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bottlebrush grass, eastern bottle-brush grass, glumeless wlldrye

slender wheatgrass, slender wild rye, wheatgrass

Habit Plants cespitose, not rhizomatous, occasionally glaucous, particularly the spikes. Plants usually cespitose, sometimes weakly rhizomatous.
Culms

50-140 cm, usually erect, occasionally geniculate below;

nodes 4-8, exposed or concealed, glabrous.

30-150 cm, ascending to erect;

nodes usually glabrous.

Leaves

evenly distributed;

sheaths usually glabrous, occasionally pilose, often purplish;

auricles usually present, 0.5-3 mm, brown to black;

ligules 1-2(3) mm;

blades 4-16 mm wide, lax, usually deep glossy green, adaxial surfaces pilose or scabridulous.

somewhat basally concentrated;

sheaths usually glabrous, sometimes markedly retrorsely hirsute or villous;

auricles absent or to 1 mm;

ligules 0.2-0.8 mm, truncate;

blades 2-5(8) mm wide, flat to involute, usually straight and ascending, abaxial surfaces usually smooth and glabrous, sometimes hairy, adaxial surfaces usually glabrous, sometimes conspicuously hairy.

Spikes

7-20 cm long, 4-7 cm wide, more or less erect, usually with 2 spikelets per node, rarely with 3 at some nodes;

internodes (3)4-8(10) mm long, (0.1)0.2-0.3(0.4) mm thick at the thinnest sections, flexuous, usually glabrous, sometimes scabrous or hirsute, usually with green lateral bands.

4-25 cm long, 0.4-1 cm wide, erect, with 1 spikelet at all or most nodes;

internodes (4)7-9(12) mm, edges scabrous, both surfaces smooth and glabrous.

Spikelets

10-18 mm, strongly divergent to patent at maturity, with (1)2-4(6) florets, lowest florets functional;

disarticulation above the glumes, beneath each floret.

9-17(20) mm long, usually at least twice as long as the internodes, 3-6 mm wide, appressed, with 3-9 florets, lowest florets functional;

rachillas glabrous or hairy, hairs to 0.3 mm;

disarticulation above the glumes, beneath each floret.

Glumes

usually vestigial, sometimes 1-3 mm long, about 0.1 mm wide, subulate, entire, with no evident veins, occasionally to 10(20) mm long including the undifferentiated awns and differing in length by more than 5 mm, 0.1-0.2 mm wide, setaceous, tapering from the base, usually glabrous, occasionally appressed-puberulent to strigose, sometimes scabrous, usually straight, rarely somewhat curving, margins firm;

lemmas 8-11 mm, usually glabrous, occasionally appressed-puberulent to strigose, especially near the margins and apices, awns (12)20-40(47) mm, usually straight, rarely somewhat curving;

paleas 7-11 mm, obtuse or truncate, occasionally emarginate;

anthers 2.5-5 mm.

subequal, 5-17 mm long, from 3/4 as long as to longer than the adjacent lemmas, 1.8-2.3 mm wide, lanceolate to narrowly ovate, widest about midlength, usually green, purple at higher latitudes and elevations, flat or asymmetrically keeled for their full length, 3-7-veined, the keel vein usually scabrous, the others smooth or scabrous, only 1 vein extending to the apex, adaxial surfaces glabrous, margins hyaline or scarious, usually more or less equal, 0.2-0.5 mm wide, widest at or slightly beyond midlength, apices acute to awned, awns to 11 mm;

lemmas 6-13 mm, glabrous, usually smooth proximally, often scabridulous distally over the veins, apices acute, usually awned, awns to 40 mm, usually straight, sometimes weakly curved if shorter than 10 mm;

paleas subequal to the lemmas, keels straight or slightly outwardly curved below the apices, tapering to the apices, apices truncate, 0.15-0.3 mm wide, keel veins often extending beyond the intercostal region, sometimes forming teeth;

anthers (0.8)1.2-2.5 (3) mm.

Anthesis

mid-June to early July.

2n

= 28.

= 28.

Elymus hystrix

Elymus trachycaulus

Distribution
from FNA
AL; AR; CT; DC; DE; GA; IA; IL; IN; KS; KY; MA; MD; ME; MI; MN; MO; NC; ND; NE; NH; NJ; NM; NY; OH; OK; PA; RI; SC; SD; TN; VA; VT; WI; WV; MB; NB; NS; ON; QC
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from FNA
AK; AZ; CA; CO; CT; IA; ID; IL; IN; KS; KY; MA; MD; ME; MI; MN; MO; MT; NC; ND; NE; NH; NJ; NM; NV; NY; OH; OR; PA; RI; SD; TX; UT; VA; VT; WA; WI; WV; WY; AB; BC; MB; NB; NL; NS; NT; ON; PE; QC; SK; YT; Greenland
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Discussion

Elymus hystrix grows in dry to moist soils in open woods and thickets, especially on base-rich slopes and small stream terraces. It grows throughout most of temperate eastern North America, extending west to Manitoba and Oklahoma, but is absent from the southern portion of the coastal plain.

Plants with pubescent lemmas have been recognized as Elymus hystrix var. bigelovianus (Fernald) Bowden. These occur infrequently north of a line from South Dakota through Kentucky to New Jersey, and are often mixed with the typical variety; uniform populations are known in the northeastern United States. Plants with pubescent blades are also more prevalent to the north. Elymus hystrix hybridizes with most eastern species of Elymus. Introgression may account for the considerable variation in glume development and spikelet appression among these species. Lack of glumes may be a recessive character, with even slight glume development indicating introgression (Church 1967b). Plants with relatively well-developed, subequal glumes are presumed to be of hybrid origin. Such plants include most material from the Carolina piedmont region, where E. glabriflorus (p. 296) is the most likely source of introgression. The relatively frequent hybrids with E. virginicus (p. 298) are usually sterile, but Church (1967b) made crosses through three segregating generations. Within the ranges of E. diversiglumis (p. 316), E. svensonii (p. 314), and E. churchii (p. 314;, there appear to be frequent introgressants between these species and E. hystrix. Further east, especially in the Appalachian regions of North Carolina, Virginia, West Virginia, and Maryland (including the shale barrens and nearby), there are scattered plants of E. hystrix with curving awns and, in a few cases, appressed spikelets (Campbell 2002). Whether these represent occasional variation within the E. hystrix gene pool, or whether they are outlying remnants of introgression with E. canadensis (p. 303) during a past eastward extension, is unknown.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Elymus trachycaulus grows from sea level to 3300 m, usually in open or moderately open areas, but sometimes in forests. Its range extends from the boreal forests of North America east through Canada to Greenland and south into Mexico. It also grows, as an introduction, in Asia and Europe. It exhibits considerable variability in the presence or absence of rhizomes, the length and density of the spike, awn development on the glumes and lemmas, and glume venation. The variability in these features has often been used to circumscribe infraspecific taxa, but most such taxa, even though locally distinctive, appear to intergrade. Some of the features appear to be strongly influenced by environmental factors. For instance, plants growing in forested areas of northwestern North America tend to be slightly rhizomatous, more gracile, and later-flowering that those in adjacent, more exposed areas; whether they constitute a distinct taxon or merely a forest ecotype is not clear. Plants growing at higher elevations tend to have glumes with more widely spaced veins and broader, often unequal margins, resembling E. violaceus in these respects. Whether this reflects ecotypic differentiation, hybridization with E. violaceus (p. 324), or greater genetic continuity than is suggested by their placement in different species is not clear.

Jozwik (1966) recognized four groups within Elymus trachycaulus. Group I comprised unawned or shortly awned specimens; group II a polymorphic assemblage of awned specimens; group III a rather homomorphic group of specimens with secund spikes and relatively long awns; and group IV a relatively homomorphic group of unawned, high-elevation specimens. He concluded that group II consists of hybrids and backcrosses between E. trachycaulus and other species of Triticeae. He based this conclusion on consideration of field observations, artificial hybrids, the polymorphism of the specimens, and the geographic distribution of the group. This last was similar to that of unawned specimens of E. trachycaulus, but the populations were highly scattered within the area concerned. Jozwik's group III is treated here as E. trachycaulus subsp. subsecundus. His group IV is treated here as E. violaceus.

Elymus trachycaulus is often confused with E. stebbinsii (p. 329). It differs in having shorter anthers, shorter internodes, and glumes that are sometimes awned. It may also be confused, particularly in the herbarium, with specimens of E. glaucus having solitary spikelets at all the spike nodes; it usually differs in having shorter anthers and less acuminate glumes. When, as is sometimes the case, the two species grow together, E. trachycaulus can be distinguished by its stiffer leaves. Elymus trachycaulus also resembles E. macrourus (p. 324) and E. alaskanus (p. 326), but its glumes are longer relative to the lemmas. It also has less hairy rachillas than most plants of those species.

C.L. Hitchcock et al. (1969) treated Elymus trachycaulus as a subspecies of E. caninus (see next); it differs consistently from the latter species in glumes that are glabrous on the adaxial (inner) surface, in a chromosome interchange, and in its molecular characteristics (Sun et al. 1998). It also tends to have a more erect spike.

Elymus trachycaulus has been implicated in several interspecific and intergeneric hybrids. Named interspecific hybrids (pp. 338-343) (and the other parent) are E. ×cayouetteorum (E. canadensis), E. ×palmerensis (E. sibiricus), E. ×pseudorepens (E. lanceolatus), and E. ×saundersii (E. elymoides). Hybrids with E. hystrix have been named ×Agroelymus dorei Bowden; the appropriate combination has not been made in Elymus. Named intergeneric hybrids are ×Elyhordeum macounii (p. 284) (Hordeum jubatum), ×Elyleymus jamesensis (p. 348) {Leymus mollis), and ×Elyleymus ontariensis (p. 346) (Leymus innovatus). Hybrids with Elymus elymoides, E. multisetus, and Hordeum jubatum have brittle rachises and tend to be awned. Others are harder to recognize.

(Discussion copyrighted by Flora of North America; reprinted with permission.)

Key
1. Lemma awns 17-40 mm long, longer than the lemma body, straight; spikes somewhat 1-sided
subsp. subsecundus
1. Lemmas unawned or with awns to 24 mm long, shorter or longer than the lemma body, straight or curved; spikes 2-sided.
→ 2
2. Lemma awns 9-24 mm long
Group II
2. Lemmas unawned or with awns to 9 mm long, the awns sometimes curved.
→ 3
3. Spike internodes 8-15 mm long; spikes 8-25 cm long; glumes unawned or with straight awns to 2 mm long; spikelet bases usually visible; lemmas unawned or with straight awns to 40 mm long
subsp. trachycaulus
3. Spike internodes 4-5 mm long; spikes 5-10 cm long; glumes awned, awns 1.8-4 mm long; spikelet bases usually concealed; lemmas awned, awns 2-3 mm long, slightly curved
subsp. virescens
Source FNA vol. 24, p. 316. FNA vol. 24, p. 321.
Parent taxa Poaceae > subfam. Pooideae > tribe Triticeae > Elymus Poaceae > subfam. Pooideae > tribe Triticeae > Elymus
Sibling taxa
E. alaskanus, E. albicans, E. arizonicus, E. bakeri, E. canadensis, E. caninus, E. churchii, E. ciliaris, E. curvatus, E. dahuricus, E. diversiglumis, E. elymoides, E. glabriflorus, E. glaucus, E. hirsutus, E. hoffmannii, E. interruptus, E. lanceolatus, E. macgregorii, E. macrourus, E. multisetus, E. pringlei, E. repens, E. riparius, E. scribneri, E. semicostatus, E. sibiricus, E. sierrae, E. stebbinsii, E. svensonii, E. texensis, E. trachycaulus, E. tsukushiensis, E. villosus, E. violaceus, E. virginicus, E. wawawaiensis, E. wiegandii, E. ×cayouetteorum, E. ×ebingeri, E. ×hansenii, E. ×palmerensis, E. ×pinalenoensis, E. ×pseudorepens, E. ×saundersii, E. ×yukonensis
E. alaskanus, E. albicans, E. arizonicus, E. bakeri, E. canadensis, E. caninus, E. churchii, E. ciliaris, E. curvatus, E. dahuricus, E. diversiglumis, E. elymoides, E. glabriflorus, E. glaucus, E. hirsutus, E. hoffmannii, E. hystrix, E. interruptus, E. lanceolatus, E. macgregorii, E. macrourus, E. multisetus, E. pringlei, E. repens, E. riparius, E. scribneri, E. semicostatus, E. sibiricus, E. sierrae, E. stebbinsii, E. svensonii, E. texensis, E. tsukushiensis, E. villosus, E. violaceus, E. virginicus, E. wawawaiensis, E. wiegandii, E. ×cayouetteorum, E. ×ebingeri, E. ×hansenii, E. ×palmerensis, E. ×pinalenoensis, E. ×pseudorepens, E. ×saundersii, E. ×yukonensis
Subordinate taxa
E. trachycaulus subsp. subsecundus, E. trachycaulus subsp. trachycaulus, E. trachycaulus subsp. virescens
Synonyms Hystrix patula var. bigeloviana, Hystrix patula, E. hystrix var. bigelovianus Agropyron trachycaulum
Name authority L. (Link) Gould
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