SPECIES: Elymus elymoides

INTRODUCTORY

SPECIES: Elymus elymoides

AUTHORSHIP AND CITATION:
Simonin, Kevin A. 2001. Elymus elymoides. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [].



ABBREVIATION:
ELYELY

SYNONYMS:
Sitanion hystrix (Nutt.) J. G. Smith [51,82,115,123,132]

NRCS PLANT CODE [193]:
ELEL5

COMMON NAMES:
bottlebrush squirreltail

TAXONOMY:
The currently accepted scientific name of bottlebrush squirreltail is Elymus elymoides (Raf.) Swezey [60,87,113] (Poaceae). Barkworth and Dewey [12] realigned Sitanion hystrix (Nuttall) J. G. Smith in the Elymus genus as Elymus elymoides. Realignment of the Elymus genus is based upon morphological and genomic characters [12,56].

The following subspecies are currently recognized: Elymus elymoides ssp. brevifolius, E. e. ssp. californicus, E. e. ssp. elymoides, and E. e. ssp. hordeoides [93]. Bottlebrush squirreltail hybridizes frequently with other Elymus species and infrequently with Hordeum species [200]. Bottlebrush squirreltail also hybridizes with saline wildrye (Leymus salinus) [106].

LIFE FORM:
Graminoid

FEDERAL LEGAL STATUS:
No special status

OTHER STATUS:
No entry


DISTRIBUTION AND OCCURRENCE

SPECIES: Elymus elymoides

GENERAL DISTRIBUTION:
Bottlebrush squirreltail is found from British Columbia to Saskatchewan, south throughout the western and central United States and into Mexico [200]. The PLANTS database provides a distributional map for bottlebrush squirreltail. Elymus elymoides ssp. brevifolius occurs in the San Bernardino Mountains, Peninsular Ranges, Modoc Plateau, and Mojave Desert of California to Oregon, the Great Plains and south to northern Mexico. Elymus elymoides ssp. californicus is found in the Klamath Range, Cascade Range, Sierra Nevada, San Gabriel Mountains, San Bernardino Mountains, east Sierra Nevada of California to Washington, Montana and Utah. Elymus elymoides ssp. elymoides is found in the Transverse Ranges, San Jacinto Mountains, and Great Basin floristic Province from California to Washington, Wyoming and Colorado. Elymus elymoides ssp. hordeoides occurs in Klamath Range from California to Washington and Nevada.

ECOSYSTEMS [80]:
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES23 Fir-spruce
FRES26 Lodgepole pine
FRES28 Western hardwoods
FRES29 Sagebrush
FRES30 Desert shrub
FRES32 Texas savanna
FRES33 Southwestern shrubsteppe
FRES34 Chaparral-mountain shrub
FRES35 Pinyon-juniper
FRES36 Mountain grasslands
FRES38 Plains grasslands
FRES40 Desert grasslands
FRES44 Alpine

STATES:
AZ CA CO ID
IL KS KY MI
MT NE NV NM
ND OK OR SD
TX UT WA WY

AB BC MB SK

MEXICO

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BLM PHYSIOGRAPHIC REGIONS [22]:
3 Southern Pacific Border
4 Sierra Mountains
5 Columbia Plateau
6 Upper Basin and Range
7 Lower Basin and Range
8 Northern Rocky Mountains
9 Middle Rocky Mountains
10 Wyoming Basin
11 Southern Rocky Mountains
12 Colorado Plateau
13 Rocky Mountain Piedmont
14 Great Plains
15 Black Hills Uplift

KUCHLER [121] PLANT ASSOCIATIONS:
K005 Mixed conifer forest
K007 Red fir forest
K008 Lodgepole pine-subalpine forest
K010 Ponderosa shrub forest
K011 Western ponderosa forest
K012 Douglas-fir forest
K016 Eastern ponderosa forest
K017 Black Hills pine forest
K018 Pine-Douglas-fir forest
K019 Arizona pine forest
K020 Spruce-fir-Douglas-fir forest
K021 Southwestern spruce-fir forest
K022 Great Basin pine forest
K023 Juniper-pinyon woodland
K024 Juniper steppe woodland
K026 Oregon oakwoods
K030 California oakwoods
K031 Oak-juniper woodland
K032 Transition between K031 and K037
K037 Mountain-mahogany-oak scrub
K038 Great Basin sagebrush
K039 Blackbrush
K040 Saltbush-greasewood
K041 Creosotebush
K046 Desert: vegetation largely lacking
K050 Fescue-wheatgrass
K052 Alpine meadows and barren
K053 Grama-galleta steppe
K054 Grama-tobosa prairie
K055 Sagebrush steppe
K056 Wheatgrass-needlegrass shrubsteppe
K057 Galleta-threeawn shrubsteppe
K059 Trans-Pecos shrub savanna
K060 Mesquite savanna
K064 Grama-needlegrass-wheatgrass
K065 Grama-buffalo grass
K066 Wheatgrass-needlegrass
K085 Mesquite-buffalo grass
K086 Juniper-oak savanna

SAF COVER TYPES [71]:
66 Ashe juniper-redberry (Pinchot) juniper
68 Mesquite
207 Red fir
209 Bristlecone pine
210 Interior Douglas-fir
217 Aspen
218 Lodgepole pine
220 Rocky Mountain juniper
229 Pacific Douglas-fir
237 Interior ponderosa pine
238 Western juniper
239 Pinyon-juniper
241 Western live oak
242 Mesquite
243 Sierra Nevada mixed conifer
244 Pacific ponderosa pine-Douglas-fir
246 California black oak
250 Blue oak-foothills pine
256 California mixed subalpine

SRM (RANGELAND) COVER TYPES [176]:
101 Bluebunch wheatgrass
102 Idaho fescue
104 Antelope bitterbrush-bluebunch wheatgrass
105 Antelope bitterbrush-Idaho fescue
106 Bluegrass scabland
107 Western juniper/big sagebrush/bluebunch wheatgrass
108 Alpine Idaho fescue
109 Ponderosa pine shrubland
110 Ponderosa pine-grassland
201 Blue oak woodland
207 Scrub oak mixed chaparral
210 Bitterbrush
211 Creosotebush scrub
212 Blackbush
301 Bluebunch wheatgrass-blue grama
310 Needle-and-thread-blue grama
314 Big sagebrush-bluebunch wheatgrass
315 Big sagebrush-Idaho fescue
318 Bitterbrush-Idaho fescue
320 Black sagebrush-bluebunch wheatgrass
321 Black sagebrush-Idaho fescue
322 Curlleaf mountain-mahogany-bluebunch wheatgrass
401 Basin big sagebrush
403 Wyoming big sagebrush
405 Black sagebrush
406 Low sagebrush
407 Stiff sagebrush
408 Other sagebrush types
409 Tall forb
410 Alpine rangeland
412 Juniper-pinyon woodland
413 Gambel oak
414 Salt desert shrub
415 Curlleaf mountain-mahogany
416 True mountain-mahogany
417 Littleleaf mountain-mahogany
501 Saltbush-greasewood
502 Grama-galleta
503 Arizona chaparral
504 Juniper-pinyon pine woodland
509 Transition between oak-juniper woodland and mahogany-oak association
607 Wheatgrass-needlegrass
611 Blue grama-buffalo grass
612 Sagebrush-grass
614 Crested wheatgrass
615 Wheatgrass-saltgrass-grama
704 Blue grama-western wheatgrass
712 Galleta-alkali sacaton
713 Grama-muhly-threeawn
714 Grama-bluestem
715 Grama-buffalo grass
718 Mesquite-grama
727 Mesquite-buffalo grass
735 Sideoats grama-sumac-juniper

HABITAT TYPES AND PLANT COMMUNITIES:
Bottlebrush squirreltail is a common component of sagebrush (Artemisia spp.)/grass communities of the Intermountain shrubsteppe [109,212].

Within rangelands of Utah, Nevada, southern Idaho, and western Wyoming, bottlebrush squirreltail commonly grows under and adjacent to shadscale (Atriplex confertifolia), black greasewood (Sarcobatus vermiculatus), and green rabbitbrush (Chrysothamnus viscidiflorus) [161]. Bottlebrush squirreltail is a common component of pinyon-juniper (Pinus spp.-Juniperus spp.) communities of the Great Basin. It occurs mostly within the mountain ranges of Nevada and Utah, and to a lesser extent in California and Idaho [191].

Arizona:
Bottlebrush squirreltail occurs in northern desert shrub communities [137] and ponderosa pine (Pinus ponderosa) forests [39,137]. Within aspen (Populus tremuloides)-bunchgrass communities of northern Arizona, bottlebrush squirreltail commonly occurs with Arizona fescue (Festuca arizonica), mountain muhly (Muhlenbergia montana), western yarrow (Achillea millefolium), lupine (Lupinus spp.), fleabane (Erigeron spp.) and American vetch (Vicia americana) [86]. Bottlebrush squirreltail is occasionally found in openings and under shrub canopies within shrub live oak (Quercus turbinella)-mixed shrub communities [173].

California:
Bottlebrush squirreltail is native to California's central valley and is commonly associated with purple tussock grass (Nassella pulchra), nodding tussock grass (N. cernua), smallflower tussockgrass (N. lepida), and shooting star (Dodecatheon spp.) [14]. It is a minor component of blue oak (Quercus douglasii) [28,126] and interior live oak (Q. wislizenii) habitats [126].

Within sagebrush scrub of the White Mountains, prairie Junegrass (Koeleria macrantha), muhly (Muhlenbergia spp.) and timberline bluegrass (Poa glauca spp. rupicola) are common associates [130]. Within alluvial fans of desert shrub communities, bottlebrush squirreltail is commonly associated with Indian ricegrass (Achnatherum hymenoides) and desert needlegrass (A. speciosum). Shrub associates of bottlebrush squirreltail include California jointfir (Ephedra californica), goldenfleece (Ericameria arborescens), white burrobrush (Hymenoclea salsola), and purple sage (Salvia dorrii) [135].

Bottlebrush squirreltail occasionally occurs in alpine flora of the Sierra Nevada eastern slope [40]. It is also an occasional understory species of California red fir (Abies magnifica) forests in the Sierra Nevada [155].

Colorado:
Bottlebrush squirreltail is a member of north-central Colorado's short-grass prairie [45,57,175]. Within the short-grass prairie, bottlebrush squirreltail is commonly associated with western wheatgrass (Pascopyrum smithii), purple threeawn (Aristida purpurea), sideoats grama (Bouteloua curtipendula), blue grama (Bouteloua gracilis), buffalo grass (Buchloe dactyloides), and needle-and-thread grass (Hesperostipa comata) [57]. It occurs in blue grama ranges of Colorado along with sun sedge (Carex heliophila), sand dropseed (Sporobolus cryptandrus), and purple threeawn [139].

Within Colorado pinyon-Utah juniper (Pinus edulis-Juniperus osteosperma) habitats of Colorado, bottlebrush squirreltail is associated with Gambel oak (Quercus gambelii), Utah serviceberry (Amelanchier utahensis), true mountain-mahogany (Cercocarpus montanus), fendlerbush (Fendlera rupicola), banana yucca (Yucca baccata), and antelope bitterbrush (Purshia tridentata). Grass associates include cheatgrass (Bromus tectorum), Indian ricegrass, mutton grass (Poa fendleriana), and brome grasses (Bromus spp.) [67].

Bottlebrush squirreltail is an occasional associate of Rocky Mountain bristlecone pine (Pinus aristata) [162].

Idaho:
Bottlebrush squirreltail is a dominant species in shadscale communities of south-central Idaho [174], along with Indian ricegrass [189].

Montana:
Bottlebrush squirreltail generally occurs as scattered plants on rangelands [150]. In the eastern plains of Montana, bottlebrush squirreltail is a dominant species of saline rangelands in association with alkali sacaton (Sporobolus airoides), western wheatgrass (Pascopyrum smithii), thickspike wheatgrass (Elymus lanceolatus), inland saltgrass (Distichlis stricta), Sandberg bluegrass (Poa secunda), and basin wildrye (Leymus cinereus). Shrub associates include black greasewood and Nuttall's saltbush (Atriplex nuttallii).

Nevada:
Bottlebrush squirreltail occurs in gray low sagebrush (Artemisia arbuscula ssp. arbuscula) and big sagebrush (A. tridentata) communities. Principal grass associates include cheatgrass [24], Sandberg bluegrass [24,25], bluebunch wheatgrass (Pseudoroegneria spicata), and Thurber needlegrass (Achnatherum thurberianum). Common shrub associates include green rabbitbrush and gray horsebrush (Tetradymia canescens). Forb associates include bird's-beak (Cordylanthus ramosus), mourning milkvetch (Astragalus atratus), woollypod milkvetch (Astragalus purshii), desert yellow fleabane (Erigeron linearis), lava aster (Lonactis alpina), Heerman's buckwheat (Eriogonum heermanii), tail cup lupine (Lupinus caudatus) and phlox (Phlox longifolia) [25].

In northeastern Nevada bottlebrush squirreltail is commonly found with black sagebrush (Artemisia nova) [107,219], shadscale, Nevada ephedra (Ephedra nevadensis), Sandberg bluegrass and Indian ricegrass [107].

Bottlebrush squirreltail also occurs in shadscale communities [24,145]. Common shrub associates include green molly (Kochia americana), winterfat (Krascheninnikovia lanata), budsage (Artemisia spinescens) and spiny hopsage (Grayia spinosa ) [145]. Common grass associates are cheatgrass, Indian ricegrass [24] and galleta (Pleuraphis jamesii) [145]. Several common forb associates are salt lover (Halogeton glomeratus), Adonis blazingstar (Mentzelia multiflora) and gooseberryleaf globemallow (Sphaeralcea grossulariifolia). Bottlebrush squirreltail is also common to juniper (Juniperus spp.) and greasewood communities [24].

Bottlebrush squirreltail is found in Wyoming big sagebrush (Artemisia tridentata ssp. wyomingensis) rangelands of Nevada [23].

Oregon:
Eastern Oregon grass associates of bottlebrush squirreltail include bluebunch wheatgrass, prairie Junegrass [31], Idaho fescue (Festuca idahoensis), Thurber needlegrass [31,59], Sandberg bluegrass and cheatgrass. Common forbs include Hood's phlox (Phlox hoodii) and maiden blue eyed Mary (Collinsia parviflora) [59]. Bottlebrush squirreltail occurs in lodgepole pine stands in the Cascades of Oregon [61], along with western needlegrass (Achnatherum occidentale) and Ross' sedge (Carex rossii) [65].

Texas:
In western Texas bottlebrush squirreltail occurs in Pinchot juniper (Juniperus pinchotii) rangelands with sideoats grama, buffalo grass, slim tridens (Tridens muticus), awnless bushsunflower (Simsia calva) and plains fleabane (Erigeron modestus) [143].

Utah:
Bottlebrush squirreltail is common in salt-desert shrub ranges along with the shrubs shadscale (Atriplex confertifolia), mat saltbush (A. corrugata), fourwing saltbush (A. canescens), valley saltbush (A. cunneata), greasewood, winterfat, spiny hopsage, budsage, black sagebrush and green rabbitbrush. Common grasses include Indian ricegrass, Sandberg bluegrass, galleta, alkali sacaton, sand dropseed, and blue grama. Bottlebrush squirreltail is also found in pinyon-juniper woodlands [29], ponderosa pine savannas [148] along with dry Douglas-fir (Pseudotsuga menziesii) and mixed conifer habitats [29].

Wyoming:
Bottlebrush squirreltail commonly occurs in big sagebrush steppe along with aspen, big sagebrush, mountain snowberry (Symphoricarpos oreophilus), Utah serviceberry, rose (Rosa spp.), Scouler's willow (Salix scouleriana), and Oregon-grape (Mahonia repens). Common forb associates include rosy pussytoes (Antennaria microphylla), arrowleaf buckwheat (Eriogonum compositum), pineywoods geranium (Geranium caespitosum) and northern bedstraw (Galium boreale). Graminoid associates include needle-and-thread grass, Columbia needlegrass (Achnatherum nelsonii), and elk sedge (Carex geyeri) [36].

Classifications describing plant communities in which bottlebrush squirreltail is a dominant species are as follows:

Idaho [97]
California [218] Colorado [119]
New Mexico [94,146]
Nevada [24,25,107,125,192,221]
Oregon [59,101,197]
Utah [151,203]
Wyoming [186]


MANAGEMENT CONSIDERATIONS

SPECIES: Elymus elymoides

IMPORTANCE TO LIVESTOCK AND WILDLIFE:
Bottlebrush squirreltail is a dietary component of several wildlife species. It is a minor component of bison and cattle summer diets within sagebrush rangelands of southern Utah [195]. Although of little importance, bottlebrush squirreltail may provide forage for mule deer [122,124]. Pronghorn of western Utah feed upon bottlebrush squirreltail [16]. Townsend's ground squirrels [211], Nuttall's cottontails [111,127], and black-tailed jackrabbits [5,72,112,127] all feed upon bottlebrush squirreltail.

In southeastern Oregon salt desert-shrub ranges, bottlebrush squirreltail is an important component of domestic livestock seasonal diets. Winter months show greatest use [83,140].

PALATABILITY:
Bottlebrush squirreltail is a very palatable winter forage for domestic sheep of Intermountain ranges. Domestic sheep relish the green foliage [104]. Overall, bottlebrush squirreltail is considered moderately palatable to livestock.

When present, the long sharp awns of bottlebrush squirreltail greatly reduce its palatability [150]. Mature awns may penetrate flesh around the mouth of grazing animals, producing inflammation [51,115]. Eye and ear injury may also occur [51].

NUTRITIONAL VALUE:
Clary [44] compared chemical constituents (%) of bottlebrush squirreltail within open and timbered ponderosa pine overstories in Arizona. Greater digestibility and significantly (p<0.05) higher crude protein were found in open versus timbered overstories:

Open Timbered
Crude protein (%) 16.0 9.7
Phosphorus (%) 0.25 0.26
Ash (%) 12.3 13.7
Digestibility (%) 66.7 61.0

Bottlebrush squirreltail nutrient levels fluctuate throughout the growing season. Levels of S, P, and K usually drop from March to October. Amounts of Mg and Ca stay relatively the same with high points in spring, late summer, and early fall [152]. Overall, bottlebrush squirreltail is a poor source of phosphorus, carotene, and digestible protein, but a good source of energy [48]. The average chemical composition (%) of bottlebrush squirreltail in Great Basin desert ranges is summarized below [47]:

Composition (%)
Ether Extract 2.6
Total protein 4.5
Ash 17.1
Lignin 8.7
Cellulose 37.5
Other Carbohydrates 29.6
Phosphorus 0.07
Gross energy 1730 (kcal/lb)
Carotene 0.5 (mg/lb)

COVER VALUE:
The degree of environmental protection provided by bottlebrush squirreltail for wildlife species is as follows [58]:

  Utah Wyoming
Pronghorn Poor Poor
Elk Poor Poor
Mule deer Poor Poor
White-tailed deer ---- Poor
Small mammals Good Good
Small nongame birds Fair Good
Upland game birds Fair Fair
Waterfowl Poor Fair

VALUE FOR REHABILITATION OF DISTURBED SITES:
Bottlebrush squirreltail is tolerant of disturbance [133]. The Bureau of Land Management, U.S. Department of the Interior, identifies bottlebrush squirreltail as a high priority species for restoring native plant diversity in the Great Basin and the Columbia River Plateau [90]. Bottlebrush squirreltail naturally colonizes disturbed sites in Yellowstone National Park and is a component of seed mixtures used for restoration of lodgepole pine communities [129]. Brown and Amacher [34] recommend bottlebrush squirreltail for use in restoration of disturbed arid to semi-arid, desert shrub and pinyon-juniper systems. Bottlebrush squirreltail is well adapted for seeding of Wyoming, black and low sagebrush communities of the Intermountain West, receiving 9 to 13 inches (229-330 mm) annual precipitation. Bottlebrush squirreltail grows well under rabbitbrush canopies in south-central Idaho rangelands [149].

Bottlebrush squirreltail inhabits xeric sandy soils (73.9% sand, 16.8% silt, 9.2% clay, 1.3% organic matter) of a 50-year-old abandoned tailings pond from a Pb-Zn-processing mill [41], and is recommended for seed mixtures used to reclaim strip mines in southeastern Montana [64].

The large ecological amplitude of bottlebrush squirreltail lends to ecotypic differentiation. Phenological differences in growth rate, root:shoot ratios, leaf area, and overall plant size exist between subspecies of bottlebrush squirreltail. Differences are directly related to subspecies distribution [100]. Bottlebrush squirreltail seed source should be considered when implementing revegetation projects. Arredondo and others [9] observed a higher root length-to-leaf area ratio in plants grown from seed collected from different environments. Differences in phenology between individuals of different habitats are common (see: SEASONAL DEVELOPMENT within the Botanical and Ecological Characteristics section for further information).

Bottlebrush squirreltail seed is available commercially [103,104,134]. The United States Department of Agriculture (USDA), Utah Division of Wildlife Resources, in conjunction with the Intermountain Research Station, Forest Service, USDA, established bottlebrush squirreltail seed quality standards. Seed quality standards as of 1990 are summarized below [181]:

Seed unit1 Acceptable purity (%)2 Acceptable viability (%)2
spikelet with or without awns 90 85

1 Reproductive structure marketed as seed.
2 Purity (%) and germination (%) expected using seed quality testing rules in seeds of commercial quality.

Germinability of primed bottlebrush squirreltail seed significantly (p<0.05) decreases when dried and stored [89].

Competition with invasive weeds:
The persistence of bottlebrush squirreltail in areas invaded by exotic weeds is well recognized. Bottlebrush squirreltail persists in areas infested with cheatgrass [9,18,99,100,103,188], medusahead (Taeniatherum caput-medusae) [9,96,169,213,216], and Japanese brome (Bromus japonicus) [166].

Bottlebrush squirreltail naturally invades rangelands dominated by cheatgrass and medusahead [9]. However, mechanisms behind bottlebrush squirreltail's ability to occupy weed-infested areas are not completely understood. Several studies have evaluated the persistence of bottlebrush squirreltail within cheatgrass infested ranges. Beckstead [18] found recently harvested bottlebrush squirreltail seeds from mountain brush and meadow sites to possess lower levels of dormancy than cheatgrass at higher temperatures, 68/86 degrees Fahrenheit (20/30 C), whereas the opposite was true of lower temperatures, 41/59 degrees Fahrenheit (5/15 C). Bottlebrush squirreltail at lower elevations (4,100 feet (1,250 m)) have a greater probability of autumn germination than cheatgrass [2]. Established bottlebrush squirreltail plants generally initiate growth before the rosettes of cheatgrass in desert rangelands of Nevada [188]. Beckstead [18] suggests fall seeding of bottlebrush squirreltail into cheatgrass infested rangelands.

Early spring growth and ability to grow at low temperatures contribute to the persistence of bottlebrush squirreltail among cheatgrass dominated ranges [100]. Bottlebrush squirreltail seedlings have the ability to grow roots at low soil temperatures, allowing for soil penetration similar to medusahead and cheatgrass in the northern regions of the Great Basin. Root development at low temperatures promotes bottlebrush squirreltail seedling establishment and effective competition with medusahead [96].

Bottlebrush squirreltail has potential to outcompete medusahead. Management goals often concentrate on protecting bottlebrush squirreltail seedlings from livestock and rabbits, along with maintaining a natural supply of seed [169]. Hironaka and Sindelar [98] evaluated bottlebrush squirreltail growth under greenhouse conditions, when closely associated with medusahead. Bottlebrush squirreltail plants (10 plants) were observed in combination with 0, 4, 12, 36, 108, and 324 medusahead/foot2. Bottlebrush squirreltail growth was not affected by medusahead until 5 weeks old, grown under densities of 108 and 324 medusahead/foot2. Although stunted, no bottlebrush squirreltail mortality was seen at all densities tested, whereas a large amount of medusahead mortality was observed in the 324 medusahead/foot2 level. Bottlebrush squirreltail acquired greater root carbohydrate reserves than medusahead under competitive conditions. Under proper management, Hironaka [96] suggests a successional sequence of cheatgrass to medusahead to bottlebrush squirreltail dominated sites for northern Great Basin areas receiving greater than 11 inches (279 mm) precipitation.

Rome and Eddelman [166] compared bottlebrush squirreltail seedling growth in competition with Japanese brome at densities of 0, 50, 100, 200, 400 Japanese brome/m2. Observations were made in Missoula, Montana at 23, 42, 56, 82, and 97 days following an 8 April seeding of bottlebrush squirreltail and Japanese brome. Bottlebrush squirreltail averaged 85% survival in areas without Japanese brome, compared to an average of 66% survival from areas with 100 to 400 Japanese brome/m2 (p<0.05). Overall, bottlebrush squirreltail under competition with Japanese brome showed the greatest competitive ability at 100 Japanese brome/m2.

Martlette and Anderson [131] observed poor bottlebrush squirreltail seed dispersal into adjacent crested wheatgrass (Agropyron cristatum) stands. Plant cover acted as a barrier restricting the dispersal capabilities of bottlebrush squirreltail.

Under greenhouse conditions, Schlatterer and Tisdale [172] found sagebrush leaf litter to significantly (p<0.05) decrease bottlebrush squirreltail germination compared to moss and rabbitbrush (Chrysothamnus spp.) litter. The average number of bottlebrush squirreltail seeds (20 seeds/pot) germinating under different litter treatments is summarized below:

Big sagebrush Moss Rabbitbrush No litter
11.25 18.75 18.25 18.25

Bottlebrush squirreltail will readily establish in pinyon-juniper tree litter when a fermentation layer is not present [69].

Robertson [165] observed seeded bottlebrush squirreltail within a big sagebrush habitat at 5,200 feet (1,585 m) in northern Nevada to be short lived, persisting for 5 years. Bottlebrush squirreltail persisted for 30 years following direct seeding within a big sagebrush/bluebunch wheatgrass site in south-central Idaho [148].

OTHER USES AND VALUES:
No entry

OTHER MANAGEMENT CONSIDERATIONS:
The addition of nitrogen to disturbed sagebrush communities in Colorado [141] and mountain meadows of Nevada [62] had no positive effect on bottlebrush squirreltail establishment. Bottlebrush squirreltail decreased after the addition of nutrients in the form of stabilized sewage sludge [78].

Bottlebrush squirreltail reproductive potential is adversely affected by jointworm larvae. Spears and Barr [179] found culm length, seed weight, germination (%), and germination rate all significantly lower (p<0.01) on bottlebrush squirreltail infested with jointworms compared to noninfested plants. Results are summarized below :

Variable Infested Uninfested
Culm length (cm) 30.0 33.7
Leaf length (cm) 22.0 23.1
# of spikelets 5 9
Seed weight (mg 25 seeds) 108.2 162.5
Germination (%) 20.0 66.0
Growth rate (Seedlings day -1 100 seeds -1) 2.8 7.8

Rangelands:
Bottlebrush squirreltail is a valuable winter range plant in the Great Basin [48], with leaves remaining green and succulent through the winter.

Bottlebrush squirreltail's total available root carbohydrate reserves are lowest in early spring (approximately 3rd leaf stage), and at the beginning of fall regrowth. Total available carbohydrates are highest after anthesis [50]. By the 4th leaf stage, bottlebrush squirreltail has replaced the carbohydrate reserves found in roots at the beginning of the growing season [20]. Wright [206] found bottlebrush squirreltail most tolerant to herbage removal at time of seed maturity, declining slightly after maturity before fall regrowth. In eastern Oregon, bottlebrush squirreltail is resistant to late season defoliation [31]

Bottlebrush squirreltail generally increases in abundance when moderately grazed or protected on the foothills of intermountain winter ranges [104]. Moderate trampling by livestock in big sagebrush rangelands of central Nevada enhanced bottlebrush squirreltail seedling emergence compared to untrampled conditions. Heavy trampling destroys germination sites and significantly (p<0.05) reduces germination, whereas moderate trampling may enhance germination [63].

Bottlebrush squirreltail is tolerant of grazing in big sagebrush rangelands of southeastern Idaho [4]. In sagebrush rangelands of western Utah, Cook and Child [46] found winter harvesting to have a minor effect on crown cover, whereas early spring (April 1, May 1) harvest greatly reduced bottlebrush squirreltail cover.

Bottlebrush squirreltail vegetative vigor was evaluated over 25 years within a sagebrush rangeland of southeastern Oregon excluded from grazing. Vigor of bottlebrush squirreltail increased significantly over the 25 year period, with the 1st decade showing slower growth than the 2nd. The average annual precipitation over the 25 years equaled 8.3 inches (210 mm) with 40% falling during April, May, and June. Winters were cold with snow cover from December to March. Summers were hot, occasionally exceeding 100 degrees Fahrenheit (38 °C) [3].

Bottlebrush squirreltail is commonly found in heavily grazed and browsed (cattle and deer) aspen stands of big sagebrush steppe in Wyoming [36].

McPherson and Wright [144] observed significantly (p<0.01) greater coverage of bottlebrush squirreltail on ungrazed versus grazed Pinchot juniper rangelands in western Texas. Within the ponderosa pine bunchgrass ranges of the central Rocky Mountains, bottlebrush squirreltail production is greatest under light and moderate grazing regimes [52]. Bottlebrush squirreltail is tolerant of heavy grazing in the ponderosa pine zone of the Coconino Plateau, Arizona, since its long, sharp awns are usually present to discourage grazing [8].

On shortgrass ranges of the central plains bottlebrush squirreltail is very tolerant of light to moderate grazing [118].

Silviculture:
Climax western juniper stands are of mixed age, consisting of 1st year seedlings to trees several hundred years old. Seral stands are composed of predominately younger aged trees. In central Oregon, Vaitkus and Eddleman [194] observed significantly greater (p<0.05) bottlebrush squirreltail production when associated with large (older) trees compared to small trees. Production of bottlebrush squirreltail was also significantly greater (p<0.05) under juniper canopies compared to intercanopy zones. McPherson and others [143] observed significantly greater (p<0.01) bottlebrush squirreltail under Pinchot juniper canopies and at canopy edges compared to areas beyond canopy, within grazed and relict grasslands of western Texas. Evaluations by Tueller and Platou [190] lend supporting evidence (see: SUCCESSION within the Botanical and Ecological Characteristics section).

Bottlebrush squirreltail does not reduce ponderosa pine seedling growth. Two-year-old pine seedlings that were planted the 1st postfire spring, after a June wildfire in northern Arizona, were not affected in height or diameter by competition with bottlebrush squirreltail [66]. In Arizona ponderosa pine forests, seedlings normally gain dominance over bottlebrush squirreltail within 5 years [8].

Bottlebrush squirreltail drastically increased 4 years after a clear-cut within a lodgepole pine forest of northeastern Utah at 8,800 feet (2,700 m). Bottlebrush squirreltail showed the largest increase in vegetative production out of all grasses present [10]:

1976 (kg/ha) 1980 (kg/ha)
Ross' sedge 56.8 42.0
elk sedge 2.1 4.4
Poa spp. 10.2 40.7
bottlebrush squirreltail 3.3 47.7
5 others 0.0 13.9

Bottlebrush squirreltail was an early colonizer after the clear-cut of a ponderosa pine forest in north-central California [138].

Bottlebrush squirreltail populations were greatest 11 to 25 years after clearcuts of a red fir forest in the Sierra Nevada, California [73].

Everett and Sharow [70] found bottlebrush squirreltail seed production was less under singleleaf pinyon (Pinus monophylla)-Utah juniper woodland canopies than in clearcut areas (1 and 2 postharvest years).


BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Elymus elymoides

GENERAL BOTANICAL CHARACTERISTICS:
Bottlebrush squirreltail is a cool season, [8] perennial bunchgrass native to the Intermountain West [18]. It is solitary [200], possessing solid, mostly flowering culms [210], with flat leaf blades. The inflorescence is a spike 0.8 to 6.7 inches (2-17 cm) long [82,150,200]. Ecotypic variation is common among bottlebrush squirreltail populations [9].

Reynolds and Fraley [164] found bottlebrush squirreltail roots to achieve depths of 39.4 inches (100 cm) below the soil surface. Depths below 39.4 inches (100 cm) were not seen due to a subsurface layer of basalt, suggesting rooting depths greater than 39.4 inches (100 cm) are possible. A lateral root extension of 16 inches (40 cm) was observed at 9.8, 20, 24 and 39.4 inch (25, 50, 60 and 100 cm) depths.

RAUNKIAER [163] LIFE FORM:
Hemicryptophyte

REGENERATION PROCESSES:
Bottlebrush squirreltail regenerates from surviving root crown [29,201] and seed [18]. Vegetative propagation is nonexistent [18]. Bottlebrush squirreltail has the ability to produce large numbers [99,214] of highly germinable seeds, with relatively rapid germination [214] when exposed to the correct environmental cues. Plants are self-fertilizing [55]. Seeds are readily dispersed by wind [15,99] a few days following maturation [18]. Dispersal is a function of bottlebrush squirreltail's long reflexed awns and disarticulating, mature inflorescence [99,131,148]. Seeds are dispersed when the spike inflorescence is carried along the ground by wind catching the long awns [131].

Although bottlebrush squirreltail has the potential for long distance seed dispersal, Martlette and Anderson [131] found natural plant cover to act as a barrier to dispersal. Wind dispersal of bottlebrush squirreltail seed did not exceed 131 feet (40 m), with viable seed remaining relatively close to mature bottlebrush squirreltail plants.

Dormancy protects bottlebrush squirreltail seeds from germinating during seasonal dry periods. Dry seeds require a period of afterippening, which widens environmental conditions conducive to germination [18]. Allen and others [2] found germination rate increased and dormancy levels decreased as the duration of dry storage increased. Desert bottlebrush squirreltail seed commonly show higher levels of dormancy than seed from mountain populations [18].

Bottlebrush squirreltail seeds may germinate without a period of afterippening, showing a partial state of dormancy. However mean germination time for recently harvested seeds is longer than for afterippened seeds.

Beckstead [19] evaluated the germination temperature requirements of recently harvested bottlebrush squirreltail seeds obtained from mountain and desert habitats. The greatest germination occurred primarily at 50/68 degrees Fahrenheit (10/20 °C) and 59/77 degrees Fahrenheit (15/25 °C), with higher temperatures of 68/86 degrees Fahrenheit (20/30 °C) inhibiting germination.

Environmental conditions and timing of phenological events greatly affect the probability of recently harvested bottlebrush squirreltail seed germination. Temperatures of 50/68 degrees Fahrenheit (10/20 °C) and 59/77 degrees Fahrenheit (15/25 °C) are unlikely to occur during summer months in desert habitats. In higher, mountain habitats, summer temperatures of 50/68 degrees Fahrenheit (10/20 °C) and 59/77 degrees Fahrenheit (15/25 °C) may occur; however, bottlebrush squirreltail usually ripens later at higher elevations [19]. In general, recently harvested bottlebrush squirreltail seeds at lower elevations have a much greater probability of fall germination than seeds from higher elevations [2].

Chabet and Billings [40] observed germination of bottlebrush squirreltail seeds from alpine sites (10,793 feet (3,290 m)) in the Sierra Nevada. The greatest germination (%) occurred at day/night temperatures of 81/73 degrees Fahrenheit (27/23 °C (96%)) and 90/82 degrees Fahrenheit (32/28 °C (92%)).

SITE CHARACTERISTICS:
Bottlebrush squirreltail has wide ecological amplitude [161], but it most commonly occurs in disturbed areas of deserts, valleys, foothills, and mountain meadows [18].

Regional:
Bottlebrush squirreltail is found throughout Colorado on dry hills, plains, and rocky slopes, and within open woods and meadows [92]. In Montana, bottlebrush squirreltail occurs in dry, open habitats, from valley to timberline [123]. Throughout the western Great Plains, bottlebrush squirreltail is commonly found on dry soils of pastures and roadsides [82]. In Utah, bottlebrush squirreltail prefers dry to moist vegetation types, from salt desert shrub to alpine grassland [200]. Plains, rocky hills, or montane slopes are common sites in New Mexico [132]. In Arizona, open sandy ground, rocky hills, and open pine woods are common sites [115]. Bottlebrush squirreltail is common to dry rangeland areas of Kansas [154]. In central Washington, bottlebrush squirreltail prefers disturbed sites. Within these sites plant density is negatively correlated with individual plant size [153]. In California, bottlebrush squirreltail is found in scattered stands at high elevations on dry, gravelly soils. It is also common to hillsides and brush associations [168].

Soils:
Bottlebrush squirreltail inhabits a wide variety of soil types and is tolerant of saline [108] and alkaline soils [168]. It is widely distributed in salt-desert shrub ranges of the west, on dry, gravelly soils, or within alkaline conditions. Bottlebrush squirreltail is found on clayey soils of northeastern California sagebrush communities [27]. Throughout Montana it occurs predominantly on dry, gravelly soils, in saline or alkaline areas [150]. Within alpine areas of Olympic National Park, Washington, bottlebrush squirreltail prefers well-drained, undifferentiated, disturbed, shallow and stony soils [21]. Passey and Hugie [158] found bottlebrush squirreltail to achieve better growth on Newdale silt loam soils than on Brunt silt loam, in areas with similar climate, slope, and exposure. Bottlebrush squirreltail may also occur on loose, ashy soil [11].

Bottlebrush squirreltail is not common within wet areas such as river lowlands and soil along irrigation canals [153].

Elevation by state:
Arizona 2,000 to 11,500 (609-3,505 m) [115]
west-central Montana 7,000 to 9,200 feet (2,135-2,805 m) [123]
New Mexico 4,500 to 11,500 feet (1,372-3,505 m) [132]
Utah 3,510 to 11,400 feet (1,070-3,500 m) [200]

SUCCESSIONAL STATUS:
Depending upon habitat type, bottlebrush squirreltail may occur as an early, mid-, or late successional species.

Shrub rangelands:
Bottlebrush squirreltail is generally a dominant component of seral big sagebrush/bunchgrass communities [217]. Bottlebrush squirreltail is represented in early seral and climax stages of big sagebrush/bluebunch wheatgrass associations in Nevada. Tueller and Platou observed the most pronounced bottlebrush squirreltail during early and climax stages of big sagebrush/bluebunch wheatgrass associations in Nevada [190]. Bottlebrush squirreltail is found within seral and climax stages of big sagebrush rangelands in southeastern Idaho [4]. It is a component of climax big sagebrush communities in Idaho [205] and is a member of climax big sagebrush/western wheatgrass communities of Colorado [183]. Within shrub-steppe ecosystems of western Colorado, bottlebrush squirreltail is an early seral species [117]. Bottlebrush squirreltail also occurs in climax shadscale communities [100].

Pinyon-juniper communities:
Bottlebrush squirreltail is common in mid-seral and climax pinyon-juniper communities of Mesa Verde, Colorado [67,68]. Bottlebrush squirreltail is a component of seral and climax western juniper (Juniper occidentalis) communities of the Pacific Northwest [54].

Ponderosa pine communities:
Bottlebrush squirreltail is a member of interior ponderosa pine climax communities within the central and southern Rocky Mountains [209].

Prior to invasion of nonnative annuals in the Snake River Plain, Idaho, bottlebrush squirreltail occupied a mid to late seral status, suppressing the early seral fescues, sixweeks fescue (Vulpia octoflora), and foxtail fescue (Vulpia myuros) [160].

SEASONAL DEVELOPMENT:
The wide ecological amplitude of bottlebrush squirreltail leads to differential timing of phenological events between individuals of differing habitats [43,109]. Flowering generally occurs in spring or early summer [18,57]. Lower elevation populations (that is, cold desert, salt desert habitats) usually mature early June with higher elevation populations (that is, mountain brush, mountain meadows) reaching maturity in late July [18]. Hironaka and Tisdale observed phenological differences between the subspecies Elymus elymoides ssp. elymoides and ssp. californicum. In a common garden experiment E. e. ssp. elymoides developed 10 to 14 days earlier than ssp. californicum [100].

Between 1960 and 1969, Murray and others evaluated bottlebrush squirreltail phenology in southern Idaho. Growth began from mid-March to mid-April. Flower stalks began to form late-April to mid-May, with anthesis occurring in early to mid-June. Plants were dormant from the middle of July to the end of August with fall regrowth occurring through October [152].

Clary [43] evaluated bottlebrush squirreltail phenology and rate of growth from different environments using a transplant garden and growth chamber. The timing of bottlebrush squirreltail phenological events and overall growth rate was closely related to homesite environmental conditions. Bottlebrush squirreltail individuals from higher elevations were limited by cold temperatures whereas individuals from lower elevations were limited by water availability and warm temperatures. Under the same environmental constraints, bottlebrush squirreltail from areas with low moisture stress and cool climates showed higher growth rates, attaining maximum height earlier than individuals from warmer drier sites. Bottlebrush squirreltail requires the longest time to flower in areas of relatively moderate temperature and moisture regimes:

Time to flowering in days for bottlebrush squirreltail individuals from different habitats is shown below. Plants were grown at 6,490 feet (1,980 m) on a clay loam with an annual precipitation of 21.4 inches (544 mm) and annual temperature of 49 degree Fahrenheit (9.5 oC).

 
Bottlebrush squirreltail collection site description Days to flower
7,410 feet (2,260 m), silt loam, ponderosa pine 205.5 
4,990 feet (1,520 m), stony clay loam, ponderosa pine 201.2 
7,200 feet (2,200 m), loam, pinyon-juniper 193.8 
7,810 feet (2,380 m), clay loam, ponderosa pine 192.5 
9,780 feet (2,980 m), gravelly loam, spruce-fir 172.5 
9,320 feet (2,840 m), gravelly sandy loam, mountain grassland 166.8 
4,530 feet (1,380 m), loamy fine sand, short grass 165.8 
4,720 feet (1,440 m), cobble clay, pinyon-juniper 162.2 
4,990 feet (1,520 m), stony clay loam, ponderosa pine 159.5 
5,510 feet (1,680 m), silty clay loam, sagebrush-greasewood 158.0 
4,530 feet (1,380 m), stony loam, oak savannah 153.5 

Bottlebrush squirreltail is responsive to fall rains in northern areas of the Great Basin, allowing for fall regrowth. Fall regrowth uses the majority of total available root carbohydrates partitioned during the summer [50]. The optimal soil temperature for root and shoot growth occurs at approximately 77 degrees Fahrenheit (25 °C). However, bottlebrush squirreltail shows continuous root growth down to 41 degrees Fahrenheit (5 °C) soil temperature [100].


FIRE ECOLOGY

SPECIES: Elymus elymoides

FIRE ECOLOGY OR ADAPTATIONS:
Bottlebrush squirreltail's small size, coarse stems, and sparse leafy material aid in its tolerance of fire [31]. Postfire regeneration occurs from surviving root crowns and from on- and off-site seed sources [29]. Frequency of disturbance greatly influences postfire response of bottlebrush squirreltail. Undisturbed plants within a 6 to 9 year age class generally contain large amounts of dead material, increasing bottlebrush squirreltail's susceptibility to fire [210].

Koniak [120] found bottlebrush squirreltail to be a major component of postfire pinyon-juniper communities of the Great Basin at any time during succession. Greatest occurrence and coverage of bottlebrush squirreltail are generally achieved during mid-seral stages.

Successional stage Occurrence (%) Percent of areas achieving > 5% cover
Early (1 year old) 43 3
Early-mid (4-8 years old) 58 15
Mid (15-17 years old) 49 28
Late-mid (22-60 years old) 90 0
Late > 60 years old 44 0


Fire regimes for plant communities in which bottlebrush squirreltail occurs are summarized below. For further information regarding fire regimes and fire ecology of communities and ecosystems where bottlebrush squirreltail is found see "The Fire Ecology and Adaptations" section of the FEIS species summary for the plant community or ecosystem dominants listed below.

Community or Ecosystem Dominant Species Fire Return Interval Range (years)
silver fir-Douglas-fir Abies amabilis-Pseudotsuga menziesii var. menziesii > 200
sagebrush steppe Artemisia tridentata/Pseudoroegneria spicata 20-70 [33]
basin big sagebrush A. t. var. tridentata 12-43 [170]
mountain big sagebrush A. t. var. vaseyana 20-60 [7,37]
Wyoming big sagebrush A. t. var. wyomingensis 10-70 (40)** [196,215]
saltbush-greasewood Atriplex confertifolia-Sarcobatus vermiculatus < 35 to < 100
desert grasslands Bouteloua eriopoda and/or Pleuraphis mutica 5-100
plains grasslands Bouteloua spp. < 35
blue grama-needle-and-thread grass-western wheatgrass B. g.-Hesperostipa comata-Pascopyrum smithii < 35
blue grama-buffalo grass B. g.-Buchloe dactyloides < 35
grama-galleta steppe B. g.-Pleuraphis jamesii < 35 to < 100
blue grama-tobosa prairie B. g.-P. mutica < 35 to < 100
cheatgrass Bromus tectorum < 10
mountain-mahogany-Gambel oak scrub Cercocarpus ledifolius-Quercus gambelii < 35 to < 100
western juniper Juniperus occidentalis 20-70
Rocky Mountain juniper J. scopulorum < 35
Sierra lodgepole pine* Pinus contorta var. murrayana 35-200
Rocky Mountain ponderosa pine* P. ponderosa var. scopulorum 2-10
Arizona pine P. p. var. arizonica 2-10
galleta-threeawn shrubsteppe Pleuraphis jamesii-Aristida purpurea < 35 to < 100
mesquite-buffalo grass Prosopis glandulosa-Buchloe dactyloides < 35
Texas savanna P. g. var. glandulosa < 10 [33]
mountain grasslands Pseudoroegneria spicata 3-40 (10)** [6]
Rocky Mountain Douglas-fir* Pseudotsuga menziesii var. glauca 25-100
interior live oak Quercus wislizenii < 35 [33]
*fire return interval varies widely; trends in variation are noted in the species summary
**(mean)

POSTFIRE REGENERATION STRATEGY [182]:
Crown residual colonizer (on-site, initial community)
Secondary colonizer (on-site or off-site seed sources)


FIRE EFFECTS

SPECIES: Elymus elymoides

IMMEDIATE FIRE EFFECT ON PLANT:
Although bottlebrush squirreltail is generally top-killed by fire, its small size and low density of coarse fuel per unit basal area make it relatively fire tolerant [31,198,208]. Low density of above ground plant tissue produces a quick, "hot" flame, transferring little heat to growing points below the soil surface [208,210]. The solid culms of bottlebrush squirreltail do not readily burn, compared to those of perennial grass associates [210].

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
No entry

PLANT RESPONSE TO FIRE:
Bottlebrush squirreltail sprouts from surviving root crown [29,201] and colonizes from seed [29].

Seasonal trends in bottlebrush squirreltail root carbohydrate reserves greatly affect postfire response. Burning is generally harmful during late spring and early summer [30,208] coinciding with low points in carbohydrate reserves [20]. Bottlebrush squirreltail is most tolerant of late summer (anthesis) or mid-fall (before regrowth) fires [30,49,79], coinciding with relatively high carbohydrate reserves [20]:



A difference in phenological traits of surviving postfire individuals may exist between small (1 to 3 inch (2.5-7.6 cm) crown diameter) and large (>3.5 inches (8.9 cm) crown diameter) bottlebrush squirreltail plants. Wright [210] found large plants to produce significantly (p<0.01) higher numbers of flowering stalks than small plants after fire.

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
Wright [208] compared bottlebrush squirreltail response to burning and clipping near Boise, Idaho. Heat was applied by propane burner for 20 to 30 seconds to raise soil surface temperature to 400 or 800 degrees Fahrenheit. The 800 degree Fahrenheit treatment killed 25% of bottlebrush squirreltail plants during July and August. No other treatment caused mortality. Burning and clipping during all seasons reduced yields 1 year after treatment. Burning reduced yield most during May. Average herbage weight (in grams) per bottlebrush squirreltail plants in relation to season and treatment at 1 postfire year is summarized below:

Season 400 F 800 F Clipped Control
May 3.94a 5.48 7.41 22.58
June 6.96 8.50 7.26 22.09
July 8.51a 4.32ab 13.25 14.01
August 7.50a 9.42 11.58 16.61
September 10.44 6.11ab 10.21 21.97

a Differences from clipped treatment significant at p<0.05
b Differences from 400 F treatment significant at p<0.05

Wright [207] conducted time/temperature evaluations of bottlebrush squirreltail mortality on 5 dates between 19 May and 21 September, at temperatures between 120 to 200 degrees Fahrenheit (48.9-93.3 C). Time required to kill bottlebrush squirreltail tissue at all temperatures within the test range increased as burning date increased. The greatest change occurred between 10 June and 21 July.

Time (minutes) required to kill bottlebrush squirreltail tissue at 172 degrees Fahrenheit (78 C) [208]:

19 May 10 June 21 July 20 August 21 September
4.00 5.50 18.50 28.00 33.50

Fox [77] found a direct association between postfire response of bottlebrush squirreltail and ponderosa pine tree density and canopy cover. The greatest cover (%) of bottlebrush squirreltail was achieved in areas with larger (>4 inches (10 cm) diameter) trees and less dense tree canopies.

Blank and others [26] grew bottlebrush squirreltail under greenhouse conditions in soil from a July wildfire site and adjacent unburned areas within a big sagebrush habitat of Nevada. Bottlebrush squirreltail had greater aboveground biomass and more total N, P and K, along with greater silica content, when grown in soil collected from wildfire sites.

Spring:
Early spring fire (May) within sagebrush ecosystems of eastern Oregon greatly reduced bottlebrush squirreltail basal area [31,32]. Basal area decreased an average of 47 % the 2nd postfire year. Britton and others [31] compared bottlebrush squirreltail postfire response in eastern Oregon to clipping. Yield (1 postfire year) after a May fire was less than yield from clipping (down to 0.4 inch (1 cm) stubble). Results the 2nd year showed no significant difference.

Bottlebrush squirreltail populations increased after a "moderate" spring (May 1972) wildfire in a ponderosa pine forest on limestone-sandstone derived soils, near flagstaff Arizona. The area observed was logged 2 years before, averaging 16,875 board feet/acre (6,750 board feet/ha). Number of bottlebrush squirreltail stems per hectare in 1972 and 1974 is summarized below in thousands/ha [17]:

Moderate burn 1972 Severe burn 1972 Control (logged, not burned) 1972
7.2 0 0
Moderate burn 1974 Severe burn 1974 Control (logged, not burned) 1974
18.1 0.1 1.1

Although frequency of bottlebrush squirreltail was too low for statistical analysis, Champlin [42] reported no damage to bottlebrush squirreltail basal cover and height 2 postfire years after a spring fire in a big sagebrush community of northern California. Bottlebrush squirreltail vigor increased the 1st and 2nd postfire growing season in central Oregon, following a spring fire within a sagebrush-bitterbrush/bunchgrass plant community [1].

Summer:
Bottlebrush squirreltail increased following an August wildfire in a big sagebrush community with an understory dominated by cheatgrass and Lyall's milkvetch (Astragalus lyallii) [95]. Significantly (p<0.01) greater biomass was achieved 1 postfire year after a 19 July prescribed fire in Oregon. At time of burn, bottlebrush squirreltail had entered summer quiescence with no green shoot material evident. Mean shoot biomass of burned plants was greater per unit crown area, compared to control. Burned plants also averaged 49% higher root biomass per unit crown area, producing a shoot:root biomass ratio of 1.73 compared to control plots at 0.43 shoot:root biomass. Burning also increased the proportion of reproductive culms; 74.8% of all shoots of burned plants produced reproductive culms compared to 14.3% for unburned plants [220].

Bottlebrush squirreltail showed a negative postfire response to summer (July) wildfire within a sagebrush rangeland in Utah, for the 2nd and 3rd postfire years compared to control [202]. Bottlebrush squirreltail decreased in abundance 1 postfire year after a summer (July) prescribed fire and after a lightning fire within a mountain mahogany-big sagebrush community [187].

Fall:
Bottlebrush squirreltail maintained previous levels of production (kg/ha) 1 postfire year after an October fire in an aspen-bunchgrass community of northern Arizona. Although total vegetative production remained constant, percent cover and density of bottlebrush squirreltail were significantly higher. The October fire resulted in a large bottlebrush squirreltail population consisting of small individuals whose combined vegetative biomass equaled or exceeded preburn levels. Associated dominants, Arizona fescue and mountain muhly, decreased [86].

For further information on bottlebrush squirreltail response to fire, see Fire Case Studies, Lyon's Research Paper (Lyon 1971), and the following Research Project Summaries:

FIRE MANAGEMENT CONSIDERATIONS:
Humphrey and Schupp [103] compared bottlebrush squirreltail seedling emergence within burned and unburned cheatgrass dominated areas of the Great Basin, Utah. Greater seedling emergence (April) occurred on seeded burned areas compared to unseeded, within loamy fine sand (85% sand) sites. On a dune site with sandy soil (95% sand), seedling emergence occurred in March with no significant difference between burned and unburned sites. However, a significantly greater proportion of bottlebrush squirreltail seedlings survived on burned dune areas compared to unburned.

Seeding:
Aerially applied seed mixture of mutton grass, prairie Junegrass, Indian ricegrass, slender wheatgrass (Elymus trachycaulus) and bottlebrush squirreltail aided in the reestablishment of bottlebrush squirreltail after a summer (August) wildfire within Mesa Verde National Park, Colorado [74]. Bottlebrush squirreltail was an important component 1, 2, [76] and 3 postfire years [75] in seeded areas, whereas no bottlebrush squirreltail was observed in unseeded areas [74].

Postfire recovery of bottlebrush squirreltail occurred after a summer (June 1956) wildfire in Arizona chaparral, aerially seeded with weeping lovegrass (Eragrostis curvula) and crested wheatgrass. Results shown that percent frequency of bottlebrush squirreltail within 9.6 foot (2.9 m) square plots increased steadily for 4 years postfire [157]:

1956 1957 1958 1960 1961
bottlebrush squirreltail 0 2.5 4.0 10.5 21.5
crested wheatgrass 0 14.0 20.5 17.5 13.0
weeping lovegrass 0.5 2.0 1.5 4.0 6.0

Seeding postfire pinyon-juniper communities of the Great Basin with desert wheatgrass (Agropyron desertorum), intermediate wheatgrass (Thinopyrum intermedium), and smooth brome (Bromus inermis) inhibits establishment of bottlebrush squirreltail [120].

Pinyon-juniper communities:
Four years after a late summer (July-August) wildfire in pinyon-juniper woodlands of Mesa Verde, Colorado, Erdman [67] found bottlebrush squirreltail as an important component. Bottlebrush squirreltail, along with Indian rice grass and mutton grass, assumed dominance after a 3-year annual grass/forb stage. At 25 postfire years, bottlebrush squirreltail is a member of climax stands.

Within pinyon-juniper ranges of west-central Utah, bottlebrush squirreltail is an important native perennial species at 5 to 6 postfire years [13].

A fire return interval less than 10 to 25 years should increase abundance of bottlebrush squirreltail in newly expanded (young) western juniper stands (Juniperus occidentalis) receiving greater than 14 inches (350 mm) precipitation, at elevations higher than 4,900 feet (1,500 m), in southwestern Idaho [35].

FIRE CASE STUDIES:

SPECIES: Elymus elymoides

FIRE CASE STUDY CITATION:
Simonin, Kevin, compiler. 2001. Bottlebrush squirreltail postfire response in a ponderosa pine forest of Arizona. In: Elymus elymoides. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ [ ].

REFERENCE:
Vose, James M.; White, Alan S. 1991. Biomass response mechanisms of understory species the first year after prescribed burning in an Arizona ponderosa pine community. Forest Ecology and Management. 40(3/4): 175-187. [199].

SEASON/SEVERITY CLASSIFICATION:
This was a late October fire. The fire smoldered for several days and consumed the entire litter layer, with a total heat yield of 115,830 calories/foot2 (1600 kJ/m2).

STUDY LOCATION:
The prescribed fire took place within open sawtimber sites, pole sites, and sapling sites within a ponderosa pine (Pinus ponderosa)/Arizona fescue (Festuca arizonica) habitat type on the Fort Valley Experiment Forest near Flagstaff.

PREFIRE VEGETATIVE COMMUNITY:
Saw timber sites 296 trees/acre (120 trees/ha)
Pole sites average tree diameter of 5.9 inches (15cm) at 1,730 trees/ha
Sapling sites average tree diameter 1.8 inches (4.5 cm) at 10,070 trees/ha


TARGET SPECIES PHENOLOGICAL STATE:
No-entry

SITE DESCRIPTION:
ponderosa pine/Arizona fescue habitat type

FIRE DESCRIPTION:
The day of initiation, average temperature was between 57.2 and 64.4 degrees Fahrenheit (14-18 C) and relative humidity was 21%.

FIRE EFFECTS ON TARGET SPECIES:
Vegetative biomass within open saw timber sites, including surviving bottlebrush squirreltail plants and new seedlings, was approximately 3 times greater (P<0.05) on burn plots compared to control, 247 lbs (112 kg/ha) and 88 lbs (40 kg/ha) respectively. Surviving bottlebrush squirreltail plants within burned, open saw timber areas had more than twice the vegetative production. Seedling recruitment within open saw timber sites was also greater on burned than control plots. Burned open saw timber plots produced an average seedling biomass 15 times greater (p<0.05) than on control plots. Bottlebrush squirreltail in pole sites and sapling sites showed a more negative response to fire than in open sawtimber sites. Results for bottlebrush squirreltail are summarized below:

Burn biomass (kg/ha) Control biomass (kg/ha) Density (plants/m2) Control density (plants/m2 )
Open saw timber sites
all plants 112.45 40.06 9.06 5.74
residual plants 96.00 40.23 4.34 4.56
seedlings 15.84 1.03 4.98 0.57
Pole sized sites
all plants 18.97 21.86 2.27 5.56
residual plants 18.23 18.64 2.05 3.41
seedlings 0.78 3.17 0.28 1.99
Sapling sites
all plants 14.08 11.72 2.91 4.18
residual plants 12.77 10.64 1.96 2.97
seedlings 1.36 0.77 1.14 0.52


No significant phenological differences were found between all burned areas and control.

FIRE MANAGEMENT IMPLICATIONS:
Overall response of bottlebrush squirreltail within pole and sapling stands was less than in open saw timber areas; however, the response of surviving plants in pole and sapling stands remained strong. Seedlings are generally more vulnerable to environmental changes than established plants. The postfire response of surviving bottlebrush squirreltail plants may result in an increased presence of bottlebrush squirreltail within later postfire stages.


Elymus elymoides: References


1. Adams, Glenn R. 1980. Results of range/wildlife prescribed burning on the Fort Rock Ranger District in central Oregon. R-6 Fuels Management Notes. September 24, 1980. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region, Aviation and Fire Management. 6 p. [292]

2. Allen, Phil S.; Debaene-Gill, Susan B.; Meyer, Susan E. 1994. Regulation of germination timing in facultatively fall-emerging grasses. In: Monsen, Stephen B.; Kitchen, Stanley G., compilers. Proceedings--ecology and management of annual rangelands; 1992 May 18-22; Boise, ID. Gen. Tech. Rep. INT-GTR-313. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 215-219. [24284]

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