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Research Article
Phylogeny of Muhlenbergia subg. Pseudosporobolus, including M. spatha (Poaceae, Chloridoideae, Cynodonteae, Muhlenbergiinae) now found in Zacatecas, Mexico
expand article infoPaul M. Peterson‡, Yolanda Herrera Arrieta§, Konstantin Romaschenko‡
‡ National Museum of Natural History, Smithsonian Institution, Washington, United States of America
§ Instituto Politécnico Nacional, Durango, Mexico

Corresponding author: Paul M. Peterson ( peterson@si.edu )

Academic editor: Stephen Boatwright
© 2018 Paul M. Peterson, Yolanda Herrera Arrieta, Konstantin Romaschenko.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation: Peterson PM, Arrieta YH, Romaschenko K (2018) Phylogeny of Muhlenbergia subg. Pseudosporobolus, including M. spatha (Poaceae, Chloridoideae, Cynodonteae, Muhlenbergiinae) now found in Zacatecas, Mexico. PhytoKeys 103: 83-94. https://doi.org/10.3897/phytokeys.103.26162
Open Access

Abstract

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

Resumen

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

Keywords

grasses, ITS, Mexico, Muhlenbergia , plastid DNA sequences, Schaffnerella gracilis , systematics, taxonomy

Introduction

Muhlenbergia spatha Columbus is a small (usually less than 20 cm tall) annual first collected in the mountains of San Miguelito, in the valley of San Luis Potosí by J.G. Schaffner in 1876 (Bentham 1882). There are few collections of this species (only two known numbers collected by Schaffner with at least 14 duplicates) and it was thought to be extirpated until it was rediscovered near the type locality by J.T. Columbus along the Río Potosino in the Sierra de San Miguelito in 2001 (Columbus et al. 2001). Muhlenbergia spatha can be separated from other species in the genus by its possessing small, few-branched condensed panicles that are partially included in a spatheolate sheath with each branch containing 1–5 sessile spikelets.

Bentham (1882) thought Schaffnerella gracilis (Benth.) Nash (≡ Schaffnera gracilis Benth. ≡ Muhlenbergia spatha) was related to other members of the Zoysieae (e.g. Zoysia Willd. and Aegopogon P. Beauv = Muhlenbergia Schreb.), but he also suggested the inflorescences were nearer to members of the Andropogoneae or Pappophoreae. Beal (1896), Nash (1912), and Conzatti (1988) followed Bentham and included the monotypic Schaffnerella Nash in the Zoysieae. There are no modern treatments of Schaffnerella in Mexico, although a fairly complete genus and species description is given in Nash (1912) and, later, much abbreviated in Conzatti (1988). Pilger (1956) placed Schaffnerella in the tribe Lappagineae Link, again near Zoysia and Aegopogon. Clayton and Renvoize (1986) indicated Schaffnerella was “an isolated genus apparently related to Opizia” J. Presl (= Bouteloua Lag.; Peterson et al. 2015) and placed it in the tribe Cynodonteae. Currently, the Cynodonteae includes 25 subtribes, 95 genera and 859 species (Peterson et al. 2010a, 2016, 2017; Soreng et al. 2017). Molecular DNA sequence studies support placement of Schaffnerella in the Cynodonteae, aligning the genus within the monogeneric subtribe Muhlenbergiinae Pilg. in Muhlenbergia Schreb. subg. Pseudosporobolus (Parodi) P.M. Peterson (Columbus et al. 2010; Peterson et al. 2010a, 2010b). There are at least 27 species of Muhlenbergia that align within M. subg. Pseudosporobolus.

In this paper, we report two new collections of Muhlenbergia spatha in Zacatecas, include a complete updated description and illustration and present a molecular phylogeny of species in M. subg. Pseudosporobolus using sequence data from two plastid markers (rpl32-trnL and rps16 intron) and a single nuclear marker (ITS). Our new molecular phylogeny of M. subg. Pseudosporobolus was included to characterise evolutionary relationships of M. spatha, since earlier analyses for this species were based on few markers (Columbus et al. 2010; Peterson et al. 2010b). Based on a single sequence (rpl32-trnL), M. spatha paired with M. alopecuroides (Griseb.) P.M. Peterson & Columbus in a plastid tree and it paired with M. wrightii Vasey ex J.M. Coult. in a combined (plastid and ITS) tree (Peterson et al. 2010b).

Materials and methods

Phylogenetic analyses. Detailed methods for DNA extraction, amplification, sequencing and phylogenetic analysis are given in Peterson et al. (2010a, 2010b, 2014, 2015, 2016). In brief, we estimated the phylogeny amongst members of Muhlenbergia based on the analysis of three molecular markers (nuclear ribosomal ITS 1&2; plastid rpl32-trnL and rps16 intron DNA sequences). For this study we included a sampling of species within the five subgenera of Muhlenbergia, M. ramulosa (Kunth) Swallen and the outgroups Distichlis scoparia (Nees ex Kunth) Arechav. (Cynodonteae, Monanthochloinae), Sporobolus indicus (L.) R. Br. (Zoysieae, Sporobolinae) and Willkommia sarmentosa Hack. (Cynodonteae, Traginae) [Peterson et al. 2010b, 2016]. Voucher information and GenBank numbers for all 41 samples used in the analysis are given in Table 1.

Table 1.
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Taxon voucher (collector, number and where the specimen is housed), country of origin and GenBank accession for DNA sequences of rps16 intron, rpl32-trnL and ITS regions (bold indicates new accession); a dash (–) indicates missing data.

N Taxon Voucher Country rps16 intron rpl32-trnL ITS
OUTGROUP
1 Distichlis scoparia var. erinacea (Nees ex Kunth) Arechav. Peterson 17475, Soreng & Refulio-Rodriguez (US) Argentina, Neuquen GU360477 GU359803 GU359334
2 Sporobolus indicus (L.) R. Br. Peterson 22025 & Saarela (US) Mexico, Chihuahua GU360355 GU359913 GU359209
3 Willkommia sarmentosa Hack. Schweickerdt 2181 (US) South Africa, GU360343 GU359924 GU359194
MUHLENBERGIA
4 Muhlenbergia ramulosa (Kunth) Swallen Peterson 22447 & Saarela (US) Mexico, Durango GU360406 GU359953 GU359115
subg. Bealia
5 Muhlenbergia arenicola Buckley Peterson 19947 & Lara-Contreras (US) Mexico, Coahuila GU360413 GU359960 GU359166
6 Muhlenbergia tricholepis (Torr.) Columbus Peterson 22099 & Saarela (US) Mexico, Chihuahua GU360305 GU359853 GU359278
subg. Clomena
7 Muhlenbergia durangensis Y. Herrera Peterson 13644, Knowles, Dietrich, Braxton & Gonzalez-Elizondo (US) Mexico, Durango HM143552 HM143162 HM143060
8 Muhlenbergia montana (Nutt.) Hitchc. Peterson 22234 & Saarela (US) Mexico, Sinaloa GU360417 GU359964 GU359162
subg. Muhlenbergia
9 Muhlenbergia glauca (Nees) B.D. Jacks. Peterson 21023, Saarela, Lara Contreras & Reyna Alvarez (US) Mexico, Coahuila HM143563 HM143173 HM143072
10 Muhlenbergia pereilema P.M. Peterson Peterson 22191 & Saarela (US) Mexico, Sinaloa GU360282 GU359993 GU359131
subg. Pseudosporobolus
11 Muhlenbergia alopecuroides (Griseb.) P.M. Peterson & Columbus Peterson 20960, Saarela, Lara Contreras & Reyna Alvarez (US) Mexico, GU360426 GU359976 GU359152
12 Muhlenbergia alopecuroides (Griseb.) P.M. Peterson & Columbus Peterson 22008 & Saarela (US) Mexico, Chihuahua GU360425 GU359975 GU359153
13 Muhlenbergia arenacea (Buckley) Hitchc. Peterson 10624 & Annable (US) Mexico, Coahuila GU360414 GU359961 GU359165
14 Muhlenbergia asperifolia (Nees & Meyen ex Trin.) Parodi Peterson 15452, Soreng, Finot & Judziewicz (US) Chile, Region III (Atacama) HM143539 HM143149 HM143048
15 Muhlenbergia atacamensis Parodi Peterson 19626, Soreng, Salariato, & Panizza, (US) Argentina, Jujuy GU360489 GU359879 GU359344
16 Muhlenbergia cuspidata (Torr. ex Hook.) Rydb. Hill 35331 (US) USA HM143546 HM143156 HM143055
17 Muhlenbergia decumbens Swallen Columbus 3653 (RSA) Mexico – – EF153029
18 Muhlenbergia fastigiata (J. Presl) Henrard Peterson 21512, Soreng, LaTorre & Rojas Fox (US) Peru, Ancash HM143556 HM143166 HM143064
19 Muhlenbergia implicata (Kunth) Trin. Peterson 22266, Saarela (US) Mexico, Oaxaca HM143568 HM143179 HM143077
20 Muhlenbergia jaime-hintonii P.M. Peterson & Valdés-Reyna Peterson 15841 & Valdés-Reyna (US) Mexico, Nuevo León HM143569 HM143181 HM143079
21 Muhlenbergia ligulata (E. Fourn.) Scribn. & Merr. Peterson 22416 & Saarela (US) Mexico, Durango GU360440 GU359863 GU359273
22 Muhlenbergia monandra Alegría & Rúgolo Peterson 17990 & Refulio-Rodriguez (US) Peru, Lima – – GQ397891
23 Muhlenbergia multiflora Columbus Peterson 7845 & Annable (US) USA, Colorado GU360289 GU359985 GU359138
24 Muhlenbergia palmirensis Grignon & Lægaard Peterson 9317 & Judziewicz (US) Ecuador, Chimborazo HM143586 HM143200 HM143098
25 Muhlenbergia paniculata (Nutt.) Columbus Peterson 12070 & Annable (US) USA, Colorado GU360375 GU359936 GU359201
26 Muhlenbergia phleoides (Kunth) Columbus Peterson 24452, Romaschenko & Valdés-Reyna (US) Mexico, Nuevo León – MH400231 MH400228
27 Muhlenbergia phleoides (Kunth) Columbus Peterson 24799, Romaschenko, Rodriguez Avalos, Herrera-Simoni, & Garcia Rodriguez (US) Mexico, Aguascalientes – MH400232 MH400229
28 Muhlenbergia pungens Thurb. ex A. Gray Ricketson 4642 (MO) USA, Arizona MH508106 MH508102 MH508098
29 Muhlenbergia repens (J. Presl) Hitchc. Peterson 7900 & Annable (US) USA, New Mexico HM143596 HM143212 HM143110
30 Muhlenbergia richardsonis (Trin.) Rydb. Peterson 7832 & Annable (US) USA, Colorado. HM143598 HM143214 HM143112
31 Muhlenbergia seatonii Scribn. Peterson 9946 Mexico, Puebla MH508107 MH508103 MH508099
32 Muhlenbergia spatha Columbus Schaffner 134 (US) Mexico, – GU359981 MH400230
33 Muhlenbergia subbiflora Hitchc. Peterson 21158, Saarela, Rosen & Reid (US) Mexico, Durango GU360439 GU359877 GU359318
34 Muhlenbergia tenuissima (J. Presl) Kunth Peterson 4751 & Annable Mexico, Jalisco MH508108 MH508104 MH508100
35 Muhlenbergia uniflora (Muhl.) Fernald Peterson 13212, Annable, Pizzolato, Gordon, Frett, Frick, Morrone & Griner (US) USA, New Jersey HM143616 HM143232 HM143130
36 Muhlenbergia uniflora (Muhl.) Fernald Peterson 20862 & Saarela (US) USA, New York GU360275 GU359994 GU359119
37 Muhlenbergia utilis (Torr.) Hitchc. Peterson 24869 & Romaschenko (US) Mexico, San Luis Potosí – MH508105 MH508101
38 Muhlenbergia villiflora Hitchc. Peterson 15811 & Valdés-Reyna (US) Mexico, Nuevo León HM143620 HM143236 HM143133
39 Muhlenbergia wrightii Vasey ex J.M. Coult. Peterson 20964, Saarela, Lara Contreras & Reyna Alvarez (US) Mexico, Coahuila HM143623 HM143240 HM143137
subg. Trichochloa
40 Muhlenbergia rigens (Benth.) Hitchc. Peterson 22129 & Saarela (US) Mexico, Chihuahua GU360357 GU359951 GU359117
41 Muhlenbergia longiligula Hitchc. Peterson 15224 & Cayouette (US) USA, Arizona HM143574 HM143187 HM143085

Results and discussion

Phylogeny. A total of 16 new sequences from Muhlenbergia phleoides (Kunth) Columbus, M. pungens Thurb. ex A. Gray, M. seatonii Scribn., M. spatha, M. tenuissima (J. Presl) Kunth and M. utilis (Torr.) Hitchc. are reported in GenBank (Table 1). Total aligned characters for individual regions and other parameters are noted in Table 2. In Figure 1, we combined the plastid–ITS sequences in our analysis since there was little incongruence between these data sets.

Table 2.
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Characteristics of the three regions, rpL32-trnL, rps16 intron and ITS and parameters used in Bayesian analyses indicated by Akaike Information Criterion (AIC).

rpL32-trnL rps16 intron Combined plastid data ITS Overall
Total aligned characters 996 1088 2084 761 2845
Likelihood score (-lnL) 4758.44 3429.94 9569.67
Number of substitution types 6 6 − 6 −
Model for amongst-site rate variation gamma gamma − gamma −
Substitution rates
rAC 1.1071 1.2315 − 1.5168 −
rAG 1.8768 1.2968 2.8806
rAT 0.4688 0.4669 1.9244
rCG 1.2702 1.0243 0.7054
rCT 1.4748 2.4545 5.3115
rGT 1.0000 1.0000 1.0000
Character state frequencies
fA 0.3827 0.3860 − 0.2585 −
fC 0.1189 0.1105 0.1967
fG 0.1241 0.1772 0.2539
fT 0.3740 0.3261 0.2907
Proportion of invariable sites 0.1608 0.3844 − 0.3790 −
Substitution model TVM+G TVM+G − GTR+I+G −
Gamma shape parameter (α) 0.9290 0.9303 − 0.7988 −
Figure 1. 

Maximum-likelihood tree inferred from combined plastid (rpl32-trnL, rps16 intron) and ITS sequences. Thick branches indicate posterior probabilities of 1; numbers above branches are posterior probabilities less than 1 but greater than 0.50; Scale bar: 2%.

The maximum-likelihood tree from the combined analysis of ITS and two plastid regions (rpL32-trnL and rps16 intron) is well resolved, with strong support for the monophyly of Muhlenbergia, including M. ramulosa and five subgenera, Bealia, Clomena, Muhlenbergia, Pseudosporobolus and Trichochloa [Fig. 1; posterior probability (PP) = 1 for thick branches]. Within Muhlenbergia subg. Pseudosporobolus, M. spatha is strongly supported (PP = 1) as sister to M. phleoides and M. alopecuroides. Muhlenbergia decumbens Swallen, M. monandra Alegría & Rúgolo, M. pungens, M. seatonii and M. tenuissima are newly reported as occurring in M. subg. Pseudosporobolus, bringing the total to 27 species for this subgenus.

It is interesting but not surprising that Muhlenbergia phleoides, a close relative of M. spatha, was found as an associated species at both collection sites in Zacatecas (Rosales 3490 & Herrera Arrieta; Peterson 25544 & Herrera Arrieta). Muhlenbergia spatha, M. alopecuroides, M. phleoides and M. phalaroides Kunth (a presumed member of this clade, not yet sampled and not included in our tree) all have plumbeous-mottled spikelets that disarticulate as a unit (below the glumes) leaving a small cuplike tip and glumes with 2–5 recurved awns (Reeder 2003; Peterson et al. in prep.). Plumbeous-mottled spikelets and well-developed sclerenchyma girders in the primary vascular bundles are additional traits shared by members of M. subg. Pseudosporobolus (Watson and Dallwitz 1992; Peterson and Herrera Arrieta 2001; Peterson et al. 2010b).

Taxonomy

Muhlenbergia spatha Columbus, Aliso 28: 66. 2010, non Muhlenbergia gracilis (Kunth) Trin. (1824).

Fig. 2A–D

Schaffnera gracilis Benth., Hooker’s Icon. Pl. 14: 59, t. 1378. 1882. Schaffnerella gracilis (Benth.) Nash, N. Amer. Fl. 17(2): 141. 1912, non Schaffneria Fée ex T. Moore (1857). Muhlenbergia columbi P.M. Peterson, Amer. J. Bot. 97(9): 1543. 2010, nom. illeg. superfl. Type: Mexico, San Luis Potosí, mountains of San Miguelita, Aug 1876, J.G. Schaffner 1070 (holotype: K-000309066 [image!]; isotypes PH-00022592 [image!], US-397116!, YU-063983 [image!]).

Description

Delicate annuals, loosely caespitose. Culms 5–20(–30) cm tall, erect, geniculate below; nodes scaberulous, branching at lower and middle nodes; internodes 2.0–4.5(–8.0) cm long, strongly 4–6 ribbed. Leaves caulescent and basal; ligules 0.8–1.8 mm long, membranous, decurrent, apex obtuse, minutely erose; sheaths 0.8–1.5 cm long, much shorter than the internodes, oblong, open, chartaceous, strongly ribbed with 7–9-veins, sometimes keeled, margins hyaline; blades 0.5–4 cm long, 0.5–1.5 mm wide, flat or folded, adaxially scattered pubescent near base, the hairs antrorse leaning, apex obtuse. Inflorescences compound, fasciculate, composed of terminal and axillary condensed panicles, these branched near the base, the basal-most branch usually with a sterile floret at the base consisting of two scales, the entire panicle partially included in a spatheolate sheath; sterile florets 2–4 mm long, 0.8–1 mm wide, linear-apiculate, flat, hyaline, 1-veined; racemose branches each bearing 1–5 fertile sessile spikelets, the spikelets separated by 1–4 mm on each branch; rachis angular, 3 or 4-ribbed. Spikelets 4.4–6 mm long, lanceolate, laterally compressed, solitary, composed of one fertile floret, plumbeous-mottled; rachilla not extended; callus short, blunt, pubescent, located just below the glumes where disarticulation occurs leaving a small cuplike tip; glumes dimorphic; lower glumes absent or obscure; upper glumes 3.5–5(–6) mm long, about as long as the lemma, oblong, chartaceous, firmer than fertile lemma (excluding the awns), 7–9-veined, lateral veins ribbed, pubescent along the veins on lower ½, apex deeply bifid and 3(5)–awned, the awns 5–7 mm long, scabrous, recurved, arising between the bifid apex; lemmas (4–)4.8–6 mm long, lanceolate, membranous, 3-veined, keeled, midvein scaberulous, lateral veins ribbed, apex acute to acuminate, minutely bifid, 1-awned, the awn 3–5 mm long arising from between the teeth; paleas 3.7–5.5 mm long, shorter than the lemmas, oblong, hyaline to membranous, tightly involute, 2-veined, apex obtuse, unawned; stamens 3; anthers 2–2.5 mm long, yellowish; ovary glabrous; caryopses 1.8–2 mm long, 0.5 mm wide, narrowly fusiform, straw coloured.

Figure 2. 

Muhlenbergia spatha: A Habit B Panicle partially enclosed by the leaf sheath C Basal panicle branch with a lower sterile floret and small cuplike structures (callus remains) where the fertile spikelets were inserted D Spikelet, side view E Upper glume showing three recurved awns, dorsal view. (Peterson 25544 & Herrera Arrieta, CIIDIR).

Distribution

The species is known only from the type locality in San Luis Potosí and Municipio Villanueva, Zacatecas.

Ecology

Muhlenbergia spatha was found by the authors growing on flat table rock in open areas near arroyos associated with Bouteloua hirsuta Lag., B. curtipendula (Michx.) Torr., Schizachyrium sanguineum (Retz.) Alston, Quercus, Juniperus, Muhlenbergia implicata (Kunth) Trin., M. phleoides, M. rigida (Kunth) Kunth, Chloris submutica Kunth, Digitaria ternata (A. Rich.) Stapf, Microchloa kunthii Desv., Aristida adscensionis L., A. divaricata Humb. & Bonpl. ex Willd., Enneapogon desvauxii P. Beauv., Piptochaetium fimbriatum (Kunth) Hitchc., Eragrostis intermedia Hitchc., E. pectincea (Michx.) Nees and Eleusine multiflora Hochst. ex A. Rich.

Habitat

At the southern edge of the Sierra Madre Occidental Range in the Sierra Fría de Aguascalientes, we recently found Muhlenbergia spatha in two localities south of the city of Zacatecas in a corridor located east of Villanueva: 1) 17.4 km east of Villanueva on flat table rock just above Arroyo “El Muerto” in open areas and 2) about 1 km northeast of the small village Palomas Viejas along Arroyo Juan Manuel on natural grasslands near cultivated fields. These two sites are approximately 204 km W (air distance) from the type locality southwest of San Luis Potosí in the Sierra de San Miguelito along the Río Potosino (Columbus et al. 2001). According to Instituto Nacional de Estadística, Geografía e Informática [INEGI] (2003, 2005), the Sierra San Miguelito is placed in the Mesa del Centro Province in the Sierra Madre Oriental. Near Villanueva, Zacatecas, the soils are Haplic Phaeozem (forming sodium carbonate, Na2CO3) over extrusive igneous rock (acidic) in a semi-dry climate with an annual mean temperature of 16°C and an annual mean precipitation of 60 cm (INEGI 2008a, 2008b). In the Sierra de San Miguelito along the Rio Potosino, the soils are Calcic Regosol (forming calcium carbonate, CaCO3) of medium texture over extrusive igneous rock (acidic) in a semi-dry climate with an annual mean temperature of 16 °C and an annual mean precipitation of 40 cm (INEGI 2008a, 2008b). Without field verification, it is uncertain whether the soil differences among these sites are significant, but we are reasonably certain that all are alkaline with an elevated pH.

Conservation status

The species is rare in Mexico and is known from only three recent collections. Since it is a diminutive, short-lived annual, the species is easily overlooked and the main concern seems to be loss of habitat via human impact, i.e. agriculture, dam and road construction.

Specimens examined

Mexico. San Luis Potosí: 1876, J.G. Schaffner 134 (GOET-006918 [image!], US-825687); Sierra de San Miguelito, Río Potosino, 22°04'55"N, 101°03'51"W, 1980 m, 2 Oct 2001, J.T. Columbus 4040 (RSA [image! in Columbus et al. 2001]). Zacatecas: Mpio. Villanueva, Arroyo Juan Manuel ± 1 km NE of Palomas Viejas near Villanueva, 22°24'36"N, 102°43'01"W, 2112 m, 22 Oct 2006, O. Rosales 3490 & Y. Herrera Arrieta (CIIDIR); Mpio Villanueva, 10.8 mi [17.4 km] E of Villanueva, just above arroyo “El Muerto”, 22°22'47.1"N, 102°43'31.5"W, 2083 m, 2 Oct 2015, P.M. Peterson 25544 & Y. Herrera Arrieta (CIIDIR, US).

Comments

The hand written script (verified by J. Rzedowski, per. comm., also see Rzedowski 1959) on the label of the holotype (K) says: “Müehlenbergia gracilis mihi”, indicating that Jose Guillermo Schaffner thought he had collected a new species of Muhlenbergia. Bentham (1882) agreed with Schaffner that it was a new species but thought it had enough unique morphological features (spatheolate sheathed panicles, sessile spikelets and 3(5)-awned upper glumes) to warrant description of a new genus.

Acknowledgements

We thank the Smithsonian Institution’s Restricted Endowment Fund, the Scholarly Studies Program, Research Opportunities, Atherton Seidell Foundation, Biodiversity Surveys and Inventories Program, Small Grants; the National Geographic Society for Research and Exploration (Grant No. 8848-10, 8087-06), Instituto Politécnico Nacional (Grants: SIP-20160729, SIP-20180988), Comisión Nacional para el Conocimiento y Uso de la Biodiversidad (Grants: V024, EE014); Flor Isela Rentana Rentaría and Alice R. Tangerini for preparing the plant illustration; Robert J. Soreng for discussions pertinent to the manuscript; and Neil Snow and Jeffery M. Saarela for suggesting improvements to the manuscript.

References

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