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852 POACEAE/BRACHYELYTRUM<br />

(e.g., Stephenson ined.) treating all three taxa as parts <strong>of</strong> a single species and o<strong>the</strong>rs considering<br />

<strong>the</strong>m worthy <strong>of</strong> recognition as separate species (Stephenson 1971; Tucker 1988; Kartesz 1999).<br />

We are following a recent revision <strong>of</strong> <strong>the</strong> genus (Saarela et al. 2003), based on morphological<br />

and molecular data, in recognizing <strong>the</strong> variation at <strong>the</strong> specific level.<br />

The disjunct e Asia–e North America distribution pattern exhibited by this and o<strong>the</strong>r genera has complex<br />

origins and is <strong>of</strong> interest <strong>to</strong> plant geographers. In <strong>the</strong> geologic past, dispersal between <strong>the</strong> Eurasian and<br />

North American continents was possible across both <strong>the</strong> Bering and North Atlantic land bridges, and <strong>the</strong><br />

combined area is considered a single “Holarctic” biogeographic region. The fossil record shows that many<br />

<strong>plants</strong> had extensive distributions across <strong>the</strong> Nor<strong>the</strong>rn Hemisphere—for example, temperate forests with<br />

tropical elements occurred very broadly and reached <strong>the</strong>ir maximum extension in <strong>the</strong> mid-Tertiary (<strong>the</strong><br />

Tertiary extended from 65 million years ago <strong>to</strong> 1.8 mya). This widespread flora has been referred <strong>to</strong> as <strong>the</strong><br />

Arc<strong>to</strong>-Tertiary flora, <strong>the</strong> Tertiaro-mesophytic flora, <strong>the</strong> boreotropical flora, or a mixed mesophytic forest.<br />

Geohis<strong>to</strong>rical events from <strong>the</strong> mid-Tertiary <strong>to</strong> <strong>the</strong> present have included alterations in <strong>the</strong> shapes <strong>of</strong> <strong>the</strong><br />

nor<strong>the</strong>rn land masses, fluctuations in sea levels, plate tec<strong>to</strong>nic movements, mountain building, glaciation,<br />

and o<strong>the</strong>r changes in <strong>the</strong> climate. As a result, <strong>the</strong>re have been great changes in both <strong>the</strong> composition and<br />

<strong>the</strong> disposition <strong>of</strong> <strong>the</strong> flora, and <strong>the</strong> ranges <strong>of</strong> many <strong>plants</strong> have been greatly restricted (e.g., eliminated<br />

from Europe and w North America). A significant number <strong>of</strong> <strong>plants</strong> that were once much more widespread<br />

now survive in only two areas, e North America and e Asia. Though this disjunct distribution pattern<br />

has complex and multiple origins, with similar present day distributions differing in time and manner<br />

<strong>of</strong> origin, <strong>the</strong> consensus is that <strong>the</strong> e Asia–e North America pattern is in general a relict <strong>of</strong> <strong>the</strong><br />

maximum development <strong>of</strong> Nor<strong>the</strong>rn Hemisphere temperate forests (with tropical elements) in <strong>the</strong> Tertiary,<br />

with greater survival in e Asia and e North America and higher rates <strong>of</strong> extinction in Europe and w<br />

North America. According <strong>to</strong> Wen (1999), ca. 65 genera <strong>of</strong> seed <strong>plants</strong> have this disjunct distribution. The<br />

genus Brachyelytrum is an example (o<strong>the</strong>r East TX examples include, but are not limited <strong>to</strong>, Aletris,<br />

Ampelopsis, Apios, Campsis, Carya, Diarrhena, Halesia, Hamamelis, Lindera, Ly onia, Menispermum, Nyssa,<br />

Par<strong>the</strong>nocissus, Penthorum, Phryma, Podophyllum, Sassafras, Saururus, Stewartia, Tipularia, Trachelospermum,<br />

Triosteum, and Zizania) (Li 1952; Little 1970; Graham 1972; Boufford & Spongberg 1983; Hamil<strong>to</strong>n<br />

1983; Hsü 1983; Wu 1983; Ying 1983; Tiffney 1985a, 1985b; Cox & Moore 1993; Graham 1993a, 1999;<br />

Xiang et al. 1998; Wen 1999, 2001; Dilcher 2000; Donoghue et al. 2001; Xiang & Soltis 2001).<br />

This genus has been variously thought <strong>to</strong> be in ei<strong>the</strong>r <strong>the</strong> Pooideae (Festucoideae) (e.g., Hitchcock<br />

1951) or Bambusoideae (e.g., Campbell et al. 1986; Tucker 1988), but recent evidence confirms<br />

its place in <strong>the</strong> Pooideae (Grass Phylogeny Working Group 2001). (Greek: brachys, short,<br />

and elytron, cover, husk, sheath, presumably in reference <strong>to</strong> <strong>the</strong> small glumes) (subfamily<br />

Pooideae, tribe Brachyelytreae)<br />

REFERENCES: Babel 1943; Koyama & Kawano 1964; Stephenson 1971, ined.; Macfarlane & Watson<br />

1980; Nixon et al. 1980a; Campbell 1983b; Campbell et al. 1986; Tucker 1988; Saarela et al. 2003.<br />

Brachyelytrum erectum (Schreb.) P. Beauv., (upright), BEARDED SHORT HUSK, LONG-AWNED WOOD<br />

GRASS. Perennial with short knotty rhizomes; culms 50–90(–100) cm tall, erect <strong>to</strong> ascending,<br />

usually with retrorse pubescence near <strong>the</strong> nodes; leaves cauline, 2–6; ligule a scale 1–3.5 mm<br />

long; leaf blades 8–17.5 cm long, 5–20 mm wide, flat; inflorescence a narrow panicle, <strong>of</strong>ten appearing<br />

racemose, 7–17 cm long; spikelets 8–13 mm long (excluding awns), with 1 fertile floret,<br />

disarticulating above <strong>the</strong> glumes; lower (first) glume absent or rudimentary (� 1 mm long); upper<br />

glume 1–4(–8) mm long; lemma 8–13 mm long, strongly antrorsely hispid, tapering in<strong>to</strong> an<br />

awn 12–25(–30) mm long; palea slightly shorter than lemma; rachilla prolonged beyond palea<br />

as a slender awn-like bristle 5–8 mm long; stamens 3, <strong>the</strong> an<strong>the</strong>rs ca. 5–6 mm long. Mesic forests,<br />

beech-hardwood slopes; Nacogdoches, Sabine (BRIT), San Augustine, Shelby (TOES 1993),<br />

Polk, Rusk, Tyler (Turner et al. 2003), and Jasper (J. Singhurst, pers. comm.) cos. in <strong>the</strong><br />

Pineywoods; se Canada and e U.S. from ME s <strong>to</strong> FL w <strong>to</strong> MN and TX. May–Sep. This species was<br />

first reported for TX from Nacogdoches Co. by Nixon et al. (1980a). Vegetatively and in habitat<br />

preference, it superficially resembles two o<strong>the</strong>r woodland species, Bromus pubescens and<br />

Chasmanthium latifolium; however, Brachyelytrum can be easily distinguished by its spikelets<br />

with only 1 floret (versus 4–many florets). (TOES 1993: IV) �

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