Muhlenbergia asperifolia
Poaceae subfam. chloridoideae
alkali muhly, or alkali muhly, scratch grass
10-60(100) cm, decumbent-ascending, bases somewhat compressed-keeled;
internodes glabrous, shiny below the nodes.
usually annual, sometimes becoming somewhat woody, internodes solid or hollow.
glabrous, margins hyaline;
ligules 0.2-1 mm, firm, truncate, ciliate, without lateral lobes;
blades 2-7(11) cm long, 1-2.8(4) mm wide, flat, occasionally conduplicate, smooth or scabridulous abaxially, scabridulous adaxially, margins and midveins not conspicuously thickened, greenish, apices acute, not sharp.
sometimes conspicuously distichous;
sheaths usually open;
auricles absent;
abaxial ligules usually absent, sometimes present as a line of hairs;
adaxial ligules membranous, often ciliate with cilia longer than the membranous base, sometimes not ciliate;
blades not pseudopetiolate;
mesophyll usually radiate;
adaxial palisade layer not present;
fusoid cells absent;
arm cells absent;
Kranz anatomy present;
midrib simple;
adaxial bulliform cells present;
stomatal subsidary cells dome-shaped or triangular;
bicellular microhairs present, usually with a short, wide apical cell;
papillae sometimes present.
6-21 cm long, 4-16 cm wide, broadly ovoid, open;
primary branches 3-12 cm, capillary, lower branches spreading 30-90° from the rachises, never appearing fascicled;
pedicels 3-14 mm, longer than the spikelets.
ebracteate, paniculate, racemose, or spicate (occasionally a single spikelet), if paniculate, often with spikelike branches;
disarticulation usually beneath the florets, sometimes at the base of the panicle branches.
1.2-2.1 mm, occasionally with 2 or 3 florets.
usually bisexual, usually laterally compressed, with 1-60 florets, distal florets often reduced.
equal, 0.6-1.7 mm, purplish, scabridulous, particularly on the veins, 1-veined, apices acute;
lemmas 1.2-2.1 mm, lanceolate to oblong-elliptic, somewhat plumbeous, glabrous, usually smooth, occasionally scabridulous near the apices, apices acute, unawned or mucronate, mucros to 0.3 mm;
paleas 1.2-2.1 mm, lanceolate, glabrous, acute;
anthers 1-1.3 mm, greenish-yellow to purplish at maturity.
usually 2, shorter or longer than the lemmas, sometimes exceeding the distal florets, lower or both glumes occasionally missing;
lemmas lacking uncinate hairs, sometimes awned, awns single or, if multiple, the bases not fused into a single column;
anthers 1-3;
ovaries glabrous;
styles 2, separate throughout, bases close.
0.8-1 mm, fusiform, brownish.
often with a free or loose pericarp;
hila short;
endosperm hard, without lipid;
starch grains simple or compound;
haustorial synergids absent;
embryos usually large relative to the endosperm, not waisted;
epiblasts usually present;
scutellar cleft present;
mesocotyl internode elongate;
embryonic leaf margins usually meeting, rarely overlapping, x = (7, 8,) 9, 10 (12).
= 20, 22, 28.
Muhlenbergia asperifolia
Poaceae subfam. chloridoideae
Muhlenbergia asperifolia grows in moist, often alkaline meadows, playa margins, and sandy washes, on grassy slopes, and around seeps and hot springs, at elevations of 55-3000 m. Its geographic range includes northern Mexico. Muhlenbergia asperifolia is morphologically similar to the southeastern M. torreyana, but differs in having glabrous, weakly compressed culms and more widely divergent panicle branches.
The caryopses of Muhlenbergia asperifolia are frequently infected by a smut, Tilletia asperifolia Ellis & Everhart, which produces a globose body filled with blackish-brown spores.
(Discussion copyrighted by Flora of North America; reprinted with permission.)
The subfamily Chloridoideae is most abundant in dry, tropical and subtropical regions. In the Flora region, it reaches its greatest diversity in the southwestern United States (Barkworth and Capels 2000). Almost all its members, and all those in the Flora region, have C4 photosynthesis. Most employ the NAD-ME or PCK pathways, but Pappophorum utilizes the NADP-ME pathway.
The subfamily has been recognized, with essentially the same limits as here, for some time, although reservations have been expressed concerning its monophyly (Campbell 1985; Jacobs 1987; Kellogg and Campbell 1987). More recent studies, both morphological (Van den Borre and Watson 1997, 2000) and molecular (Soreng and Davis 1998; Hilu et al. 1999; Hsaio et al. 1999; Grass Phylogeny Working Group 2001; Hilu and Alice 2001) support its recognition as a monophyletic unit. There is less agreement concerning the subfamily's closest relative, some studies pointing to the Arundinoideae (Grass Phylogeny Working Group 2001) and some to the Danthonioideae (Barker et al. 1995; Hilu and Esen 1993; Hilu and Alice 2001).
There is considerable disagreement concerning the tribal treatment within the Chloridoideae, the number of tribes recognized varying from two (Prat 1936) to eight (Gould and Shaw 1983). Hilu and Wright (1982, p. 28) concluded, on the basis of their morphological study, that "... the boundaries between most of the tribes in this subfamily are not pronounced." They noted that Savile (1979) reached the same conclusion from considering the host specificity of various pathogenic fungi.
More recent work supports Hilu and Wright's conclusion. Van den Borre and Watson (1997, 2001) recognized eight informal groups within the subfamily. Five of the groups were large, the smallest including around 133 species and the largest around 380. The other three groups, which correspond to the Orcuttieae, Pappophoreae, and subtribe Triodiinae, include 9, 42, and 54 species, respectively. The difference in size is of no concern; the fact that all three of the small groups are embedded within one of the five large groups, the Pappophoreae and Triodiinae in a group than includes Eragrostis subg. Eragrostis and the Orcuttieae in the group that includes Muhlenbergia, is disturbing. Van den Borre and Watson noted that part of the problem was that that Eragrostis, and probably some of the other large genera, are not monophyletic.
Hilu and Alice (2001) recognized four clades within the Chloridoideae. Like Van den Borre and Watson, they found the Orcuttieae and Triodiinae to be monophyletic, although their placement within the subfamily was not clear. Unlike Van den Borre and Watson, Hilu and Alice found Pappophorum, and hence the Pappophoreae, to be polyphyletic.
The treatment presented here is conservative in recognizing the Orcuttieae and Pappophoreae as distinct tribes. It departs from most other treatments in merging all other North American taxa into a single tribe, the Cynodonteae. Consensus on how the Cynodonteae sensu lato should be broken up is unlikely to be reached until the generic limits of its members have been more thoroughly examined.
(Discussion copyrighted by Flora of North America; reprinted with permission.)
1. Leaves with little or no distinction between the sheath and blade; ligules not present; plants annual, viscid | Orcuttieae |
1. Leaves clearly differentiated into sheath and blade; ligules present; plants annual or perennial, not viscid. | → 2 |
2. Lemmas 5-13-veined, all the veins extending into awns, often alternating with hyaline lobes or teeth | Pappophoreae |
2. Lemmas 1-11-veined, unawned or with 1 or 3 awns, sometimes with hyaline lobes on either side of the central awns | Cynodonteae |
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