SPECIES: Calamagrostis rubescens

INTRODUCTORY

SPECIES: Calamagrostis rubescens
AUTHORSHIP AND CITATION:
Matthews, Robin F. 2000. Calamagrostis rubescens. 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:
CALRUB

SYNONYMS:
No entry

SCS PLANT CODE:
CARU

COMMON NAMES:
pinegrass
pine reedgrass

TAXONOMY:
The currently accepted scientific name of pinegrass is Calamagrostis rubescens Buckl. (Poaceae) [69,71,76,82,154]. There are no recognized infrataxa.

LIFE FORM:
Graminoid

FEDERAL LEGAL STATUS:
No special status

OTHER STATUS:
No entry

DISTRIBUTION AND OCCURRENCE

SPECIES: Calamagrostis rubescens
GENERAL DISTRIBUTION:
Pinegrass is found from Manitoba west through British Columbia and south through Washington, Idaho, Montana, Oregon, Wyoming, Utah, Colorado, and into California [69,70,71,82,153,154]. The Natural Resources Conservation Service provides a map of pinegrass' distribution in the United States (http://plants.usda.gov/plants/cgi_bin/topics.cgi).

ECOSYSTEMS [52]:
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES22 Western white pine
FRES23 Fir-spruce
FRES25 Larch
FRES26 Lodgepole pine
FRES29 Sagebrush
FRES34 Chaparral-mountain shrub
FRES36 Mountain grasslands
FRES44 Alpine

STATES:
CA CO ID MT NV OR UT WA WY
AB BC MB

BLM PHYSIOGRAPHIC REGIONS [15]:
1 Northern Pacific Border
2 Cascade Mountains
3 Southern Pacific Border
5 Columbia Plateau
6 Upper Basin and Range
8 Northern Rocky Mountains
9 Middle Rocky Mountains
10 Wyoming Basin
11 Southern Rocky Mountains

KUCHLER [80] PLANT ASSOCIATIONS:
K002 Cedar-hemlock-Douglas-fir forest
K010 Ponderosa shrub forest
K011 Western ponderosa forest
K012 Douglas-fir forest
K013 Cedar-hemlock-pine forest
K014 Grand fir-Douglas-fir forest
K015 Western spruce-fir forest
K018 Pine-Douglas-fir forest
K037 Mountain-mahogany-oak scrub
K047 Fescue-oatgrass
K050 Fescue-wheatgrass
K051 Wheatgrass-bluegrass
K052 Alpine meadows and barren
K055 Sagebrush steppe
K063 Foothills prairie

SAF COVER TYPES [46]:
205 Mountain hemlock
206 Engelmann spruce-subalpine fir
208 Whitebark pine
210 Interior Douglas-fir
211 White fir
212 Western larch
213 Grand fir
215 Western white pine
217 Aspen
218 Lodgepole pine
219 Limber pine
224 Western hemlock
226 Coastal true fir-hemlock
227 Western redcedar-western hemlock
228 Western redcedar
229 Pacific Douglas-fir
230 Douglas-fir-western hemlock
237 Interior ponderosa pine
238 Western juniper
244 Pacific ponderosa pine-Douglas-fir
245 Pacific ponderosa pine

SRM (RANGELAND) COVER TYPES [123]:
101 Bluebunch wheatgrass
102 Idaho fescue
104 Antelope bitterbrush-bluebunch wheatgrass
105 Antelope bitterbrush-Idaho fescue
107 Western juniper/big sagebrush/bluebunch wheatgrass
108 Alpine Idaho fescue
109 Ponderosa pine shrubland
110 Ponderosa pine-grassland
302 Bluebunch wheatgrass-Sandberg bluegrass
304 Idaho fescue-bluebunch wheatgrass
309 Idaho fescue-western wheatgrass
311 Rough fescue-bluebunch wheatgrass
312 Rough fescue-Idaho fescue
314 Big sagebrush-bluebunch wheatgrass
315 Big sagebrush-Idaho fescue
316 Big sagebrush-rough fescue
317 Bitterbrush-bluebunch wheatgrass
318 Bitterbrush-Idaho fescue
319 Bitterbrush-rough fescue
322 Curlleaf mountain-mahogany-bluebunch wheatgrass
409 Tall forb 410 Alpine rangeland
415 Curlleaf mountain-mahogany
420 Snowbrush
421 Chokecherry-serviceberry-rose

HABITAT TYPES AND PLANT COMMUNITIES:
Throughout its range, pinegrass is an important and often dominant understory species in Douglas-fir (Pseudotsuga menziesii) [73,97,113,133], subalpine fir (Abies lasiocarpa)-Engelmann spruce (Picea engelmannii) [73,93,113,130], grand fir (A. grandis) [51,73,131], ponderosa pine (Pinus ponderosa) [30,60,73,156], lodgepole pine (P. contorta) [29,113,130], and quaking aspen (Populus tremuloides) forests [3,104,157].

Pinegrass is the most abundant grass in forested areas of interior southern British Columbia, where it is a dominant understory species in the Douglas-fir and sub-boreal pine-spruce zones. It is common in drier zones of the sub-boreal spruce and montane spruce zones as well. Pinegrass also occurs in drier portions of the interior western redcedar-western hemlock (Thuja plicata-Tsuga heterophylla) and subalpine fir-Engelmann zones and at lower elevations and in moist areas of the ponderosa pine and bunchgrass zones [30].

Douglas-fir/pinegrass habitat types are widespread and are recognized from southern British Columbia southeast through Montana and Wyoming and south to the eastern foothills of the Cascade Range and the Wallowa and Blue Mountains of Oregon [36,97,113,130,133,146,155]. According to Mauk and Henderson [93], the southern extent of this habitat type probably occurs in northern Utah's Uinta range. Douglas-fir/pinegrass appears to be the most widely distributed habitat type in the Douglas-fir series within the Northern Rockies [51,113,133,155]. A publication describing the type and outlining succession and management in the Douglas-fir/pinegrass habitat type in central Idaho is available [133].

Some classifications identifying plant communities in which pinegrass is described as a dominant understory species follow:

Classification of the forest vegetation of Wyoming [2]
Forest vegetation of the Medicine Bow National Forest in southeastern Wyoming: a habitat type classification [3]
Vegetation of two drainages in Eagle Cap Wilderness, Wallowa Mountains, Oregon [29]
Forest habitat types of northern Idaho: a second approximation [31]
Forest vegetation of eastern Washington and northern Idaho [37]
Natural vegetation of Oregon and Washington [51]
Plant communities of the Blue Mountains in eastern Oregon and southeastern Washington [60]
Coniferous forest habitat types of northern Utah [93]
Plant communities of the Similkameen Valley, British Columbia [97]
Aspen community types of the Intermountain Region [104]
Forest habitat types of Montana [113]
A preliminary description of plant communities found on the Sawtooth, White Cloud, Boulder, and Pioneer Mountains [118]
Forest habitat types of eastern Idaho-western Wyoming [130]
The Douglas-fir/pinegrass habitat type in central Idaho: succession and management [133]
Forested plant associations of the Okanogan National Forest [155]

Some common plant associates of pinegrass in Douglas-fir habitats include ninebark (Physocarpus malvaceus), oceanspray (Holodiscus discolor), baldhip rose (Rosa gymnocarpa), white spirea (Spiraea betulifolia), common snowberry (Symphoricarpos albus), mountain snowberry (S. oreophilus), Scouler willow (Salix scouleriana), Saskatoon serviceberry (Amelanchier alnifolia), Rocky Mountain maple (Acer glabrum), bearberry (Arctostaphylos uva-ursi), chokecherry (Prunus virginiana), pachistima (Pachistima myrsinites), Oregon-grape (Mahonia repens), heartleaf arnica (Arnica cordifolia), big-leaved sandwort (Arenaria macrophylla), western yarrow (Achillea millefolium), mountain sweet-cicely (Osmorhiza chilensis), tailcup lupine (Lupinus caudatus), wood strawberry (Fragaria vesca), and elk sedge (Carex geyeri) [2,26,54,73].

Common plant associates of pinegrass in subalpine fir-Engelmann spruce habitats include menziesia (Menziesia ferruginea), grouse whortleberry (Vaccinium scoparium), other huckleberry species (Vaccinium spp.), bearberry, beargrass (Xerophyllum tenax), twinflower (Linnaea borealis), pachistima, heartleaf arnica, queencup beadlily (Clintonia uniflora), bunchberry (Cornus canadensis), twisted stalk (Streptopus amplexifolium), and elk sedge [73,93].

Some common plant associates of pinegrass in quaking aspen (Populus tremuloides) habitats include russet buffaloberry (Shepherdia canadensis), mountain snowberry, rose (Rosa spp.), bearberry, Oregon-grape, shrubby cinquefoil (Dasiphora floribunda), western yarrow, strawberry (Fragaria spp.), showy aster (Aster conspicuus), northern bedstraw (Galium boreale), Richardson's geranium (Geranium richardsonii), sticky geranium (G. viscosissimum), Fendler meadowrue (Thalictrum fendleri), lupine (Lupinus spp.), and elk sedge [2,78,104,157].

MANAGEMENT CONSIDERATIONS

SPECIES: Calamagrostis rubescens
IMPORTANCE TO LIVESTOCK AND WILDLIFE:
Because of its abundant herbage production and widespread distribution, pinegrass is considered an important or principal forage species over much of its range [30,45,70,98,106,111,144,149].

Pinegrass is primary forage on southern British Columbia summer ranges, providing over half of the weight of forage for cattle in the Douglas-fir, lodgepole pine, and fir-spruce zones [1,30,98]. Cattle readily consume pinegrass from spring through midsummer but tend to avoid it by August [13,98]. Pinegrass is grazed by domestic sheep as well [146,149].  Pinegrass is sensitive to overgrazing, and its forage value decreases over the course of the season [1,13,30,56,98,149]. 

Pinegrass provides forage or other value to various wildlife species. Grizzly bears forage on pinegrass in Yellowstone National Park, Wyoming, and in the Bitterroot Mountains of southwestern Montana, especially in the spring [16,38]. Pinegrass is also utilized by black bear, white-tailed deer, mule deer, bighorn sheep, and elk [,24,45,59,77,81,105,146]. Pinegrass-dominated ledges are often utilized by Rocky Mountain goats in Glacier National Park, Montana, for feeding and bedding from May through September [25]. Douglas-fir/pinegrass and subalpine fir/pinegrass habitat types are used by boreal owls for nesting and roosting in the northern Rocky Mountains [67].

PALATABILITY:
Pinegrass is most palatable in the spring but declines over the growing season.  It is considered 1 of the least valuable of the important western range grasses [13,30,97,146,147,149]. Pinegrass appears to be less palatable when growing in shaded conditions [98]; on some clearcuts in Montana it was preferred to rough fescue (Festuca altaica) and bluebunch wheatgrass (Pseudoroegneria spicata) early in the season [13]. Pinegrass palatability is improved by burning [110,150]. Overall, the palatability of pinegrass to livestock and wildlife has been rated as follows:

cattle and horses poor to good  [30,41,59,103,146,147,149]
domestic sheep poor to fair  [41,103,144,149]
elk poor to good
mule deer poor to fair  [41,103]
white-tailed deer poor 
pronghorn poor
small mammals fair  [41]

NUTRITIONAL VALUE:
In general, pinegrass is moderate quality forage, but its overall value declines throughout the growing season [30,98]. It is rated moderate in crude protein value in the spring and early summer which then declines, and moderate in crude fiber value in early summer.  Crude fiber increases toward fall [30,75,98]. The following crude protein (%) and digestible dry matter (%) values for elk were reported in the beginning of July and the end of September on burned (postfire year 3) and unburned quaking aspen sites [24]:

  Crude Protein - July Crude Protein - Sept. IVDMD* - July IVDMD -    Sept.
Burned 14.1 8.8 66.0 56.6
Unburned 18.0 9.8 68.5 57.7
*in-vitro dry matter digestibility

In other studies comparing nutritive values in similar situations, the increased flowering and seed production on the burned aspen sites had apparently caused the decrease in forage quality (decreases in  percent IVDMD, crude protein,  phosphorus, and calcium) [40].

Other authors have also found higher protein values and higher in-vitro digestible dry matter values for pinegrass growing under a forest canopy than in open or thinned areas [98,145]. Hedrick and others [68], however, found crude protein values higher in openings versus under heavy shade in grand-fir-mixed conifer forests in northeastern Oregon.

Pinegrass has higher protein content than timothy (Phleum pratense) or orchard grass (Dactylis glomerata) (both of which are used to seed burned Douglas-fir sites) from late June to mid-September [94,134].

Johnston and Bezeau [75] have listed seasonal nutrient composition of pinegrass in Alberta as follows:

Stage Dry Matter Protein Crude Fat Crude Fiber Ash  Ca P
Leaf 91.85 14.90 3.55 28.00 12.25 0.22 0.19
Head 92.37 8.03 2.30 32.73 10.00 0.28 0.17
Ripe Seed 91.70 8.40 3.60 29.00 9.00 0.53 0.17
Cured 92.30 3.33 2.50 29.70 15.70 0.52 0.08
Weathered 95.47 4.40 2.23 35.23 7.77 0.60 0.06


Trace element content analysis of pinegrass shows that copper and molybdenum levels are satisfactory for livestock most of the growing season, whereas zinc is below the minimum requirements for livestock [30].

COVER VALUE:
Because of its low stature, pinegrass provides some cover only to small mammals and birds [41].

VALUE FOR REHABILITATION OF DISTURBED SITES:
Pinegrass frequently exhibits vigorous growth following disturbance [27,84,85,107,134,155]. Its rhizomes and dense network of fine roots form a thick sod, which holds up under trampling [28,147,149]. In addition, the thick sod provides control of surface soil erosion for watershed protection [30,147]. Pinegrass sod may also play a role in rehabilitation by excluding the establishment of more competitive species [30]. Pinegrass has been included in seed mixtures developed for restoration of disturbed western redcedar-western hemlock sites in Glacier National Park [92].

OTHER USES AND VALUES:
No entry

OTHER MANAGEMENT CONSIDERATIONS:
Pinegrass abundance and productivity are best on open sites [27,68,145]. In grand fir-mixed conifer types in Oregon, pinegrass production decreased from 260 pounds per acre (291 kg/ha) in unlogged open areas to 60 pounds per acre (67 kg/ha) under closed canopies. In logged areas, pinegrass produced 380 pounds per acre (425 kg/ha) on open sites and 70 pounds per acre (78 kg/ha) on shaded sites [44,68]. Other authors have also reported increases in pinegrass following overstory removal of grand fir in central Idaho [131]. Thinning of ponderosa pine increased pinegrass production by 47% in Oregon [44]; pinegrass cover increased 40% in thinned ponderosa pine in eastern Washington where spacings were greater than 16.5 feet (5 m) [30]. Thinning of lodgepole pine in eastern Oregon resulted in an average increase of 90% in pinegrass [145]; canopy removal increased pinegrass abundance in subalpine forests in British Columbia [97] and in Douglas-fir/pinegrass types in various locations [97,133]. However, on some drier sites overstory removal may cause a decrease in abundance. Pinegrass decreased following logging on low elevation, open Douglas-fir/fescue (Festuca spp.) sites in the Similkameen Valley, British Columbia [97], and in bluebunch wheatgrass phases of the Douglas-fir/pinegrass types in Montana [113].

Pinegrass is a major competitor with conifer seedlings throughout its range [14,50,66,86,108,112,134]. Its ability to aggressively compete for soil moisture is often detrimental to conifer seedling establishment and growth [13,30,84]. This aggressiveness may be useful in areas where overstocking of lodgepole pine is a problem [30]. Due to the lack of a favorable conifer seedbed, seedling establishment is also hindered where thick pinegrass sod occurs [114,122,126,127,134]. Extensive scarification is often required to reduce competition for seedling establishment [113,155]; a 2-by-2-foot (60 × 60 cm) scalp may not be adequate to reduce pinegrass' competitive edge [127,134]. In general, profuse pinegrass flowering can be prevented on conifer/pinegrass sites if some tree cover is left during harvesting [86].  Planting artificial regeneration early, before pinegrass becomes well established, also favors conifer establishment [30,86,114,122,155]. Herbicide control and application rates for pinegrass have been reported by various authors [27,30,100,136]. 

Light to medium soil disturbance from mechanical site preparation or other means is favorable to pinegrass, while heavy disturbance may retard pinegrass growth and reduce its cover by displacing the rhizomes and roots [27,30,53]. Anchor chain drag scarification can increase pinegrass abundance and cover. Disc trenching or patch scarification will initially reduce pinegrass, but the control will be short-lived [27,30]. On Douglas-fir sites in central Idaho, pinegrass underwent a major increase in vegetative growth in response to clearcutting with no additional site preparation. A minor decrease in cover was reported following shelterwood cuts and clearcuts followed by mechanical scarification. A major increase in cover due to vegetative expansion and some establishment of seedlings occurred in clearcuts followed by burning and after wildfires [134].

Pinegrass is susceptible to heavy grazing [27,86,98,159]. This is partially due to the fact that large clumps can easily be pulled out of the ground [30,98,143]. Environmental conditions, especially summer rainfall, affect the response of pinegrass to grazing or clipping, and year-to-year fluctuations in growth because of variability in conditions can exceed variation from grazing treatments [30]. Simulated grazing studies have revealed that pinegrass will decrease in yield and vigor after 1 season of heavy grazing (clipping to 2 inches (5 cm) every 2 weeks), with successive years of grazing causing increasing deterioration. Effects of clipping were less severe in years of high rainfall and/or cool temperatures. It has been estimated after heavy grazing for 4 consecutive years, pinegrass would require 20 years to fully recover. Clipping to 6 inches (15 cm) did not affect pinegrass vigor, and pinegrass was most sensitive to herbage removal in July, when growth was slowing down and summer dormancy was setting in [1,64,98,143]. It has been recommended that pinegrass range be utilized early in the season or in the spring and again in late summer and fall. Grazing pinegrass throughout the growing season is not recommended [1,64,79,98,143]. According to McLean [98], deterioration in pinegrass range is hard to detect in the early stages.

Pinegrass has increased dramatically in abundance and cover following applications of nitrogen-based fertilizers [27,30].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Calamagrostis rubescens
GENERAL BOTANICAL CHARACTERISTICS:
Pinegrass is a native, perennial, rhizomatous grass. Culms are hollow and can reach up to 44 inches (110 cm) in height, but more typically grow 11 to 20 inches (30-50 cm) tall [65,146,154]. Dense panicles are 3 to 6 inches (7-15 cm) long. Pinegrass may produce moderate-sized clumps rather than single stems  and is referred to in the literature as a sod-former [30,147,149,154], particularly when the overstory is removed [135]. Rhizomes grow mostly in the top 2 inches (5 cm) of mineral soil [30,53,72]. Seed-producing plants usually grow in the open and  are somewhat tufted and erect, differing in appearance from the stemless, drooping form that usually grows in shade [147].

RAUNKIAER [115] LIFE FORM:
Hemicryptophyte
Geophyte

REGENERATION PROCESSES:
Pinegrass reproduces primarily through the lateral extension of rhizomes [30,143,150]. Pinegrass rarely flowers except in open areas or after removal of the forest canopy [34,36,82,120,147,154]. Seedlings reach reproductive age in 1 to 2 years [72]. Seeds are wind-dispersed and apparently do not persist in the soil [30,72,134].  Seeds germinate best on mineral soil [134]; stratification or scarification are not required, which may indicate that pinegrass seeds can germinate in the fall when an adequate supply of moisture is available and other conditions are suitable  [30]. 

SITE CHARACTERISTICS:
Pinegrass grows in open areas and under forest canopies from valley floors to subalpine zones. It tolerates mesic to dry sites and is found on all aspects [30,42,70,71,82]. Pinegrass is often found growing in extensive stands [82]. Some elevational ranges have been reported in the following states and province; 8,052 to 9,058 feet (2440-2745 m) in Utah [154], 2,700 to 7,800 feet (818-2363 m) in Montana [113], and 1,980 to 6,105 feet (600-1850 m) in the Similkameen Valley, British Columbia [97].

Pinegrass is found on a wide variety of parent materials and soils within its geographic range. Most often, soils are well drained, loamy to coarse-textured Luvisols and Brunisols [30]. In the Similkameen Valley of southern British Columbia, Douglas-fir/pinegrass habitats are wide ranging and have developed mostly on glacial till, glacial outwash, and alluvium. These soils are chiefly loams to sandy loams that are well drained and are rapidly permeable [97]. On pinegrass-dominated sites in the Garnet Mountains of western Montana, pinegrass cover is highest on limestone-derived soils, and the grass is found more often on limestone than on granitic parent materials. No significant difference was found between limestone- and quartzite-derived soils [55].

Throughout interior British Columbia, pinegrass occurs on sites with nutrient regimes that range from very poor to very rich. Soils supporting dense stands of pinegrass are often low in nitrogen. Pinegrass often grows on sites that experience at least a moderate moisture deficit during the growing season. Pinegrass is most abundant in the interior Douglas-fir zone, where it occupies extremely xeric to hygric sites [30].

SUCCESSIONAL STATUS:
Pinegrass is shade tolerant [34,98,149,154] and is persistent throughout all successional stages [35,51,86,113]. In many situations, pinegrass can maintain moderate cover under moderate shade conditions, but with increased light will acquire new vigor and dominate the herb layer [27,131]. Pinegrass generally produces seedlings only in natural forest openings or in openings created by disturbance [,34,36,114,131,143]. Pinegrass is an aggressive competitor for soil moisture [13,84,86] due to the fact that it undergoes rapid growth early in the season and has high soil water usage [84,124]. It is also capable of  photosynthesis at very low leaf water potentials (-3.5 mP) [13,124]. Pinegrass is resistant to disturbance and readily invades or resprouts on sites following logging or fire, often with an increase over predisturbance canopy cover [86,102,126,142].

Pinegrass is classified by various authors as a seral to climax species throughout its range. For example, pinegrass is considered seral in Engelmann spruce/queencup beadlily habitat types in Glacier National Park [5], in some Douglas-fir stands in interior British Columbia [146], in the Douglas-fir/Rocky mountain maple type in central Idaho [132], in many lodgepole pine stands [60,113,130], and in the subalpine fir/big huckleberry (Vaccinium membranaceum)-beargrass [126] and western hemlock habitat types in Idaho [158].

Over much of its range, pinegrass occurs as a climax understory dominant with Douglas-fir [31,37,51,60,83,97,109,113,134,146], and to a somewhat lesser extent with subalpine fir [94,98,114,131,] grand fir [51,131,135], and ponderosa pine [156]. Some quaking aspen/pinegrass communities in the Intermountain Region are considered climax on sites suitable for quaking aspen dominance, or may be seral to Douglas-fir/pinegrass or subalpine fir/pinegrass habitats, especially in the absence of fire [104,157].

SEASONAL DEVELOPMENT:
Pinegrass generally initiates growth in late April or early May and typically goes into dormancy by early August [1,98]. In a study on the growth and development of pinegrass in interior British Columbia, new stem initiation was completed by mid-May, and no new stem development occurred until fall. All measures of growth (leaf number, stem height, stem weight, and leaf blade area) indicated a cessation of growth in July [143]. Regrowth takes place in late August or early September, depending on site and timing of fall rains [1,98]. Although flowering is infrequent [143,147,154], it generally takes place July and August [71,110,144].

The following phenological observations were reported for pinegrass in the northern Rocky Mountains (dates given are means; east of Continental Divide includes Montana and Yellowstone National Park; west of Continental Divide includes Montana and northern Idaho) [119]:

Location Growth starts Leaves fully grown Flowering starts Flowering stops Seeds ripe Seed fall starts Leaf fall begins Leaf fall ends 1st frost injury
East of Divide May 6 June 22 July 2 July 22 Aug. 6 Aug. 20 Aug. 21 Oct. 5 Sept. 3
West of Divide April 19 May 30 May 30 July 2 July 14 July 28 Aug. 28 Sept. 26 Sept. 13

FIRE ECOLOGY

SPECIES: Calamagrostis rubescens
FIRE ECOLOGY OR ADAPTATIONS:
Pinegrass sprouts from rhizomes following fire [27,74,139,150]. Rhizomes are buried in the top 2 inches (5 cm) of mineral soil [18,72], allowing pinegrass to survive fires that do not completely consume the duff layer [34,50,114,128,150]. Pinegrass is seldom, if ever, eliminated from a site even after severe wildfires [141].  Throughout its range, repeated fires promote increased cover in pinegrass and often result in its early postfire dominance [17,27,30,91,130,155].

Pinegrass seedlings establish on burned sites from off-site sources, and pinegrass undergoes mass flowering in the years immediately following fire [34,72,114,142,151], allowing for rapid postfire colonization [72,74,150].

Historically, pinegrass stands have perpetuated surface fires that maintained open stands of ponderosa pine. Ponderosa pine is now being replaced by Douglas-fir, white fir, or grand fir in the Blue and Wallowa Mountains of Oregon, in central Idaho, and along the east side of the Cascade Range [61,62,134,152]. Pinegrass stands have also perpetuated low-intensity, high frequency fires in the interior Douglas-fir zone of British Columbia, maintaining an open stand structure [56]. Estimated fire return intervals for several pinegrass-dominated habitats follow:

Habitat type Location Estimated fire return interval
Ponderosa pine/pinegrass Blue Mts., OR 10 yrs [73]
Douglas-fir/pinegrass interior British Columbia 13 yrs [56]
Douglas-fir/pinegrass Forest-grassland ecotones,  southwestern MT 22 yrs [8]
moist Douglas-fir/ pinegrass Bitterroot NF, MT 28 yrs [6]
dry Douglas-fir/pinegrass Lolo NF, MT 10 yrs [39]

Other habitat types in which pinegrass is named as a dominant understory species have been placed in "Fire Groups" in certain regions. Descriptions of these "Fire Groups", and the importance of pinegrass within them, along with fire history information and fire management considerations are available by region in the literature: eastern Idaho and western Wyoming [17]; northern Idaho [128]; central Idaho [32]; Utah [18]; Montana (east of the Continental Divide) [50]; and western Montana [39,49].

Estimated fire return intervals for some ecosystems and communities in which pinegrass occurs follow:

Community or Ecosystem Dominant Species Fire Return Interval Range in Years
silver fir-Douglas-fir Abies amabilis-Pseudotsuga menziesii var. menziesii > 200 
grand fir A. grandis 35-200 
sagebrush steppe Artemisia tridentata/Pseudoroegneria spicata 20-70 
California montane chaparral Ceanothus and/or Arctostaphylos spp. 50-100 [23]
curlleaf mountain-mahogany* Cercocarpus ledifolius 13-1000 [9,121]
western juniper Juniperus occidentalis 20-70 
western larch Larix occidentalis 25-100 
Engelmann spruce-subalpine fir Picea engelmannii-Abies lasiocarpa 35 to > 200 
whitebark pine* Pinus albicaulis 50-200 [23]
Rocky Mountain lodgepole pine* P. contorta var. latifolia 25-300+ [7,117]
western white pine* P. monticola 50-200 
Pacific ponderosa pine* P. ponderosa var. ponderosa 1-47 
Rocky Mountain ponderosa pine* P. ponderosa var. scopulorum 2-10 [23]
quaking aspen (west of the Great Plains) Populus tremuloides 7-120 [23,57,99]
mountain grasslands Pseudoroegneria spicata 3-40 (10)** [7]
Rocky Mountain Douglas-fir* Pseudotsuga menziesii var. glauca 25-100 [23]
coastal Douglas-fir* P. menziesii var. menziesii 40-240 [23,101,116]
western redcedar-western hemlock Thuja plicata-Tsuga heterophylla > 200 
mountain hemlock* T. mertensiana 35 to > 200 [23]
*fire return interval varies widely; trends in variation are noted in the species summary
**(mean)

POSTFIRE REGENERATION STRATEGY [139]:
Rhizomatous herb, rhizome in soil
Initial off-site colonizer (off-site, initial community)
Secondary colonizer - on-site seed

FIRE EFFECTS

SPECIES: Calamagrostis rubescens
IMMEDIATE FIRE EFFECT ON PLANT:
Pinegrass is usually top-killed by fire [95]. Fires that consume the duff layer may kill pinegrass rhizomes [18,96,114]. In the northern Rocky Mountains, pinegrass is seldom, if ever, eliminated from a site even after severe wildfire [141].

DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
No entry

PLANT RESPONSE TO FIRE:
Pinegrass sprouts from rhizomes and establishes from seed following fire. It may bloom profusely for the 1st 2 or 3 postfire years, allowing rapid colonization of burned areas. It can also invade burned areas from off-site sources. Pinegrass generally increases in response to fire, often exceeding preburn levels [20,27,32,72,74,128,138,139,142].

DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
Following the Pattee Canyon Wildfire in a Douglas-fir type near Missoula, Montana, pinegrass bloomed profusely in the 1st postfire summer. Subsequent seedling establishment was observed in the following 2 years, along with vegetative spread from rhizomes. Pinegrass responded rapidly to the July wildfire. Even on severely burned sites it became the most abundant native species. Pinegrass cover increased from 0.6 to 8.5% on upland sites, and from 1.0 to 1.3% on ravine sites from postfire year 1 to postfire year 3 [33,34].

Johnson [74] summarized pinegrass response to several fires in the mountains of northeastern Oregon. Pinegrass responded quickly and positively to low, moderate, and high severity fires in grand fir, Douglas-fir, ponderosa pine, and in ninebark-common snowberry shrublands. In all cases, pinegrass sprouted from rhizomes and reproduced from seed, often with dramatic increases over preburn cover. Pinegrass recovery was most delayed following severe fires in ponderosa pine types.

In western Montana, pinegrass cover increased following the high severity Sleeping Child Fire in seral Rocky Mountain lodgepole pine communities within subalpine fir/beargrass and subalpine fir/grouse whortleberry habitat types. Cover values (means from 11 transects) were 0.3% in postfire year 1; 11.6% in postfire year 12; and 19% in postfire year 21 [89,90].

The following coverage class values (2=6% to 25% cover; 3=26% to 50% cover; 4=51% to 75% cover) were reported for pinegrass following moderately severe prescribed fires in a western quaking aspen and quaking aspen-mixed conifer type in the Caribou National Forest, Idaho [20]:

Type Untreated Postfire Yr 2 Postfire Yr 4
Quaking aspen-mixed conifer 4 4 3
Pure quaking aspen 3 3 2

In these study areas, pinegrass produced more biomass, uniformly flowered, and produced seed on severely burned sites. Pinegrass also produced many new culms in burned areas, but few were produced in unburned areas [40].

Pinegrass followed a general trend (increase in frequency and/or cover) in the first 17 years following the Waterfalls Canyon Fire in Grand Teton National Park. The fire occurred in a subalpine fir-Engelmann spruce dominated area, with some lodgepole pine contributing to the overstory. Cover and frequency were temporarily reduced in postfire year 1 on moderately burned and severely burned sites, as compared to an adjacent stand that had burned 43 years earlier and an adjacent unburned stand. Pinegrass increased by continued sprouting and seedling establishment from abundant postfire seed production, with greater cover in burned stands compared to unburned stands during most of the successive 17-year period. Mean percent frequency and cover, respectively, for pinegrass during the course of the study follow (frequencies were not provided after postfire year 1) [10,43]:

Site Postfire Yr 1 Postfire Yr 2 Postfire Yr 3 Postfire Yr 9 Postfire Yr 17
43-yr-old Burn 78 (12) ---  --- ---  --- ---  --- ---  ---
Unburned 42 (7) ---/(7) ---/(9) ---/(5) ---/(5)
Moderate Burn 7 (1) ---/(2) ---/(3) ---/(13) ---/(9)
Severe Burn 15 (1) ---/(4) ---/(3) ---/(27) ---/(9)

On ponderosa pine and Douglas-fir communities in the Blue Mountains of northeastern Oregon, pinegrass cover and frequency were higher on sites that had been thinned 6 years previously than on prescribed burned, thinned-and-burned, or control sites. Pinegrass was determined to be an indicator species for thinned sites (P≤0.05). For further information on the effects of thinning and burning treatments on pinegrass and 48 other species, see the Research Project Summary of Youngblood and others' [160] study.

For further information on prescribed fire use and postfire response of plant species, including pinegrass, see Fire Case Studies, Lyon's Research Paper, and the following Research Project Summaries:

FIRE MANAGEMENT CONSIDERATIONS:
Fire suppression in the Pacific Northwest has caused a downward trend in range condition where pinegrass has historically played an important role. The invasion of Douglas-fir and grand fir into ponderosa pine communities has caused an overall increase in crown cover which in turn has caused a decrease in pinegrass production. The highest densities of pinegrass in the region are found in fire-maintained ponderosa pine stands [63].

According to Johnson [74], low- and moderate-severity fires are best for pinegrass enhancement in Douglas-fir/pinegrass associations of the Blue Mountains and Wallowa Mountains in Oregon.

Prescribed fire guidelines have been developed for Douglas-fir/pinegrass habitat types in Montana that are adjacent to grasslands where shrub enhancement for wildlife is important [58].

Stout and Brooke [143] developed an equation for determining the relationship between culm height and leaf area for pinegrass. Total leaf blade area (y) was predicted from culm height (x): y=0.39375 + 0.051604x + 0.00419223x² (R²=0.97). This may be useful in predicting fuel loading. Others have evaluated the National Fire Danger Rating System grass fuel models [129], compiled photo guides for appraising fuel loading in Rocky Mountain habitat types with pinegrass indicators [47,48], estimated fuel weights for pinegrass [21], and developed surface area to volume ratio models as indicators of flammability [19,22]. Fuel loads have been estimated for the Douglas-fir/pinegrass habitat type in northwestern Montana [4].

FIRE CASE STUDIES

SPECIES: Calamagrostis rubescens
1st CASE STUDY:
FIRE CASE STUDY CITATION:
Matthews, Robin F., compiler. 2000. Pinegrass development following prescribed burning of Douglas-fir in south-central Idaho. In: Calamagrostis rubescens. 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/ [ ].

REFERENCES:
Lyon, L. Jack. 1966. Initial vegetal development following prescribed burning of Douglas-fir in south-central Idaho. Res. Pap. INT-29. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 17 p. [87].

Lyon, L. Jack. 1971. Vegetal development following prescribed burning of Douglas-fir in south-central Idaho. Res. Pap. INT-105. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 30 p. [88].

SEASON/SEVERITY CLASSIFICATION:
August/high

STUDY LOCATION:
The study took place in Neal Canyon, a small drainage off of Eagle Creek and the Big Wood River, located within the Sawtooth National Forest. The area is approximately 6 miles north of Ketchum, Idaho (NE1/4, Sec.13, T3N, R17E, Boise Meridian).

PREFIRE VEGETATIVE COMMUNITY:
Preburn frequencies based on occurrence in 25 2-by-2-foot (60 cm × 60 cm) quadrats located in the study area follow ("present" indicates species was present in the stand but not detected in quadrats): Douglas-fir (Pseudotsuga menziesii), 20%; subalpine fir (Abies lasiocarpa), present; Engelmann spruce (Picea engelmannii), present; lodgepole pine (Pinus contorta), present; limber pine (Pinus flexilis), present; quaking aspen (Populus tremuloides), present; Rocky Mountain maple (Acer glabrum), 12%; bristly black currant (Ribes lacustre), 20%; (Ribes viscosissimum), 4%; (Penstemon fruticosus), 8%; mountain snowberry (Symphoricarpos oreophilus), 12%; pinegrass, 24%; bluebunch wheatgrass (Pseudoroegneria spicata), 8%; (Poa nervosa), 20%; elk sedge (Carex geyeri), 52%; (Trisetum spicatum), 12%; heartleaf arnica (Arnica cordifolia), 48%; showy aster (Aster conspicuus), 8%; (Castilleja miniata), 8%; bull thistle (Cirsium vulgare), 12%; (Collinsia parviflora), 8%; fireweed (Epilobium angustifolium), 20%; giant frasera (Frasera speciosa), 16%; cinquefoil (Potentilla spp.), 20%; (Pyrola secunda), 12%; (Senecio cymbalarioides), 44%; (Viola adunca), 12%.

The stand had been logged twice in the previous 13 years and conifers remaining on the site were mostly mistletoe (Arceuthobium spp.)-deformed saplings and poles.

TARGET SPECIES PHENOLOGICAL STATE:
Although not stated, pinegrass would have been entering summer dormancy at the time the study site was burned.

SITE DESCRIPTION:
The 120-acre (48-ha) study area was located at elevations of 6,500 to 7,000 feet (1970-2120 m) and general aspect was north-northeast. Annual rainfall at the time of the study at 2 nearby weather stations was 14 to 17 inches (355-432 mm).  Total snowfall was 85 to 120 inches (2160-3050 mm). Soils in Neal Canyon are of limestone origin, rocky, poorly developed, and highly stable. Analysis at the site showed at least 50% gravel with a shift from sandy loams on the surface to a sandy clay loam below 14 inches (35.6 cm). Since some parts of the area had been heavily disturbed during previous years of logging, a specific 20-acre (8 ha) site of "less disturbed" terrain on a lower, middle slope was selected for examination. True aspect at this location was N 10-20 degrees.  Slope was 64% and elevation was 6,500 feet (1970 m).

FIRE DESCRIPTION:
Basic objectives of the burn were sanitation and site preparation of the area for silvicultural purposes. Moisture content of fuel moisture sticks examined each day in the week preceding the burn had been a relatively constant 5 to 6%. During the firing period, temperatures rose from the mid-50 degrees Fahrenheit to nearly 80 degrees Fahrenheit (12.8-26.6 oC). Relative humidity dropped from approximately 50 to 10% or less. Surface winds were mostly under 5 miles per hour, with fire-induced gusts to 30 miles per hour. The fire was started with a flame thrower through the middle of the site and on the lower edge. Fire swept through tree crowns almost immediately. Within the site all litter and herbaceous material, all dead material less than 3 inches (7.6 cm), and all live stems less than 2 inches (5.1 cm) at the base were completely consumed. Fire intensity was measured with water can integrating devices described by Beaufait [12], resulting in a mean heat energy flux at the surface of the cans of 200 calories/second for about 45 minutes.  It was inferred by the author that the water loss was comparable to losses during a severe broadcast burn in deep, dry slash.

FIRE EFFECTS ON TARGET SPECIES:
The fire consumed all aboveground portions of pinegrass. Mean frequency (%) of pinegrass in the prefire and postfire communities, as well as in an adjacent stand that burned 30 years earlier, follow ("x" represents present in stand but not detected in quadrats):

Preburn Postfire Yr 1 Postfire Yr 2 Postfire Yr 3 Postfire Yr 4 Postfire Yr 5 Postfire Yr 6 Postfire Yr 7 Adj. 30-yr-old Wildfire
24 x 20 28 24 24 64 52 32

FIRE MANAGEMENT IMPLICATIONS:
As the findings of the study indicate, pinegrass appeared early in the postfire stand at frequencies only slightly different than the preburn community. Lyon's notes suggested an aggregate rather than random distribution pattern of pinegrass individuals. He predicted no major expansion of pinegrass, except for an increase in cover in the years to follow.

2nd CASE STUDY:
FIRE CASE STUDY CITATION:
Matthews, Robin F., compiler. 2000. Pinegrass response to spring and fall burning for wildlife habitat improvement, west-central Montana. In: Calamagrostis rubescens. 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:
Noste, Nonan V. 1982. Vegetation response to spring and fall burning for wildlife habitat improvement. In: Baumgartner, David M., compiler & editor. Site preparation and fuels management on steep terrain: Proceedings of a symposium; 1982 February 15-17; Spokane, WA. Pullman, WA: Washington State University, Cooperative Extension: 125-132. [107].

SEASON/SEVERITY CLASSIFICATION:
spring (April)/low
fall (October)/high

STUDY LOCATION:
The two study sites were located in the O'Keefe Creek area, approximately 10 miles (16 km) north of Missoula, Montana, within the Lolo National Forest.

PREFIRE VEGETATIVE COMMUNITY:
The prefire vegetation in the study area consisted of a senescent seral shrub community within a Douglas-fir (Pseudotsuga menziesii)/ninebark (Physocarpus malvaceus) habitat type. Wildfire had burned the O'Keefe Creek area 34 years earlier.

Volumes expressed as feet³/acre for some shrubs, herbs, and graminoids in the prefire community follow:

Species Spring Burn Fall Burn
Saskatoon serviceberry (Amelanchier alnifolia) 37,753  18,551
Snowbrush ceanothus (Ceanothus velutinus) 27,991  35,948 
Oregon-grape (Mahonia repens) 238  0
Ninebark 51,547  45,041 
Scouler willow (Salix scouleriana) 55,786  8,183 
Common snowberry (Symphoricarpos albus) 4,295 15,438 
Bearberry (Arctostaphylos uva-ursi) 119  0
Arrowleaf balsamroot (Balsamorhiza sagittata) 89  0
Elk sedge (Carex geyeri) 3,528  3,217 
Orchard grass (Dactylis glomerata) 253  554
Pinegrass (Calamagrostis rubescens) 164 208

TARGET SPECIES PHENOLOGICAL STATE:
Although not specifically stated, pinegrass would have probably initiated spring growth prior to the April burn, and would have been undergoing regrowth after fall rains during the October burn.

SITE DESCRIPTION:
The fall burn was conducted within a 50-acre (20-ha) area; the spring burn comprised 125 acres (50 ha). Aspect within the 2 burned areas was generally southeast; elevations were below 4,950 feet (1500 m). Slope averaged 30% with a maximum of 50%. No additional information was given on moisture regime or soils in the area.

FIRE DESCRIPTION:
The study was initiated to compare spring and fall prescribed fire treatments to improve wildlife habitat in a deteriorated seral shrub community. Although there was a higher fuel loading on the fall than on the spring burn sites, loadings on both sites were low. The amount of fine fuels on both sites was marginal for carrying the fires. Weather conditions during the fires follow:

Transect Temperature Relative humidity Wind velocity
deg F (oC) % miles/hr (km/hr)
Fall upper slope 59 (15) 30 6 to 8 (10)
Fall mid-slope 69 (20) 23 8 (13)
Fall lower slope 70 (21) 18 7 to 12 (11)
Spring upper slope 65 (18) 36 5 to 7 (8)
Spring mid-slope 57 (14) 30 6 to 10 (10)
Spring lower slope 62 (17) 37 6 to 8 (10)

Rate of spread ranged from 660 to 1,188 feet per hour (200-360 m/hr) on the spring burn, contrasted with 3,366 to 4,620 feet per hour (1020-1400 m/hr) spread rates on the fall burn. Flame lengths were correspondingly higher on the fall burn.

FIRE EFFECTS ON TARGET SPECIES:
Volumes of pinegrass in the pre- and postfire communities, given as ft³/acre, follow:

Prefire Postfire Yr 1 Postfire Yr 2
Fall Burn 208  0 12,654
Spring Burn 164  0 923 

FIRE MANAGEMENT IMPLICATIONS:
Pinegrass recovered rapidly and showed a dramatic increase in volume over preburn levels following both the fall and spring burns.

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