Chloris virgata Peterson 21468 VIR GAY ORT PIL AND LAT CUC MOS PIL PRI CRU VIR HAL BAR PYC GAY SUB EKM 1 . Allen s. n Chloris andropogonoides Cory 52467 Chloris andropogonoides Swallen 10091 Chloris andropogonoides Swallen 10232 Chloris pycnothrix Bidgood 3170 Chloris pycnothrix Peterson 24262 Chloris andropogonoides Peterson 18839 ITS 65 Chloris x subdolichostachya Chloris virgata Peterson 249 1 Chloris pycnothrix Peterson 24172 Chloris virgata Peterson 24789 Chloris virgata Peterson 24659 Chloris virgata Peterson 24954 Chloris pycnothrix Peterson 24184 Chloris virgata Peterson 21468 0.64 Chloris virgata Peterson 24854 Chloris pycnothrix Peterson 22278 Chloris prieurii Laegaard 17863 Chloris cruciata Correll 50431 Chloris pycnothrix Harley 17042 Chloris pycnothrix Pedersen 6378 Chloris cruciata Correll 48667 Chloris prieurii Laegaard 17061 89 Chloris barbata Saarela 1830 Chloris virgata Peterson 24286 Chloris halophila Peterson 13970 Chloris halophila Laegaard 71450 Chloris sagraeana Chloris orthonoton Harley 16629 Chloris barbata Saarela 1740 Chloris barbata Peterson 23795 Chloris barbata Bentley 102 Chloris barbata Peterson 22255 Chloris barbata Ernst 1507 Chloris barbata King 4038 Ochthochloa compressa Gloves 485 Ochthochloa compressa USDA 154 Chloris cucullata Crutchfield 5475 72 Chloris cucullata Isely 10647 0.99 Chloris gayana Peterson 23972 87 73 Chloris ekmanii Liogier 12416 Chloris gayana Peterson 23997 Chloris rufescens Chloris ekmanii Liogier 15884 Chloris pycnothrix Peterson 24070 Chloris latisquamea Fleetwood s.n. 1.00 Chloris pycnothrix Peterson 24305 Chloris pycnothrix Peterson 23949 61 0.99 Chloris pycnothrix Peterson 23974 Chloris latisquamea Stanford s.n. Chloris verticillata Chloris pilosa Peterson 23919 99 Chloris gayana Batty 125 100 1.00 91 Chloris cubensis Chloris submutica Peterson 22393 Chloris texana 0.99 1.00 Chloris gayana Balleza 1788 Chloris submutica McGregor 706 Chloris submutica Peterson 24560 Chloris submutica Peterson 24939 Chloris submutica Peterson 24842 74 Chloris pilosa Brenan 9601 1.00 0.99 Chloris pilosa 72 0.98 89 73 77 Chloris submutica Reeder 4845 0.94 83 0.89 1.00 0.91 95 100 90 1.00 Chloris gayana Peterson 24253 Chloris divaricata var. cynodontoides Chloris divaricata 1.00 Ochthochloa compressa Rechinger 29410 100 1.00 66 87 1.00 Chloris pectinata Chloris castilloniana 1.00 1.00 67 Chloris mossambicensis Peterson 23803 0.93 98 87 0.89 Chloris amethystea 1.00 0.95 52 100 54 1.00 0.96 0.54 0.86 99 98 94 84 93 1.00 1.00 Chloris orthonoton Pedersen 5817 1.00 1.00 100 1.00 1.00 100 100 1.00 1.00 0.53 A B C D 100 1.00 100 1.00 Chloris mossambicensis Oakes 1211 Chloris mossambicensis Peterson 24219 98 Leptochloa digitata Risler 476 1.00 Lintonia nutans 0.68 96 1.00 95 1.00 95 1.00 66 0.75 Enteropogon macrostachyus Peterson 24228 59 0.55 0.02 100 1.00 ica f r North America South America A Australia + Pacific Chloris

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these differ little genetically from the two other subspecies free pericarps and inflorescences with biseriate racemes (2 or of D. fusca. Contrary to our best guess in Snow & Peterson 3 in Austrochloris and 1 in Astrebla; Watson & Dallwitz, 1992; (2012b), we report that Dinebra divaricatissima aligns within Clayton & al., 2006–). Geography seems to have played a large the Diplachne clade, again with very little genetic variation role in the evolution of these grasses since the Austrochloris- between it and D. fusca. Astrebla lineage is an Australian endemic clade. We include for the first time a full complement of DNA Our tree depicts Schoenefeldia paraphyletic with a clade markers for Enteropogon mollis (Columbus & al., 2007) and of two samples of Afrotrichloris martinii embedded within. Chloris exilis, and confirm their placement in Leptochloa s.str. At this time it would be premature to submerge Afrotrichlo­ Anderson (1974) mentioned that “Chloris mollis Nees (= Entero­ ris martinii (type) within Schoenefeldia because we have not pogon mollis) is unique in the genus [Chloris] in having sterile sampled A. hyaloptera Clayton and we have no support for the florets with a prominently pubescent callus and long awns”, and Schoenefeldia gracilis (type)–S. transiens–A. martinii clade that it has many characters similar to Gymnopogon P.Beauv. in the ITS tree where Schoenefeldia appears paraphyletic (see Possession of long-awned (4–12 mm) florets is a common fea­ Electr. Suppl.: Fig. S1). However, we have moderate support ture of Leptochloa s.str., although this trait was only recently (85/–, see Fig. 1A) for the Schoenefeldia gracilis–S. tran­ found important after viewing a molecular phylogeny (Peterson siens–A. martinii clade in the plastid tree. Further study may & al., 2012). Only the Australian endemic, Leptochloa digitata, indicate hybridization and introgression with members of this which is sister to the rest of Leptochloa, has entire to aristu­ clade and other species in the Eleusininae. late lemmas (Nightingale & al., 2005b). Anderson (1974) was We found strong support for a monophyletic Enteropogon correct in assuming Chloris mollis was misplaced in Chloris in its current circumscription, which includes eight species, but the characters he mentions as being intermediate between contrary to earlier reports of polyphyly of the genus (Hilu Gymnopogon and Chloris, i.e., short, stiff leaf blades and sterile & Alice, 2001; Columbus & al., 2007; Peterson & al. 2010a; florets, seem to have arisen in parallel, since the Gymnopogo­ GPWG, 2011). None of these previous studies included more ninae is far removed from Eleusininae (Peterson & al., 2014a). than two species of the genus, whereas Columbus & al. (2007) Renvoize (1984) indicated that Chloris exilis may be confused studied two species, Enteropogon chlorideus and E. mollis, with Enteropogon mollis, although the latter has 2(–3)-flowered which are here treated in Tetrapogon and Leptochloa, respec­ spikelets and strongly dorsally compressed lower lemmas. In tively. Peterson & al. (2010a) depicted a clade with two species our tree these two species are in a grade between Leptochloa of Enteropogon (E. macrostachyus, E. ramosus B.K.Simon) digitata (basal) and L. chloridiformis (Hack.) Parodi–L. crinita treated here in the genus and one species of Eustachys. Tra­ (Lag.) P.M.Peterson & N.Snow–L. pluriflora (E.Fourn.) P.M.​ ditionally, Enteropogon was separated from Chloris by hav­ Peterson & N.Snow–L. virgata (L.) P.Beauv. (derived). We also ing a single, rarely two, spikelike racemes and fertile lemmas verify that Leptochloa chloridiformis is included in Leptochloa that are strongly dorsally compressed with raised midveins s.str., as previously suggested (Peterson & al., 2012), bringing (Clayton, 1967; Anderson, 1974). However, Enteropogon acicu­ the genus to seven species. laris (Lindl.) Lazarides from Australia and E. dolichostachyus This study corroborates the strong support found by Peter­ (Lag.) Keng ex Lazarides from Southeast Asia have many son & al. (2010a) for a monophyletic Astrebla sister to Austro­ (3–14) digitate to subdigitately arranged spikelike racemes and chloris dichanthioides (Everist) Lazarides, and we confirm that dorsally compressed lemmas with raised midveins, and our in our study using three species of Astrebla, including A. pecti­ data confirm these are part of Enteropogon. Tateoka (1962) nata (Lindl.) F.Muell. ex Benth. (type). Jozwik (1969) prepared found simple, angular starch grains in Enteropogon while an excellent morphological, anatomical, and hybridization study Chloris had compound starch grains, and both E. acicularis of the four species of Astrebla but there is no mention of their and E. dolichostachyus have simple, angular starch grains affinity with other genera, other than Tateoka (1957) placing the (Anderson, 1974). Delimitation of E. macrostachyus is problem­ genus in the Chlorideae based on chromosome number. Clayton atic (Fig. 2B) where there are two separate clades of E. macro­ & Renvoize (1986) indicate Astrebla has “no obvious relatives stachyus, implying a polyphyletic origin (or one of these clades … but some resemblance to Tetrapogon”. Lazarides (1972), the may represent an undescribed taxon). Diploids and tetraploids author of Austrochloris, felt that Chloris was somehow related have been documented in individuals of E. macrostachyus to A. dichanthioides but noted that dorsally compressed florets (Roodt & Spies, 2002, 2003a, b). There is little ITS variation and caryopses in Austrochloris were important features not among all accessions of E. barbatus C.E.Hubb, E. macro­ found in Chloris. In our tree the Austrochloris-Astrebla clade stachyus, E. monostachyus (Vahl) K.Schum., and E. sechel­ lies between Leptochloa (basal) and Disakisperma (derived), lensis (Baker) T. Durand & Schinz (see Electr. Suppl.: Fig. S1) two splits (Afrotrichloris-Schoenefeldia and Enteropogon) suggesting homogenization or simply recent, rapid radiation removed from the derived Chloris clade. Astrebla and Aus­ (i.e., they form a clade). However, moderate genetic variation trochloris have dorsally compressed to terete caryopses with occurs among the plastid markers in these four species, and their

◄ Fig. 3. A comparison (tanglegram) of the ITS and combined plastid phylograms for species of Chloris s.str. (see Fig. 2). Numbers above branches are bootstrap values; numbers below branches are posterior probabilities; major clades are labeled A–D; abbreviation for species uses the first three letters; dashed lines indicate changed positions of species between and within clades; arrow indicates the Chloris clade; color indicates native distribution (see legend); scale bars = 2% (ITS) and 0.3% (plastid).

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alignment appears to be congruent with geography (a clade of (2006) reconsidered and placed D. boivinii back in Chloris. Our three accessions of E. macrostachyus from southern Africa results confirm that D. boivinii is not related to Chloris s.str. and a clade of E. barbatus and two accessions of E. macro­ nor Cynodon but is sister to Lepturus. stachyus from east Africa; see Electr. Suppl.: Fig. S2). Tradi­ Brachyachne, Chrysochloa, Cynodon, Eustachys, Harpo- tional characters used to differentiate among the species of chloa, Micrachne, Microchloa, Oxychloris, and Stapfochloa. Enteropogon, such as leaf sheaths margins (glabrous versus — Within the Microchloa clade considerable genetic varia­ hairy and strongly keeled versus not strongly keeled), callus tion exists among individuals of each of the three species, hair length (< 2 mm versus 3–4 mm), and lemma awn length as M. altera (Rendle) Stapf and M. caffra Nees appear in grades, (1–5 mm versus 10–18 mm) are inconsistent (Clayton, 1967). and M. kunthii Desv. forms a clade with strongly supported For instance, E. macrostachyus is characterized by having gla­ substructure. Launert (1966) revised Microchloa because brous leaf sheaths that are not strongly keeled but Oakes 1250 “specimens were often indiscriminately attributed to any of from South Africa has hairy sheaths that are strongly keeled the described taxa”. Currently, six species are placed in Micro­ like E. sechellensis (Renvoize, 1974a; Phillips, 1995) whereas chloa since Rendlia altera Rendle (= Microchloa altera) with Smook 9026 and Laegaard 15902 have glabrous sheaths. 2 or 3-flowered spikelets (spikelets 1-flowered in Microchloa We sampled four species traditionally treated in Tetra­ s.str.) was subsumed in Microchloa (Clayton, 1967; Clayton pogon and lack only T. ferrugineus (Renvoize) S.M.Phillips, & Renvoize, 1986; Cope, 1999). Harpochloa falx is sister to known from Ethiopia, Kenya, and Somalia (Phillips, 1995). We our moderately supported Microchloa clade (85/1.00), differing have moderate support (85/1.00, Fig. 2B) for these four species from the latter genus by having shorter glumes not completely previously included in Tetrapogon plus Chloris roxburghiana. enclosing the florets (Cope, 1999). Clayton (1967) suggested Tetra ­pogon includes species with 2–5 fertile florets per spikelet that since Harpochloa, Microchloa, and Rendlia share many whereas Chloris generally has a single fertile floret (Renvoize, morphological similarities, the “three genera might well be 1974b; Phillips, 1995; Cope, 1999). Chloris roxburghiana has united”. Our data clearly support this idea since these three spikelets with a single perfect floret with 2 or 3 progressively taxa all lie in a moderately supported clade (76/1.00, Fig. 2C). smaller, reduced or sterile florets above but is unusual in Chlo­ Sister to the Harpochloa-Microchloa clade is an Australian ris in possessing 30–50 or more (1–3 in Tetrapogon), spikelike endemic, Oxychloris scariosa, although this relationship is racemes, racemosely inserted along a short axis (Anderson, poorly supported. Clayton (1967), Lazarides (1972), and Ander­ 1974; Renvoize, 1974c). Sister to Tetrapogon (including C. rox­ son (1974) suggested the unique features, i.e., broadly winged burghiana ) is a moderately supported clade with Enteropogon fertile lemmas with a sharp, elongated callus and upper reduced brandegeei and E. chlorideus (71/0.99; Fig. 2B). These two sterile florets, of Chloris scariosa F.Muell. (= Oxychloris scari­ species have never been attributed to Tetrapogon but together osa) might warrant generic status (Lazarides, 1985), which our have at one time been placed in seven other genera (Chloris, results support. Dinebra, Diplachne, Eutriana, Gouinia, Gymnopogon, Lepto­ Brachyachne fulva Stapf, B. obtusifolia (Benth.) C.E.Hubb., chloa). It seems most parsimonious to recognize E. brande­ and B. patentiflora (Stent & J.M.Rattray) C.E.Hubb. form a geei, E. chlorideus, and Saugetia fasciculata (type) in a strongly strongly supported clade that we are describing as a new genus supported (100/1.00) Tetrapogon s.l. Clayton & Renvoize (1986), below since the type, B. convergens (F.Muell.) Stapf, lies within who published a designation (invalid name) for Enteropogon Cynodon. Micrachne gen. nov. differs from Cynodon in hav­ fasciculatus and, never formally, made a combination for this ing golden brown to bronze spikelets (at least in herbarium or the sister species, Saugetia pleiostachya Hitchc. & Ekman specimens) bearing a rachilla extension and glumes that com­ (not sampled here), both of which are treated in Enteropogon in pletely enclose the floret with the lower glume being slightly their online World Grass Flora (Clayton & al., 2006–) as forsan asymmetrical. Although not sampled in our study, we include (“perhaps” in Latin). Similar to Chloris roxburghiana, E. bran­ a Zambia endemic, B. simonii Kupicha & Cope and B. pilosa degeei, E. chlorideus, S. fasciculata, and S. pleiostachya have Van der Veken from the Democratic Republic of the Congo spikelets with a single fertile floret and a single sterile floret in Micrachne, and provide a complete description of all five (Hitchcock & Chase, 1917; Anderson, 1974). However, the inflo­ African species below (Van der Veken, 1958; Kupicha & Cope, rescences of E. brandegeei and E. chlorideus have 3–20 spike­ 1985). Cope (1999) in the Flora Zambesiaca included a key to like racemes, racemosely to subverticillately arranged along easily differentiate these four species (excluding B. pilosa). the peduncle, whereas S. fasciculata and S. pleiostachya have a We sampled 11 species of Eustachys and there is a strongly single raceme (Watson & Dallwitz, 1992; Clayton & al., 2006–). supported American clade (100/1.00, Fig. 2C) containing two The alternative hypothesis would be to place E. brandegeei separate subclades, each with a geographic component: (1) and E. chlorideus in a new genus and retain Saugetia. one comprising five species from North America (82/0.99) We include the monotypic, Daknopholis boivinii for the and (2) the other comprising four species from South America first time in our analysis, which is strongly supported as sister (73/0.94). The American Eustachys clade is sister to (in reverse to four species of Lepturus. This is surprising since no mor­ order of divergence) the African E. tenera (J.Presl) A.Camus phological characters suggest a link between these two genera. and the Australian E. paspaloides (Vahl) Lanza & Matte. Nash Clayton (1967) separated Daknopholis boivinii from Chloris (1898) was one of the first agrostologists to treat Eustachys at by possessing a naked rachilla extension and suggested it was the generic level rather than a subgenus of Chloris, recognizing linked to Cynodon (Clayton & Renvoize, 1986). Clayton & al. the strongly flattened, folded, equitant leaf sheaths; bilobed

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upper glumes with a short awn borne between the lobes; and appear to be undergoing rapid speciation since many morpho­ fertile florets that are commonly pale to dark brown or black logical forms do not always segregate in a shared lineage. We with awnless or short-awned lemmas (Anderson, 1974). agree with Cope (1999), who stated in the Flora Zambesiaca, The strongly supported Chrysochloa clade, consisting “The taxonomy of Cynodon is the most difficult problem of three species, is sister to the Stapfochloa-Cynodon clade among southern African Cynodonteae.” (Fig. 2C). Even though Chrysochloa annua C.E.Hubb. has been Chloris. — In the plastid tree Ochthochloa compressa is placed as a synonym of C. hindsii C.E.Hubb. by Zon (1992) and sister to the remaining species of Chloris. This species was Clayton & al. (2006), we retain these accessions as a different initially described in Panicum L. by Forsskål (1775), trans­ species since they are genetically variable, but either circum­ ferred to Eleusine by Ascherson & Schweinfurth (1922) scription is consistent with our data. before being placed in a monotypic genus by Hilu (1981). The Chloris ciliata complex as recognized by Anderson Even though O. compressa has 3–6 fertile florets per spikelet (1974) included five species that have elliptic, ovate or lanceo­ (versus 1 or 2 fertile florets in Chloris) and caryopses with late fertile lemmas with densely ciliate margins (usually along free pericarps (occasionally found in Chloris) we place this the entire length). As pointed out earlier by Anderson (1974), enigmatic taxon in Chloris and make the new combination the African Chloris lamproparia (= Stapfochloa lamproparia) below. Likewise, Lintonia nutans has never been associated is the only Old World member of the Chloris ciliata complex. with Chloris but it is clearly a member in our ITS and plastid The observation of Scholz & Müller (2004) that “their [species tree (Fig. 3). Lintonia nutans has 2–5 fertile florets per spikelet included in the Chloris ciliata complex] striking resemblance with 5–11-veined (confluent at the base), cartilaginous lem­ with C. lamproparia in lemma form and indument is surely a mas (versus 3-veined and membranous to cartilaginous) with result of parallel evolution and does not reflect close genetic clavicorniculate hairs and dorsally flattened caryopses with relationship” is not supported by our tree (Fig. 2C). Anderson free pericarps (versus usually subterete or trigonous caryopses; (1974) mentioned that the Chloris ciliata complex “presents Watson & Dallwitz, 1992). Clavicorniculate lemmatal hairs more interspecific taxonomic problems that any other group”, have arisen twice in the Eleusininae since they appear in Lin­ which we confirm based on high genetic variability among tonia and Disakisperma. individuals of C. ciliata and C. elata. Stapfochloa has priority, The major phylogenetic problem in Chloris, however, is the and we make the necessary new combinations for the American presence of the C-D clade in the ITS tree that encompasses spe­ species and emend the generic description below. cies representing two distant plastid lineages. The most prob­ In our tree three of the four species of Brachyachne (B. cili­ able scenario underlying the formation of the C-D clade in the aris (Kuntze) C.E.Hubb., B. convergens, B. tenella (R.Br.) ITS tree could be ancient hybridization between the lineages C.E.Hubb.) that are embedded in Cynodon are endemic to in the early stages of speciation with subsequent alternation Australia, whereas B. ambigua Ohwi is native to Java, Lesser and homogenization of the ITS signal. Sunda Islands, and New Guinea (Nightingale & al., 2005a). The subgeneric classification of Chloris by Varadarajan Traditionally, Brachyachne was separated from Cynodon by & Gilmartin (1983a, b) based on phenetic and cladistic anal­ having glumes that are longer than the floret (versus at least yses of morphological features does not coincide with our one glume shorter that the floret) and lemmas that are long work. Their Chloris subg. Phacelaria (Steud.) Varadarajan hairy on the veins or all over (versus hairy or not only along & Gilmartin included species we place in Leptochloa, Micr­ the midvein; Nightingale & al., 2005a). Clearly this poorly sup­ achne, and Stapfochloa (Peterson & al., 2012) and their Chlo­ ported clade (k) arose within Cynodon, the most derived clade ris subg. Chloris includes species we place in Tetrapogon in the Eleusininae. Few morphological differences separate the (Peterson & al., 2012). Within Chloris we found four separate species in Cynodon s.l., and several species do not form mono­ strongly supported clades in the ITS tree (A–D, Fig. 3), which phyletic clades. For instance, C. aethiopicus Clayton & Harlan correspond to four clades in the plastid tree, two of which are (Smook 4140) from South Africa is placed in an unresolved strongly supported (A and C), one is moderately supported (D), clade (clearly differentiated from a clade with three accessions and one is poorly supported (B, PP =0.69). of C. aethiopicus) with five other accessions of C. nlemfuensis Clades A and C contain species in the Chloris inflata Link Vanderyst from South Africa, United States, Costa Rica, and complex (= C. barbata, see Kartesz & Gandhi, 1992), where two from Ecuador. Morphological features used to differenti­ most species have single-celled microhairs, Chloris-type silica ate C. aethiopicus from C. nlemfuensis are: lemma keel densely cells (cuboidal to saddle-shaped), and fertile lemmas with a tuft pubescent (former) versus glabrous or only with a few hairs of hairs on the upper margins (Anderson, 1974). Chloris prieu­ (latter); plants robust, often woody and coarse, 60–100 cm tall rii, which is sister to C. barbata in clade A (ITS and plastid versus plants fairly slender to robust, not woody, 30–60 cm tall tree), and C. orthonoton, which is included in clade C (plastid (Gibbs Russell & al., 1991). Likewise, many species have been tree only), lacks a tuft of hairs on the upper margins of the placed as synonyms or as varieties of C. dactylon (L.) Pers., fertile lemma but instead has a row of clavate glandular hairs such as C. aristiglumis­ Caro & E.A.Sánchez, C. erectus J.Presl, along the sides of the lemma (Anderson, 1974). Chloris rufe­ and C. pascuus Nees (Salariato & Zanotti, 2012), and these scens (attributed by Anderson to the C. verticillata complex) appear in a large clade at the base of the tree with C. trans­ also occurs in the plastid tree only and it has been indicated vaalensis Burtt Davy, a morphologically distinct species with as a close relative of C. orthonoton but differs in having short- strong support (97/1.00; Fig. 2C). The species of Cynodon s.l. ciliate less than 1 mm long ligules (versus long ciliate, about

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2 mm long), shorter hairs along the lemma margins (versus Chloris nutans (Stapf) P.M.Peterson, comb. nov. ≡ Linto­ hairs > 1.5 mm), and lemmas without clavate glandular hairs nia nutans Stapf in Hooker’s Icon. Pl. 30: t. 2949. 1911 (Anderson, 1974). – Holotype: KENYA. Nairobi, A. Linton 193 (K barcode Clade B (Fig. 3) contains Chloris amethystea, C. andro­ K000365107 [image!]). pogonoides, C. cucullata, C. latisquamea, C. × subdolicho­ stachya, C. verticillata, and C. mossambicensis, although two Cynodon ambiguus (Ohwi) P.M.Peterson, comb. nov. ≡ individuals of the latter species are sister to Lintonia nutans in the Brachyachne ambigua Ohwi in Bull. Tokyo Sci. Mus. 18: ITS tree, suggesting a past hybridization event. Anderson (1974, 11. 1947 – Holotype: JAVA. Gersik, E Java, on low rocky 1975) referred to most members of this clade as the C. verticil­ hill, Dec 1924, J.D. Dorzelo 3169 (BO; isotype: L). lata complex and suggested that C. andropogonoides, C. cucul­ lata, C. verticillata, and C. × subdolichostachya, form a hybrid­ Cynodon prostratus (C.A.Gardner & C.E.Hubb.) P.M.Peterson, ization complex that “seem to blend and recombine in various comb. nov. ≡ Brachyachne prostrata C.A.Gardner & ​C.E.​ ways”. Anderson (1974) found that the lemmatal awn length and Hubb. in Hooker’s Icon. Pl. 34: t. 3362. 1938 – Holotype: the sterile lemma width are best to differentiate among the three AUSTRALIA. Western Australia, Eremean Prov., 1 mi N of former (see preceding sentence) species while the latter taxon Meekatharra, 2 Aug 1927, C.A. Gardner s.n. (PERTH; iso­ is perhaps a hybrid of C. cucullata crossed with C. verticillata types: K barcodes K000885870 & K000885871 [images!]). or C. andropogonoides (Brown, 1969). Anderson (1974) included Chloris cubensis, C. ekmanii, Cynodon simonii P.M.Peterson, nom. nov. ≡ Cynodon ciliaris C. halophila, and C. sagraeana in his C. radiata (L.) Sw. com­ Benth., Fl. Austral. 7: 610. 1878, non (L.) Raspail in Ann. plex, all of which appear in clade D in both the ITS and plastid Sci. Nat., Bot. 5: 302. 1825 ≡ Capriola ciliaris Kuntze, trees (Fig. 3). These species have comparatively narrow spike­ Revis. Gen. Pl. 2: 764. 1891 ≡ Microchloa ciliaris (Kuntze) lets, reduced sterile florets, and relatively long-awned lemmas Domin in Biblioth. Bot. 85: 364. 1915 ≡ Brachyachne cili­ (Anderson, 1974). In addition to these core species our trees aris (Kuntze) C.E.Hubb. in Bull. Misc. Inform. Kew 1934: include C. cruciata (mentioned by Anderson as having similar 448. 1934 – Holotype: AUSTRALIA. Northern Territory, overall spikelet structure to the C. radiata complex), C. divari­ central Australia, Charlotte Waters, W.E.P. Giles s.n. (K cata, C. pectinata, C. pycnothrix, C. submutica, and C. texana. barcode K000885872 [image!]; isotype: MEL). Chloris texana is a possible hybrid (listed as a synonym of Etymology. – Named in honor of Bryan Kenneth Simon what we now call Stapfochloa ciliata ≡ C. ciliata; Anderson, (1943–2014), an outstanding Australian agrostologist. 1974; Soreng & al., 2014). In the ITS tree, C. texana is found in a weakly supported clade with C. halophila, C. orthonoton, Diplachne divaricatissima (S.T.Blake) P.M.Peterson, comb. and C. rufescens and then is an unresolved member in a poorly nov. ≡ Leptochloa divaricatissima S.T.Blake in Contr. supported C. pycnothrix clade in the plastid tree, possibly indi­ Queensland Herb. 14: 8, fig. 5. 1972 ≡ Dinebra divaricatis­ cating hybridization. sima (S.T.Blake) P.M.Peterson & N.Snow in Phytoneuron 2012-71: 1. 2012 – Holotype: AUSTRALIA. Queensland, Goondiwindi, S.T. Blake 10517 (BRI No. 131665-6; iso­ TAXONOMY types: CANB, K barcodes K000899894 & K000899895 [images!], MO, NSW, PRE). Because our molecular analysis renders Chloris s.l. and Brachyachne polyphyletic, we describe a new genus, Micr­ Leptochloa anisopoda (Scribn. ex B.L.Rob.) P.M.Peterson, achne, found in a separate, strongly supported clade with five comb. nov. ≡ Chloris anisopoda Scribn. ex B.L.Rob. in species and emend Stapfochloa to include six species. In addi­ Proc. Amer. Acad. Arts 38: 118. 1902 – Lectotype (desig­ tion, we provide new combinations in Chloris (2), Cynodon nated by Hitchcock in N. Amer. Fl. 17: 596. 1939): ECUA­ (2), Diplachne (1), Leptochloa (2), Micrachne (5), Stapfochloa DOR. Galapagos Islands, Charles (Santa Maria) Island, (5), and Tetrapogon (5); provide a new name in Cynodon; and Jun 1891, G. Baur 333 (GH barcode GH00023358 [image!]; lectotypify Eleusine flagellifera. isolectotype: US No. 2830885! fragm.) = Gymnopogon mollis Nees in Martius & al., Fl. Bras. Enum. Pl. Chloris flagellifera (Nees) P.M.Peterson, comb. nov. ≡ 2(1): 427. 1829 ≡ Chloris mollis (Nees) Swallen in N. Amer. Eleusine flagellifera Nees in Linnaea 16(2): 220. 1842 – Fl. 17: 596. 1939 ≡ Enteropogon mollis (Nees) Clayton in Lectotype (designated here): SAUDI ARABIA. In pla­ Kew Bull. 37: 419. 1982, non Leptochloa mollis Kunth, nitie deserti prope Dscheddam, 10 Jan 1836, W. Schimper Révis. Gramin. 2: 443, t. 135. 1831. 800 (US No. 3298791!; isolectotypes: REG barcode 000227 = Gymnopogon rupestris Ridl. in J. Linn. Soc., Bot. 27: 73. 1890. [image!], US Nos. 88428! & 1535370!). ≡ Chloris rupestris (Ridl.) Hitchc., Man. Grasses W. Ind.: = Panicum compressum Forssk., Fl. Aegypt.-Arab.: 18. 1775 ≡ 126. 1936, non Leptochloa rupestris C.E.Hubb. in Bull. Eleusine compressa (Forssk.) Asch. & Schweinf. ex C.Chr. Misc. Inform. Kew 1941(2): 195–196. 1941. in Dansk Bot. Ark. 4(3): 12. 1922 ≡ Ochthochloa compressa (Forssk.) Hilu in Kew Bull. 36(3): 560. 1981, non Chloris Leptochloa exilis (Renvoize) P.M.Peterson, comb. nov. ≡ compressa DC., Cat. Pl. Horti Monsp.: 94. 1813. Chloris exilis Renvoize in Kew Bull. 37(2): 323, fig. 2.

458 Version of Record TAXON 64 (3) • June 2015: 445–467 Peterson & al. • Phylogeny and classification of the Eleusininae

1982 – Holotype: BRAZIL. 49 km N of Senhor do Bonfim Brachyachne patentiflora (Stent & J.M.Rattray) C.E.Hubb. on BA Hwy 130 to Juazeiro, W 40°13′ S 10°05′, 500 m, 26 in Bull. Misc. Inform. Kew 1933: 503. 1933 – Holotype: Feb 1974, R.M. Harley 16363, Renvoize, Erskine, Brighton ZIMBABWE. Harare, Dec 1919, F. Eyles 1955 (BM; iso­ & Pinheiro (CEPEC; isotypes: K barcode K000003260 types: EA barcode EA000000464 [image!], K barcodes [image!], US No. 2955111!). K000365039, K000365040 & K000365041 [images!], PRE barcode PRE0664092-0 [image!], SRGH barcode Micrachne P.M.Peterson, Romasch. & Y.Herrera, gen. nov. SRGH0106276 [image!]). – Type: Micrachne fulva (Stapf) P.M.Peterson (≡ Brachy­ achne fulva Stapf). Micrachne pilosa (Van der Veken) P.M.Peterson, comb. nov. Diagnosis. – Micrachne differs from Cynodon sect. ≡ Microchloa pilosa Van der Veken in Bull. Jard. Bot. Brachyachne Benth. in having golden brown to bronze spikelets État. Bruxelles 24(1): 84. 1958 – Holotype: DEMOCRATIC (at least in herbarium specimens) bearing a rachilla extension, REPUBLIC OF THE CONGO. District de Haut-Katanga, glumes that completely enclose the floret, and the lower glume Parc National de l’Upemba, region de Kanongo, petit ruis­ being slightly asymmetrical. sellement tributaire de la riv. Lukanga, sur larges dalles Description. – Plants perennial, occasionally annual, caes­ de laterite, Feb 1949, Van Meel in de G.F. Witte 5458 (BR pitose to geniculately ascending to erect. Culms 10–60(–70) cm barcode BR0000008756367 [image!]; isotype K barcode tall. Leaf sheaths glabrous above, occasionally densely pilose K000365034 [image!]). below, often forming a cushion of fibers near base; ligule a short ciliate membrane; blades 0.5–15(–25) cm long, 0.5–3 mm Micrachne simonii (Kupicha & Cope) P.M.Peterson, comb. wide, flat or involute, glabrous or sparsely hairy, strait or nov. ≡ Brachyachne simonii Kupicha & Cope in Kew recurved, occasionally setaceous and rigid. Inflorescences Bull. 40(1): 89. 1985 – Holotype: ZAMBIA. 3 km E of paniculate usually with a single terminal spikelike branch or Chizela, 27 Dec 1969, B.K. Simon & Williamson 2068 (K sometimes with (2–)3–5(–7) digitately inserted branches; the barcode K000365038 [image!]; isotypes: BM, BR barcode 1-sided spikelike branches 1.5–12 cm long, erect or somewhat BR0000005628681 [image!], MO No. 2154714-1660917 curved and bearing imbricate spikelets. Spikelets 1-flowered, [image!], PRE barcodes PRE0592099-0 & PRE0664091-0 laterally compressed, golden brown to bronze (at least in her­ [images!], SRGH). barium specimens), the single fertile floret bearing a rachilla extension; glumes 2–5.3 mm long, longer than the floret, Stapfochloa H.Scholz – Type: Stapfochloa lamproparia (Stapf) 1-veined, chartaceous to thinly coriaceous, glabrous or sparsely H.Scholz (≡ Chloris lamproparia Stapf). hairy, completely enclosing the floret, the lower glume slightly Diagnosis. – Stapfochloa differs from Chloris in having asymmetrical, apex obtuse to acute; lemmas 1.6–4 mm long, elliptic, ovate or lanceolate fertile lemmas with lemma margins 3-veined, hyaline to membranous, delicate, oblong-elliptic, that are densely ciliate, usually along the entire length. ovate to oblong-lanceolate, folded along the midvein, hairy Description. – Plants perennial, occasionally annual, along the midvein and marginal veins, apex obtuse to truncate caespitose, rhizomatous or stoloniferous. Culms 5–80 cm tall. or slightly emarginated, often short mucronate; paleas a little Leaf sheaths glabrous; ligule a short ciliate crown or lacking; shorter than the lemmas, hairy or glabrous along the veins; blades 1–28 cm long, 1–8 mm wide, flat or folded, glabrous stamens 3, anthers 0.8–2.3 mm long. Caryopses ellipsoid. to pilose. Inflorescences paniculate with 1–12(–28) terminal Distribution and habitat. – The five species are distrib­ spikelike branches, usually digitately or subdigitately inserted; uted in central and eastern Africa, often on rock outcrops with the spikelike branches 3–20 cm long, erect to widely spreading thin soils, sandy washes, laterite pans to open woodlands; and bearing spikelets in two rows on one side of the branch. 700–2130 m. Spikelets with 1 basal fertile floret with 2 or 3 reduced, sterile florets above; fertile lemmas 1.3–4.2 mm long, 0.5–1.7 mm Micrachne fulva (Stapf) P.M.Peterson, comb. nov. ≡ Brachy- wide, elliptic, ovate or lanceolate, with a single apical or subapi­ ­achne fulva Stapf in Hooker’s Icon. Pl. 31: t. 3099. cal awn 1.4–6 mm long, the keel and margins with spreading 1922 – Holotype: ZAIRE. Katanga, Homble 21 (K bar­ ciliate hairs usually along the entire length, the hairs 0.5–3 mm code K000365033 [image!]; isotypes: BR barcodes long. Caryopses 1–2 mm wide, 0.5–1 mm wide, ovoid to ellip­ BR0000008756374 & BR0000008756411 [images!]). soid to trigonous. Distribution and habitat. – Five species are distributed in Micrachne obtusiflora (Benth.) P.M.Peterson, comb. nov. the New World and a single species, S. lamproparia, is native ≡ Microchloa obtusiflora Benth., Niger Fl.: 565. 1849 ≡ to Africa. Plants occur in open grasslands to wet savannahs, Brachyachne obtusiflora (Benth.) C.E.Hubb. in Bull. Misc. often on rocky slopes. Inform. Kew 1933: 503. 1933 – Holotype: NIGERIA. Vogel 189 (K barcode K000365032 [image!]). Stapfochloa berroi (Arechav.) P.M.Peterson, comb. nov. ≡ Chloris berroi Arechav. in Anales Mus. Nac. Montevideo Micrachne patentiflora (Stent & J.M.Rattray) P.M.Peterson, 1: 388, t. 44. 1896 – Holotype: URUGUAY. Dept. Soriano, comb. nov. ≡ Microchloa patentiflora Stent & J.M.Rattray Estancia Vera, 5 Jan 1895, M.B. Berro 962 (MVFA barcode in Proc. & Trans. Rhodesia Sci. Assoc. 32: 59. 1933 ≡ MVFA0000272 [image!]; isotypes: BAA No. 655-00001714

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[image!], CORD No. 879335, K barcode K000308975 Pl. 1: 280. 1833 ≡ Chloris chloridea (J.Presl) Hitchc. in [image!], W No. 19160034138 [image!]). Proc. Biol. Soc. Wash. 41: 162. 1928 ≡ Enteropogon chlo­ rideus (J.Presl) Clayton in Kew Bull. 37: 419. 1982 – Holo­ Stapfochloa canterae (Arechav.) P.M.Peterson, comb. nov. ≡ type: MEXICO. Haenke s.n. (PR; isotypes: MO, US No. Chloris canterae Arechav. in Anales Mus. Nac. Monte­ 865870!). video 1: 385. 1896 – Lectotype (designated by Rosengurtt & Izaguirre in Bol. Soc. Argent. Bot. 12: 124. 1968): URU­ Tetrapogon fasciculatus (Hitchc. & Chase) P.M.Peterson, GUAY. Paysandú, J. Arechavaleta s.n. (MVM; isolecto­ comb. nov. ≡ Saugetia fasciculata Hitchc. & Chase types: BAA barcode BAA00001717 [image!], CORD bar­ in Contr. U.S. Natl. Herb. 18(7): 378. 1917 – Holotype: code CORD00001558 [image!], US No. 879336! fragm. CUBA. Pinar del Río, in small wood savanna San Julián, ex MVM). S of Guane, 28 Dec 1916, Leon 6901 & M. Roca (US No. 950204!; isotypes: BAA barcode BAA00002812 [image!], Stapfochloa ciliata (Sw.) P.M.Peterson, comb. nov. ≡ Chlo­ NY barcode 00071123 [image!]). ris ciliata Sw., Prodr.: 25. 1788 ≡ Cynodon ciliatus (Sw.) Raspail in Ann. Sci. Nat. (Paris) 5: 303. 1825 – Holotype: Tetrapogon pleiostachyus (Hitchc. & Ekman) P.M.Peterson, JAMAICA. O.P. Swartz s.n. (S No. 06-646 [image!]; iso­ comb. nov. ≡ Saugetia pleiostachya Hitchc. & Ekman in types: BAA No. 661-00001719 [image!], US No. 80821! Hitchcock, Man. Grasses W. Ind.: 123. 1936 – Holotype: fragm. ex S). CUBA. Prov. Pinar del Río, Mendoza Municipio, Cerro de Mendoza, 26 Jun 1920, E.L. Ekman 11483 (US No. 1296127!; Stapfochloa elata (Desv.) P.M.Peterson, comb. nov. ≡ Chloris isotype: NY barcode 00071124). elata Desv. in Mém. Soc. Agric. Angers 1: 177. 1831 – Holo­ type: BRAZIL. s.coll., s.n. (P). Tetrapogon roxburghiana (Schult.) P.M.Peterson, comb. nov. = Chloris dandyana C.D.Adams in Phytologia 21: 408. 1971. ≡ Chloris roxburghiana Schult., Mant. 2: 339. 1824 ≡ Chlo­ ris polystachya Roxb., Fl. Ind. 1: 332. 1820, nom. illeg., Stapfochloa lamproparia (Stapf) H.Scholz in Willdenowia non Lag., Gen. Sp. Pl. 4. 1816 – Holotype: INDIA. Based 34: 131, fig. 1–2. 2004 ≡ Chloris lamproparia Stapf in on a fairly complete description and Roxburgh painting Mém. Soc. Bot. France 2, mém. 8: 220. 1912 – Holotype: (K No. 2023). CHAD. Baguirmi, emplacement de la ville de Massenia, Aug 1903, A.J.B. Chevalier 9633 bis (P barcode P00439460 [image!]; isotype: K barcode K000365096 [image!]). ACKNOWLEDGEMENTS

Stapfochloa parvispicula (Caro & E.A.Sánchez) P.M.Peterson, We thank the National Geographic Society Committee for comb. nov. ≡ Chloris parvispicula Caro & E.A.Sánchez in Research and Exploration (Grant No. 8848-10, 8087-06) for field and Kurtziana 6: 227, fig. 3. 1971 – Holotype: ARGENTINA. La laboratory support; the Smithsonian Institution’s Restricted Endow­ Rioja, Los Llanos, Los Cerrillos, 15 Feb 1940, A. Castel­ ments Fund, the Scholarly Studies Program, Research Opportunities, lanos s.n. (BAA No. 33416-00000082 [image!]; isotype: Atherton Seidell Foundation, Biodiversity Surveys and Inventories LIL No. 42941-000028 [image!]). Program, Small Grants Program, and the Laboratory of Analytical Biology; and the United States Department of Agriculture, all for Tetrapogon brandegeei (Vasey) P.M.Peterson, comb. nov. ≡ financial support. We would also like to acknowledge Gabriel Johnson Diplachne brandegeei Vasey in Proc. Calif. Acad. Sci., ser. for help in the laboratory; Robert J. Soreng and Jeffery M. Saarela for 2: 213. 1889 (“brandegei”) ≡ Gouinia brandegeei (Vasey) accompanying the first author on numerous field expeditions; Maria Hitchc. in Bull. Bur. Pl. Industr. U.S.D.A. 33: 21. 1903 Vorontsova for finding type images at K; Robert J. Soreng for many (“brandegei”) ≡ Leptochloa brandegeei (Vasey) Hitchc. in extended discussions pertinent to the manuscript; Jeffery M. Saarela Bull. Bur. Pl. Industr. U.S.D.A. 33: 21. 1903 (“brandegei”) and Neil Snow for providing helpful comments on the manuscript; and ≡ Chloris brandegeei (Vasey) Swallen in Amer. J. Bot. 22: Franz Stadler for critical review during production. 41. 1935 (“brandegei”) ≡ Enteropogon brandegeei (Vasey) Clayton in Kew Bull. 37: 419. 1982 (“brandegei”) – Lecto­ type (designated by Hitchcock in Contr. U.S. Natl. Herb. LITERATURE CITED 17: 352. 1913): MEXICO. Baja California Sur, Magdalena Island, 18 Jan 1889, T.S. Brandegee 11 (US No. 78812!). Anderson, D.E. 1974. Taxonomy of the genus Chloris (Gramineae). Sci. Etymology. – Originally named by Vasey to honor Bull. Brigham Young Univ., Biol. Ser. 19(2): 1–133. Townshend Stith Brandegee (1843–1925) and is therefore to Anderson, D.E. 1975. 73. Chloris Swartz. Pp. 316–333 in: Gould, F.W. be spelled “brandegeei” as per Art. 60.12 and Rec. 60C.1(a). (ed.), The grasses of Texas. College Station: Texas A&M Univer­ sity Press. Ascherson, P.F.A. & Schweinfurth, G.A. 1922. Index to Forsskål: Tetrapogon chlorideus (J.Presl) P.M.Peterson, comb. nov. ≡ Flora aegyptiaco-arabica 1775 with a revision of Herbarium Dinebra chloridea J.Presl in C. Presl, Reliq. Haenk. 1(4–5): Forskålii contained in the Botanical Museum of the University of 291. 1830 ≡ Eutriana chloridea (J.Presl) Kunth, Enum. Copenhagen. Dansk Bot. Ark. 4(3): 12. 1922

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A Gibbs Russell, G.E., Watson, L., Koekemoer, M., Smook, L., Barker, molecular phylogeny and classification of Leptochloa (Poaceae: N.P., Anderson, H.M. & Dallwitz, M.J. 1991. Grasses of southern Chloridoideae: Chlorideae) sensu lato and related genera. Ann. Africa. Mem. Bot. Surv. S. Africa 58: 1–437. Bot. (Oxford) 109: 1317–1329. GPWG (Grass Phylogeny Working Group) II 2011. New grass phy­ http://dx.doi.org/10.1093/aob/mcs077 logeny resolves deep evolutionary relationships and discovers C4 Peterson, P.M., Romaschenko, K. & Herrera Arrieta, Y. 2014a. A origins. New Phytol. 193: 304–312. molecular phylogeny and classification of the Cteniinae, Farrag­ http://dx.doi.org/10.1111/j.1469-8137.2011.03972.x ininae, Gouiniinae, Gymnopogoninae, Perotidinae, and Tricho­ Hilu, K.W. 1981. Taxonomic status of the disputable Eleusine compressa neurinae (Poaceae: Chloridoideae: Cynodonteae). Taxon 63: (Gramineae). Kew Bull. 36: 559–563. 275–286. http://dx.doi.org/10.12705/632.35 http://dx.doi.org/10.2307/4117586 Peterson, P.M., Romaschenko, K., Herrera Arrieta, Y. & Saarela, Hilu, K.W. & Alice, L.A. 2001. A phylogeny of Chloridoideae (Po­aceae) J.M. 2014b. A molecular phylogeny and new subgeneric classifica­ based on matK sequences. Syst. Bot. 26: 386–405. tion of Sporobolus (Poaceae: Chloridoideae: Sporobolinae). Taxon Hitchcock, A.S. & Chase, A. 1917. Grasses of the West Indies. Contr. 63: 1212–1243. http://dx.doi.org/10.12705/636.19 U.S. Natl. Herb. 18(7): 261–494. Phillips, S.M. 1972. A survey of the genus Eleusine Gaertn. (Gra­ http://dx.doi.org/10.5962/bhl.title.53799 mineae) in Africa. Kew Bull. 27: 251–270. Huelsenbeck, J.P. & Ronquist, F.R. 2001. MRBAYES: Bayesian infer­ http://dx.doi.org/10.2307/4109450 ence of phylogenetic trees. Bioinformatics 17: 754–755. Phillips, S.M. 1995. Poaceae (Gramineae). Pp. 1–420 in: Hedberg, I. http://dx.doi.org/10.1093/bioinformatics/17.8.754 & Edwards, S. (eds.), Flora of Ethiopia and Eritrea, vol. 7. Addis Jozwik, F.X. 1969. Some systematics aspects of Mitchell grasses Ababa: Addis Ababa University; Uppsala: Uppsala University. (Astrebla F. Muell.). Austral. J. Bot. 17: 359–374. Posada, D. 2008. jModelTest model averaging. Molec. Biol. Evol. 25: http://dx.doi.org/10.1071/BT9690359 1253–1256. http://dx.doi:10.1093/molbev/msn083 Kartesz, J.T. & Gandhi, K.N. 1992. Chloris barbata Sw. and C. elata Renvoize, S.A. 1974a. Enteropogon. Pp. 331–335 in: Polhill, R.M. (ed.), Desvaux (Poaceae), the earlier names from C. inflata Link and Flora of tropical East Africa, Gramineae, part 2. London: Crown C. dandeyana Adams. Rhodora 94: 135–140. Agents for Oversea Governments and Administrations.

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Renvoize, S.A. 1974b. Tetrapogon. Pp. 347–351 in: Polhill, R.M. (ed.), Snow, N., Peterson & Romaschenko, K. 2013. Systematics of Disaki­ Flora of tropical East Africa, Gramineae, part 2. London: Crown sperma (Poaceae, Chloridoideae, Chlorideae). PhytoKeys 26: Agents for Oversea Governments and Administrations. 21–70. http://dx.doi.org/10.3897/phytokeys.26.5649 Renvoize, S.A. 1974c. Chloris. Pp. 337–347 in: Polhill, R.M. (ed.), Flora Soreng, R.J., Davidse, G., Peterson, P.M., Zuloaga, F.O., Judziewicz, of Tropical East Africa, Gramineae, part 2. London: Crown Agents E.J., Filgueiras, T.S., Morrone, O. & Romaschenko, K. 2014. A for Oversea Governments and Administrations. World -wide phylogenetic classification of Poaceae (Gramineae): Renvoize, S.A. 1984. The grasses of Bahia. Richmond: Royal Botanic căo (草), capim, çayır, çimen, darbha, ghaas, ghas, gish, gramas, Gardens, Kew. graminius, gräser, grasses, gyokh, he-ben-ke, hullu, kasa, kusa, Romaschenko, K., Garcia-Jacas, N., Peterson, P.M., Soreng, R.J., nyasi, pastos, pillu, pullu, zlaki, etc. http://www.tropicos.org/pro­ Vilatersana, R. & Susanna, A. 2014. Miocene–Pliocene specia­ jectwebportal.aspx?pagename=ClassificationNWG&projectid=10 tion, introgression, and migration of Patis and Ptilagrostis (Poa­ (accessed Oct 2014). ceae: Stipeae). Molec. Phylogen. Evol. 70: 244–259. Soreng, R.J., Peterson, P.M., Romaschenko, K., Davidse, G., http://dx.doi.org/10.1016/j.ympev.2013.09.018 Zuloaga, F.O., Judziewicz, E.J., Filgueiras, T.S., Davis, J.I. & Ronquist, F. & Huelsenbeck, J.P. 2003. Mr Bayes 3: Bayesian phyloge­ Morrone O. 2015. A worldwide phylogenetic classification of the netic inference under mixed models. Bioinformatics 19: 1572–1574. Poaceae (Gramineae). J. Syst. Evol. 53: 117–137. http://dx.doi.org/10.1093/bioinformatics/btg180 http://dx.doi.org/10.1111/jse.12150 Roodt, R. & Spies, J.J. 2002. Chromosome studies on African plants. Tateoka, T. 1957. Notes on some grasses. V. Bot. Mag., Tokyo 70: 115–17. 18. The subfamily Chloridoideae. Bothalia 32(2): 240–249. Tateoka, T. 1962. Starch grains of endosperm in grass systematics. Bot. Roodt, R. & Spies, J.J. 2003a. Chromosome studies in the grass sub­ Mag., Tokyo 75: 377–383. family Chloridoideae. I. Basic chromosome numbers. Taxon 52: Van der Veken, P. 1958. Graminées nouvelles ou peu connues de 557–566. http://dx.doi.org/10.2307/3647454 l’Afrique Centrale II. Bull. Jard. Bot. État. Bruxelles 24(1): 84–91. Roodt, R. & Spies, J.J. 2003b. Chromosome studies in the grass sub­ 1958. family Chloridoideae. II. An analysis of polyploidy. Taxon 52: Varadarajan, G.S. & Gilmartin, A.J. 1983a. Phenetic and cladistic 736–746. http://dx.doi.org/10.2307/3647348 analyses of North American Chloris (Poaceae). Taxon 32: 380–386. Salariato, D.L. & Zanotti, C.A. 2012. Cynodon Rich., nom. cons. Pp. http://dx.doi.org/10.2307/1221494 100–106 in: Zuloaga, F.O., Rúgolo, Z.E. & Anton, A.M. (eds.), Varadarajan, G.S. & Gilmartin, A.J. 1983b. Subgeneric classification Flora vascular de la Republica Argentina, vol. 3, tomo 1. Cordoba: of Chloris Sw. (Poaceae). Taxon 32: 471–473. Graficamente Ediciones. http://dx.doi.org/10.2307/1221518 Scholz, H. & Müller, J. 2004. Stapfochloa (Poaceae: Cynodonteae: Watson, L. & Dallwitz, M.J. 1992. The grass genera of the World. Chloridinae), a new genus from Africa. Willdenowia 34: 129–33. Wallingford, U.K.: CAB International. http://dx.doi.org/10.3372/wi.34.34111 Zon, A.P.M. 1992. Graminées du Cameroun. Wageningen Agric. Univ. Snow, N. & Peterson, P.M. 2012a. Systematics of Trigonochloa Pap. 92-1(2): 1–557. (Po ­aceae, Chloridoideae, Chlorideae). PhytoKeys 13: 25–38. Zwickl, D.J. 2006. Genetic algorithm approaches for the phylogenetic http://dx.doi.org/10.3897/phytokeys.13.3355 analysis of large biological sequence datasets under the maxi­ Snow, N. & Peterson, P.M. 2012b. Nomenclatural notes on Dinebra, mum likelihood criterion. Ph.D. dissertation, University of Texas Diplachne, Disakisperma and Leptochloa (Poaceae: Chlorid­ at Austin, U.S.A. oideae). Phytoneuron 2012-71: 1–2.

Appendix 1. List of specimens sampled. Taxon voucher (collector, number, and where the specimen is housed), country of origin, and GenBank accession for DNA sequences of rpl32-trnL, ndhA, rps16, rps16-trnK and ITS regions (bold indicates new accession); a dash (–) indicates missing data; an asterisk (*) indicates sequences not generated in our lab. OUTGROUP: Aristida gypsophila Beetle, Peterson 15839 & Valdes-Reyna (US), Mexico, GU359977, GU359386, GU360286, GU360570, GU359267; Chas­ manthium latifolium (Michx.) H.O.Yates, Peterson 22463 (US), U.S.A., GU359891, GU359379, GU360438, GU360517, GU359319; Danthonia compressa Austin, Peterson 21986 & Levine (US), U.S.A., GU359865, GU359370, GU360483, GU360521, GU359345; Capeochloa cincta subsp. sericea (N.P.Barker) N.P.Barker & H.P.Linder, Barker 1545 (GRA), South Africa, JF729173, JF729163, JF729181, JF729074, –; Rytidosperma pallidum (R.Br.) A.M.Humphreys & H.P.Linder, Peterson 19685, Saarela & Sears (US), U.S.A., GU359984, GU359518, GU360291, GU360671, GU359183; CENTROPODIEAE: Ellisochloa rangei (Pilg.) P.M.Peterson & N.P.Barker, Barker 960 (BOL), Namibia, –, JF729166, JF729184, JF729079, JQ345167; TRIRAPHIDEAE: Neyraudia reynau­ diana (Kunth) Keng ex Hitchcock, Soreng 5318, Peterson & Sun Hang (US), China, GU360003, GU359397, GU360272, –, GU359124; Triraphis ramosissima Hack., Seydel 4278 (US), South Africa, GU359931, GU359541, GU360338, GU360651, GU359188; ERAGROSTIDEAE: Cottea pappophoroides Kunth, Peterson 21463, Soreng, LaTorre & Rojas Fox (US), Peru, GU359842, GU359363, GU360456, GU360600, GU359237; Ectrosia scabrida C.E.Hubb., Lazarides 4772 (US), Australia, GU359799, GU359476, GU360459, GU360497, GU359317; Enneapogon desvauxii P.Beauv., Peterson 21999 & Saarela (US), Mexico, GU359796, GU359474, GU360486, GU360495, GU359339; Entoplocamia aristulata (Hack. & Rendle) Stapf, Seydel 187 (US), South Africa, GU359793, GU359469, GU360468, GU360492, GU359342; Eragrostis desertorum Domin, Peterson 14358, Soreng & Rosenberg (US), Australia, GU359787, GU359471, GU360462, GU360545, GU359289; Eragrostis nigricans (Kunth) Steud., Peterson 21623, Soreng, LaTorre & Rojas Fox (US), Peru, GU359790, GU359491, GU360398, GU360520, GU359299; Harpachne harpachnoides (Hack.) B.S.Sun & S.Wang, Soreng 5288, Peterson & Sun Hang (US), China, GU359815, GU359435, GU360382, GU360611, GU359113; Psammagrostis wiseana C.A.Gardner & C.E.Hubb., Peterson 14345, Soreng & Rosenberg (US), Australia, GU359986, GU359533, GU360288, GU360703, GU359137; Tetrachne dregei Nees, Jarman 120 (US), South Africa, GU359904, GU359513, GU360365, GU360622, GU359218; Uniola condensata Hitchc., Peterson 9342 & Judziewicz (US), Ecuador, GU359927, GU359534, GU360340, GU360649, GU359191; Calamovilfa longifolia (Hook.) Hack. ex Scribn. & Southw., Hatch 5738 & Bearden (US), U.S.A., GU359880, GU359357, GU360441, GU360548, GU359300; Crypsis aculeata (L.) Aiton, Soreng 7940, Johnson, Johnson, Dzyubenko, Dzyubenko & Schilnikov (US), Russia, JQ345316, JQ345205, JQ345275, JQ345233, JQ345163; Spartina densiflora Brongn., Peterson 19154, Soreng, Salariado & Panizza (US), Argentina, GU359916, GU359510, GU360352, JQ345271, GU359206; Sporobolus indicus (L.) R.Br., Peterson 22025 & Saarela (US), Mexico, GU359913, GU359504, GU360355, GU360630, GU359209; Sporobolus lasiophyllus Pilg., Peterson 21879, Soreng & Sanchez Vega (US), Peru, GU359912, GU359505, GU360356, GU360629, GU359210; Sporobolus virginicus (L.) Kunth, Peterson 15683 & Soreng (US), Chile, GU359892, GU359502, GU360362, GU360610, GU359215; Zoysia macrantha subsp. walshii M.E.Nightingale, Loch 435 (US), Australia, GU359922, GU359548, GU360345, GU360642, GU359197; CYNODONTEAE: Tripogoninae: Tripogon multiflorus Miré & H.Gillet,

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Appendix 1. Continued. Spellenberg 7441 (MO), Yemen, JQ345360, JQ345232, JQ345315, JQ345274, JQ345204; Tripogon trifidus Munro ex Hook.f., Laegaard 21668 & Norsangsri (MO), Thailand, KP873641, KP873957, KP874094, KP873822, KP873422; Eragrostiella leioptera (Stapf) Bor, Chand 7961 (US), India, GU359827, GU359486, JQ345280, GU360529, GU359305; Oropetium capense Stapf, Venter 9939 & Venter (MO), South Africa, KM010692, KM010533, KM010917, KM011120, KM010324; Melanocenchris abyssinica (R.Br. ex Fresen.) Hochst., DeWilde 6912 (MO), Ethiopia, JQ345355, JQ345228, JQ345310, JQ345268, JQ345198; Desmostachya bipinnata (L.) Stapf, Dwyer 13120 (MO), Saudi Arabia, JQ345319, JQ345207, JQ345277, JQ345235, JQ345165; CYNODONTEAE: Pappo- phorinae: Neesiochloa barbata (Nees) Pilg., Swallen 4491 (US), Brazil, GU360005, –, GU360279, GU360724, GU359122; Pappophorum pappiferum (Lam.) Kuntze, Peterson 21689, Soreng, La Torre & Rojas Fox (US), Peru, GU359996, GU359402, GU360276, GU360700, GU359128; Tridens flavus var. chapmanii (Small) Shinners, McCauley 438 (MO), U.S.A., KF827689, KF827615, KF827751, KF827817, KF827568; CYNODONTEAE: Traginae: Monelytrum luederit­ zianum Hack., Smook 10031 (US), South Africa, GU359969, GU359459, GU360421, GU360682, GU359158; Tragus berteronianus Schult., FLSP 457 (US), Peru, GU359898, GU359503, GU360370, GU360616, GU359224; Willkommia sarmentosa Hack., Schweickerdt 2181 (US), South Africa, GU359924, GU359545, GU360343, GU360645, GU359194; CYNODONTEAE: Scleropogoninae: Blepharidachne benthamiana (Hack.) Hitchc., Melix 570 & Cherobini (US), Argentina, GU359857, –, –, GU360579, GU359275; Scleropogon brevifolius Phil., Peterson 19280, Soreng, Salariado & Panizza (US), Argentina, GU359919, GU359530, –, GU360635, GU359203; CYNODONTEAE: Hilariinae: Hilaria cenchroides Kunth, Peterson 22339 & Saarela (US), Mexico, GU359813, GU359424, GU360380, GU360697, GU359143; Pleuraphis rigida Thurber, Boyd 11566 (BRY), U.S.A., JQ345356, JQ345229, JQ345311, JQ345269, JQ345200; CYNODONTEAE: Monanthochloinae: Distichlis eludens (Soderstr. & H.F.Decker) H.L.Bell & Columbus, Peterson 24864 & Romaschenko (US), Mexico, KP873561, –, –, –, KP873345; Distichlis eludens (Soderstr. & H.F.Decker) H.L.Bell & Columbus, Reeder 6430 & Reeder (US), Mexico, JQ345320, –, GU360290, GU360701, GU359139; Distichlis humilis Phil., Peterson 19362, Soreng, Salariato & Panizza (US), Argentina, GU359835, GU359430, GU360478, GU360502, GU359333; Distichlis littoralis (Engelm.) H.L.Bell & Columbus, Moran 10570 (US), Mexico, GU359970, GU359460, GU360422, GU360699, GU359157; CYNODONTEAE: Boutelouinae: Bouteloua simplex Lag., Peterson 21289, Saarela & Flores Villegas (US), Mexico, GU359834, GU359406, GU360297, GU360607, GU359231; CYNODONTEAE: Muhlenbergiinae: Muhlenbergia japonica Steud., Soreng 5301, Peterson & Sun Hang (US), China, HM143184, HM143389, HM143572, HM143668, HM143082; Muhlenbergia schreberi J.F.Gmel., Peterson 19443, Soreng, Salariato & Panizza (US), Argentina, GU359950, GU359456, GU360404, GU360679, GU359161; CYNODONTEAE: Aeluropodinae: Aeluropus lagopoides (L.) Trin. ex Thwaites, Weinert s.n. & Mosawi (US), Iraq, GU360013, GU359391, GU360284, GU360576, GU359261; Aeluropus littoralis (Gouan) Parl., Ferguson 634 (US), Greece, GU360018, GU359390, GU360308, GU360575, GU359262; Aeluropus pungens (M.Bieb.) K.Koch, Yunatov 124, Li Shyin & Yuan Y-fen (US), China, GU360014, GU359389, GU360319, GU360574, GU359263; CYNODONTEAE: Triodiinae: Triodia intermedia Cheel, Peterson 14384, Soreng & Rosenberg (US), Australia, GU359941, GU359563, GU360327, GU360661, GU359179; CYNODONTEAE: Orcuttiinae: Orcuttia tenuis Hitchc., Stone 771 (US), U.S.A., GU360001, GU359398, GU360271, GU360727, JQ345199; Tuctoria greenei (Vasey) Reeder, Reeder 6656 & Reeder (US), U.S.A., GU359928, –, –, –, GU359190; CYNODONTEAE: Gouiniinae: Gouinia paraguayensis (Kuntze) Parodi, Peterson 11526 & Annable (US), Argentina, GU359817, GU359437, GU360384, GU360504, GU359314; Triplasis purpurea (Walter) Chapm., Peterson 14238, Weakley & LeBlond (US), U.S.A., GU359921, GU359536, GU360347, GU360656, GU359184; Vaseyochloa mul­ tinervosa (Vasey) Hitchc., Swallen 10041 (US), U.S.A., GU359925, GU359544, GU360342, GU360646, GU359193; CYNODONTEAE: Cteniinae: Ctenium brevispicatum J.G.Sm., Filgueiras 3251 & Oliviera (US), Brazil, JQ345317, JQ345206, JQ345276, JQ345234, JQ345164; CYNODONTEAE: Trichoneurinae: Trichoneura eleusinoides (Rendle) Ekman, Seydel 448 (US), South Africa, GU359988, GU359522, GU360277, GU360705, GU359135; CYNODONTEAE: Farragoinae: Craspedorhachis rhodesiana Rendle, Strohbach 5699 (US), Namibia, KF827623, KF827574, KF827696, KF827760, KF827504; Farrago rac­ emosa Clayton, Peterson 23851, Soreng, Romaschenko & Abeid (US), Tanzania, KF827634, KF827582, KF827706, KF827772, KF827516; CYNODONTEAE: Perotidinae: Mosdenia leptostachys (Ficalho & Hiern) Clayton, Schweickerdt 1542 (US), South Africa, GU359967, GU359458, GU360420, GU360681, GU359159; Perotis arenacea (Judz.) P.M.Peterson, Phillipson 4117 & Raharilala (MO), Madagascar, JQ345358, –, JQ345313, JQ345272, JQ345202; Perotis hordeiformis Nees, Soreng 5717, Peterson & Sun Hang (US), China, GU359991, GU359520, GU360283, GU360708, GU359132; Trigonochloa uniflora (Hochst. ex A.Rich.) P.M.Peterson & N.Snow, Greenway 14075 (MO), Tanzania, JQ345347, –, –, JQ345261, –; Trigonochloa uniflora (Hochst. ex A.Rich.) P.M.Peterson & N.Snow, Snow 6978, Burgoyne & Gumbi (MO), South Africa, JQ345348, –, –, –, JQ345192; CYNODONTEAE: Gymnopogoninae: Bewsia biflora (Hack.) Gooss., Peterson 23905, Soreng, Romaschenko & Abeid (US), Tanzania, KF827617, KF827570, KF827692, KF827754, KF827498; Dignathia hirtella Stapf, McCallum-Webster 5251 (US), Kenya, –, GU359368, GU360481, GU360490, GU359316; Gymnopogon grandiflorus Roseng., B.R.Arill. & Izag., Peterson 16642 & Refulio-Rodriguez (US), Peru, GU359816, GU359436, GU360383, GU360581, GU359200; Leptocarydion vulpiastrum (De Not.) Stapf, Mboya 565 (MO), Tanzania, KF827658, KF827593, KF827723, KF827790, KF827537; Leptothrium rigidum Kunth, Davidse 3281 (MO), Jamaica, KF827662, KF827596, KF827727, KF827794, KF827541; Lophacme digitata Stapf, Smook 1453 (MO), South Africa, JQ345354, –, JQ345309, JQ345267, JQ345197; CYNODONTEAE: Eleusininae: Acrachne racemosa (B.Heyne ex Roem. & Schult.) Ohwi, Jackson 1408 (US), Malawi, KP873423, KP873823, –, –, KP873206; Acrachne rac­ emosa (B.Heyne ex Roem. & Schult.) Ohwi, Pappi 5990 (US), Eritrea, –, –, –, –, KP873207; Acrachne racemosa (B.Heyne ex Roem. & Schult.) Ohwi, Rattray 1303 (US), Zimbabwe, KP873424, –, KP873958, –, KP873208; Acrachne racemosa (B.Heyne ex Roem. & Schult.) Ohwi, Strohbach 1308 (US), Namibia, KP873425, KP873824, KP873959, KP873642, KP873209; Acrachne racemosa (B.Heyne ex Roem. & Schult.) Ohwi, Zavattari 1195 (FT), Ethiopia, KP873426, KP873825, KP873960, KP873643, KP873210; Acrachne racemosa (B.Heyne ex Roem. & Schult.) Ohwi, Zavattari 1196 (FT), Ethiopia, KP873427, KP873826, KP873961, KP873644, KP873211; Afrotrichloris martinii Chiov., Hemming 3407 (FT), Somalia, KP873428, KP873827, KP873962, KP873645, KP873212; Afrotrichloris martinii Chiov., Herlocker 459 (MO), Somalia, –, –, –, –, KP873213; Apochiton burttii C.E.Hubb., Greenway 11513 & Polhill (US), Tanzania, –, –, –, GU360571, GU359266; Apochiton burttii C.E.Hubb., Peterson 24163, Soreng, Romaschenko & Abeid (US), Tanzania, KP873429, KP873828, KP873963, KP873646, KP873214; Astrebla elymoides F.Muell. ex F.M.Bailey, Hubbard 7976 (US), Australia, KP873430, GU359414, GU360313, –, GU359269; Astrebla lappacea (Lindl.) Domin, McKinlay s.n. (US), Australia, GU360009, GU359395, GU360312, GU360568, GU359270; Astrebla pectinata (Lindl.) F.Muell. ex Benth., Chalmers 5 (US), Australia, GU359861, GU359421, GU360311, GU360567, GU359286; Austrochloris dichanthioides (Everist) Lazarides, Anson s.n. (US), Australia, GU359860, GU359420, GU360310, GU360566, GU359272; Brachyachne ambigua Ohwi, Leach 3996 (MEL), Australia, KP873431, KP873829, KP873964, KP873647, KP873215; Brachyachne ciliaris (Kuntze) C.E.Hubb., Latz 14224 (MEL), Australia, KP873432, KP873830, KP873965, KP873648, KP873216; Brachyachne ciliaris (Kuntze) C.E.Hubb., Mullen s.n. (US), Australia, –, –, KP873966, KP873649, –; Brachyachne convergens (F.Muell.) Stapf, Adams 851 (US), Australia, GU359885, GU359349, –, GU360586, GU359252; Brachyachne fulva Stapf, Robson 1122 (US), Mozambique, GU359884, GU359348, GU360444, GU360585, GU359253; Brachyachne obtusiflora (Benth.) C.E.Hubb., Laegaard 16295 (US), Zimbabwe, GU359883, GU359374, GU360458, GU360584, GU359254; Brachyachne obtusiflora (Benth.) C.E.Hubb., Schweinfurth 2150 (US), South Sudan, KP873433, –, KP873967, KP873650, KP873217; Brachyachne patentiflora (Stent & J.M.Rattray) C.E.Hubb., Bogdan 7075 (US), Kenya, GU359862, GU359350, GU360295, GU360605, GU359233; Brachyachne tenella (R.Br.) C.E.Hubb., Lazarides 4281 (US), Australia, GU359882, GU359376, GU360442, GU360583, GU359255; Chloris amethystea Hochst., Archer 8650 (US), Ethiopia, KP873434, KP873831, KP873968, KP873651, KP873218; Chloris andropogonoides E.Fourn., Cory 52467 (US), U.S.A., KP873435, –, –, –, KP873219; Chloris andropogonoides E.Fourn., Peterson 18839, Valdés-Reyna & Sifuentes (US), Mexico, KP873436, KP873832, KP873969, KP873652, KP873220; Chloris andropogonoides E.Fourn., Swallen 10091 (US), U.S.A., KP873437, KP873833, KP873970, KP873653, KP873221; Chloris andro­ pogonoides E.Fourn., Swallen 10232 (US), U.S.A., –, –, KP873971, KP873654, KP873222; Chloris barbata Sw., Bentley 102 (US), Ecuador, KP873438, –, KP873972, –, KP873223; Chloris barbata Sw., Ernst 1507 (US), Dominica, KP873439, KP873834, KP873973, KP873655, KP873224; Chloris barbata Sw., King 4038 (US), Mexico, KP873440, KP873835, KP873974, KP873656, KP873225; Chloris barbata Sw., Peterson 22255 & Saarela (US), Mexico, GU359873, GU359377, GU360435, GU360514, GU359320; Chloris barbata Sw., Peterson 23795, Soreng & Romaschenko (US), Tanzania, KP873441, KP873836, KP873975, KP873657, KP873226; Chloris barbata Sw., Saarela 1740, Peterson, Soreng & Judziewicz (CAN), Australia, KP873442, KP873837, KP873976, KP873658, KP873227; Chloris barbata Sw., Saarela 1830, Peterson, Soreng & Judziewicz (CAN), Australia, KP873443, KP873838, KP873977, KP873659, KP873228;

Version of Record 463 Peterson & al. • P hylogeny and classification of the Eleusininae TAXON 64 (3) • June 2015: 445–467

Appendix 1. Continued. Chloris canterae var. grandiflora (Roseng. & Izag.) D.E.Anderson, Izaguirre de Artucio 2642 (US), Uruguay, –, –, –, KP873660, KP873229; Chloris canterae Arechav., Pedersen 9597 (US), Paraguay, KP873444, KP873839, KP873978, KP873661, KP873230; Chloris canterae Arechav., Schinini 5321 (US), Argen­ tina, –, KP873840, KP873979, –, –; Chloris canterae Arechav., Smith 01 (US), U.S.A., KP873445, KP873841, KP873980, KP873662, KP873231; Chloris castilloniana Lillo & Parodi, Gallinal H. 5542, Aragone, Bergalli, Campal & Rosengurtt (US), Uruguay, KP873446, –, KP873981, KP873663, KP873232; Chloris ciliata Sw., Lundell 14872 (US), U.S.A., KP873447, KP873842, KP873982, KP873664, KP873233; Chloris ciliata Sw., Peterson 24622 & Romaschenko (US), Mexico, KP873448, –, –, –, KP873234; Chloris ciliata Sw., Peterson 24632 & Romaschenko (US), Mexico, KP873449, –, –, –, KP873235; Chloris cruciata (L.) Sw., Correll 48667 & Proctor (US), Bahamas, KP873450, –, –, –, KP873236; Chloris cruciata (L.) Sw., Correll 50431 & Correl (US), Bahamas, KP873451, –, KP873983, KP873665, KP873237; Chloris cubensis Hitchc. & Ekman, Box 35 (US), Antigua and Barbuda, KP873452, –, –, –, KP873238; Chloris cucullata Bisch., Crutchfield 5475 & Johnston (US), Mexico, KP873453, KP873843, –, KP873666, KP873239; Chloris cucullata Bisch., Isely 10647 (US), U.S.A., KP873454, KP873844, –, KP873667, KP873240; Chloris divaricata var. cynodontoides (Balansa) Lazarides, Greenwood 226C (US), Fiji, KP873455, –, –, KP873668, KP873241; Chloris divaricata R.Br., Swallen 10572 (US), U.S.A., KP873456, –, –, KP873669, KP873242; Chloris ekmanii Hitchc., Liogier 12416 (US), Dominican Republic, KP873457, KP873845, –, KP873670, KP873243; Chloris ekmanii Hitchc., Liogier 15884 (US), Dominican Republic, KP873458, KP873846, –, KP873671, KP873244; Chloris elata Desv., Acosta-Solis 12667 (US), Ecuador, KP873459, –, –, –, KP873245; Chloris elata Desv., Beck 4454 (US), Bolivia, KP873460, KP873847, –, KP873672, KP873246; Chloris elata Desv., Spiaggi 40 (US), Bolivia, KP873461, KP873848, –, KP873673, KP873247; Chloris elata Desv., Swallen 14461 (US), U.S.A., KP873462, KP873849, –, KP873674, KP873248; Chloris elata Desv., Webster 10789, Samuel & Williams (US), Bahamas, KP873463, –, –, KP873675, KP873249; Chloris exilis Renvoize, Anderson 37049, Stieber & Kirkbride (US), Brazil, KP873464, –, –, KP873676, KP873250; Chloris exilis Renvoize, Anderson 37190, Sticber & Kirkbride (US), Brazil, KP873465, –, –, KP873677, KP873251; Chloris gayana Kunth, Balleza 1788 (US), Mexico, KP873466, –, –, KP873678, KP873252; Chloris gayana Kunth, Batty 125 (US), U.S.A., KP873467, KP873850, –, KP873679, KP873253; Chloris gayana Kunth, Peterson 23972, Soreng, Romaschenko & Abeid (US), Tanzania, KP873468, –, KP873984, KP873680, KP873254; Chloris gayana Kunth, Peterson 23997, Soreng, Romaschenko & Abeid (US), Tanzania, KP873469, KP873851, KP873985, KP873681, KP873255; Chloris gayana Kunth, Peterson 24253, Soreng, Romaschenko & Mbago (US), Tanzania, KP873470, KP873852, KP873986, KP873682, KP873256; Chloris halophila Parodi, Laegaard 71450 (US), Ecuador, KP873471, KP873853, –, KP873683, KP873257; Chloris halophila Parodi, Peterson 13970 & Refulio-Rodriguez (US), Peru, KP873472, KP873854, –, KP873684, KP873258; Chloris lamproparia Stapf, Sherif 3945 (K), Sudan, KP873473, –, –, –, KP890054; Chloris latisquamea Nash, Fleetwood s.n. (US), U.S.A., KP873474, KP873855, –, KP873685, KP873259; Chloris latisquamea Nash, Stan­ ford s.n., Lauber & Taylor (US), Mexico, KP873475, KP873856, –, KP873686, KP873260; Chloris mossambicensis K.Schum., Oakes 1211 (US), South Africa, GU359903, –, –, GU360621, GU359219; Chloris mossambicensis K.Schum., Peterson 23803, Soreng & Romaschenko (US), Tanzania, KP873476, KP873857, KP873987, KP873687, KP873261; Chloris mossambicensis K.Schum., Peterson 24219, Soreng, Romaschenko & Mbago (US), Tanzania, KP873477, KP873858, KP873988, KP873688, KP873262; Chloris orthonoton Döll, Harley 16629, Renvoize, Erskine, Brighton & Pinheiro (US), Brazil, KP873478, –, –, KP873689, KP873263; Chloris orthonoton Döll, Pedersen 5817 (US), Argentina, KP873479, –, –, KP873690, KP873264; Chloris pectinata Benth., Saarela 1699, Peter­ son, Soreng & Judziewicz (CAN), Australia, KP873480, KP873859, KP873989, KP873691, KP873265; Chloris pilosa Schumach., Allen s.n. & Baumgartner (US), France, KP873481, KP873860, –, KP873692, KP873266; Chloris pilosa Schumach., Brenan 9601 (US), Nigeria, KP873482, –, –, KP873693, KP873267; Chloris pilosa Schumach., Peterson 23919, Soreng, Romaschenko & Abeid (US), Tanzania, KP873483, KP873861, KP873990, KP873694, KP873268; Chlo­ ris prieurii Kunth, Laegaard 17061 & Traore (US), Senegal, KP873484, KP873862, –, KP873695, KP873269; Chloris prieurii Kunth, Laegaard 17863 & Traore (US), Senegal, KP873485, –, –, KP873696, KP873270; Chloris pycnothrix Trin., Bidgood 3170 & Vollesen (US), Tanzania, KP873486, KP873863, –, KP873697, KP873271; Chloris pycnothrix Trin., Harley 17042 (US), Brazil, KP873487, KP873864, –, KP873698, KP873272; Chloris pycnothrix Trin., Pedersen 6378 (US), Argentina, KP873488, –, –, KP873699, KP873273; Chloris pycnothrix Trin., Peterson 22278 & Saarela (US), Mexico, GU359872, GU359366, GU360434, GU360513, GU359321; Chloris pycnothrix Trin., Peterson 23949, Soreng & Romaschenko (US), Tanzania, KP873489, KP873865, KP873991, KP873700, KP873274; Chloris pycnothrix Trin., Peterson 23974, Soreng, Romaschenko & Abeid (US), Tanzania, KP873490, –, KP873992, KP873701, KP873275; Chloris pycnothrix Trin., Peterson 24070, Soreng, Romaschenko & Abeid (US), Tanzania, KP873491, KP873866, KP873993, KP873702, KP873276; Chloris pycnothrix Trin., Peterson 24172, Soreng, Romaschenko & Abeid (US), Tanzania, KP873492, KP873867, KP873994, KP873703, KP873277; Chloris pycnothrix Trin., Peterson 24184, Soreng, Romaschenko & Mbago (US), Tanzania, KP873493, KP873868, KP873995, KP873704, KP873278; Chlo­ ris pycnothrix Trin., Peterson 24262, Soreng, Romaschenko & Mbago (US), Tanzania, KP873494, KP873869, KP873996, KP873705, KP873279; Chloris pycnothrix Trin., Peterson 24305, Soreng, Romaschenko & Mbago (US), Tanzania, KP873495, KP873870, KP873997, KP873706, KP873280; Chloris rox­ burghiana Schult., Peterson 23812, Soreng & Romaschenko (US), Tanzania, KP873496, KP873871, KP873998, KP873707, KP873281; Chloris roxburghiana Schult., Peterson 24194, Soreng, Romaschenko & Mbago (US), Tanzania, KP873497, KP873872, KP873999, KP873708, KP873282; Chloris roxburghiana Schult., Peterson 24225, Soreng, Romaschenko & Mbago (US), Tanzania, KP873498, KP873873, KP874000, KP873709, KP873283; Chloris rufescens Lag., McVaugh 19517 (US), Mexico, KP873499, KP873874, –, KP873710, KP873284; Chloris sagraeana A.Rich., McKee 10539 (US), Guadeloupe, KP873500, KP873875, –, KP873711, KP873285; Chloris subdolichostachya C.Müll., Trouart 17 (US), U.S.A., KP873501, KP873876, –, KP873712, KP873286; Chlo­ ris submutica Kunth, McGregor 706, Harms, Robinson, Rosarie & Seagal (US), Mexico, KP873502, KP873877, –, KP873713, KP873287; Chloris submutica Kunth, Peterson 22393 & Saarela (US), Mexico, GU359871, GU359375, GU360471, GU360512, GU359322; Chloris submutica Kunth, Peterson 24560 & Romaschenko (US), Mexico, KP873503, –, –, –, KP873288; Chloris submutica Kunth, Peterson 24842 & Romaschenko (US), Mexico, KP873504, –, –, –, KP873289; Chloris submutica Kunth, Peterson 24939, Romaschenko & González-Elizondo (US), Mexico, KP873505, –, –, –, KP873290; Chloris submutica Kunth, Reeder 4845 & Reeder (US), Mexico, KP873506, KP873878, –, KP873714, KP873291; Chloris texana (Vasey) Nash, Silvens 423 (US), U.S.A., KP873507, –, –, KP873715, KP873292; Chloris verticillata Nutt., Reeder 5787 & Reeder (US), U.S.A., KP873508, –, –, KP873716, KP873293; Chloris virgata Sw., Peterson 21468, Soreng, LaTorre & Rojas Fox (US), Peru, GU359869, GU359373, GU360445, GU360510, GU359324; Chloris virgata Sw., Peter­ son 24286, Soreng, Romaschenko & Mbago (US), Tanzania, KP873509, KP873879, KP874001, KP873717, KP873294; Chloris virgata Sw., Peterson 24659 & Romaschenko (US), Mexico, KP873510, –, –, –, KP873295; Chloris virgata Sw., Peterson 24789, Romaschenko, Rodriguez Avalos, Herrera-Simoni & Garcia Rodriguez (US), Mexico, KP873511, –, –, –, KP873296; Chloris virgata Sw., Peterson 24854 & Romaschenko (US), Mexico, KP873512, –, –, –, KP873297; Chloris virgata Sw., Peterson 24911 & Romaschenko (US), Mexico, KP873513, –, –, –, KP873298; Chloris virgata Sw., Peterson 24954 & Romaschenko (US), Mexico, KP873514, –, –, –, KP873299; Chrysochloa annua C.E.Hubb., Arkrok 20417 (US), Ghana, KP873515, KP873880, KP874002, KP873718, KP873300; Chrysochloa hindsii C.E.Hubb., Reekmans 11068 (US), Burundi, GU359868, –, GU360485, GU360509, GU359325; Chrysochloa orientalis (C.E.Hubb.) Swallen, Burtt 5666 (US), Tanzania, KP873516, –, KP874003, KP873719, KP873301; Coelachyrum lagopoides (Burm.f.) Senaratna, Saldanha 15334 (US), India, GU359844, –, –, GU360602, –; Coelachyrum poiflorum Chiov., Burger 2915 (US), Ethiopia, GU359843, –, GU360457, GU360601, GU359236; Cynodon aethiopicus Clayton & J.R.Harlan, Peterson 24268, Soreng, Romaschenko & Mbago (US), Tanzania, KP873517, KP873881, KP874004, KP873720, KP873302; Cynodon aethiopicus Clayton & J.R.Harlan, Peterson 24318, Soreng, Romaschenko & Mbago (US), Tanzania, KP873518, KP873882, KP874005, KP873721, KP873303; Cynodon aethiopicus Clayton & J.R.Harlan, Smook 4140 (US), South Africa, KP873519, KP873883, KP874006, KP873722, KP873304; Cynodon aethiopicus Clayton & J.R.Harlan, Vesey-Fitzgerald 6475 (US), Tanzania, KP873520, –, –, –, –; Cynodon affinis Caro & E.A.Sánchez, Booth 1251 (US), U.S.A., KP873521, KP873884, KP874007, KP873723, KP873305; Cynodon affinis Caro & E.A.Sánchez, Cabrera 5149 (US), Argentina, KP873522, KP873885, KP874008, KP873724, KP873306; Cynodon affinis Caro & E.A.Sánchez, Collins 63 (US), U.S.A., –, KP873886, KP874009, KP873725, KP873307; Cynodon aristiglumis Caro & E.A.Sánchez, Beetle 2731 (US), U.S.A., KP873523, KP873887, KP874010, KP873726, KP873308; Cynodon aristulatus Caro & E.A.Sánchez, Egler 42-314 (US), Guatemala, KP873524, KP873888, KP874011, KP873727, KP873309; Cynodon aristulatus Caro & E.A.Sánchez, Fassett 26000 (US), Colombia, KP873525, KP873889, KP874012, KP873728, KP873310; Cynodon aristulatus Caro & E.A.Sánchez,

464 Version of Record TAXON 64 (3) • June 2015: 445–467 Peterson & al. • Phylogeny and classification of the Eleusininae

Appendix 1. Continued. Killip 31600 (US), U.S.A., KP873526, KP873890, KP874013, KP873729, KP873311; Cynodon dactylon var. longiglumis Caro & E.A.Sánchez, Dunkle 8366 (US), U.S.A., KP873527, –, –, –, –; Cynodon dactylon var. longiglumis Caro & E.A.Sánchez, Dunkle 8652 (US), U.S.A., KP873528, –, –, –, –; Cynodon dac­ tylon var. pilosus Caro & E.A.Sánchez, Marsh 141 (US), U.S.A., KP873529, –, –, –, –; Cynodon dactylon var. longiglumis (L.) Pers., Peterson 19701, Saarela & Sears (US), U.S.A., KP873530, –, –, –, –; Cynodon dactylon (L.) Pers., Peterson 22000 & Saarela (US), Mexico, GU359836, GU359359, GU360453, GU360580, GU359243; Cynodon dactylon var. pilosus (L.) Pers., Peterson 24626 & Romaschenko (US), Mexico, KP873531, –, –, –, KP873312; Cynodon dactylon var. pilosus (L.) Pers., Saarela 1606, Peterson, Soreng & Judziewicz (CAN), Australia, KP873532, KP873891, KP874014, KP873730, KP873313; Cynodon erectus J.Presl, Asplund 10905 (US), Peru, –, –, KP874015, KP873731, KP873314; Cynodon erectus J.Presl, Martin 1814 (US), U.S.A., KP873533, KP873892, KP874016, KP873732, KP873315; Cynodon hirsutissimus (Litard. & Maire) Caro & E.A.Sánchez, Parodi 15066 (US), Argentina, –, –, –, –, KP873316; Cynodon hirsutus Stent, Smook 6616 (US), South Africa, GU359876, GU359358, GU360452, GU360594, GU359229; Cynodon incompletus Nees, Oakes 663 (US), South Africa, KP873534, –, –, –, –; Cynodon incompletus Nees, Smook 10152 (US), South Africa, GU359847, GU359347, GU360451, GU360609, GU359245; Cynodon incompletus Nees, Smook 2408 & Gibbs-Russell (US), South Africa, GU359849, KP873893, GU360450, KP873733, GU359246; Cyno­ don maritimus Kunth, Howard 10214 & Howard (US), Bahamas, GU359889, GU359365, GU360448, GU360591, GU359248; Cynodon maritimus var. vagini­ florus Caro, Martindale s.n. (US), U.S.A., KP873535, –, –, –, –; Cynodon mucronatus Caro & E.A.Sánchez, Fosberg 273 & Batts (US), U.S.A., KP873536, KP873894, KP874017, KP873734, KP873317; Cynodon mucronatus Caro & E.A.Sánchez, Hermann 8502 (US), U.S.A., KP873537, KP873895, KP874018, KP873735, KP873318; Cynodon nitidus Caro & E.A.Sánchez, Macbride 524 & Featherstone (US), Peru, –, –, KP874019, KP873736, KP873319; Cynodon nlemfuensis Vanderyst, Hansen 13002 (US), U.S.A., KP873538, KP873896, KP874020, KP873737, KP873320; Cynodon nlemfuensis Vanderyst, Laegaard 19845 (US), Ecuador, KP873539, KP873897, KP874021, KP873738, KP873321; Cynodon nlemfuensis Vanderyst, Laegaard 19876 (US), Ecuador, KP873540, KP873898, KP874022, KP873739, KP873322; Cynodon nlemfuensis Vanderyst, Lobo 2099, Cascante, Quesada & Prieto (CR), Costa Rica, KJ768976, KP873899, KP874023, KP873740, KJ768881; Cynodon nlemfuensis Vanderyst, Molina 25803 (US), Honduras, –, GU359353, –, GU360589, –; Cynodon nlemfuensis Vanderyst, Peterson 23791, Soreng & Romaschenko (US), Tanzania, KP873541, –, –, KP873741, KP873323; Cynodon nlemfuensis Vanderyst, Peterson 24058, Soreng, Romaschenko & Abeid (US), Tanzania, KP873542, KP873900, KP874024, KP873742, KP873324; Cynodon nlemfuensis Vanderyst, Peterson 24113, Soreng, Romaschenko & Abeid (US), Tanzania, KP873543, KP873901, KP874025, KP873743, KP873325; Cynodon nlemfuensis Vanderyst, Peterson 24207, Soreng, Romaschenko & Mbago (US), Tanzania, KP873544, KP873902, KP874026, KP873744, KP873326; Cynodon pascuus Nees, Moran 12518 (US), Mexico, GU359888, GU359354, GU360447, GU360590, GU359249; Cynodon plectostachyus (K.Schum.) Pilg., Troupin 11610 (US), Rwanda, GU359890, GU359356, GU360449, GU360592, GU359247; Cynodon radiatus Roth, Clayton 5836 (US), Sri Lanka, GU359837, GU359360, GU360454, GU360596, GU359257; Cynodon radiatus Roth, Laegaard 21611 (US), Thailand, KP873545, KP873903, KP874027, KP873745, KP873327; Cynodon transvaalensis Burtt Davy, Laegaard 20560 (US), Ecuador, –, –, KP874028, KP873746, KP873328; Cynodon transvaalensis Burtt Davy, Smook 6710 (US), South Africa, GU359887, GU359352, GU360446, GU360588, GU359250; Daknopholis boivinii (A.Camus) Clayton, Ranaivojaona 1441, Andrianjafy, Phillipson & Lubke (MO), Madagascar, KP873548, KP873907, KP874032, KP873750, KP873332; Dinebra aquatica (Scribn. & Merr.) P.M.Peterson & N.Snow, Soderstrom 650 (US), Mexico, KP873549, KP873908, KP874033, KP873751, KP873333; Dinebra caudata (K.Schum.) P.M.Peterson & N.Snow, Kuchar 24265 (MO), Tan­ zania, JQ345324, JQ345210, JQ345282, JQ345239, JQ345170; Dinebra chinensis (L.) P.M.Peterson & N.Snow, Snow 6909 (MO), Botswana, JQ345325, –, –, JQ345240, JQ345171; Dinebra coerulescens (Steud.) P.M.Peterson & N.Snow, Harris 289 & Fay (MO), Central African Republic, –, –, JQ345283, –, JQ345172; Dinebra decipiens subsp. asthenes (Roem. & Schult.) P.M.Peterson & N.Snow, Snow 7327 (MO), Australia, JQ345326, –, JQ345284, JQ345241, JQ345173; Dinebra decipiens subsp. decipiens (R.Br.) P.M.Peterson & N.Snow, Snow 7328 & Simon (MO), Australia, JQ345327, JQ345211, JQ345285, JQ345242, JQ345174; Dinebra decipiens subsp. asthenes (Roem. & Schult.) P.M.Peterson & N.Snow, Snow 7355 (MO), Australia, JQ345328, JQ345212, JQ345286, JQ345243, JQ345175; Dinebra decipiens subsp. peacockii (Maiden & Betche) P.M.Peterson & N.Snow, Snow 7361 & Simon (MO), Australia, JQ345329, –, JQ345287, JQ345244, JQ345176; Dinebra decipiens subsp. decipiens (R.Br.) P.M.Peterson & N.Snow, Waterhouse 5948 (MO), Australia, JQ345330, –, JQ345288, JQ345245, JQ345177; Dinebra divaricatissima (S.T.Blake) P.M.Peterson & N.Snow, Brown 253 (MEL), Australia, KP873550, KP873909, –, KP873752, KP873334; Dinebra haareri (Stapf & C.E.Hubb.) P.M.Peterson & N.Snow, Peterson 24231, Soreng, Romaschenko & Mbago (US), Tanzania, KP873551, KP873910, KP874034, KP873753, KP873335; Dinebra ligulata (Lazarides) P.M.Peterson & N.Snow, Snow 7402 (MO), Australia, JQ345336, JQ345218, JQ345294, JQ345251, JQ345183; Dinebra marquisensis (F.Br.) P.M.Peterson & N.Snow, Wood 10145 (MO), French Polynesia, JQ345337, JQ345219, JQ345295, JQ345252, JQ345184; Dinebra nealleyi (Vasey) P.M.Peterson & N.Snow, Snow 5806 (MO), U.S.A., JQ345338, JQ345220, JQ345296, JQ345253, JQ345185; Dinebra nee­ sii (Thwaites) P.M.Peterson & N.Snow, Snow 7380 & Simon (MO), Australia, JQ345339, JQ345221, JQ345297, JQ345254, JQ345186; Dinebra panicea subsp. brachiata (Steud.) P.M.Peterson & N.Snow, Peterson 20086, Alvarez Marvan & Dejesos (US), Mexico, JQ345341, –, –, –, –; Dinebra panicea subsp. brachiata (Steud.) P.M.Peterson & N.Snow, Peterson 22185 & Saarela (US), Mexico, GU359810, GU359431, GU360389, –, GU359146; Dinebra panicea subsp. mucronata (Michx.) P.M.Peterson & N.Snow, Peterson 9546, Annable & Herrera (US), Mexico, JQ345342, JQ345222, JQ345299, JQ345256, JQ345188; Dinebra panicoi­ des (J.Presl) P.M.Peterson & N.Snow, Snow 5810a (MO), U.S.A., JQ345343, –, JQ345300, JQ345257, –; Dinebra retroflexa (Vahl) Panz., Ndegwa 610 (US), Kenya, GU359778, GU359355, GU360479, GU360503, GU359332; Dinebra retroflexa (Vahl) Panz., Snow 6950 & Burgoyne (MO), Swaziland, KP873552, –, –, –, KP873336; Dinebra scabra (Nees) P.M.Peterson & N.Snow, Snow 5788 (MO), U.S.A., JQ345344, JQ345223, JQ345301, JQ345258, JQ345189; Dinebra somalensis (Stapf) P.M.Peterson & N.Snow, Faden 74-991 & Faden (MO), Kenya, JQ345321, –, JQ345278, JQ345236, JQ345166; Dinebra southwoodii (N.Snow & B.K.Simon) P.M.Peterson & N.Snow, Snow 7362 (MO), Australia, JQ345345, JQ345224, JQ345302, JQ345259, JQ345190; Dinebra squarrosa (Pilg.) P.M.Peterson & N.Snow, Kayombo 5196 (MO), Tanzania, JQ345346, JQ345225, JQ345303, JQ345260, JQ345191; Dinebra viscida (Scribn.) P.M.Peterson & N.Snow, Peterson 22184 & Saarela (US), Mexico, GU359808, GU359429, GU360430, GU360693, GU359148; Dinebra viscida (Scribn.) P.M.Peterson & N.Snow, Snow 6528a& Prinzie (MO), Mexico, JQ345352, –, JQ345307, JQ345265, –; Dinebra xerophila (P.M.Peterson & Judz.) P.M.Peterson & N.Snow, Perlman 18438 (MO), French Polynesia, JQ345353, JQ345227, JQ345308, JQ345266, JQ345196; Diplachne fusca var. muelleri (Benth.) J.M.Black, Badman 1282 (MO), Aus­ tralia, JQ345334, JQ345216, JQ345292, JQ345249, JQ345181; Diplachne fusca var. uninervia (J.Presl) P.M.Peterson & N.Snow, Peterson 20786, Soreng, Romaschenko & Gonzalez-Elizondo (US), Peru, JQ345335, JQ345217, JQ345293, JQ345250, JQ345182; Diplachne fusca var. uninervia (J.Presl) P.M.Peterson & N.Snow, Peterson 21305, Saarela & Flores Villegas (US), Mexico, GU359809, GU359461, GU360391, GU360694, GU359147; Diplachne fusca var. fusca (L.) P.Beauv. ex Roem. & Schult., Peterson 24322, Soreng, Romaschenko & Mbago (US), Tanzania, KP873553, KP873911, KP874035, KP873754, KP873337; Diplachne fusca var. fusca (L.) P.Beauv. ex Roem. & Schult., Walsh 6558 (MEL), Australia, KP873554, KP873912, KP874036, KP873755, KP873338; Disakisperma dubium (Kunth) P.M.Peterson & N.Snow, Peterson 22334 & Saarela (US), Mexico, GU359811, GU359442, GU360416, GU360695, GU359145; Disakisperma dubium (Kunth) P.M.Peterson & N.Snow, Peterson 24453, Romaschenko & Valdés-Reyna (US), Mexico, KP873555, –, –, –, KP873339; Disaki­ sperma dubium (Kunth) P.M.Peterson & N.Snow, Peterson 24554 & Romaschenko (US), Mexico, KP873556, –, –, –, KP873340; Disakisperma dubium (Kunth) P.M.Peterson & N.Snow, Peterson 24566, Romaschenko & Valdés-Reyna (US), Mexico, KP873557, –, –, –, KP873341; Disakisperma dubium (Kunth) P.M.Peterson & N.Snow, Peterson 24665 & Romaschenko (US), Mexico, KP873558, –, –, –, KP873342; Disakisperma dubium (Kunth) P.M.Peterson & N.Snow, Peterson 24845 & Romaschenko (US), Mexico, KP873559, –, –, –, KP873343; Disakisperma dubium (Kunth) P.M.Peterson & N.Snow, Peterson 24905 & Romaschenko (US), Mexico, KP873560, –, –, –, KP873344; Disakisperma dubium (Kunth) P.M.Peterson & N.Snow, Peterson 8105 & Annable (US), Mexico, JQ345332, JQ345214, JQ345290, JQ345247, JQ345179; Disakisperma eleusine (Nees) P.M.Peterson & N.Snow, Snow 6982 (MO), South Africa, JQ345333, JQ345215, JQ345291, JQ345248, JQ345180; Disakisperma obtusiflorum (Hochst.) P.M.Peterson & N.Snow, Belsky 527 (MO), Kenya, JQ345340, –, JQ345298, JQ345255, JQ345187; Disakisperma obtusiflorum (Hochst.) P.M.Peterson & N.Snow, Peterson 24198, Soreng, Romaschenko & Mbago (US), Tanzania, KF574411, KF574405, KF574417, KF574423, KF574400; Disakisperma obtusiflorum (Hochst.) P.M.Peterson & N.Snow, Peterson 24211, Soreng, Romaschenko & Mbago (US), Tanzania, KF574412, KF574406, KF574418, KF574424, KF574401; Disakisperma obtusiflorum (Hochst.) P.M.Peterson & N.Snow, Peterson 24243,

Version of Record 465 Peterson & al. • P hylogeny and classification of the Eleusininae TAXON 64 (3) • June 2015: 445–467

Appendix 1. Continued. Soreng, Romaschenko & Mbago (US), Tanzania, KF574413, KF574407, KF574419, KF574425, KF574402; Disakisperma yemenicum (Schweinf.) P.M.Peterson & N.Snow, Peterson 24254, Soreng, Romaschenko & Mbago (US), Tanzania, KF574414, KF574408, KF574420, KF574426, KF574403; Disakisperma yemeni­ cum (Schweinf.) P.M.Peterson & N.Snow, Verdcourt 3275 (US), Kenya, KF574415, KF574409, KF574421, KF574427, KF574404; Eleusine africana Kenn.- O’Byrne, Kayombo 424 (MO), Tanzania, KP873562, KP873913, KP874037, KP873756, KP873346; Eleusine africana Kenn.-O’Byrne, Peterson 24072, Soreng, Romaschenko & Abeid (US), Tanzania, KP873563, KP873914, KP874038, KP873757, KP873347; Eleusine africana Kenn.-O’Byrne, Qing Liu 090 (US), Mexico, HQ202625, HQ202648, HQ202640, HQ202632, –; Eleusine africana Kenn.-O’Byrne, Seydel 2798 (US), Namibia, KP873564, –, –, –, KP873348; Eleusine africana Kenn.-O’Byrne, Seydel 3889 (US), Namibia, KP873565, –, –, –, KP873349; Eleusine coracana (L.) Gaertn., Peterson 24112, Soreng, Romaschenko & Abeid (US), Tanzania, KP873566, KP873915, KP874039, KP873758, KP873350; Eleusine coracana (L.) Gaertn., Qing Liu 093 (US), Ethiopia, KP873567, –, –, –, KP873351; Eleusine coracana (L.) Gaertn., Qing Liu 103 (US), Ethiopia, KP873568, –, –, –, KP873352; Eleusine floccifolia (Forssk.) Spreng., Hilu s.n.(VPI), Ethiopia, –, –, –, –, AY515208*; Eleusine indica (L.) Gaertn., Brown 628 & Clebsch (US), U.S.A., KP873569, –, –, –, KP873353; Eleusine indica (L.) Gaertn., Chase 9922 (US), U.S.A., KP873570, –, –, –, KP873354; Eleusine indica (L.) Gaetrn., Peterson 21362, Saarela & Flores Villegas (US), Mexico, GU359797, GU359473, GU360472, GU360496, GU359338; Eleusine indica (L.) Gaetrn., Peterson 23802, Soreng & Romaschenklo (US), Tan­ zania, KP873571, KP873916, KP874040, KP873759, KP873355; Eleusine indica (L.) Gaertn., Peterson 24429, Romaschenko & Knapp (US), U.S.A., KP873572, –, KP874041, KP873760, KP873356; Eleusine jaegeri Pilg., Hilu 203 (VPI), Kenya, –, –, –, –, AY515213*; Eleusine jaegeri Pilg., Kindeketa 762 (MO), Tanzania, KP873573, KP873917, KP874042, KP873761, KP873357; Eleusine jaegeri Pilg., Peterson 24299, Soreng, Romaschenko & Mbago (US), Tanzania, KP873574, KP873918, KP874043, KP873762, KP873358; Eleusine kigeziensis S.M.Phillips, Purseglove 3384 (K), Uganda, –, –, –, –, AY515216*; Eleusine multiflora Hochst. ex A.Rich., Davidse 30264 & Davidse (MO), Mexico, KP873575, –, –, –, KP873359; Eleusine multiflora Hochst. ex A.Rich., Peterson 24272, Soreng, Romaschenko & Mbago (US), Tanzania, KP873576, KP873919, KP874044, KP873763, KP873360; Eleusine multiflora Hochst. ex A.Rich., Qing Liu 100 (US), Kenya, HQ202630, HQ202653, HQ202646, HQ202638, –; Eleusine tristachya (Lam.) Lam., Beck 3227 (US), Bolivia, KP873577, –, –, –, KP873361; Eleusine tristachya (Lam.) Lam., Hilu 2464 (VPI), Uruguay, –, –, –, –, AY515224*; Eleusine tristachya (Lam.) Lam., Qing Liu 107 (US), Mexico, HQ202631, HQ202654, HQ202647, HQ202639, –; Eleusine tristachya (Lam.) Lam., Zardini 35791 & Tillería (MO), Paraguay, KP873578, KP873920, KP874045, KP873764, KP873362; Enteropogon acicularis (Lindl.) Lazarides, Cochrane 521B (MEL), Australia, KP873579, KP873921, KP874046, KP873765, KP873363; Enteropogon barbatus C.E.Hubb., Kabuye 575, Luke, Robertson, Mungai & Mathenge (US), Kenya, KP873580, –, KP874047, KP873766, KP873364; Entero­ pogon brandegeei (Vasey) Clayton, Carter 2897 & Kellogg (US), Mexico, KP873581, KP873922, KP874048, KP873767, KP873365; Enteropogon chlorideus (J.Presl) Clayton, Gould 12679 (US), Mexico, KP873582, KP873923, KP874049, KP873768, KP873366; Enteropogon chlorideus (J.Presl) Clayton, Leaven­ worth 1900 (US), Mexico, KP873583, –, KP874050, KP873769, KP873367; Enteropogon chlorideus (J.Presl) Clayton, Meyer s.n. (US), U.S.A., KP873584, –, KP874051, KP873770, KP873368; Enteropogon chlorideus (J.Presl) Clayton, Sohns 384 (US), Mexico, KP873585, –, –, –, KP873369; Enteropogon doli­ chostachyus (Lag.) Keng ex Lazarides, Blake 23488 (MEL), Australia, KP873586, –, –, –, KP873370; Enteropogon dolichostachyus (Lag.) Keng ex Lazarides, Gould 13680 & Cooray (US), Sri Lanka, KP873587, –, KP874052, KP873771, KP873371; Enteropogon dolichostachyus (Lag.) Keng ex Lazarides, Lazarides 7416 (US), Thailand, KP873588, –, –, KP873772, KP873372; Enteropogon macrostachyus (Hochst. ex A.Rich.) Munro ex Benth., Laegaard 15902 (US), Zimbabwe, GU359795, GU359472, GU360470, GU360494, GU359340; Enteropogon macrostachyus (Hochst. ex A.Rich.) Munro ex Benth., Oakes 1250 (US), South Africa, KP873589, KP873924, KP874053, KP873773, KP873373; Enteropogon macrostachyus (Hochst. ex A.Rich.) Munro ex Benth., Peterson 24197, Soreng, Romaschenko & Mbago (US), Tanzania, KP873590, KP873925, KP874054, KP873774, KP873374; Enteropogon macrostachyus (Hochst. ex A.Rich.) Munro ex Benth., Peterson 24228, Soreng, Romaschenko & Mbago (US), Tanzania, KP873591, KP873926, KP874055, KP873775, KP873375; Enteropogon macrostachyus (Hochst. ex A.Rich.) Munro ex Benth., Smook 9026 (US), Namibia, KP873592, KP873927, KP874056, KP873776, KP873376; Enteropogon mollis (Nees) Clayton, Proosdij 813, van de Riet & Zauder (US), Aruba, KP873593, KP873928, –, KP873777, KP873377; Enteropogon mollis (Nees) Clayton, Steyermark 29192 (US), Guatemala, KP873594, –, –, KP873778, KP873378; Enteropogon monostachyus (Vahl) K.Schum., Barber 2607 (US), India, KP873595, KP873929, KP874057, KP873779, KP873379; Enteropogon monostachyus (Vahl) K.Schum., Clayton 5954 (US), Sri Lanka, KP873596, –, KP874058, KP873780, KP873380; Enteropogon ramosus B.K.Simon, Peterson 14367, Soreng & Rosenberg (US), Australia, GU359794, GU359470, GU360469, GU360493, GU359341; Enteropogon ramosus B.K.Simon, Saarela 1663, Peterson, Soreng & Judziewicz (CAN), Australia, KP873597, KP873930, KP874059, KP873781, KP873381; Enteropogon rupestris (J.A.Schmidt) A.Chev., Bogdan 4026 (US), Kenya, KP873598, –, KP874060, KP873782, KP873382; Enteropogon rup­ estris (J.A.Schmidt) A.Chev., Gillet 13082 (US), Kenya, KP873599, KP873931, KP874061, KP873783, KP873383; Enteropogon rupestris (J.A.Schmidt) A.Chev., McKinnon 124 (US), Somalia, KP873600, –, –, KP873784, –; Enteropogon rupestris (J.A.Schmidt) A.Chev., Vesey-Fitzgerald 4973 (US), Kenya, KP873601, KP873932, KP874062, KP873785, KP873384; Enteropogon sechellensis (Baker) T.Durand & Schinz, Fosberg 49814 (US), Seychelles, –, KP873933, KP874063, KP873786, KP873385; Enteropogon sechellensis (Baker) T.Durand & Schinz, Peterson 23815, Soreng & Romaschenko (US), Tan­ zania, KP873602, KP873934, KP874064, KP873787, KP873386; Enteropogon sechellensis (Vahl) K.Schum., Peterson 23833, Soreng, Romaschenko & Abeid (US), Tanzania, KP873603, KP873935, KP874065, KP873788, KP873387; Eustachys calvescens (Hack.) Caro & E.A.Sánchez, Chase 10971 (US), Brazil, KP873604, KP873936, KP874066, KP873789, KP873388; Eustachys calvescens (Hack.) Caro & E.A.Sánchez, Rojas 6633 (US), Paraguay, KP873605, –, KP874067, KP873790, KP873389; Eustachys caribaea (Spreng.) Herter, Renvoize 4022 & Cope (US), Bolivia, KP873606, KP873937, KP874068, KP873791, KP873390; Eustachys distichophylla (Lag.) Nees, Harris s.n. (US), U.S.A., KP873607, KP873938, –, KP873792, KP873391; Eustachys floridana Chapm., Kral 44355 (US), U.S.A., KP873608, KP873939, –, KP873793, KP873392; Eustachys glabrescens (Hack.) Caro & E.A.Sánchez, Pedersen 6121 (US), Argen­ tina, –, –, KP874069, KP873794, KP873393; Eustachys glabrescens (Hack.) Caro & E.A.Sánchez, Pedersen 6307 (US), Argentina, KP873609, KP873940, KP874070, KP873795 , KP873394; Eustachys glauca Chapm., Swallen 8221 (US), U.S.A., KP873610, KP873941, –, KP873796, KP873395; Eustachys neglecta (Nash) Nash, Chase 4499 (US), U.S.A., KP873611, –, –, KP873797, KP873396; Eustachys paspaloides (Vahl) Lanza & Mattei, Ndegway 741 (US), Kenya, GU359819, GU359439, GU360386, GU360522, GU359312; Eustachys petraea (Sw.) Desv., Strong 3124 (US), U.S.A., GU359833, GU359438, GU360385, GU360637, GU359313; Eustachys petraea (Sw.) Desv., Taylor 12539 (US), Mexico, –, –, –, –, KP873397; Eustachys retusa (Lag.) Kunth, Krapovickas 22799 & Cristobal & Quarin (US), Brazil, –, –, –, –, KP873398; Eustachys tenera (J.Presl) A.Camus, Strong 10564 & Clemens (US), Papua New Guinea, KP873612, –, KP874071, KP873798, KP873399; Harpochloa falx (L.f.) Kuntze, Zietsman Peyper, Avenant & Van der Walt (MO), South Africa, KP873613, –, KP874072, KP873799, KP873400; Leptochloa chloridiformis Parodi, Myndel-Pedersen 2662 (US), Argentina, KP873614, –, KP874073, KP873800, KP873401; Lepto­ chloa chloridiformis Parodi, Myndel-Pedersen 3741 (US), Argentina, KP873615, –, –, KP873801, KP873402; Leptochloa crinita (Lag.) P.M.Peterson, Renvoize 1987 (US), Bolivia, JQ345359, JQ345231, JQ345315, JQ345273, JQ345203; Leptochloa digitata (R.Br.) Domin, Risler 476 (MO), Australia, JQ345331, JQ345213, JQ345289, JQ345246, JQ345178; Leptochloa pluriflora (E.Fourn.) P.M.Peterson & N.Snow, Peterson 15048 & Refulio-Rodriguez (US), Peru, GU359905, GU359554, GU360334, GU360623, GU359212; Leptochloa pluriflora (E.Fourn.) P.M.Peterson & N.Snow, Sohns 1258 (US), Mexico, GU359934, GU359553, GU360374, GU360653, GU359214; Leptochloa pluriflora (E.Fourn.) P.M.Peterson & N.Snow, Swallen 10100 (US), U.S.A., KP873616, KP873942, KP874074, KP873802, KP873403; Leptochloa virgata (L.) P.Beauv., Peterson 15088 & Refulio-Rodriguez (US), Peru, JQ345349, –, JQ345304, JQ345262, JQ345193; Leptochloa virgata (L.) P.Beauv., Rimachi 8359 (US), Peru, JQ345350, –, JQ345305, JQ345263, JQ345194; Leptochloa virgata (L.) P.Beauv., Vargas 2710 (US), Bolivia, JQ345351, JQ345226, JQ345306, JQ345264, JQ345195; Lepturus gasparricensis Fosberg, Herbst 9687 (US), U.S.A. Wake Isl., GU359807, GU359477, GU360429, GU360692, GU359149; Lepturus geminatus C.E.Hubb., Morton 1711 (MEL), Australia, KP873617, KP873943, KP874075, KP873803, KP873404; Lepturus radicans (Steud.) A.Camus, Gilleopie 38 (US), Kenya, GU359806, –, –, –, –; Lepturus repens (G.Forst.) R.Br., Peterson 23835, Soreng, Romaschenko & Abeid (US), Tanzania, KP873618, KP873944, KP874076, KP873804, KP873405; Lepturus repens R.Br., Whistler 9853 (US), Chagos Archipelago, Diego Garcia Isl., GU359893, GU359427, GU360428, GU360691, GU359150; Lintonia nutans Stapf, Mwasumbi 14374 (US), Tanzania, GU359980, GU359426, GU360427, GU360690, GU359151; Microchloa altera (Rendle) Stapf, Brand 315, Kay, Logie & Logie (MO), South Africa, KP873619, KP873945, KP874077,

466 Version of Record TAXON 64 (3) • June 2015: 445–467 Peterson & al. • Phylogeny and classification of the Eleusininae

Appendix 1. Continued. KP873805, KP873406; Microchloa altera (Rendle) Stapf, Malaisse 16572 (MO), Congo (Kinshasa) Dem. Rep., KP873620, KP873946, KP874078, KP873806, KP873407; Microchloa altera (Rendle) Stapf, Robinson 6362 (US), Zambia, KP873621, KP873947, KP874079, KP873807, KP873408; Microchloa caffra Nees, Peterson 24288, Soreng, Romaschenko & Mbago (US), Tanzania, KP873622, KP873948, KP874080, KP873808, KP873409; Microchloa caffra Nees, Smook 10441 (US), South Africa, GU359972, GU359453, GU360424, GU360670, GU359155; Microchloa kunthii Desv., Hagerup 455 & Olufsen (US), Mali, KP873623, KP873949, KP874081, KP873809, –; Microchloa kunthii Desv., Peterson 22152 & Saarela (US), Mexico, GU359971, GU359434, GU360423, GU360684, GU359141; Microchloa kunthii Desv., Peterson 24048, Soreng, Romaschenko & Abeid (US), Tanzania, KP873624, KP873950, KP874082, KP873810, KP873410; Microchloa kunthii Desv., Peterson 24255, Soreng, Romaschenko & Mbago (US), Tanzania, KP873625, KP873951, KP874083, KP873811, KP873411; Microchloa kunthii Desv., Peterson 24716, Romaschenko & Zamudio Ruíz (US), Mexico, KP873626, –, –, –, KP873412; Ochthochloa compressa (Forssk.) Hilu, Gloves 48 & Gilliband (US), Somalia, KP873628, –, –, –, KP890055; Ochthochloa compressa (Forssk.) Hilu, Rechinger 29410 (US), Pakistan, KP873629, –, KP874084, –, KP890056; Ochthochloa compressa (Forssk.) Hilu, USDA 154-57 (US), Pakistan, KP873630, –, KP874085, –, KP890057; Oxychloris scariosa (F.Muell.) Lazarides, Forster 20737 & Holland (MEL), Australia, KP873631, KP873952, KP874086, KP873813, KP873414; Saugetia fasciculata Hitchc. & Chase, Ekman s.n. (US), Dominican Republic, GU359982, GU359528, GU360317, GU360638, GU359156; Schoenefeldia gracilis Kunth, Pauwels 7413 (MO), Benin, KJ768981, –, KP874087, KP873814, KJ768886; Schoenefeldia transiens (Pilg.) Chiov., Greenway 9781 (US), Kenya, GU360007, KP873953, GU360349, GU360636, GU359202; Schoenefeldia transiens (Pilg.) Chiov., Peterson 24216, Soreng, Romaschenko & Mbago (US), Tanzania, KP873632, KP873954, KP874088, KP873815, KP873415; Schoenefeldia transiens (Pilg.) Chiov., Spjut 3920 (BRY), Kenya, JQ345357, JQ345230, JQ345312, JQ345270, JQ345201; Tetrapogon bidentatus Pilg., Bogdan 3903 (US), Kenya, KP873633, –, KP874089, KP873816, KP873416; Tetra­ pogon bidentatus Pilg., Greenway 12778 & Kanuri (US), Kenya, KP873634, –, KP874090, –, KP873417; Tetrapogon cenchriformis (A.Rich.) Clayton, BN 7701 (US), Ethiopia, KP873635, –, KP874091, KP873817, –; Tetrapogon cenchriformis (A.Rich.) Clayton, Greenway 8844 (US), Kenya, KP873636, –, –, –, –; Tetrapogon cenchriformis (A.Rich.) Clayton, Peterson 24192, Soreng, Romaschenko & Mbago (US), Tanzania, KP873637, –, –, KP873818, KP873418; Tetrapogon cenchriformis (A.Rich.) Clayton, Peterson 24240, Soreng, Romaschenko & Mbago (US), Tanzania, KP873638, KP873955, KP874092, KP873819, KP873419; Tetrapogon tenellus (J.Koenig ex Roxb.) Chiov., Peterson 23794, Soreng & Romaschenko (US), Tanzania, KP873639, KP873956, KP874093, KP873820, KP873420; Tetrapogon tenellus (J.Koenig ex Roxb.) Chiov., Peterson 24244, Soreng, Romaschenko & Mbago (US), Tanzania, KP873640, –, –, KP873821, KP873421; Tetrapogon villosus Desf., Ash 2561 (US), Ethiopia, GU359902, –, GU360366, GU360620, GU359220; Tetrapogon villosus Desf., Johannes s.n. (US), Canary Islands, GU359901, GU359514, GU360367, GU360619, GU359221; Incertae sedis: Pogononeura biflora Napper, Greenway 10091 (US), Tanzania, KM010694, –, –, –, –; Neobouteloua lophostachya (Griseb.) Gould, Peterson 11515 & Annable (US), Argentina, GU360004, GU359396, GU360273, GU360725, GU359123; Neobouteloua pauciracemosa M.G.López & Biurrun, Porter 11968 & Columbus (US), Argentina, KP873627, –, –, KP873812, KP873413; Dactyloctenium geminatum Hack., Peterson 23808, Soreng & Romaschenko (US), Tanzania, KP873546, KP873904, KP874029, KP873747, KP873329; Dactyloctenium robecchii Chiov., Puccioni 626 & Stefanini (FT), Somalia, KP873547, KP873905, KP874030, KP873748, KP873330; Dactyloctenium scindicum Boiss., Hemming 2358 (US), Somalia, –, KP873906, KP874031, KP873749, KP873331.

Version of Record 467 Vol. 64 (3) • June 2015

International Journal of Taxonomy, Phylogeny and Evolution

Electronic Supplement to

A molecular phylogeny and classification of the Eleusininae with a new genus, Micrachne (Poaceae: Chloridoideae: Cynodonteae) Paul M. Peterson, Konstantin Romaschenko, & Yolanda Herrera Arrieta

Taxon : – Fig. S1. Maximum-likelihood tree from analysis of ITS sequences. Numbers above branches are posterior probabilities; numbers below branches are bootstrap values.

Chasmanthium latifolium P22463 1.00 Danthonia compressa P21986 100 Rytidosperma penicellatum P19685 Aristida gypsophila P15839 89 1.00 Neyraudia reynaudiana RJS5318 Triraphis ramosissima Sey4278 57 100 1.00 Cottea pappophoroides Enneapogon desvauxii P21999 1.00 98 1.00 Uniola condensata P9342 95 53 1.00 Entoplocamia aristulata Sey187 0.98 51 Tetrachne dregei Jarm120 61 1.00 Eragrostis desertorum AU 1.00 0.91 86 Psammagrostis wiseana P14345 100 0.98 Eragrostis nigricans P21623 51 0.95 Harpachne harpachnoides RJS5288 53 Ectrosia scabrida Laz4772 Zoysia macrantha walshii Loch435 0.96 1.00 Sporobolus indicus P22025 63 1.00 0.87 Crypsis aculeata Soreng7940 95 100 1.00 60 Sporobolus lasiophyllus P21879 93 1.00 Sporobolus virginicus P15683 78 1.00 Calamovilfa longifolia Ha5738 100 Spartina densiflora P19154 0.56 Demostachya bipinnata Dwyer13120 1.00 63 Melanocenchris abyssinica Wilde6912 100 1.00 Eragrostiella leioptera Chand7961 100 1.00 Oropetium capense Veuter9939 75 1.00 Tripogon multiflorus Spellenberg7441 100 Tripogon trifidus Laegaard21668 0.53 1.00 Tragus racemosus 457 0.91 Monelytrum luederitzianum Sm10031 52 100 72 Willkommia sarmentosa Sch2181 1.00 Muhlenbergia japonica RJS5301 0.59 100 Muhlenbergia schreberi P19443 Neesiochloa barbata Reed6247 1.00 1.00 57 63 1.00 Pappophorum pappiferum P21689 1.00 100 Tridens flavus chapmanii McCanley438 1.00 73 Blepharidachne benthamiana Me570 0.74 77 Scleropogon brevifolius P19280 1.00 Hilaria cenchroides2 P22339 0.99 100 Pleuraphis rigida Boyd11566 0.63 Bouteloua simplex P21289 1.00 Distichlis littoralis Mor10570 Distichlis humilis P19362 99 0.81 51 0.81 Distichlis eludens P24864 52 Distichlis eludens Reed6430 1.00 Neobouteloua paucirracemosa PorterJM 11968 99 Neobouteloua lophostachya P11515 Acrachne racemosa JacksonG 1408 Acrachne racemosa StrohbachB 1308 1.00 1.00 Acrachne racemosa ZavattariE 1195 97 1.00 100 Acrachne racemosa ZavattariE 1196 97 Acrachne racemosa PappiA 5990 1.00 Acrachne racemosa RattrayJM 1303 100 1.00 Dactyloctenium robecchii NPuccioni626 100 1.00 Dactyloctenium geminatum P23808 100 Dactyloctenium scindicum HemmingCF 2358

1.00 Triplasis purpurea P14238 1.00 Gouinia paraguayensis P11526 84 90 Vaseyochloa multinervosa Sw10041 1.00 1.00 Ctenium brevispicatumFilgueiras 71 61 Trichoneura eleusinoides Sey448 1.00 Leptothrium rigidum Davidse3281 58 1.00 Dignathia hirtella CW5251 Leptocarydion vulpiastrum Mboya565 63 0.80 Bewsia biflora Dav6471 0.86 86 1.00 Gymnopogon grandiflorus P16642 1.00 97 70 0.72 Lophachme digitata Smook1453 1.00 57 Craspedorhachis rhodesiana Strohbach 5699 0.66 100 Farrago racemosa P23851a Trigonochloa uniflora Greenway14075 1.00 Mosdenia phleoides Sch1542 100 1.00 100 1.00 Perotis hordeiformis RJS5717 100 Perotis arenacea Phillipson4117 Triodia intermedia P14384 1.00 Orcuttia tenuis 771 100 Tuctoria greenei Ree6656 0.98 Aeluropus lagopoides Weinert 0.60 1.00 71 100 1.00 Aeluropus littoralis Ferg634 65 92 Aeluropus pungens Yunat124 0.55 Dinebra chinensis Snow6909 1.00 Dinebra marquisensis Wood10145 75 Dinebra xerophila Perlman18438 1.00 1.00 Dinebra panicea brachiata P22185 0.98 100 85 0.53 98 Dinebra panicea mucronata P9546 0.90 Dinebra squarrosa Kayombo5196 65 0.96 Dinebra retroflexa Nd610 1.00 83 Dinebra retroflexa SnowN BurgoyneP 6950 100 0.64 Dinebra aquatica Soderstrom650 1.00 55 Dinebra coerulescens Harris 289 95 0.52 Dinebra caudata Kuchar 2426524 52 Dinebra viscida P22184 0.88 1.00 98 1.00 Dinebra nealteyi Snow5806 100 Dinebra scabra Snow5788 1.00 1.00 Dinebra haareri P24231 96 100 1.00 Dinebra somalensis Faben74 991 Dinebra neesii Snow7380 98 1.00 1.00 Dinebra ligulata Snow7402 98 100 Dinebra southwoodii Snow7362 1.00 Dinebra decipiens peacockii Snow7361 88 0.52 1.00 Dinebra decipiens asthenes Snow7327 98 Dinebra decipiens asthenes Snow7355 1.00 Dinebra decipiens decipiens Snow7328 100 Dinebra decipiens decipiens Waterhouse5948 Apochiton burttii P24163 1.00 Coelachyrum poiflorum Bur2915 100 0.93 Eleusine multiflora P24272 1.00 84 1.00 Eleusine jaegeri Kindeketa762 98 100 Eleusine jaegeri P24299 57H11 Eleusine africana SeydelR 2798 1.00 1.00 Eleusine africana SeydelR 3889 95 Eleusine coracana africana Kayombo424 Eleusine P24072 1.00 Eleusine indica P24429 100 1.00 Eleusine caracana P24112 Eleusine coracana LiuQ 093 89 1.00 Eleusine coracana LiuQ 103 96 0.96 Eleusine indica BrownF ClebschA 628 93 Eleusine indica ChaseVH 9922 1.00 Eleusine indica P23802 Eleusine indica P21362 93 0.99 84 1.00 Eleusine tristachya BeckStG 3227 97 Eleusine tristachya Zardini35791 Diplachne divaricatissima BrownAJ 253 1.00 Diplachne uninervia P21305 100 Diplachne fusca P24322 0.88 66 Diplachne fusca unineura Peterson20786 0.94 Diplachne fusca fusca WalshNG 6558 70 Diplachne fusca muelleri Badman1282 Leptochloa digitata Risler476 1.00 Chloris exilis AndersonWR 37049 1.00 0.52 100 Chloris exilis AndersonWR 37190 98 1.00 Chloris mollis ProosdijASJ 813 100 Chloris mollis SteyenmarkJA 29192 1.00 0.65 1.00 Leptochloa pluriflora Swallen 10100 89 59 100 0.61 Leptochloa crinita Renv1987 1.00 73 Leptochloa pluriflora P15048 100 Leptochloa virgata RimachiY8359 0.58 1.00 Leptochloa chloridiformis MyndelPedersen2662 74 99 Leptochloa chloridiformis MyndelPedersen3741 1.00 Leptochloa virgata Peterson15088 99 Leptochloa virgata Vargas2710 Schoenfeldia gracilis Pauwels7413 1.00 Austrochloris dichanthioides Anson 0.74 Astrebla lappacea Mckin 62 92 1.00 100 0.59 Astrebla elymoides Hub7976 54 Astrebla pectinata Cha5 Disakisperma yemenicum P24254 1.00 Disakisperma eleusine Snow6982 100 Disakisperma obtusiflora Belsky527 0.84 Disakisperma obtusiflora P24211 53 1.00 1.00 Disakisperma obtusiflora P24198 100 68 Disakisperma obtusiflora P24243 1.00 Disakisperma dubia P24554 92 Disakisperma dubia P8105 1.00 Disakisperma dubia P22334 1.00 100 Disakisperma dubia P24566 0.93 80 Disakisperma dubia P24665 79 Disakisperma dubia P24845 0.84 0.97 Disakisperma dubia P24453 55 69 Disakisperma dubia P24905 0.98 Enteropogon ramosus Saarela1663 1.00 69 Enteropogon acicularis CochraneAC 521B 93 Enteropogon dolichostachyus GouldFW 13680 1.00 Enteropogon dolichostachyus LazaridesM 7416 87 Enteropogon dolichostachyus BlakeST 23488 1.00 Enteropogon barbatus KabuyeC 575 100 Enteropogon macrostachyus SL15902 Enteropogon macrostachyus OakesAJ 1250 Enteropogon macrostachyus SmookL 9026 1.00 Enteropogon monostachyus africanus P23833 Enteropogon sechellensis FosbergFR 49814 97 Enteropogon sechellensis P23815 0.90 Enteropogon macrostachyus P24197 64 Enteropogon macrostachyus P24228 0.97 Enteropogon monostachyus BarberCA 2607 62 Enteropogon monostachyus ClaytonD 5954 1.00 Enteropogon somalensis BogdanA 4026 94 0.90 Enteropogon rupestris GilletJB 13082 74 Enteropogon rupestris VeseyFitzgeraldD 4973 1.00 Afrotrichloris martinii HemmingCF 3407 1.00 100 Afrotrichloris martinii Herlocker459 99 1.00 Schoenefeldia transiens P24216 Schoenfeldia transiens2 Spjut3920 1.00 100 90 Daknopholis boivinii Ranaivojaona1441 1.00 Lepturus repens P23835 100 1.00 Lepturus repens Whi9853 100 0.54 Lepturus gasparricensis Herb9687 53 Lepturus geminatus MortonA 1711 Saugetia fasciculata Ekm 1.00 Tetrapogon bidentatus BogdanA 3903 98 Tetrapogon bidentatus Greenway 12778 1.00 Tetrapogon tenellus P23794 97 Tetrapogon tenellus P24244 1.00 0.99 Chloris roxburghiana P24194 100 88 1.00 Chloris roxburghiana P23812 72 Chloris roxburghiana P24225 0.98 Tetrapogon cenchriformis P24192 0.99 84 Tetrapogon cenchriformis P24240 78 1.00 Tetrapogon villosus Ash2561 83 Tetrapogon villosus Johan 0.84 Chloris brandegeei CarterAM 2897 Enteropogon chlorideus GouldFW 12679 65 1.00 Enteropogon chlorideus MeyerFG 24 10 1958 98 0.63 1.00 Enteropogon chlorideus LeavenworthWC 1900 97 Enteropogon chlorideus SohnsER 384 0.91 Ochthochloa compressa Gloves 485 1.00 77 Ochthochloa compressa USDA 154 0.75 100 Ochthochloa compressa Rechinger 29410 66 1.00 Chloris mossambicensis Oakes 1211 1.00 98 Chloris mossambicensis P24219 96 Lintonia nutans Mwa14374 1.00 Chloris prieurii Laegaard 17061 0.55 99 Chloris prieurii Laegaard 17863 1.00 59 1.00 Chloris barbata 22255 82 100 Chloris barbata BentleyPS 102 Chloris barbata P23795 0.89 Chloris inflata ErnstWR 1507 89 1.00 Chloris inflata KingRM 4038 95 Chloris Saarela1740 Chloris Saarela1830 Chloris amethystea ArcherWA 8650 1.00 Chloris mossambicensis P23803 100 0.53 1.00 Chloris cucullata CrutchfieldJ 5475 1.00 100 Chloris cucullata IselyD 10647 93 1.00 Chloris latisquamea FleetwoodIF 24 02 1959 0.54 100 Chloris latisquamea StanfordLR 54 Chloris verticillata Reeder 5787 1.00 Chloris subdolichostachya TrouartJE 17 95 0.99 Chloris andropogonoides P18839 1.00 910.99 Chloris andropogonoides CoryVL 52467 100 910.99 Chloris andropogonoides Swallen 10091 89 Chloris andropogonoides Swallen 10232 1.00 Chloris castilloniana GallinalH 5542 100 Chloris orthonoton PedersenTM 5817 Chloris gayana P24253 0.68 1.00 Chloris gayana BallezaJJ 1788 99 1.00 Chloris gayana BattyR 125 98 1.00 Chloris gayana P23972 1.00 1.00 73 Chloris gayana P23997 100 100 0.95 Chloris pilosa AllenP 19 10 1967 87 Chloris pilosa BrenanJPM 9601 1.00 Chloris pilosa P23919 98 0.89 Chloris virgata P24286 67 0.99 Chloris virgata P24954 0.94 Chloris virgata P21468 72 72 1.00 Chloris virgata P24854 1.00 Chloris virgata P24911 95 87 1.00 Chloris virgata P24659 73 Chloris virgata P24789 Chloris cynodontoides GreenwoodW 226C 1.00 Chloris divaricata Swallen 10572 94 Chloris Saarela1699 Chloris submutica Reeder 4845 1.00 Chloris cruciata CorrellDS 48667 100 Chloris cruciata CorrellDS 50431 Chloris cubensis BoxHE 35 1.00 0.86 1.00 Chloris ekmanii LiogierAH 12416 100 90 Chloris ekmanii LiogierAH 15884 Chloris sagrana McKeeHS 10539 Chloris submutica P24842 Chloris submutica P24939 1.00 Chloris submutica McGregorRL 706 87 1.00 Chloris submutica P22393 84 Chloris submutica P24560 Chloris orthonoton HarleyRM 16629 0.93 Chloris rufescens MaVaughR 19517 66 Chloris texana SilveusWA 423 0.98 Chloris halophila LaegaardS 71450 0.96 74 Chloris halophila P13970 52 Chloris pycnothrix P23949 Chloris pycnothrix P23974 1.00 Chloris pycnothrix P24070 83 Chloris pycnothrix P24305 Chloris pycnothrix P24172 Chloris pycnothrix P24184 Chloris pycnothrix P24262 0.99 Chloris pycnothrix BidgoodS 3170 61 Chloris pycnothrix HarleyRM 17042 Chloris pycnothrix PedersenTM 6378 Chloris pycnothrix P22278 Oxychloris scariosa ForsterPI 20737 Harpochloa flax Zletsmam4015 Microchloa altera Brand315 1.00 1.00 Microchloa altera RobinsonEA6362 82 1.00 100 Microchloa altera Malaisse6572 Microchloa caffra Sm10441 100 1.00 Microchloa kunthii P24288 99 1.00 0.94 100 1.00 Microchloa kunthii P24255 59 83 0.87 Microchloa kunthii P24048 66 Microchloa kunthii P24716 Eustachys paspaloides Nd741 1.00 Eustachys tenera StrongClemensM 10564 100 Eustachys distichophylla HarrisJP 30 04 1951 1.00 1.00 Eustachys glabrescens PedersenTM 6121 99 1.00 99 Eustachys glabrescens PedersenTM 6307 78 1.00 0.57 Eustachys retusa KrapovickasA 22799 98 52 1.00 Eustachys caribaea Renvoize 4022 100 1.00 Eustachys calvescens ChaseA 10971 99 Eustachys calvescens RojasT 6633 Eustachys glauca Swallen 8221 1.00 Eustachys petraea TaylorJ 12539 1.00 97 Eustachys floridana KralR 44355 100 0.76 Eustachys neglecta ChaseA 4499 72 Eustachys petraea St3124 1.00 Brachyachne fulva Rob1122 100 1.00 Brachyachne chrysolepis Bog7075 100 0.68 Brachyachne obtusiflora Schweinfurth 2150 75 Brachyachne patentiflora SL16295 1.00 Chrysochloa orientalis BurttBD 5666 0.78 100 1.00 Chrysochloa annua ArkrokJO 20417 55 99 Chrysochloa hindsii Ree11068 Chloris canterae PedersenTM 9597 Chloris canterae SmithJE 01 1.00 1.00 Chloris grandiflora Izaguirre 2642 100 95 Chloris elata AcostaSolisM 12667 Chloris elata Swallen 14461 0.98 Chloris elata WebsterGL 10789 74 0.96 Chloris elata BeckStG 4454 63 Chloris elata SpiaggiL 40 1.00 Chloris ciliata LundellCL 14872 96 0.99 Chloris ciliata P24622 82 Chloris ciliata P24632 1.00 Cynodon maritimus Hov10214 1.00 94 Cynodon transvaalensis Laegaard20560 82 Cynodon transvaalensis Smook6710 1.00 Cynodon radiatus Laegaard21611 100 1.00 Brachyachne ambigua LeachGJ 3996 100 Brachyachne convergens Ad851 Brachyachne ciliaris LatzPK 14224 Brachyachne tenella Laz4281 0.56 Cynodon hirsutus Smook 6616 1.00 Cynodon aethiopicus P24268 100 Cynodon aethiopicus P24318 Cynodon nlemfuensis P24058 1.00 Cynodon nlemfuensis P24113 98 Cynodon nlemfuensis P24207 Cynodon nlemfuensis HansenB 13002 1.00 Cynodon nlemfuensis Laegaard 19876 100 Cynodon nlemfuensis Laegaard 19845 98 Cynodon nlemfuensis Lobo2099 1.00 Cynodon aethiopicus Smook4140 68 Cynodon plectostachyus Tro11610 0.75 Cynodon aristulatus EglerFE 42 314 Cynodon aristulatus FassettNC 26000 1.00 Cynodon aristulatus KillipEP 31600 91 Cynodon dactylon pilosus P24626 Cynodon dactylon pilosus Saarela1606 Cynodon affinis BoothWE 1251 Cynodon affinis CabreraAL 5149 Cynodon affinis CollinsRW 63 Cynodon aristiglumis BeetleAA 2731 Cynodon nlemfuensis P23791 0.94 Cynodon erectus AsplundE 10905 86 Cynodon erectus MartinRF 1814 Cynodon hirsutissimus ParodiLR 15066 Cynodon mucronatus FosbergFR 273 Cynodon mucronatus HermannFJ 8502 Cynodon nitidus MacbrideJF 524 Fig. S2. Maximum-likelihood tree from analysis of 4 plastid markers (rpl32-trnL, ndhA intron, rps16 intron, rps16-trnK). Numbers above branches are posterior probabilities; numbers below branches are bootstrap values.

Chasmanthium latifolium P22463 Aristida gypsophila P15839 1.00 Capeochloa cincta subsp. sericea Barker1545 100 1.00 Danthonia compressa P21986 100 Rytidosperma penicellatum P19685 Ellisochloa rangei Barker960 1.00 Neyraudia reynaudiana RJS5318 1.00 100 Triraphis ramosissima Sey4278 78 1.00 Cottea pappophoroides 1.00 100 Enneapogon desvauxii 1.00 97 1.00 Uniola condensata P9342 100 100 1.00 Tetrachne dregei Jarm120 1.00 100 Entoplocamia aristulata Sey187 100 Eragrostis desertorumAU 1.00 0.83 Eragrostis nigricans P21623 100 1.00 Harpachne harpachnoides RJS5288 59 1.00 100 Ectrosia scabrida Laz4772 100 1.00 Zoysia macrantha walshii Loch435 100 Sporobolus indicus P22025 0.68 1.00 Sporobolus lasiophyllus P21879 54 1.00 93 Sporobolus virginicus P15683 1.00 83 1.00 Crypsis aculeata Soreng7940 100 98 1.00 Calamovilfa longifolia Ha5738 97 Spartina densiflora P19154 Triodia intermedia P14384 1.00 Aeluropus lagopoides Weinert 1.00 100 Aeluropus littoralis Ferg634 100 Aeluropus pungens Yunat124 1.00 Neesiochloa barbata Reed6247 1.00 82 Pappophorum pappiferum P21689 100 Tridens flavus chapmanii McCanley438

1.00 Desmostachya bipinnata Dwyer13120 1.00 95 Melanocenchris abyssinica Wilde6912 100 1.00 Eragrostiella leioptera Chand7961 1.00 100 Tripogon multiflorus Spellenberg7441 100 Tripogon trifidus Laegaard21668 0.68 1.00 Pogononeura biflora Greenway10091 isoT 1.00 Willkommia sarmentosa Sch2181 96 1.00 Monelytrum luederitzianum Sm10031 99 1.00 1.00 92 Tragus berteronianus 1.00 83 Blepharidachne benthamiana Me570 97 99 Scleropogon brevifolius P19280 0.98 1.00 Muhlenbergia japonica RJS5301 59 100 Muhlenbergia schreberi P19443 0.51 0.92 1.00 Hilaria cenchroides P22339 100 Pleuraphis rigida Boyd11566 59 Bouteloua simplex P21289 1.00 Distichlis humilis P19362 83 1.00 97 1.00 Distichlis littoralis Mor10570 92 0.81 Distichlis eludens P24864 69 Distichlis eludens Reed6430 1.00 Neobouteloua lophostachya 100 Neobouteloua paucirracemosa 1.00 77 0.52 Dactyloctenium robecchii NPuccioni626 29bG01 1.00 Dactyloctenium scindicum BoulosL9431 29bG06 0.97 98 Dactyloctenium geminatum P23808 58H07 59 1.00 1.00 Orcuttia tenuis 1.00 80 Tuctoria greenei Ree6656 97 63 0.88 Triplasis purpurea P14238 61 1.00 Gouinia paraguayensis P11526 100 Vaseyochloa multinervosa Sw10041 1.00 1.00 Craspedorhachis rhodesiana Strohbach 5699 85 100 Farrago racemosa P23851a 1.00 Trichoneura eleusinoides Sey448 88 1.00 Leptothrium rigidum Davidse3281 Dignathia hirtella CW5251 0.68 56 0.62 68 1.00 Leptocarydion vulpiastrum Mboya565 98 1.00 Gymnopogon grandiflorus P16642 99 0.67 Bewsia biflora Dav6471 59 Lophachme digitata Smook1453 Ctenium brevispicatumFilgueiras 0.62 1.00 Trigonochloa uniflora Greenway14075 1.00 89 Trigonochloa uniflora Snow6978 92 1.00 Mosdenia phleoides Sch1542 95 0.97 Perotis hordeiformis RJS5717 85 Perotis arenacea Phillipson4117 0.87 Diplachne fusca fusca WalshNG 6558 1.00 62 Diplachne fusca P24322 0.79 Diplachne fusca muelleri Badman1282 100 Diplachne divaricatissima BrownAJ 253 50 Diplachne fusca unineura Peterson20786 Diplachne uninervia P21305 Acrachne racemosa JacksonG 1408 Acrachne racemosa StrohbachB 1308 1.00 Acrachne racemosa ZavattariE 1195 100 Acrachne racemosa ZavattariE 1196 Acrachne verticillata PappiA 5990 Dinebra coerulescens Harris 289 Dinebra chinensis Snow6909 1.00 1.00 Dinebra squarrosa Kayombo5196 0.82 97 96 Dinebra retroflexa Nd610 1.00 0.87 97 Dinebra retroflexa SnowN BurgoyneP 6950 96 84 1.00 Dinebra marquisensis Wood10145 0.92 100 Dinebra xerophila Perlman18438 68 1.00 Dinebra panicea mucronata P9546 97 1.00 Dinebra panicea brachiata P20086 96 Dinebra panicea brachiata P22185 0.53 Dinebra aquatica Soderstrom650 0.85 75 Dinebra caudata Kuchar 98 1.00 Dinebra panicoides Snow5810a 100 1.00 Dinebra viscida P22184 0.64 77 Dinebra viscida Snow6528a 1.00 Dinebra haareri P24231 100 Dinebra somalensis Faben74 991 0.90 1.00 Dinebra nealteyi Snow5806 71 100 Dinebra scabra Snow5788 0.98 Dinebra neesii Snow7380 1.00 92 1.00 1.00 Dinebra ligulata Snow7402 67 84 0.86 Dinebra southwoodii Snow7362 53 Dinebra decipiens peacockii Snow7361 1.00 Dinebra decipiens decipiens Snow7328 98 1.00 Dinebra decipiens decipiens Waterhouse5948 75 Dinebra decipiens asthenes Snow7327 Dinebra decipiens asthenes Snow7355 Coelachyrum lagopoides 1.00 Apochiton burttii P24163 93 1.00 Coelachyrum poiflorum Bur2915 1.00 Eleusine jaegeri Kindeketa762 98 0.98 100 Eleusine jaegeri P24299 81 Eleusine multiflora P24272 1.00 96 0.73 Eleusine indica P21362 17B02 0.58 84 0.59 Eleusine tristachya BeckStG 3227 58 65 Eleusine tristachya Zardini35791 Eleusine africana SeydelR 2798 1.00 Eleusine africana SeydelR 3889 97 Eleusine africana P24072 Eleusine coracana africana Kayombo424 Eleusine africana P24072 1.00 Eleusine caracana P24112 88 Eleusine coracana LiuQ 093 1.00 Eleusine coracana LiuQ 103 Eleusine indica P23802 70 Eleusine indica P24429 1.00 Eleusine indica BrownF ClebschA 628 65 Eleusine indica ChaseVH 9922 Leptochloa digitata Risler476 1.00 1.00 Chloris mollis ProosdijASJ 813 83 100 Chloris mollis SteyenmarkJA 29192 1.00 1.00 Chloris exilis AndersonWR 37049 93 94 Chloris exilis AndersonWR 37190 1.00 Leptochloa chloridiformis MyndelPedersen2662 0.64 Leptochloa chloridiformis MyndelPedersen3741 97 51 0.64 Leptochloa virgata RimachiY8359 56 Leptochloa pluriflora Swallen 10100 1.00 1.00 Leptochloa virgata Peterson15088 75 87 Leptochloa virgata Vargas2710 1.00 Leptochloa crinita Renv1987 87 Leptochloa pluriflora P15048 0.98 Austrochloris dichanthioides Astrebla lappacea 62 1.00 Astrebla pectinata 100 Astrebla elymoides Hub7976 1.00 Disakisperma yemenicum P24254 60 1.00 Disakisperma eleusine Snow6982 100 0.91 Disakisperma obtusiflorum Belsky527 92 0.78 Disakisperma obtusiflorum P24211 59 0.90 Disakisperma obtusiflorum P24198 1.00 87 Disakisperma obtusiflorum P24243 97 Disakisperma dubium P22334 17C06 Disakisperma dubium P24453 78F08 Disakisperma dubium P24554 78F03 1.00 Disakisperma dubium P24566 78F09 Disakisperma dubium P24665 77H11 Disakisperma dubium P24845 77E05 Disakisperma dubium P24905 77D01 Disakisperma dubium P8105 24G02 Schoenfeldia gracilis Pauwels7413 0.86 Afrotrichloris martinii Herlocker459 85 0.93 1.00 Schoenefeldia transiens1 Gre9781 81 Schoenefeldia transiens P24216 100 0.62 Schoenfeldia transiens2 Spjut3920 1.00 Enteropogon ramosus Saarela1663 100 Enteropogon acicularis CochraneAC 521B Enteropogon melicoides BarberCA 2607 Enteropogon monostachyus ClaytonD 5954 1.00 0.73 Enteropogon monostachyus africanus P23833 91 57 0.97 Enteropogon sechellensis P23815 64 Enteropogon macrostachyus OakesAJ 1250 1.00 1.00 Enteropogon macrostachyus SL15902 97 91 Enteropogon macrostachyus SmookL 9026 Enteropogon sechellensis FosbergFR 49814 Enteropogon dolichostachyus BlakeST 23488 1.00 Enteropogon rupestris GilletJB 13082 86 Enteropogon somalensis McKinnonAS 124 1.00 Enteropogon dolichostachyus GouldFW 13680 100 Enteropogon dolichostachyus LazaridesM 7416 1.00 0.67 Enteropogon rupestris VeseyFitzgeraldD 4973 92 Enteropogon somalensis BogdanA 4026 1.00 Enteropogon barbatus KabuyeC 575 Enteropogon macrostachyus P24197 92 Enteropogon macrostachyus P24228 Ochthochloa compressa GlovesPE485 1.00 Ochthochloa compressa RechingerKH 29410 100 Ochthochloa compressa USDA 154 57 40E5 Lintonia nutans Mwa14374 1.00 Chloris prieurii Laegaard 17061 88 Chloris prieurii Laegaard 17863 1.00 Chloris barbata 22255 91 Chloris barbata BentleyPS 102 Chloris barbata P23795 1.00 Chloris barbata Saarela1740 98 Chloris barbata Saarela1830 1.00 Chloris inflata ErnstWR 1507 100 1.00 Chloris inflata KingRM 4038 99 1.00 Chloris gayana BallezaJJ 1788 1.00 95 Chloris gayana P23997 0.91 100 Chloris gayana BattyR 125 1.00 Chloris gayanaP24253 69 97 1.00 Chloris gayana P23972 100 1.00 Chloris pilosa AllenP 19 10 1967 95 Chloris pilosa BrenanJPM 9601 1.00 Chloris orthonoton HarleyRM 16629 Chloris orthonoton PedersenTM 5817 0.89 100 0.86 Chloris pilosa P23919 62 70 Chloris virgata P24286 Chloris rufescens MaVaughR 19517 Chloris virgata P24659 Chloris virgata P24854 0.97 Chloris virgata P24911 69 Chloris virgata P24789 0.96 Chloris virgata P24954 66 Chloris virgata P21468 Chloris castilloniana GallinalH 5542 0.77 Chloris amethystea ArcherWA 8650 1.00 Chloris mossambicensis P23803 99 Chloris mossambicensis P24219 0.69 0.78 Chloris andropogonoides CoryVL 52467 Chloris andropogonoides Swallen 10091 0.98 52 Chloris andropogonoides Swallen 10232 62 Chloris cucullata CrutchfieldJ 5475 0.99 0.98 Chloris subdolichostachya TrouartJE 17 53 68 Chloris verticillata Reeder 5787 Chloris latisquamea FleetwoodIF 24 02 1959 0.80 Chloris andropogonoides P18839 53 0.80 0.99 Chloris cucullata IselyD 10647 52 71 Chloris latisquamea StanfordLR Chloris halophila LaegaardS 71450 Chloris halophila P13970 Chloris submutica P24842 Chloris submutica P24939 0.83 Chloris submutica McGregorRL 706 Chloris submutica P24560 1.00 Chloris submutica Reeder 4845 87 Chloris submutica P22393 Chloris cynodontoides GreenwoodW 226C 1.00 Chloris divaricata Swallen 10572 95 Chloris pectinata Saarela1699 Chloris cruciata CorrellDS 50431 1.00 Chloris ekmanii LiogierAH 12416 82 Chloris ekmanii LiogierAH 15884 Chloris sagrana McKeeHS 10539 Chloris pycnothrix P22278 Chloris pycnothrix P23949 Chloris pycnothrix P23974 Chloris pycnothrix P24070 Chloris pycnothrix P24172 0.78 Chloris pycnothrix P24184 Chloris pycnothrix P24262 Chloris pycnothrix P24305 Chloris pycnothrix HarleyRM 17042 Chloris texana SilveusWA 423 1.00 Chloris pycnothrix BidgoodS 3170 73 Chloris pycnothrix PedersenTM 6378 Saugetia fasciculata Enteropogon brandegeei CarterAM 2897 0.98 Enteropogon chlorideus GouldFW 12679 1.00 1.00 61 62 Enteropogon chlorideus MeyerFG 24 10 1958 94 1.00 Enteropogon chlorideus LeavenworthWC 1900 0.97 90 Enteropogon chlorideus SohnsER 384 1.00 Tetrapogon tenellus P23794 94 1.00 Tetrapogon tenellus P24244 70 Tetrapogon bidentatus BogdanA 3903 0.89 Tetrapogon bidentatus Greenway 12778 0.67 1.00 Chloris roxburghiana P23812 90 Chloris roxburghiana P24194 0.69 Chloris roxburghiana P24225 53 Tetrapogon villosus 1.00 1.00 Tetrapogon spathaceus Ash2561 67 100 0.58 51 Tetrapogon cenchriformis Greenway 8844 0.98 Tetrapogon cenchriformis BN7701 820.58 Tetrapogon cenchriformis P24192 Tetrapogon cenchriformis P24240 Daknopholis boivinii Ranaivojaona1441 1.00 89 Lepturus gasparricensis Herb9687 1.00 Lepturus radicans Gill 38 92 1.00 93 Lepturus repens P23835 Lepturus repens Whi9853 Oxychloris scariosa ForsterPI 20737 1.00 Harpochloa flax Zletsmam4015 25E07 76 0.58 0.94 Microchloa altera Brand315 64 1.00 Microchloa altera RobinsonEA6362 1.00 100 Microchloa altera Malaisse6572 81 Microchloa caffra Sm10441 1.00 Microchloa kunthii P24288 1.00 100 1.00 1.00 Microchloa abyssinica HagerupO 455 60 87 1.00 100 Microchloa kunthii P24048 97 1.00 Microchloa kunthii P24255 100 Microchloa kunthii P24716 1.00 Brachyachne fulva Rob1122 100 1.00 Brachyachne chrysolepis Bog7075 100 1.00 Brachyachne obtusiflora Schweinfurth 2150 87 Brachyachne patentiflora SL16295 Eustachys tenera StrongClemensM 10564 1.00 Eustachys paspaloides Nd741 99 0.97 Eustachys calvescens RojasT 6633 Eustachys distichophylla HarrisJP 30 04 1951 0.74 80 Eustachys floridana KralR 44355 1.00 Eustachys glauca Swallen 8221 100 Eustachys neglecta ChaseA 4499 Eustachys petraea St3124 Eustachys capensis PedersenTM 6121 0.54 Eustachys caribaea Renvoize 4022 570.70 Eustachys calvescens ChaseA 10971 59 Eustachys capensis PedersenTM 6307 1.00 Chrysochloa orientalis 94 1.00 100 1.00 Chrysochloa annua 100 Chrysochloa hindsii Chloris canterae PedersenTM 9597 Chloris canterae SmithJE 01 Chloris canterae SchininiA 5321 Chloris gaugate AcostaSolisM 12667 1.00 Chloris grandiflora Izaguirre 2642 98 Chloris elata Swallen 14461 0.53 53 Chloris elataWebsterGL 10789 1.00 1.00 Chloris elata BeckStG 4454 93 95 Chloris elata SpiaggiL 40 0.78 Chloris ciliata LundellCL 14872 65 0.95 Chloris ciliata P24622 80 Chloris ciliataP24632 Cynodon maritimus Hov10214 Cynodon maritimus vaginaeflorus MartindaleIC 0.96 Cynodon erectus AsplundE 10905 72 Cynodon nitidus MacbrideJF 524 1.00 Cynodon hirsutus Sm6616 94 1.00 Cynodon incompletus OakesAJ 663 95 Cynodon incompletus Russell 2408 1.00 Brachyachne ambigua LeachGJ 3996 98 1.00 0.55 Brachyachne convergens Ad851 Brachyachne tenella Laz4281 94 0.510.66 58 1.00 Brachyachne ciliaris MullenAM apr1906 97 Brachyachne ciliaris LatzPK 14224 Cynodon radiatus Laegaard21611 0.98 Cynodon plectostachyusTro11610 54 1.00 0.64 Cynodon aethiopicus Vesey6475 54 0.61 Cynodon aethiopicus P24268 87 0.90 Cynodon aethiopicus P24318 61 Cynodon nlemfuensis P23791 Cynodon aethiopicus Smook4140 0.86 Cynodon nlemfuensis HansenB 13002 66 Cynodon nlemfuensis Laegaard 19876 1.00 Cynodon nlemfluensis LaegaardS 19845 87 Cynodon nlemfluensis Lobo2099 Cynodon nlemfluensis P24113 Cynodon nlemfluensis OakesAJ 1602 1.00 Cynodon nlemfluensis P24058 75 Cynodon nlemfluensis P24207 Cynodon affinis CabreraAL 5149 Cynodon affinis CollinsRW 63 1.00 Cynodon aristiglumis BeetleAA 2731 78 Cynodon erectus MartinRF 1814 Cynodon mucronatus FosbergFR 273 0.57 Cynodon mucronatus HermannFJ 8502 0.76 Cynodon transvaalensis Laegaard20560 72 Cynodon transvaalensis Smook6710 0.76 Cynodon affinis BoothWE 1251 56 Cynodon aristulatus EglerFE 42 314 Cynodon aristulatus FassettNC 26000 0.62 Cynodon aristulatus KillipEP 31600 56 Cynodon dactylon P24626 Cynodon dactylon pilosus MarshEG 141 Cynodon dactylon Saarela1606 0.98 Cynodon dactylon longiglumis DunkleMB 8366 86 Cynodon dactylon longiglumis DunkleMB 8652 Cynodon dactylon P19701