|FEIS Home Page|
SPECIES: Galium boreale, G. triflorum
|northern bedstraw||fragrant bedstraw|
|© 2004 Dr. Virginia Kline
University of Wisconsin Arboretum
|© 2001 Thayne Tuason
Central Washington Native Plants
Galium boreale L., northern bedstraw
Galium triflorum Michx., fragrant bedstraw
In accordance with current taxonomic views, no infrataxa are recognized for either northern bedstraw or fragrant bedstraw in this review [131,269]. However, some systematists recognize subspecies of northern bedstraw . Throughout this review, bedstraw will refer to both of the above species. When referring to any species individually, the common names listed above will be used.LIFE FORM:
Distributional maps of bedstraw and the 2 individual species are accessible through the Plants database.ECOSYSTEMS :
Northern and fragrant bedstraw -
Canada: Both species are typical of the North American taiga.
Coniferous forests: In white spruce-balsam fir (Picea glauca-Abies balsamea) and black spruce (P. mariana) communities, bedstraw presence is normally greater in white spruce-balsam fir forests . Bedstraw occurs in nutrient-rich white spruce-black spruce-highbush cranberry (Viburnum edule) associations in British Columbia .
Northwestern U.S: A diversity of riparian, coniferous, and deciduous habitats of the northwestern U.S. include bedstraw.
Coniferous forests: In northern Idaho, bedstraw associates with subalpine fir (A. lasiocarpa), grand fir (A. grandis), mountain hemlock (Tsuga mertensiana), western hemlock (T. heterophylla), ponderosa pine (Pinus ponderosa), western redcedar (Thuja plicata), and Douglas-fir (Pseudotsuga menziesii) . In several Montana and southeastern Idaho riparian habitats bedstraw is common. In subalpine fir/red baneberry (Actaea rubra), subalpine fir/claspleaf twistedstalk (Streptopus amplexifolius), and spruce/field horsetail (Picea spp./Equisetum arvense) habitats fragrant bedstraw has greater constancy than northern bedstraw. In the subalpine fir/fragrant bedstraw habitat type, fragrant bedstraw is 100% constant, and northern bedstraw is 60% constant [100,105]. Bedstraw also occurs in Montana's spruce/ninebark (Physocarpus malvaceus) habitat type .
In western North Dakota, the 2 bedstraw species are present with almost equal frequencies in Rocky Mountain juniper (Juniperus scopulorum) communities . Ponderosa pine/Kentucky bluegrass (Poa pratensis) habitat types of the Rio Grande, San Isabel, and San Juan national forests of Colorado are also bedstraw habitat .
Deciduous forests: Both bedstraw species occur in quaking aspen (Populus tremuloides)/Kentucky bluegrass and yellow willow/beaked sedge (Salix lutea/Carex rostrata) riparian habitats of Montana , quaking aspen-paper birch (Betula papyrifera) communities of western North Dakota , narrowleaf cottonwood/Saskatoon serviceberry (P. angustifolia/Amelanchier alnifolia) communities of Colorado's White River National Forest , and bur oak (Quercus macrocarpa) communities of western North Dakota .
Shrub and grassland communities: In Utah, northern bedstraw occurs in sagebrush (Artemisia spp.), meadow, and mountain brush habitats. Mountain brush vegetation may include big sagebrush (A. tridentata), Gambel oak (Q. gambelii), bigtooth maple (Acer grandidentatum), serviceberry (Amelanchier spp.), and/or mountain-mahogany (Cercocarpus spp.) .
Classifications: Bedstraw is recognized as a dominant species in many vegetation
CO: fragrant bedstraw [3,68]
ID: fragrant bedstraw [100,250,292]
MT: fragrant bedstraw [50,105,204,250]
NM: fragrant bedstraw 
WY: northern bedstraw 
fragrant bedstraw [50,250,292]
Northern bedstraw -
Northwest: Northern bedstraw is common in the following northwestern habitat types.
Coniferous forests: Northern bedstraw is described in dry ponderosa pine  and white spruce/twinberry honeysuckle (Lonicera involucrata) vegetation types of British Columbia . In Glacier National Park, Montana, northern bedstraw is typical above 5,000 feet (1,525 m) where Engelmann spruce (Picea engelmannii), subalpine fir, alpine larch (Larix lyallii), and whitebark pine (Pinus albicaulis) dominate . In other parts of Montana, northern bedstraw maintains 85% to 100% constancy in ponderosa pine/common snowberry (Symphoricarpos albus), ponderosa pine/chokecherry (Prunus virginiana), limber pine/Idaho fescue (Pinus flexilis/Festuca idahoensis), and limber pine/common juniper (J. communis) habitat types . Engelmann spruce/subalpine fir and ponderosa pine communities are common northern bedstraw habitat in the Big Horn Mountains of Wyoming .
Deciduous and mixed forests: Northern bedstraw commonly associates with quaking aspen. In Alberta, northern bedstraw occurs in quaking aspen communities with common snowberry and Saskatoon serviceberry . Coverage of northern bedstraw decreases with stand age in quaking aspen/highbush cranberry/twinflower (Linnaea borealis) communities in the taiga of interior Alaska . In the Mackenzie Valley of Canada's Northwest Territories, northern bedstraw occurs in alder (Alnus spp.) scrub communities and in mixed white spruce-aspen (Populus spp.)-jack pine (Pinus banksiana) forests .
Northern bedstraw is typical in balsam poplar (Populus balsamifera) forests of Alberta . Spruce/red-osier dogwood (Cornus sericea ssp. sericea) riparian forests in Montana also provide northern bedstraw habitat . In eastern Montana, northern bedstraw occupies green ash (Fraxinus pennsylvanica) woodlands .
Shrub and grassland communities: Northern bedstraw is a prominent forb in several Canadian grasslands. In the high diversity fescue-oatgrass (Festuca spp.-Danthonia spp.) prairies, northern bedstraw is conspicuous but rarely has high coverage. Northern bedstraw is also present in subclimax, boreal wildrye (Leymus innovatus) shrub savannahs of Banff and Jasper national parks . Northern bedstraw is considered the most important forb in grasslands dominated by shortbristle needle and thread grass (Hesperostipa curtiseta) and California oatgrass (D. californica) in Alberta. In slender wheatgrass (Elymus trachycaulus)  and rough fescue/shrubby cinquefoil (F. altaica-Dasiphora floribunda) grasslands, northern bedstraw is also prominent . Northern bedstraw is also typical in several native wheatgrass communities of Alberta. Bluebunch wheatgrass (Pseudoroegneria spicata), slender wheatgrass, thickspike wheatgrass (E. lanceolatus), and Montana wheatgrass (E. albicans) are common here .
On steep south-facing slopes in Alaska's Yukon lowlands, northern bedstraw associates with fringed sagebrush/purple pinegrass (Artemisia frigida/Calamagrostis purpurascens) vegetation . Shrubby cinquefoil/tufted hairgrass (Deschampsia cespitosa) riparian habitat types in Montana also provide northern bedstraw habitat . In western Wyoming, mountain big sagebrush (A. tridentata ssp. vaseyana) is a northern bedstraw associate . In montane riparian sites throughout Wyoming, the aster (Aster spp.)-northern bedstraw community type is recognized .
Southwest: In the southwest, northern bedstraw occupies shrublands and forests.
Deciduous and mixed forests: Northern bedstraw is typical of white fir (Abies concolor)/bigtooth maple habitat types on cool, moist, canyon slopes throughout the southwest . Welsh and others  describe lodgepole pine (Pinus contorta), aspen, and spruce-fir (Abies spp.) overstories with northern bedstraw in Utah. In the Crested Butte area of Colorado, quaking aspen is a typical associate .
Shrub and grassland communities: In Nevada, northern bedstraw occupies sagebrush and pinyon-juniper (Pinus-Juniperus spp.) vegetation . Near Gunnison County, Colorado, northern bedstraw is 56% frequent in sagebrush communities between 8,500 and 12,000 feet (2,590-3,660 m) and 46% frequent in Thurber fescue (F. thurberi) grasslands .
North-central: Northern bedstraw is common in several deciduous forest and grassland vegetation types of the north-central U.S. and Canada.
Deciduous and mixed forests: In southern Saskatchewan, northern bedstraw was present in all wooded draws dominated by silver sagebrush (Artemisia cana), boxelder (Acer negundo), quaking aspen, Bebb willow (Salix bebbiana), chokecherry, western snowberry (Symphoricarpos occidentalis), creeping juniper (J. horizontalis), or fragrant sumac (Rhus aromatica) . In the Great Sand Hills of Saskatchewan, northern bedstraw occupies creeping juniper habitat .
Northern bedstraw is described in green ash and American elm (Ulmus americana) communities of the northern Great Plains [39,278]. Constancy of northern bedstraw is 75% or more in Rocky Mountain juniper/littleseed ricegrass (Piptatherum micranthum), quaking aspen/Oregon-grape (Mahonia repens), and quaking aspen/water birch (B. occidentalis) habitat types of the Missouri Plateau . In southwestern North Dakota, northern bedstraw has 100% frequency in green ash/chokecherry, quaking aspen/chokecherry, bur oak-chokecherry, bur oak-hazel (Corylus spp.), and paper birch/western blue virginsbower (Clematis occidentalis) habitat types .
Shrub and grassland communities: Several North Dakota grasslands include northern bedstraw. Northern bedstraw is an important associate of the bluegrass-little bluestem-needlegrass (Poa spp.-Schizachyrium scoparium-Achnatherum spp.) community type of eastern North Dakota's Oakville Prairie . In south-central North Dakota, northern bedstraw occurs in previously farmed or overgrazed Kentucky bluegrass communities, in shrubland communities dominated by silverberry (Elaeagnus commutata), and in tallgrass communities characterized by little bluestem, mat muhly (Muhlenbergia richardsonis), and switchgrass (Panicum virgatum) . In south-central North Dakota, northern bedstraw occupies the blue grama (Bouteloua gracilis)-sun sedge (Carex inops ssp. heliophila)- little bluestem vegetation type . Western Minnesota's blue grama-porcupine grass (Hesperostipa spartea), prairie dropseed (Sporobolus heterolepis)-little bluestem, big bluestem-northern reedgrass (Andropogon gerardii var. gerardii/Calamagrostis stricta ssp. inexpansa) tallgrass prairies often include northern bedstraw .
Northeast: Northeastern mixed oak woodlands are typical northern bedstraw habitat.
Northern bedstraw occurs in mixed oak woodlands in the Yale-Myers forest of Eastford, Connecticut, where eastern white pine (Pinus strobus), black oak (Q. velutina), white oak (Q. alba), northern red oak (Q. rubra), and sweet birch (Betula lenta) make up the overstory . In New York, oak (Quercus spp.), aspen, maple (Acer spp.), and beech (Fagus spp.) forests are described as northern bedstraw habitat . Northern bedstraw also occupies Mendon Ponds Park of Monroe County, New York, where water horsetail (Equisetum fluviatile), slender flatsedge (Cyperus bipartitus), and American chestnut (Castanea dentata) are typical .
Northwest: Fragrant bedstraw is a common understory species in numerous coniferous, deciduous, and mixed forests of the northwest.
Coniferous forests: In southeastern Alaska, fragrant bedstraw inhabits several Sitka spruce (Picea sitchensis), western hemlock, and mixed conifer habitat types . Fragrant bedstraw is characteristic of several productive Douglas-fir-dominated habitats of southwestern British Columbia . Fragrant bedstraw occurs in interior Douglas-fir (Pseudotsuga menziesii var. glauca), western redcedar-western hemlock, and montane spruce forests of the Kamloops Forest as well .
In Washington, fragrant bedstraw is common to several western hemlock forests. In the Gifford Pinchot National Forest, fragrant bedstraw indicates mesic sites in western hemlock/Pacific dogwood (Cornus nuttallii)/sweet after death (Achlys triphylla), western hemlock/devil's club (Oplopanax horridus)/western sword fern (Polystichum munitum), western hemlock/lady fern (Athyrium filix-femina), and western hemlock/American skunkcabbage (Lysichiton americanus) communities . In the Olympic National Forest, fragrant bedstraw is recognized in western hemlock/devil's club and western hemlock/western sword fern-threeleaf foamflower (Tiarella trifoliata) vegetation types .
In southwestern Oregon and northwestern California, fragrant bedstraw occurs in several community types characterized by the presence of Port-Orford-cedar (Chamaecyparis lawsoniana) and western hemlock or fir . Constancy of fragrant bedstraw is greater than 50% in Douglas-fir/salmonberry (Rubus spectabilis)/western sword fern, western hemlock/evergreen huckleberry (Vaccinium ovatum)/western sword fern, and Port-Orford-cedar/evergreen huckleberry/western sword fern forests of southwestern Oregon .
A diversity of overstory species associate with fragrant bedstraw in Idaho. In the Selway-Bitterroot Wilderness, fragrant bedstraw persists in 315- to 600-year-old western redcedar stands . In east-central Idaho, the presence of fragrant bedstraw identifies the Engelmann spruce/fragrant bedstraw habitat type. Other habitat types where fragrant bedstraw is important include Engelmann spruce/softleaf sedge (Carex disperma), grand fir/Rocky Mountain maple (Acer glabrum), grand fir/queencup beadlily (Clintonia uniflora), subalpine fir/claspleaf twistedstalk, and subalpine fir/queencup beadlily . The aforementioned habitat types are recognized in Montana and western Wyoming as well. Other overstory associates include, lodgepole pine, blue spruce (Picea pungens), Engelmann spruce, and subalpine fir .
Several forest types recognize fragrant bedstraw as an important understory species. From Montana to northwestern Wyoming, the Engelmann spruce/fragrant bedstraw habitat type is a topoedaphic climax on streams, seepages, benches, and swales between 6,100 and 8,200 feet (1,860-2,500 m) . Fragrant bedstraw is common in western larch (Larix occidentalis)- and whitebark pine-dominated forests of the northern Rockies . Constancy of fragrant bedstraw is between 95% and 100% in the spruce/fragrant bedstraw, subalpine fir/fragrant bedstraw, spruce/field horsetail, and subalpine fir/bluejoint reedgrass (Calamagrostis canadensis) habitat types in Montana .
Deciduous and mixed forests: Common deciduous canopy species in northwestern fragrant bedstraw habitats include aspen, poplar, alder, and dogwood (Cornus spp.) in the Northwest. In the taiga of interior Alaska, fragrant bedstraw is typical of mature balsam poplar/devil's club stands . In Alberta, researchers found fragrant bedstraw associated with and growing on decaying logs and stumps in 28-year-old, aspen-dominated boreal forests .
Fragrant bedstraw is frequent in red alder-Oregon ash/Himalayan blackberry/reed canarygrass (Alnus rubra-Fraxinus latifolia/R. discolor/Phalaris arundinacea) and California bay (Umbellularia californica)-Douglas-fir/vine maple (Acer circinatum)/western sword fern communities of the Umpqua River Valley . Atzet and others  describe fragrant bedstraw in ponderosa pine-California black oak (Q. kelloggii) and western hemlock-tanoak (Lithocarpus densiflora) vegetation of southwestern Oregon. Fragrant bedstraw occurs in riparian vegetation of the Trout Creek Mountains as well .
Riparian vegetation typical of Montana and southern Idaho includes fragrant bedstraw . Fragrant bedstraw is an important understory species in Rocky Mountain juniper/red-osier dogwood, Douglas-fir/red-osier dogwood, quaking aspen/bluejoint reedgrass, Bebb willow, and fleshy hawthorn (Crataegus succulenta) vegetation . In central and eastern Idaho, western Wyoming, and likely northern Utah, red-osier dogwood/fragrant bedstraw is a major community type at elevations below 6,595 feet (2,010 m) .
Southwest: Fragrant bedstraw is a typical understory species in several southwestern coniferous, deciduous, and mixed forest types.
Coniferous forests: Fragrant bedstraw is more than 50% constant but rarely occupies much coverage in redwood (Sequoia sempervirens)-western hemlock/evergreen huckleberry, redwood-western hemlock/salmonberry, redwood/western sword fern, and redwood-red alder/salmonberry vegetation associations in northwestern California and southwestern Oregon . In northwestern California's Klamath Mountains, fragrant bedstraw is highly visible in white fir/Pacific trillium (Trillium ovatum), white fir/American vetch (Vicia americana), and California red fir (Abies magnifica)/twinflower forest types . Fragrant bedstraw is also typical of giant sequoia (Sequoiadendron giganteum)-mixed conifer forests with white fir and incense-cedar (Calocedrus decurrens) . In California's Sacramento Ranger District, fragrant bedstraw is well represented in cold moist areas characterized by the white fir/burnet ragwort (Packera sanguisorboides) vegetation type .
In northern New Mexico and southern Colorado, fragrant bedstraw typifies the white fir/fragrant bedstraw riparian forest habitat type [3,68]. Hayward  describes fragrant bedstraw in the ponderosa pine-Douglas-fir-white fir vegetation association of Utah's Wasatch and Uinta mountains.
Deciduous and mixed forests: In southern California's montane coniferous forests, fragrant bedstraw associates with ponderosa pine, Jeffrey pine (Pinus jeffreyi), Coulter pine (P. coulteri), white fir, incense-cedar, and California black oak . Endangered walnut (Juglans spp.) forests of southern California are also fragrant bedstraw habitat. Southern California walnut (J. californica) and coast live oak (Q. agrifolia) make up the overstory and wild oat (Avena fatua) and fragrant bedstraw the understory . In the Humboldt-Toiyabe National Forest, fragrant bedstraw occupies several communities dominated by quaking aspen, red-osier dogwood, and/or willow (Salix spp.) . Kartesz  reports fragrant bedstraw in Nevada's California red fir forests.
In western Colorado, fragrant bedstraw is common to riparian montane and subalpine forests. Blue spruce-narrowleaf cottonwood/thinleaf alder (Alnus incana ssp. tenuifolia)-twinberry honeysuckle, white fir-blue spruce-narrowleaf cottonwood/Rocky Mountain maple, and subalpine fir-Engelmann spruce/thinleaf alder-twinberry honeysuckle are typical fragrant bedstraw communities . Fragrant bedstraw is a principal understory species in the blue spruce/red-osier dogwood habitat type of southern Colorado and northern New Mexico [3,68]. Fragrant bedstraw is 100% constant in the Engelmann spruce/sprucefir fleabane (Erigeron eximius) and blue spruce/sprucefir fleabane habitat types that occupy elevations of 8,000 feet (2,440 m) or more in northern Arizona's White Mountains and Plateau region .
North-central: Fragrant bedstraw is a conspicuous understory species in many forests in the northern Plains and Great Lake states.
Deciduous and mixed forests: In west-central North Dakota, fragrant bedstraw occurs in green ash-box elder forests . Fragrant bedstraw is important in Itasca Park, Minnesota, where deciduous sugar maple-basswood (Tilia americana) forests meet balsam fir-white spruce coniferous forests [36,61]. In northeastern Minnesota's hardwood forests with sugar maple, yellow birch (B. alleghaniensis), basswood, and white spruce, fragrant bedstraw frequency is 19% .
In cedar swamps of northern Wisconsin, fragrant bedstraw occupies glaciated lowland habitats where northern white-cedar (Thuja occidentalis) dominates but jack pine, black ash (Fraxinus nigra), balsam fir, paper birch, and American elm can be important. Fragrant bedstraw is also prominent in hardwood swamps in which eastern hemlock, sugar maple, and American beech (Fagus grandifolia) are most common .
Northeast: Many northeastern hardwood forests include fragrant bedstraw in the understory.
Deciduous and mixed forests: Fragrant bedstraw occurs in the central hardwood forests of southern Ohio characterized by an overstory of white oak, chestnut oak, and black oak . Lutz  describes fragrant bedstraw in northwestern Pennsylvania's hemlock-beech vegetation. In New York's Adirondack uplands, fragrant bedstraw is found in red maple, striped maple (Acer pensylvanicum), fir, yellow birch, and beech (Fagus spp.) mixed forests . Fragrant bedstraw in central Vermont occupies old-age hemlock-northern hardwood forests with sugar maple, American beech, white ash (Fraxinus americana), yellow birch, American elm, eastern hemlock, and basswood . Ross  describes fragrant bedstraw in eastern white pine forests of Strafford County, New Hampshire; eastern white pine, northern red oak, red maple, and bigtooth aspen dominate.
Similar vegetation associations are described in Canada. In Newfoundland, fragrant bedstraw is present in blackberry (Rubus spp.)/balsam fir, cinnamon fern (Osmunda cinnamomea)/black spruce, mountain alder-birch (Alnus viridis spp. crispa-Betula spp.), and blackberry/birch forest types . In the Lac des Deuz-Montagnes area of Quebec, fragrant bedstraw is important in swamp white oak (Q. bicolor) communities .
Southeast: Southeastern fragrant bedstraw habitats include hardwood and river bottom forests.Deciduous and mixed forests: Fragrant bedstraw occurs with low frequency in American beech-sugar maple and red spruce-Fraser fir (Picea rubens-Abies fraseri) communities in the southern Appalachian mountains of Tennessee and North Carolina . Fragrant bedstraw is also present in river bottom forests the Blood and Jonathan rivers in Kentucky. Overstory species in forests along the Blood River include sweetgum, overcup oak (Q. lyrata), river birch, red maple, and cherrybark oak (Q. pagoda). Forests lining the Jonathan River typically include pin oak (Q. palustris), red maple, green ash, and sycamore (Platanus occidentalis) .
|northern bedstraw||fragrant bedstraw|
|© 2005 Dan Tenaglia, missouriplants.com|
Bedstraw is a native perennial forb. Square stems and whorled leaves are characteristic [59,118]. Rhizome growth or schizocarp seed dispersal is bedstraw's method of spread [59,92,118].
Northern bedstraw: Northern bedstraw grows more erect than fragrant bedstraw, and is often between 7.9 and 31.5 inches (20-80 cm) tall. The multiple stems are mostly glabrous. Leaves are in whorls of 4 and measure 0.4 to 2.6 inches (1-6.5 cm) long by 2 to 12 mm wide [59,91,92,118]. Northern bedstraw's rhizomes are considered well developed. Fruits are typically 2 mm in diameter and glabrous to inconspicuously hairy . If hairs are present, they are short and without hooks [38,59,118]. Stevens  reports that 1,000 seeds weigh 0.6 g.
Fragrant bedstraw: Fragrant bedstraw is similar in size to northern bedstraw, but this species has weak branches that give rise to a scrambling or prostrate growth form. On the lower portion of the plant, hooked hairs concentrate at the stem angles [91,92]. Leaves are most often in whorls of 5 to 6 [59,118], but whorls of 4 are also possible . Leaves measure 0.6 to 2.6 inches (1.5-6 cm) long by 4 to 15 mm wide and smell of vanilla [59,92,118]. Rhizomes are slender . Seeds are coated with dense hooked hairs and are typically 1.5 to 2.2 mm in diameter [59,92,118].RAUNKIAER  LIFE FORM:
Breeding system: Bedstraw has perfect flowers .
Pollination: In a southeastern Minnesota pioneer cemetery site, a single collection of insects on northern bedstraw plants yielded 6 total insect species, 3 of which were unique to northern bedstraw . This study suggests that pollination of bedstraw is insect mediated.
Seed production: Northern bedstraw can produce large amounts of seed, but likely seed germination is secondary to vegetative reproduction as a means of surviving disturbances. Stevens  found 1 northern bedstraw plant produced 1,300 seeds. The focus plant was "well-developed," of "average" stature, and growing with "little competition" from other plants. Realizing the difficulty in distinguishing a single rhizomatous plant, the author evaluated a single stem for species with rhizomes .
Archibold  monitored seed inputs into burned areas of northern Saskatchewan. Seeds were trapped in trays of potting soil installed even with the ground. Trapping occurred in 1978 and 1979. Natural predation was not discouraged and trays were left for 1 year. Just 4 northern bedstraw germinated in 1979 in areas that experienced crown fires in the spring of 1977. The prefire community was dominated by white spruce, paper birch, and quaking aspen .
Seed dispersal: Fragrant bedstraw's seed with its dense covering of hooked hairs is better adapted for long-distance animal dispersal than northern bedstraw's largely glabrous seed. On an annually flooded gravel bar on a 5th order stream in Oregon's Cascade Range, researchers trapped 2.6 fragrant bedstraw seeds/m² although fragrant bedstraw occupied no coverage 3.3 feet (1 m) from trapping site .
In the Black Sturgeon boreal forests of northwestern Ontario, researchers compared the seed banks of uncut and harvested stands. Before encouraging seed banks to germinate in the greenhouse, all vegetative propagules were removed. White spruce, black spruce, balsam fir, quaking aspen, and paper birch dominated preharvest forests. Occurrence of fragrant bedstraw in collected seed rain was 0.2% in clearcut areas and 0.01% in partially-cut areas, although fragrant bedstraw did not occur in the above ground vegetation in the research area .
In northern Delaware and southern Pennsylvania, researchers calculated migration rates for fragrant bedstraw based on plant distances from an old-growth ecotone to the furthest plant or to the furthest occurrence where plants grew at 1/2 peak density. Fragrant bedstraw's migration rates were 0.87±0.55 (s x) m/year and 0.91±0.55 m/year based on the furthest 1/2 peak density and furthest individual calculations, respectively .
Seed banking: The amount of bedstraw seed recovered in seed bank studies varies with study location, collection timing, and degree of site disturbance. Most seed bank studies suggest a heavy reliance on vegetative reproduction.
Northern bedstraw: In ponderosa pine/common snowberry communities of southeastern Washington, researchers estimated from greenhouse germination trials that 50±80 (s) northern bedstraw seeds/m² were in spring seed banks and 133±103 seeds/m² were in fall seed banks. The coverage of northern bedstraw in the study area was 5% and constancy was 96% .
Following the 1988 Yellowstone fires, Clark  collected soil and seed samples from the most severely burned areas within several habitat types. Northern bedstraw was present in the postfire above-ground vegetation within the subalpine fir/pinegrass community but did not germinate in soil samples collected in the same area. These findings suggest northern bedstraw recovered vegetatively . From soil collected in central Saskatchewan's native porcupine grass and Montana wheatgrass prairies, Archibold  reported 42 northern bedstraw "root sprouts"/m².
Fragrant bedstraw: In old-growth deciduous forests of southwestern Quebec, researchers calculated a maximum of 25 fragrant bedstraw seeds/m² from soil samples collected in May. Sugar maple, striped maple, American beech, and white ash between 150 and 450 years old dominated the sites . Soil samples 4 inches (10 cm) deep from a total area of 2.3 m² were collected from early May to late August in Douglas-fir-grand fir forests of central Idaho and contained 23 total viable fragrant bedstraw seeds. The maximum seed density was 126/m². Most seed (87%) came from the top 2 inches (5 cm) of soil .
In xeric limestone prairies of Pennsylvania, researchers compared the vegetation and seed banks of forested, prairie, and prairie-edge sites. Fragrant bedstraw occurred in 2 edge plots and 3 forested plots, but no seed germinated from soil collected from any of the 3 sites .
On intermittently flooded (2- to 10-year flood intervals, typically) gravel bars, 36 fragrant bedstraw seeds/m² emerged from soil collected on 3rd order streams where coverage of fragrant bedstraw was 1%; on 5th order streams, 36.5 seeds/m² emerged from soil collected on sites with 1.4% fragrant bedstraw coverage. On annually flooded gravel bars, collected soil had 0.5 to 3.5 seeds/m² where coverage of fragrant bedstraw was 1% or less. Streams flowed in Oregon's Cascade Range .
The seed banks from undisturbed and disturbed communities in southwestern British Columbia reveal an increased density of fragrant bedstraw seed with increased disturbance levels. Characteristic species in undisturbed and slightly disturbed sites included Douglas-fir, western hemlock, western redcedar, maple, and red alder. Highly disturbed sites occurred within maintained right of ways. Fragrant bedstraw plants were present in each study site. The distribution of fragrant bedstraw along a disturbance gradient and within the soil profile is provided below .
|Site condition||Undisturbed (n=18)||
Low disturbance (n=11)
High disturbance (n=18)
|Total number of germinants||60||68||
|Mean seeds± SE/m²||20±6 (litter)||7±2
Ahlgren  compared seedling emergence from intact blocks of soil collected in late summer from severely burned and unburned sites. The author described the fire as recent, but time since fire was unclear. In the 270-year-old red pine stands of northeastern Minnesota, fragrant bedstraw occurred at 40% frequency on burned sites and 93% frequency on unburned sites. Based on greenhouse experiments, the author calculated that 10,890,000 seedlings/ha could germinate from burned soils and 218,000 seedlings/ha could emerge from unburned soils.
Germination: No literature addressed the germination rates of fragrant bedstraw, and the little literature addressing northern bedstraw germination reports different characteristics, making comparisons difficult. Seed collected in August of 1946 from Wisconsin prairie remnants had low germination percentages. Regardless of stratification, 15% of northern bedstraw seed germinated under greenhouse conditions . Unstratified northern bedstraw seed collected from remnant prairies of southern Wisconsin took 17 days to germinate and took 28 days to reach peak germination levels. A high number (likely > 2,000) of seedlings emerged per ounce of clean seed .
Seedling establishment/growth: Information regarding the early development and growth of bedstraw is lacking. One study, however, did examine underground growth of northern bedstraw. Nimlos and others  studied the rooting depths of understory species in ponderosa pine, Douglas-fir, and western larch forests near Missoula, Montana. Researchers injected radioactive iodine into the soil at known depths. The later detection of radioactive iodine in the plant suggested a rooting depth similar to the injection depth. Northern bedstraw reached rooting depths of 72 inches (1.8 m) on mesic sites. On drier sites, roots were concentrated in the top 24 inches (61 cm) of soil. The rooting depths of northern bedstraw on the 2 sites are shown below .
|Soil depth (inches)||6||12||24||36||48||60||72|
|Radioactive plants (%)||80 (n=10)||42 (n=19)||25 (n=20)||10 (n=20)||7 (n=14)||13 (n=15)||0|
|Soil depth (inches)||6||12||24||36||48||60||72|
|Radioactive plants (%)||30 (n=30)||50 (n=32)||26 (n=34)||29 (n=35)||12 (n=34)||6 (n=31)||18 (n=27)|
Asexual regeneration: Bedstraw reproduces asexually through rhizome production. After excavating multiple plants from alpine plant communities in the glaciated mountain ranges of south-central Alaska, researchers described northern bedstraw spread as "guerrilla" clonal growth. This type of asexual regeneration is characterized by daughter ramets arising from long rhizomes reaching beyond the parent plant's canopy . Stickney and Campbell  consider fragrant bedstraw's rhizomes to be more fire sensitive than those of northern bedstraw.SITE CHARACTERISTICS:
Northern bedstraw: Northern bedstraw often occupies stony slopes and meadows of Alaska and Canada  and meadows and damp slopes in the Southwest . In Michigan and Wisconsin, northern bedstraw is described in open oak, hickory, aspen woodlands, pine woodlands, fields, meadows, prairie remnants, fens, tamarack swamps, and thickets and along ditches, rivers, and lake banks [275,294]. In western Montana's mountain grasslands, northern bedstraw production was greater on southwestern exposures than on northeastern exposures .
Fragrant bedstraw: In the Intermountain West, moist woods and riparian areas are typical fragrant bedstraw habitat [59,280]. In more southwestern regions, fragrant bedstraw is restricted to mesic, shady sites [172,190]. In the Great Plains states, fragrant bedstraw rarely occupies moist prairie sites . Voss  describes fragrant bedstraw in deciduous, coniferous, and mixed forests as well as cedar swamps, fens, and river banks in Michigan. In the Gulf and Atlantic coast states, fragrant bedstraw is common to deciduous forests, fields, brush thickets, and roadsides [75,211].
Elevation: Several western regions report elevational ranges for northern and fragrant bedstraw.
|State, province, or region||Elevational range|
|Alberta||500-1,750 m |
|California||15-2,000 m |
|Colorado||1,520-3,050 m |
|Intermountain West||up to 2,700 m |
|New Mexico||1,830-3,050 m |
|Utah||1,650-3,100 m |
|State, province, or region||Elevational range|
|Adirondacks||290-900 m |
|Alberta||500-1,500 m |
|California||10-3,000 m |
|southern California||below 2,440 m |
|Colorado||1,980-2,740 m |
|Montana to northwestern Wyoming||1,860-2,500 m [50,148]|
|Montana's Gallatin National Forest (spruce/fragrant bedstraw HT)||854-2,151 m|
|central and eastern Montana (subalpine fir/fragrant bedstraw HT)||1,439-2,440 m |
|New Mexico||2,130-2,740 m |
|Utah||1,220-2,500 m |
Climate: A widely distributed species such as bedstraw must tolerate a wide range of climatic conditions. Semiarid and continental climates are typically described in conjunction with bedstraw. In the Taiga of interior Alaska, bedstraw persists in semiarid, continental climates where temperature extremes can reach lows of -60 °F (-51 °C) and highs of 100 °F (38 °C). Annual precipitation averages 11 inches (280 mm), and 70 inches (1,780 mm) of snow accumulation remains on the ground from mid-October through mid-May . In parts of northeastern Alberta, average summer temperatures are 56 °F (13.5 °C), and winter temperatures average 8.2 °F (-13 °C). A majority (9.5 inches (240 mm)) of precipitation falls in the summer with less (2.5 inches (64 mm)) precipitation in the winter months . In northeastern Oregon's bedstraw habitats, winters are cold and wet, and summers are hot and dry . In western North Dakota, temperature extremes between -49 °F (-45 °C) and 114 °F (45.5 °C) are possible, frost is typical 8 months of year, and the mean annual precipitation is 15 inches (380 mm). Rainfall in this area occurs predominantly (75%) from April through September .
Northern bedstraw: In Idaho fescue-bearded wheatgrass (Elymus caninus) grasslands of Montana's Bridger Mountain Range, coverage of northern bedstraw was greater on sites receiving increased snow levels. Sites were subjected to 6 years of snow levels measuring 23.6 inches (60 cm), 47.2 inches (120 cm), and 95 inches (240 cm). Coverage of northern bedstraw was 5.0±1.3% (s x) at snow levels of 24 inches (60 cm), 9.0±1.3% at 48 inches (120 cm), and 11.2±3.2% at 95 inches (240 cm). Flowering was delayed on sites with 95 inches (240 cm) of snow accumulation .
Soils: Bedstraw favors moist but well-drained soils and tolerates a range of acidities and textures.
Northern bedstraw: Deep mineral soils with sandy loam to loam textures are described in northern bedstraw habitats of Vancouver Island, British Columbia . In dry grasslands of Alberta, northern bedstraw soils have pH levels ranging from 4.7 at shallow depths to 8.6 at 25.6 inches (65 cm) below the soil surface . In southwestern North Dakota woodlands, soil pH ranged from 6.8 to 8.4 on sites where northern bedstraw occurred . Strausbaugh and Core  describe a rocky soil texture in northern bedstraw habitats of West Virginia.
Fragrant bedstraw: Soils described in fragrant bedstraw habitats on Vancouver Island, British Columbia, are acidic and nitrogen rich . In the subalpine fir/fragrant bedstraw habitat type of central and eastern Montana, soils range from neutral to strongly acidic . Fragrant bedstraw habitat in the Adirondacks has "higher" pH soils .SUCCESSIONAL STATUS:
A powerful windstorm in July of 1983 caused substantial tree mortality in northern pin oak- and eastern white pine-dominated forests of Anoka County, Minnesota. In the pine forest, more than 50% of the trees were removed from the canopy, and in the oak forests more than 30% of the trees were removed. After the storm, both bedstraw species showed short-lived increases in frequency. Northern bedstraw frequency was 33% in 1983, 35% in 1984, and 39% in 1985. Fragrant bedstraw frequency was 21.7% in 1983, 40.8% in 1984, and 33.3% in 1985. By 1990, frequency of both bedstraw species was lower than in 1983; frequencies were 24% and 14.2% for northern bedstraw and fragrant bedstraw, respectively .
Northern bedstraw: The following studies indicate that northern bedstraw tolerates a broad range of disturbances and persists in many communities deemed early-, mid-, or late seral. Likely the preference of certain successional staged communities relates to disturbance severity, site conditions, and/or community type.
General successional relationships: Northern bedstraw occupies 5% cover and is 96% constant in mid-successional ponderosa pine/common snowberry communities of southeastern Washington. These sites had not experienced any major disturbance in the last 90 years . In bunchberry (Cornus canadensis)-dominated sites of central Alaska, northern bedstraw was a principal species in both early and late seral communities . Northern bedstraw frequency and cover decreased with increased age of quaking aspen-dominated woodlands in the taiga of interior Alaska. In 50- to 70-year-old forests, northern bedstraw cover and frequency were 7% and 23%, respectively. In 130-year-old-stands, cover was 2% and frequency was 7% . Stringer  considers northern bedstraw common in "subclimax" boreal wildrye-dominated shrub savannahs in Banff and Jasper national parks. These high elevation communities found on steep south-facing slopes are maintained by frequent snow slides and rock falls. In subarctic northern Manitoba, researchers consider northern bedstraw typical of disturbed sites (roadsides, abandoned settlements, rights of ways, etc.) .
Different-aged river deposits of the Chena River near Fairbanks, Alaska, revealed increased frequency of northern bedstraw in younger communities. In 15-year-old willow stands and in 50- to 120-year-old balsam poplar stands, northern bedstraw cover was 3% to 4%. Northern bedstraw was not recorded in 220-year-old white spruce-black spruce forests or in "climax" black spruce/sphagnum communities. Freezing and thawing patterns were different for early and late seral communities and may have influenced northern bedstraw's distribution .
Light intensity relationships: The following information relates to northern bedstraw's light intensity preferences. Much of the following information addresses fragrant bedstraw's response to logging practices. While light intensity is indeed altered through logging operations, mechanical soil disturbances also occur and may influence findings.
Northern bedstraw coverage was greatest at intermediate light intensities, while frequency was greatest at low light intensities in red pine-dominated forests in north-central Minnesota. Study sites ranged from 5% to 95% of total sunlight, but cut-off values for intermediate and low light level categories are unknown .
Northern bedstraw persists in recently clearcut (6- to 12-year-old-stands) and mature lodgepole pine forests in the Lower Foothills of Alberta . Likewise, Crouch  reports northern bedstraw's presence on both uncut and clearcut moist sites within central Colorado's subalpine forests. In ponderosa pine/common snowberry vegetation of northeastern Oregon, northern bedstraw coverage and density significantly increased (p≤0.05) with canopy cover reductions [219,220]. However, in large clearcut areas (≥0.25 mile) of mixed conifer forests near Priest River in northern Idaho, Larsen [151,152] reports that northern bedstraw is removed from the community.
Small-scale disturbances: Northern bedstraw is well adapted to colonizing rodent mounds in prairie communities. In a northwestern Iowa big bluestem-indiangrass (Sorghastrum nutans) prairie, northern bedstraw occupied a greater proportion of Plains pocket gopher mounds than similar undisturbed quadrats. The proportion of mounds and undisturbed quadrats covered by northern bedstraw is given below .
|1 year-old-mound (n = 40)||5.85||undisturbed quadrat
(n = 49)
|2 year-old-mound (n = 40)||7.32||undisturbed quadrat
(n = 25)
In northern mixed-grass prairies of McPherson County, South Dakota, researchers compared the colonization of artificially constructed mounds in low slope, big bluestem-dominated and steep slope, little bluestem-dominated prairies. In big bluestem prairies, northern bedstraw abundance was greater on mounds 1, 3, and 5 years following mound creation. On little bluestem prairie sites, increased abundance on mounds occurred only the 1st year after mound creation. In the 3rd and 5th years, abundance of northern bedstraw was greater off mounds. Differences between mounded and nonmounded areas were not statistically significant .
Large-scale and/or multiple disturbances: Northern bedstraw commonly increases following canopy layer thinning and disturbance of soils. The same pattern exists following large-scale and/or multiple disturbances. In quaking aspen woodlands of northeastern British Columbia, the coverage of northern bedstraw was greatest in harvested and grazed areas; coverage of northern bedstraw was lowest on uncut sites. Harvesting occurred in the winter when soils were typically frozen, and the grazing treatment achieved 75% use of available forage. Results are provided below :
In northern Idaho Douglas-fir/ninebark communities, retrogressive studies compared sites with different disturbance histories. Disturbances included logging, grazing, burning, and combinations of these. The coverage of northern bedstraw was greatest on burned sites. However, samples sizes were low, time since disturbance was variable, and sites had soil type differences, so ascribing this finding to a fire effect is difficult. For more information see [43,44].
In several Canadian studies, northern bedstraw is important on burned sites. In the Selkirk Mountains of British Columbia, Shaw  lists northern bedstraw as important in the early reforestation stage following fire in western hemlock, quaking aspen, and lodgepole pine communities. Following stand-replacing fires in subalpine fir-spruce forests in northern British Columbia, northern bedstraw is among the important species in the resulting mountain grasslands . In coniferous forests of the eastern Rockies near Alberta's western border, northern bedstraw is most frequent in recently burned areas (10 to 20 years since fire). This study does not report an absence from later successional stages however .
Fragrant bedstraw: Like northern bedstraw, fragrant bedstraw tolerates early-, mid-, and late seral environmental conditions. However, many studies reveal a preference for diffuse canopy habitats and a tolerance of disturbances.
General successional relationships: The following studies describe research from various seral staged communities indicating the presence of or recent invasion by fragrant bedstraw.
After reviewing successional change and disturbance dynamics studies within western forests, McKenzie and others  classify fragrant bedstraw as a "release herb," one that responds positively to canopy removal or other disturbance. Fragrant bedstraw successfully colonized sites that were substantially disturbed in northwestern Connecticut. White pine forests were clearcut and then bulldozed to expose the mineral soil. Fragrant bedstraw seedlings identified by the presence of cotyledons likely came from seed produced by plants occupying nearby forested areas .
Researchers compared sites in Manitoba with different levels of land-use: urban, suburban, high-intensity rural (high density of crops with regular pesticide and fertilizer use), low-intensity rural (presence of forage crops without regular pesticide and fertilizer use), and relatively undisturbed sites. Fragrant bedstraw coverage decreased with increasing disturbance intensity. Undisturbed and low- and high-intensity rural sites had significantly (p<0.0001) more fragrant bedstraw coverage than urban or suburban sites .
A study of alluvial deposits along the McKenzie River in Oregon revealed the highest cover of fragrant bedstraw in the earliest seral community. On low floodplain areas dominated by red alder, fragrant bedstraw had 6% canopy cover. On high floodplains, grand fir replaced red alder after 30 to 70 years, and here fragrant bedstraw had 3% cover. In later seral stages dominated by Douglas-fir and western hemlock, fragrant bedstraw occupied 2% to 3% coverage. Coverage decreased to 1% in late seral western hemlock communities .
In north-central Idaho's western hemlock-western redcedar forests, northern bedstraw did not occur in the earliest seral community (burned 3 years prior), but was present in all others described as immature shrub to near climax communities . Similarly, in northern lower Michigan, studies in mature 2nd growth (55-82 years old) and disturbed (≤15 years old) stands revealed an association between fragrant bedstraw and disturbed mesic sites. Quaking aspen, sugar maple, and American beech dominated the mesic sites . In hybrid white spruce × Engelmann spruce forests of central British Columbia, fragrant bedstraw occurred in all forests 14 to 140 years old . Habeck  reports fragrant bedstraw in climax (315- to 600-year old stands) western redcedar forests in Idaho's Selway-Bitterroot Wilderness.
While the above studies suggest a tolerance of early-, mid-, and late seral conditions, the following studies indicate that preferences within a community type or area exist as well. In Douglas-fir forests, fragrant bedstraw frequency of occurrence was significantly (p< 0.01) greater in mature (80-195 years) forests than in old-growth (≥195 years) or young (< 80 years) forests in Oregon's Cascade Mountains. In western Washington's Douglas-fir forests, however, northern bedstraw frequency of occurrence was almost equal in mature and young forests, but was significantly lower (p< 0.01) in old-growth forests . In rich mesic forests of western Massachusetts, researchers found fragrant bedstraw frequency was significantly (p≤0.05) lower in more open sites . In central Idaho's Douglas-fir/ninebark habitat type, fragrant bedstraw is considered a major late seral species that decreases following logging and wildfire disturbances .
Light intensity relationships: Much of the following information addresses fragrant bedstraw's light intensity preference as a result of logging practices. While light intensity is indeed altered through logging operations, mechanical soil disturbances also occur and may influence findings. In general, fragrant bedstraw favors diffuse light over full sun or full shade conditions.
In mixed conifer forests of southeastern Oregon's Siskiyou Mountains, the percent cover of fragrant bedstraw was highest in sites receiving 25% to 60% full light. The range of full sunlight received and corresponding fragrant bedstraw coverage were as follows :
|Percentage of full light||0-3.5||3.5-6||6-11||11-25||25-60||60-100+|
Researchers compared old-growth, even-aged, and uneven-aged hardwood (sugar maple, basswood, yellow birch, and eastern hemlock) forests in northern Wisconsin and Michigan. Fragrant bedstraw coverage, frequency, and constancy were greater in uneven-aged forests where photosynthetically active radiation levels were significantly (α = 0.05) greater than in either other type . Likewise, in southern boreal forests of northeastern Minnesota, researchers established that on average, fragrant bedstraw occurred with similar density on postfire and postlogging sites aged 25 to 100 years. This finding suggests that canopy release was the most important factor in fragrant bedstraw occurrence within this time frame .
In boreal mixed woods of Thunder Bay, Ontario, fragrant bedstraw abundance increased in riparian areas adjacent to upland burned sites as compared to riparian areas next to undisturbed woodlands. Sites burned in the 1999 Nipigon Fire, and the study was published in 2003. The authors suggest increased light availability as the reason for increases in riparian sites dominated by red-osier dogwood and thinleaf alder . In black spruce forests of northeastern Ontario and western Quebec, fragrant bedstraw reached a high of 1.9% coverage on nutrient-rich logged sites where the stand age averaged 35.5 years and a high of 0.9% cover on nutrient-rich unlogged sites . Increases in fragrant bedstraw density were significant (p<0.01) following thinning treatments in giant sequoia groves of Tulare County, California . Frequency following the thinning reportedly decreased, however .
While fragrant bedstraw increases following canopy release predominate, Freedman and Habeck  found coverage and presence of fragrant bedstraw to be lower in treated (logged, burned, logged and burned) versus untreated Douglas-fir-, ponderosa pine-, and western larch-dominated forests in Swan Valley, Montana.
Other studies suggest that season of canopy removal and method of removal may affect the response of fragrant bedstraw. In northern Minnesota, researchers monitored understory vegetation changes for 2 years following winter and spring logging and 2 methods of harvest, full-tree logging (trees skidded intact) and tree-length logging (trees limbed and topped on site). All trees greater than 1 inch (2.5 cm) dbh were cut. On the tree-length logged site, piles were burned in July. Fire conditions included a high build-up index of 26, relative humidity of 45%, average temperature of 88 °F (31 °C), and an initial wind speed of 19 km/h with gusts of up to 48 km/h. Density of fragrant bedstraw was significantly lower (p=0.05) on untreated and tree-length burned sites in the 2nd posttreatment year. The density of fragrant bedstraw for each of the treatments is given below :
|Treatment||Control||Full-tree logging (winter)||Full-tree logging (spring)||Control||Full-tree logging (winter)||Full-tree logging (spring)||Tree-length logging (winter) & burning|
Major disturbance events: Fragrant bedstraw is often present in very early seral communities resulting from extreme weather events or volcanic activity. After the eruption of Mount St. Helens in Washington, snow and ice rapidly melted from the volcano sides. As water flowed, it collected rocks, debris, and organic materials. Massive amounts of material were deposited along and in the Muddy River. Following these events, researchers recorded fragrant bedstraw on stump bases with deposits of organic material, on root wads of uprooted trees, in moist depressions and sinks of the mudflow channel, and on sites with original soils covered with a layer of mudflow material. The frequency of fragrant bedstraw on the Muddy River was 6% .
A debris flow along a 2nd order stream in Douglas-fir, western hemlock, and red alder communities in Oregon's central Coast Range occurred in the winter of 1989 and 1990. Researchers monitored vegetation changes for 10 years following the event. Fragrant bedstraw percent constancy (or percent occurrence in all plots) was greatest the 2nd recovery year. The percent constancy of fragrant bedstraw during the succession of this area is provided below .
Bailey  revisited sites affected by the 1914-15 eruptions in northeastern
California's Lassen Volcanic National park in 1963. Fragrant bedstraw occurred
at the edge of aspen stands considered by the researcher to be "far from climax."
From areas reporting seasonal development of both northern and fragrant bedstraw, it appears that fragrant bedstraw development is slightly later than northern bedstraw's. The states or regions indicating flower or fruit set timing for bedstraw provide broad ranging dates to incorporate year-to-year variation in climate and wide regional distributions.
|State, region||Flowering dates|
|Blue Ridge Province||May-August |
|Great Plains states||June-September |
|northern Idaho||June-August |
|southeastern Illinois||flowering begins late May-late June, typically lasts 30 days |
|western Montana||1st bloom: early May-mid-July, end of blooming: late July-mid-August .|
|North Dakota||late May-early September |
|Utah's Wasatch Mountains||mid-June-early August |
|West Virginia||May-August |
Fruiting dates for northern bedstraw are between mid-July and mid-August in New England . In subarctic northern Manitoba, Staniforth and Scott  report that northern bedstraw had immature fruits as of mid-September. From data collected over 10 years in western Montana's mountain grasslands, Mueggler  indicated fruit dissemination occurred from early August through early September, and plants were dry by mid-September.
|State, region||Flowering dates|
|Atlantic and Gulf coast states||May-August |
|Blue Ridge Province||July-August |
|southern California||May-July |
|northern Idaho||June-August |
|southeastern Illinois||early June-late August |
|north-central Texas||June-July |
|West Virginia||May-September |
Fire regimes: A diversity of communities provide bedstraw habitat, and since fire regimes are dictated by the overstory community, bedstraw experiences a wide range of fire regimes. Davis and others , in a review, classify the spruce/fragrant bedstraw and subalpine fir/fragrant bedstraw habitat types as having "infrequent, severe fires with long-lasting effects." The same habitat types described on the Lolo National Forest, Montana, occupy moist environments that burn infrequently. The estimated fire return interval for these sites is 24 to 140 years . Both bedstraw species occupy northern spruce-fir forests that are typically maintained under moist conditions, and the fire return interval in these forests ranges from 35 to more than 200 years . Much shorter fire return intervals are reportedly tolerated by bedstraw as well. Northern bedstraw is typical in fescue-oatgrass mountain grasslands that are characterized by a fire return interval of less than 35 years . Fragrant bedstraw occupies southern California walnut woodlands of southern California that burn annually due to an increased presence of annual grasses .
The following table provides fire return intervals for plant communities and ecosystems where bedstraw is important. For further information, see the FEIS review of the dominant species listed below. This list may not be inclusive for all plant communities in which bedstraw occurs. Find further fire regime information for the plant communities in which these species may occur by entering the species' names in the FEIS home page under "Find Fire Regimes".
|Community or Ecosystem||Dominant Species||Fire Return Interval Range (years)|
|grand fir||Abies grandis||35-200 |
|maple-beech-birch||Acer-Fagus-Betula spp.||> 1,000|
|silver maple-American elm||Acer saccharinum-Ulmus americana||< 35 to 200|
|sugar maple||Acer saccharum||> 1,000|
|sugar maple-basswood||Acer saccharum-Tilia americana||> 1,000 |
|bluestem prairie||Andropogon gerardii var. gerardii-Schizachyrium scoparium||< 10 [142,200]|
|sagebrush steppe||Artemisia tridentata/Pseudoroegneria spicata||20-70 |
|basin big sagebrush||Artemisia tridentata var. tridentata||12-43 |
|mountain big sagebrush||Artemisia tridentata var. vaseyana||15-40 [14,37,184]|
|Wyoming big sagebrush||Artemisia tridentata var. wyomingensis||10-70 (40**) [272,291]|
|plains grasslands||Bouteloua spp.||< 35 [200,289]|
|blue grama-needle-and-thread grass-western wheatgrass||Bouteloua gracilis-Hesperostipa comata-Pascopyrum smithii||< 35 [200,227,289]|
|cheatgrass||Bromus tectorum||< 10 [202,281]|
|sugarberry-America elm-green ash||Celtis laevigata-Ulmus americana-Fraxinus pennsylvanica||< 35 to 200|
|Atlantic white-cedar||Chamaecyparis thyoides||35 to > 200|
|beech-sugar maple||Fagus spp.-Acer saccharum||> 1,000|
|black ash||Fraxinus nigra||< 35 to 200 |
|western juniper||Juniperus occidentalis||20-70|
|Rocky Mountain juniper||Juniperus scopulorum||< 35 |
|western larch||Larix occidentalis||25-350 [13,24,63]|
|yellow-poplar||Liriodendron tulipifera||< 35 |
|Great Lakes spruce-fir||Picea-Abies spp.||35 to > 200|
|northeastern spruce-fir||Picea-Abies spp.||35-200 |
|southeastern spruce-fir||Picea-Abies spp.||35 to > 200 |
|Engelmann spruce-subalpine fir||Picea engelmannii-Abies lasiocarpa||35 to > 200 |
|black spruce||Picea mariana||35-200|
|conifer bog*||Picea mariana-Larix laricina||35-200 |
|blue spruce*||Picea pungens||35-200 |
|red spruce*||Picea rubens||35-200 |
|pinyon-juniper||Pinus-Juniperus spp.||< 35 |
|whitebark pine*||Pinus albicaulis||50-200 [1,10]|
|jack pine||Pinus banksiana||<35 to 200 |
|Rocky Mountain lodgepole pine*||Pinus contorta var. latifolia||25-340 [23,24,262]|
|Sierra lodgepole pine*||Pinus contorta var. murrayana||35-200|
|Pacific ponderosa pine*||Pinus ponderosa var. ponderosa||1-47 |
|interior ponderosa pine*||Pinus ponderosa var. scopulorum||2-30 [12,19,154]|
|Arizona pine||Pinus ponderosa var. arizonica||2-15 [19,51,235]|
|red pine (Great Lakes region)||Pinus resinosa||10-200 (10**) [74,88]|
|red-white-jack pine*||Pinus resinosa-P. strobus-P. banksiana||10-300 [74,113]|
|eastern white pine||Pinus strobus||35-200|
|eastern white pine-eastern hemlock||Pinus strobus-Tsuga canadensis||35-200|
|eastern white pine-northern red oak-red maple||Pinus strobus-Quercus rubra-Acer rubrum||35-200|
|sycamore-sweetgum-American elm||Platanus occidentalis-Liquidambar styraciflua-Ulmus americana||< 35 to 200 |
|eastern cottonwood||Populus deltoides||< 35 to 200 |
|aspen-birch||Populus tremuloides-Betula papyrifera||35-200 [74,277]|
|quaking aspen (west of the Great Plains)||Populus tremuloides||7-120 [12,96,181]|
|black cherry-sugar maple||Prunus serotina-Acer saccharum||> 1,000 |
|mountain grasslands||Pseudoroegneria spicata||3-40 (10**) [11,12]|
|Rocky Mountain Douglas-fir*||Pseudotsuga menziesii var. glauca||25-100 [12,14,15]|
|coastal Douglas-fir*||Pseudotsuga menziesii var. menziesii||40-240 [12,186,221]|
|California mixed evergreen||Pseudotsuga menziesii var. menziesii-Lithocarpus densiflorus-Arbutus menziesii||< 35|
|California oakwoods||Quercus spp.||< 35 |
|oak-hickory||Quercus-Carya spp.||< 35|
|northeastern oak-pine||Quercus-Pinus spp.||10 to < 35 |
|oak-gum-cypress||Quercus-Nyssa-spp.-Taxodium distichum||35 to > 200 |
|southeastern oak-pine||Quercus-Pinus spp.||< 10 |
|coast live oak||Quercus agrifolia||2-75 |
|white oak-black oak-northern red oak||Quercus alba-Q. velutina-Q. rubra||< 35 |
|canyon live oak||Quercus chrysolepis||<35 to 200 |
|northern pin oak||Quercus ellipsoidalis||< 35 |
|Oregon white oak||Quercus garryana||< 35 |
|California black oak||Quercus kelloggii||5-30 |
|bur oak||Quercus macrocarpa||< 10 |
|oak savanna||Quercus macrocarpa/Andropogon gerardii-Schizachyrium scoparium||2-14 [200,277]|
|chestnut oak||Quercus prinus||3-8|
|northern red oak||Quercus rubra||10 to < 35|
|black oak||Quercus velutina||< 35|
|live oak||Quercus virginiana||10 to< 100 |
|little bluestem-grama prairie||Schizachyrium scoparium-Bouteloua spp.||< 35 |
|redwood||Sequoia sempervirens||5-200 [12,82,258]|
|western redcedar-western hemlock||Thuja plicata-Tsuga heterophylla||> 200 |
|eastern hemlock-yellow birch||Tsuga canadensis-Betula alleghaniensis||> 200 |
|western hemlock-Sitka spruce||Tsuga heterophylla-Picea sitchensis||> 200|
|mountain hemlock*||Tsuga mertensiana||35 to > 200 |
|elm-ash-cottonwood||Ulmus-Fraxinus-Populus spp.||< 35 to 200 [74,277]|
Northern and fragrant bedstraw: The postfire response is not always the same for northern and fragrant bedstraw in burned areas where both occur together.
Fire effects related to seasonality/severity: In the early 1960s, 17 wildfires burned in south-central New York. All but 1 of the fires burned in the spring, and sampling occurred 10 to 26 months following fire. Northern and fragrant bedstraw frequencies were significantly higher (p=0.01) on burned sites. Northern bedstraw averaged 2% frequency on unburned sites and 33% on burned sites within goldenrod (Solidago spp.)-poverty oatgrass habitats. Fragrant bedstraw averaged 2% frequency on unburned sites and 29% on burned sites in hardwood and mixed oak forests .
The postfire responses for northern and fragrant bedstraw were opposite in quaking aspen boreal forests of northeastern Alberta. Following a lightning-ignited spring wildfire, Lee  compared the immediate postfire seed banks and 2nd year postfire vegetation of unburned, "lightly" burned, and severely burned sites. Severely burned sites had all downed wood (≥ 7.9 inches (20 cm)) and the top 2.4 to 4 inches (6-10 cm) of organic material oxidized. Light burns partially oxidized small and mid-sized downed wood and just the top 0.8 inch (2 cm) of organic matter. Seed density estimates came from seedling and vegetative emergence techniques. Fragrant bedstraw rhizomes likely did not survive the fire while seed did, and the reverse was true for northern bedstraw. These findings may reflect different rhizome and seed heat tolerances for the 2 species or may indicate the occupation of different microsites where fire effects were different. Data are summarized below :
|Postfire characteristic||Mean seed density
|2nd postfire year coverage (%)|
Repeated fires: Sites in east-central Minnesota's oak savannas burned at frequencies of 0 to 19 years in a 20-year period. The coverage of fragrant bedstraw decreased with increased fire frequency; however, the r value for this relationship was -0.52 (significant at p < 0.10). Northern bedstraw coverage was not significantly changed by fire .
Northern bedstraw: Northern bedstraw recovers quickly following fire. Severe or growing season fires may result in decreased northern bedstraw coverage and/or frequency, but typically decreases are short lived.
Northern bedstraw is often mentioned as an important species in postburn communities of Canada. In the Selkirk Mountains of British Columbia, Shaw  lists northern bedstraw as a prominent herb in the early postfire reforestation of western hemlock, lodgepole pine, and quaking aspen forests. Seip and Bunnell  describe northern bedstraw in mountain grasslands resulting from stand-replacing fires in subalpine spruce forests of northern British Columbia. In coniferous forests of Alberta's eastern Rockies, northern bedstraw is most frequent in recently burned areas (10 to 20 years since fire) .
In interior Alaska's white spruce forests, Foote  visited sites burned between 6 months and 200 or more years ago. Northern bedstraw frequency was greatest but coverage was lowest on sites burned 6 months prior in a surface fire that scorched stems and killed some trees . One of the fires that was included in the previous postfire recovery chronosequence was the 1950 Porcupine River fire. Foote  investigated the postfire vegetation recovery 1, 4, 7, 10, 23, and 30 years following the fire. Northern bedstraw frequency and coverage were greatest in the 10th postfire year .
The following studies highlight fire effects that are likely a result of fire severity or seasonality. These fire characteristics are difficult to consider singly; studies listed in this section highlight severity or seasonality.
Fire effects related mainly to severity: Generally, northern bedstraw increases following low-severity fires. The postfire response of northern bedstraw to high-severity fires is less predictable. Researchers burned ponderosa pine-dominated forests in the fall on the Coeur d'Alene, Idaho, Indian Reservation. Different fire severity levels resulted. High-severity fires consumed 80% of the duff layer, and low-severity fires removed 40%. Comparisons between unburned and burned sites revealed northern bedstraw frequency and coverage were greatest on sites burned in low-severity fires and lowest on sites burned in high-severity fires. However, treatment differences were not significant (p<0.1) . See the Research Project Summary Understory recovery after low- and high-intensity fires in northern Idaho ponderosa pine forests for an extended report on this study.
In mixed oak forests of Eastford, Connecticut, researchers burned 2 sites in April. The 1st site burned in 1984, and the 2nd site burned in 1985. Fires burned under similar conditions; fuel moistures were between 18% and 28%, fire spread was slow (1m/min), and flame lengths were 11.8 inches (30 cm) or less. Within each site, portions burned more severely than others resulting in high mortality of the overstory. On the severely burned portion of site 1, 70% of the density and 60% of the basal area were removed. On the severely burned part of site 2, 95% of the density and basal area were removed. Northern bedstraw occurred only on burned sites. The density and frequency of northern bedstraw 7 to 8 years following these fires are shown below .
|Density (stems/ha)||Frequency (%)|
|Site||Dominants||intact overstory||no overstory||intact overstory||no
|1||eastern white pine
|2||northern red oak
Prefire vegetation was compared to lightly (1%-20% of litter and duff consumed and 0-few trees killed), moderately (21%-80% of litter and duff consumed and <90% of trees killed), and heavily (81%-100% of litter and duff consumed and >90% of trees killed) burned vegetation following a late August prescription fire in quaking aspen-dominated communities of northwestern Wyoming. Northern bedstraw produced less biomass before the fire than 3 years following the fire on lightly and heavily burned sites. Northern bedstraw productivity was less 12 years following the fire than before the fire [25,26]. See the Research Project Summary Vegetation recovery following a mixed-severity fire in aspen groves of western Wyoming for an extended report on this fire study.
Fall prescription fires burned quaking aspen communities of Colorado's Front Range. Fire severity was greater on plots with an understory of common juniper than on plots with an herbaceous understory. Northern bedstraw densities were significantly (p=0.05) greater 1 year following fire. Increases were greater on less severely burned plots. The differences for pre- and postburn northern bedstraw coverage and density are given below . See the Research Project Summary Vegetation changes following prescription fires in quaking aspen stands of Colorado's Front Range for an extended report on this fire study.
|Burn status||prefire (1980)||postfire (1982)||prefire (1980)||postfire (1982)|
(number of stems/0.1m)
Fire effects related mainly to seasonality: Dormant season fires (early spring or late fall) rarely cause decreases in northern bedstraw frequency and/or cover, but growing season (summer) fires may initially decrease northern bedstraw. In a central Saskatchewan rough fescue grassland, researchers compared the postfire recovery of northern bedstraw following spring (May 6), summer (June 26), and fall (October 8) prescription fires. In the 2nd postfire season, northern bedstraw density was lower for spring and summer burns than for unburned and fall burned sites . See the Research Project Summary Seasonal fires in Saskatchewan rough fescue prairie for an extended report on this fire study.
In central Alberta in 1972, almost pure, semimature quaking aspen stands burned in spring and fall prescription fires. Northern bedstraw coverage and frequency were greater on burned sites regardless of fire season or number of fires. Northern bedstraw coverage and frequency on burned sites, reburned sites, and unburned sites as of August 1978 are given below . See the Research Project Summary Understory recovery after burning and reburning quaking aspen stands in central Alberta for an extended report on this fire study.
Western snowberry-dominated communities southeast of Edmonton, Alberta, burned in spring prescription fires. Fires burned in early May of 1970 and 1971. The coverage of northern bedstraw was significantly greater on burned plots (p<0.05) 3 months following the fire. Researchers monitored vegetation for the next 2 growing seasons as well. Postfire results are below :
|Burn status||Unburned (n=125)||Burned (n=125)|
|Time since fire||0||3 months|
|Burn status||Unburned (n=23)||Burned (n=28)|
On the Namekagon River barrens of northern Wisconsin where jack pine and bur oak codominate, spring prescription fires burned. Northern bedstraw was common on both burned and unburned sites. On unburned sites, northern bedstraw averaged 89% frequency. On burned sites, frequency averaged 76%. An increased frequency of grasses following the fire may explain the slightly lower northern bedstraw frequency on burned sites .
McGee  compared early spring and late summer prescription fires in northwestern Wyoming's mountain big sagebrush communities. Two years after the fires, northern bedstraw coverage and frequency were greatest the 2nd postfire season on sites burned in the late summer. The coverage and frequency on unburned and spring burned sites were very similar .
In a fescue-oatgrass community of southern Alberta, researchers compared burned and unburned vegetation following a mid-December wildfire in 1997. The Granum wildfire burned when temperatures averaged 55 °F (13 °C), relative humidity was 17%, and winds were 19 to 25 mi/hr (30-40 km/hr) with gusts of 43.5 mi/hr (70 km/hr). Prefire fuel loads were unavailable, but nearby unburned sites had 900 kg/ha litter loadings. Postfire growing season precipitation was 46% greater than the long-term average. Northern bedstraw coverage was similar on interior burned plots and unburned plots 2 years following the fire. However, coverage was almost double on perimeter burn sites (those on the blackened side of fire line) when compared to unburned sites the 1st postfire year .
An early-spring prescription fire (May 2, 1972) stimulated northern bedstraw flowering on burned undisturbed mesic, highly-disturbed mesic, and on highly-disturbed wet to mesic prairie sites of northeastern Minnesota. Disturbances on the sites included grazing, sod production, and hay production but were discontinued approximately 15 years prior to the study. The fire occurred during periods of high humidity, virtually no wind, and wet to damp soils .
Repeated fires: The following studies report mixed postfire responses of northern bedstraw following multiple fires. Some report a tolerance of annual fires while others suggest that multiple fires followed by multiple years of rest are favored by northern bedstraw. Higgins and others  in a review suggest that northern bedstraw does not change or slightly decreases following periodic spring fires in the Northern Great Plains. In north-central South Dakota, northern bedstraw density was significantly greater (p<0.05) on northern mixed prairie plots burned annually for 3 consecutive years in the fall (October 5-17) than on unburned control plots .
In oak woodlands of east-central Minnesota's Cedar Creek Natural History Area, White  compared several burning schedules. All fires burned in the spring. However, particular overstory densities and soil series of the different sites were significantly (p≤0.05) correlated with northern bedstraw and could not be reliably related to burned sites .
Fragrant bedstraw: The following information suggests that fragrant bedstraw is not as fire tolerant as northern bedstraw. Fewer studies report increases in fragrant bedstraw following low-severity and dormant season fires than were reported for northern bedstraw. In north-central Idaho western hemlock-western redcedar habitats, fragrant bedstraw was absent 3 years following fire. Fire timing or severity are unknown. Steele and Geier-Hayes  consider fragrant bedstraw a major late-seral species in central Idaho's Douglas-fir/ninebark habitat type that decreases following logging and wildfires.
As time since fire increases however, the presence of fragrant bedstraw can decrease as well. In 1955 and 1956, Neiland  compared northwestern Oregon's mature (~300 years) western hemlock and Douglas-fir stands to sites that burned severely in 1933, 1939, and 1945. Fragrant bedstraw was absent from unburned forests but averaged 3% frequency on burned sites. In forests codominated by balsam fir, black spruce, and paper birch around Lake Duparquet, Quebec, fragrant bedstraw coverage was 1.9% on sites burned 26 years ago. On other sites that burned between 46 and 230 years ago, coverage of bedstraw varied from 0.1% to 0.5% .
The following studies highlight fire effects that are likely a result of fire severity or seasonality. These fire characteristics are difficult to consider singly; studies listed in this section highlight severity or seasonality.
Fire effects related mainly to severity: Fragrant bedstraw can survive low- and high-severity fires, but typically unburned frequencies or coverages are greater than those of burned sites. Following a "holocaustic" fire that killed all above ground vegetation, consumed all litter, and left bare mineral soil in the Pack River Valley of northern Idaho, fragrant bedstraw occurred on 5 of 18 sites and averaged 2% frequency . Following severe fires in 270-year-old red pine stands of northeastern Minnesota, fragrant bedstraw occurred at 40% frequency on burned sites and 93% frequency in unburned stands . See Seed banking for more information on this study.
In the Priest River Experimental Forest of northern Idaho, researchers compared the postfire regeneration following dry and moist prescription fires. Douglas-fir, western redcedar, and grand fir mixed forests were harvested and burned. The moist season burn occurred on June 1, 1989, when air temperatures were 69 °F to 76°F (21-24 °C), relative humidity was 43% to 50%, and winds were 1 to 8 mph. The dry season fire burned on September 13 and 14, 1989, when air temperatures were 54 °F to 77 °F (12-25 °C), relative humidity was 39% to 66%, and winds were 1 to 5 mph. Coverage of fragrant bedstraw decreased on both the moist and dry burn sites. There was no statistical analysis of the data. However, decreases were greater on dry burn sites. Coverage increased on unburned sites :
|Fire type||Prefire cover (%)||Postfire cover (%)|
Two forest sites within the Engelmann spruce-subalpine fir zone of central British Columbia were clearcut in the winter. One site burned in a low-severity prescribed fire the following fall. The coverage of fragrant bedstraw on the burned sites had not regained prefire levels by 11 years postfire. On logged unburned sites, increased fragrant bedstraw coverage lasted for 5 years following the disturbance . See the Research Project Summary Revegetation in a subalpine forest after logging and fire in central British Columbia for an extended report on this study. Cox  compared the recovery of fragrant bedstraw in clearcut and clearcut and burned Douglas-fir forests of Oregon's Coast Range. The slash burn produced a moderately severe fire (litter, duff, and woody debris consumed, but mineral soil color unchanged). No prefire data were available. Differences between burned and unburned plots 1 and 2 years following fire were negligible .
While decreases in fragrant bedstraw coverage and frequency following fire predominate, the frequency of fragrant bedstraw increased following low-severity, spring prescription fires in quaking aspen woodlands of southern Ontario. The frequency of fragrant bedstraw on unburned sites was 4%. The frequency 4 months postfire was 21% and a little over 1 year postfire was 12.5% . Following a mid-July crown fire near Missoula, Montana, fragrant bedstraw frequency had doubled from the 1st to the 2nd postfire year .
Fire effects related mainly to seasonality: Many of the following studies suggest that spring and fall fires may increase the frequency of fragrant bedstraw, while summer fires may decrease its frequency. Following spring fires in American beech-sugar maple and black oak-red maple forests in south-central New York, burned and unburned sites were compared. Fragrant bedstraw frequency was significantly higher (p=0.01) on burned sites; frequency on unburned sites was 2.4% and on burned sites was 28.6% .
In mixed conifer-hardwood forests of northeastern Minnesota, researchers assessed vegetation recovery in burned areas. Two sites dominated by black spruce, jack pine, and paper birch burned, one in late April and the other in mid-July. The late April fire occurred during high winds, leaving small unburned patches. Fragrant bedstraw frequency of occurrence on burned sites was over double that of unburned sites 3 years following the spring fire. The data collected on burned and unburned sites are summarized below :
|Fire||Unburned||Spring fire||Summer fire|
In white pine forests of Strafford County, New Hampshire, fall (1976) and spring (1977) prescription fires burned. The fires produced flame lengths of 3 to 24 inches (7.6-61 cm) and scorched trees at heights of 2 to 8 feet (0.6-2.4 m). Fragrant bedstraw was not on control plots and was not present on plots before the fire. However, it did occur following the fall and spring fires on white pine-dominated forests and following the spring fires in white pine mixed forests. Fragrant bedstraw plants on the burned plots resulted from seed germination. Plants on the spring-burned plots matured by late July and produced seed by the end of the growing season. The survival and/or development of plants on fall burned plots is unknown [42,226].
Fragrant bedstraw frequency decreased, but coverage was unchanged following a prescription fire in beetle-damaged white spruce forests of southern Alaska's Chuguch National Forest. The fire top-killed all overstory and understory vegetation in June of 1984. Prefire (1980) coverage of fragrant bedstraw was 2%, and frequency of occurrence was 24%. Seven years following the fire coverage remained 2% and the frequency of occurrence was 12% .
A prescription head fire within the Grand fir-Oregon boxwood (Paxistima myrsinites) habitat type of north-central Idaho also decreased the frequency of fragrant bedstraw. The fire burned mid-May of 1975 when temperatures were 82 °F (28 °C), relative humidity was 25%, and winds were negligible. Decreases in frequency were greater for sites that were grass seeded than unseeded sites following the fire. Statistical significance of the results was not addressed. Fragrant bedstraw frequency of occurrence is provided below :
|Time since fire||Prefire||3 months||1year||2 years|
|Burned & seeded
(# of occurrences/10 plots)
(# of occurrences/10 plots)
Repeated fires: The only study reporting fragrant bedstraw recovery following multiple fires indicates a tolerance of annual fires for up to 3 years. In southern Ohio hardwood forests, prescription fires burned some sites once and burned other sites for 3 consecutive years in March and April. Flame lengths were less than 20 inches (50 cm), and fire severity was low. The frequency of fragrant bedstraw increased by more than 10% on burned plots .
Hamilton's Research Papers (Hamilton 2006a, Hamilton 2006b) and the following Research Project Summaries provide further information on prescribed fire use and postfire response of many plant species including bedstraw:
However, burning bedstraw may increase its forage value as indicated by the following study. A tall grass prairie in eastern North Dakota burned in early May of 1966. Frequencies of northern bedstraw were the same on burned and unburned sites, but herbage production was much greater on burned sites. Statistical comparisons were not made. The results of this study are summarized below :
|Site condition||Herbage weight
|Calories/m²||% total (calories/m²)|
Northern bedstraw -
Livestock: Studies report conflicting responses of northern bedstraw to grazing. Several studies indicate an increased presence of northern bedstraw on sites grazed by livestock. The biomass of northern bedstraw was greater on grazed than ungrazed fescue grasslands of central Alberta. A decrease in grass yields was thought to facilitate northern bedstraw increases . In aspen stands of Colorado and Wyoming, northern bedstraw is constant on moderately grazed ranges, and its removal from grazed vegetation may indicate mismanagement . In rough fescue grasslands of southwestern Alberta, researchers tracked changes in the percent composition of northern bedstraw under different stocking rates and over a 32-year period. Northern bedstraw increased with length of grazing time but was relatively unaffected by stocking rates. Complete study results are shown below :
|Sampling times||1st 6 years of grazing
|Last 6 years of grazing
Others report decreases in northern bedstraw with livestock grazing, or increased utilization of northern bedstraw with increased lengths of grazing time. On Douglas-fir/ninebark habitat types near Moscow, Idaho, the production and frequency of northern bedstraw was greater on ungrazed than cattle grazed sites. Ungrazed sites were not closed to native ungulate grazing, and stocking rates in the area averaged 1 animal/13 ha for 20 years . On aspen ranges within the Black Mesa Experimental Forest of western Colorado, the utilization of northern bedstraw after 21 cattle-grazing days was 1%, after 38 days was 2%, and after 57 days was 6%. No utilization occurred after 78 days of use, but this was because most forbs on the site had senesced .
Native ungulates: The amount of northern bedstraw in elk, deer, mountain goat, and bighorn sheep diets is typically low, but season and/or stocking levels can increase utilization rates. Northern bedstraw made up a trace of winter mule deer diets in the Snowy Mountains of central Montana. After monitoring 96 feeding sites and analyzing 21 rumen samples, Kamps  found northern bedstraw constituted less than 0.5% of January diets and 1% of February diets. In a review of Rocky Mountain elk forage habits, Kufeld  considers northern bedstraw a least valuable forage plant. Least valuable forage is eaten by elk, but either makes up a small portion of the diet or is consumed in a much smaller proportion than is available. In June-collected elk feces from the Mount Saint Helens blast zone in southwestern Washington, just 0.1% of the total density was northern bedstraw .
Bentz and Woodard  consider northern bedstraw a secondary forage species for bighorn sheep in subalpine forests of southwestern Alberta. In the Sun River area of west-central Montana, 3 of 803 observed plant feeding instances by bighorn sheep were on northern bedstraw . The stomach contents of 27 mountain goats from the Crazy Mountains of Montana contained 0.9% northern bedstraw by volume (0.5% by weight) in the summer and just a trace of the volume (0.1% by weight) in the fall. These findings came from 5 stomachs collected in the summer and 18 collected in the fall .
In several Canadian National Parks, Stelfox  compared bighorn sheep diets on winter ranges from 1968 to 1970 where the frequency of northern bedstraw ranged from 80% to 100%. In Waterton Lakes National Park, northern bedstraw did not comprise any portion of sheep diets. Zero utilization was likely because ungulate stocking rates were low. In Banff, northern bedstraw made up 9.1% of bighorn sheep's diet composition and was utilized at 20% frequency. Ungulate stocking rates were moderate in Banff. In Jasper National Park, ungulate stocking rates were high, and northern bedstraw comprised 1.1% to 2.2% of sheep diets but was utilized at frequencies of 12% to 86%. Utilization was greatest in the spring in Jasper and in the summer in Banff, but some utilization occurred year round in both parks .
Omnivores: Researchers recorded high levels of northern bedstraw usage by black bears in interior Alaska. Young stems and leaves were present in the spring diet. From 23 stomach contents, northern bedstraw occurred at 17% frequency and constituted 10.2% mean volume. From 16 intestines, the frequency of northern bedstraw was 33% and mean volume was 15%. No scat samples contained northern bedstraw . Usage of northern bedstraw was less by black bears in the Rocky Mountains of southwestern Alberta, but research relied on scat samples alone. From scat collected in the summer of 1984 (n=22), northern bedstraw frequency of occurrence was 5%. Frequency was 8% in scat collected in the fall (n=13) of the same year .
Birds: Northern bedstraw may be important to breeding and ground foraging birds. Bird surveys in the Little Missouri National Grasslands of western North Dakota revealed heavy usage of ash woodlands where northern bedstraw is a prominent understory herb. Researchers conducted surveys from mid-May through mid-July in 1979, 1980, and 1981. The highest density of ground foragers and 531 nesting pairs/40 ha were in ash woodlands. Three bird species were exclusive to ash woodlands, and 6 species occurred with their highest densities in ash woodlands .
Insects: Findings from a single insect study indicate that northern bedstraw may be important to certain insect species. In a southeastern Minnesota pioneer cemetery site, a single collection of insects on northern bedstraw plants yielded 6 total insect species, 3 of which were unique to northern bedstraw. The insect species were not identified .
Fragrant bedstraw -
Native ungulates: While fragrant bedstraw has relatively low grazing value it is a valuable indicator of productive elk, deer, and moose habitat. On Vancouver Island, black-tailed deer ate the new growth of fragrant bedstraw in the spring and summer. Utilization was low given the abundance of the plant . Throughout a 3-year-long study of habitat selection by elk in western Montana, the subalpine/fragrant bedstraw habitat type was "strongly selected for." Selection described the use of a vegetation type that exceeded the availability of the type. Elk used this habitat predominantly for feeding, although fragrant bedstraw was not a utilized food source . Lonner  highlights moist sites within the same subalpine/fragrant bedstraw habitat type as very important elk summer range. In south-central Montana, the spruce/fragrant bedstraw habitat characterizes good year-round moose habitat and elk and deer winter range. The subalpine fir/fragrant bedstraw habitat in the same area receives moderate to heavy deer and elk use. Moose use valley bottom sites .
Other native mammals: Fragrant bedstraw may be an important rodent food source. In the Cascade foothills near Blue River, Oregon, 125-year-old Douglas-fir forests were logged and logged and burned. Fragrant bedstraw was an important herb in the 2nd postfire and postlogging years. The creeping vole increased in density on treated sites. The author considered increased vole densities and herbaceous understory vegetation to be related, as the vole feeds on the leaves and stems of shrubs and herbs .
Omnivores: Fragrant bedstraw identifies important grizzly bear habitat and is an important black bear food source. In the Bob Marshall Wilderness, Montana, the spruce/fragrant bedstraw habitat is ranked as the 2nd (out of 10) most important habitats for grizzly bears during the herbaceous foraging season (den emergence to July 31) and 3rd most important during the fruit foraging season (from August 1 to den entry). However, fragrant bedstraw was not listed as important grizzly bear food . In a review, Rogers and Allen  list fragrant bedstraw as 1 of several herbaceous species commonly found in the early spring black bear diets in northeastern Minnesota and Massachusetts.
Palatability/nutritional value: Few studies address the palatability and nutritional content of bedstraw. The lack of fragrant bedstraw's inclusion in nutritional studies may be due to its low palatability . Some have even suggested that bedstraw may be poisonous . Below are some specific findings regarding nutritional value of northern and fragrant bedstraw in various environments.
Northern bedstraw: Paulsen  found northern bedstraw produces 17 pounds of forage/acre in aspen communities of western Colorado. Herbage production of northern bedstraw taken from western Montana's mountain grasslands ranged from 19 to 72 kg/ha (dried). Production was greater on southwest exposures than on northeast exposures . Northern bedstraw on burned sites may have increased forage value. Following a spring fire in an eastern North Dakota tallgrass prairie, northern bedstraw herbage production was much greater on burned sites even though frequencies were the same on burned and unburned sites. The complete results of this study are summarized in the Fire Management Considerations section .
Fragrant bedstraw: Fragrant bedstraw collected in July from Hubbard Brook, New Hampshire's hardwood and boreal forests had the following nutritional composition :
|Content||2.1%||2.8%||0.2%||1.7%||0.3%||0.2%||318 ppm||109 ppm||294 ppm||20 ppm||12 ppm|
In south-central Montana, the subalpine fir/fragrant bedstraw habitat
provides important big game cover .
VALUE FOR REHABILITATION OF DISTURBED SITES:
Bedstraw may be valuable in the revegetation of abandoned mining sites. Northern bedstraw, although not directly seeded onto a coal mine spoil, was present at 20% to 75% frequency on a 31-year-old coal mine restoration site in southeastern Ohio . Fragrant bedstraw made up 1.1% of the vegetative cover on a 15- to 20-year-old abandoned coal surface mine in Campbell County, Tennessee .
Northern bedstraw is successfully transplanted using a sod relocation method.
Northern bedstraw survived when sod taken from undisturbed rough fescue grasslands
in Alberta was transplanted to a new site . During prairie restoration efforts
in northern Wisconsin, researchers found direct seeding of northern bedstraw to be
fairly successful but rated the transplanting success of seedlings in a sod form
and as 1-year-old transplants as excellent. Individual seedlings showed poor survival
in the field, but sod transplants survived even when there was no precipitation for
the 2 weeks following transplanting .
There were multiple distinct uses of the 2 bedstraw species by native people.
Northern bedstraw: The Gwich'in Athabaskan people of Fort Yukon, Alaska, used a poultice of northern bedstraw green shoots to treat general aches and pains. The same shoots in tea treated cold symptoms .
Ditidaht, indigenous people of the Pacific Northwest Coast, used fragrant bedstraw
as a rinse to enhance the thickness and luster of their hair. Fragrant bedstraw flowers
when dried were used as a perfume . This species is also used to flavor wines . In a
review, Turner and Bell  report that the Kwakiutl people of British Columbia rubbed
fragrant bedstraw on the skin to treat chest pains. Hellebore (Helleborus spp.)
roots were often applied following this preparation.
OTHER MANAGEMENT CONSIDERATIONS:
Northern bedstraw: A study of grassland sites dominated by native and nonnative species suggests that northern bedstraw may decrease in coverage on sites invaded by nonnative forbs. Tyser  found northern bedstraw coverage was 1.8% in timothy (Phleum pratense)-dominated sites, 0.6% in native fescue-dominated sites, and 0.2% on sites invaded by spotted knapweed (Centaurea maculosa). Likely the differences in coverage relate to changes in species dominance as all sites had homogeneous topography, slopes, aspects, and substrates.
Fragrant bedstraw: Several studies suggest that fragrant bedstraw can indicate environmental conditions and productive sites in several Pacific Northwest forests. In western Oregon and southwestern Washington, fragrant bedstraw is indicative of moist, well-drained sites in low to mid-elevation forests . The presence of fragrant bedstraw in riparian zones of central Oregon suggests high productivity sites for conifers . Fragrant bedstraw is also 1 of several understory species indicating productive Douglas-fir habitat in southwestern British Columbia .
An extensive study of trampling in montane grasslands and forests of western Montana, suggests fragrant has high resiliency. The trampling treatments were completed by 130- to 190-pound people wearing lug-soled boots. Seventy-five to 100 trampling passes per year reduced fragrant bedstraw's frequency of occurrence by 50% and coverage increased less than 10% from the end of the 1st trampling season (August) to the beginning of the 2nd trampling season (June). Long-term resilience was high however; fragrant bedstraw increased by more than 30% after given 3 years recovery time. Fragrant bedstraw recovered in 10 months from less than 41 trampling passes without losing more than 20% of pretrampling coverage. If trampled for 3 seasons and given a longer recovery time (3 years), fragrant bedstraw tolerates high trampling levels (≥1,200 passes). Whether or not these findings of human trampling can be related to large herbivore trampling is unknown .
1. Agee, James K. 1994. Fire and weather disturbances in terrestrial ecosystems of the eastern Cascades. Gen. Tech. Rep. PNW-GTR-320. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 52 p. (Everett, Richard L., assessment team leader; Eastside forest ecosystem health assessment; Hessburg, Paul F., science team leader and tech. ed., Volume III: assessment). 
2. Ahlgren, Clifford E. 1979. Emergent seedlings on soil from burned and unburned red pine forest. Minnesota Forestry Research Notes No. 273. St. Paul, MN: University of Minnesota, College of Forestry. 4 p. 
3. Alexander, Robert R. 1987. Classification of the forest vegetation of Colorado by habitat type and community type. Res. Note RM-478. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 14 p. 
4. Allman, Verl Phillips. 1953. A preliminary study of the vegetation in an exclosure in the chaparral of the Wasatch Mountains, Utah. Utah Academy Proceedings. 30: 63-78. 
5. Anderson, Murray L.; Bailey, Arthur W. 1979. Effect of fire on a Symphoricarpos occidentalis shrub community in central Alberta. Canadian Journal of Botany. 57: 2820-2823. 
6. Archibold, O. W. 1980. Seed input into a postfire forest site in northern Saskatchewan. Canadian Journal of Forest Research. 10: 129-134. 
7. Archibold, O. W. 1981. Buried viable propagules in native prairie and adjacent agricultural sites in central Saskatchewan. Canadian Journal of Botany. 59: 701-706. 
8. Archibold, O. W.; Ripley, E. A.; Delanoy, L. 2003. Effects of season of burning on the microenvironment of fescue prairie in central Saskatchewan. Canadian Field Naturalist. 117(2): 257-266. 
9. Armour, Charles D.; Bunting, Stephen C.; Neuenschwander, Leon F. 1984. Fire intensity effects on the understory in ponderosa pine forests. Journal of Range Management. 37(1): 44-48. 
10. Arno, Stephen F. 1976. The historical role of fire on the Bitterroot National Forest. Res. Pap. INT-187. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 29 p. 
11. Arno, Stephen F. 1980. Forest fire history in the Northern Rockies. Journal of Forestry. 78(8): 460-465. 
12. Arno, Stephen F. 2000. Fire in western forest ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 97-120. 
13. Arno, Stephen F.; Fischer, William C. 1995. Larix occidentalis--fire ecology and fire management. In: Schmidt, Wyman C.; McDonald, Kathy J., compilers. Ecology and management of Larix forests: a look ahead: Proceedings of an international symposium; 1992 October 5-9; Whitefish, MT. Gen. Tech. Rep. GTR-INT-319. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 130-135. 
14. Arno, Stephen F.; Gruell, George E. 1983. Fire history at the forest-grassland ecotone in southwestern Montana. Journal of Range Management. 36(3): 332-336. 
15. Arno, Stephen F.; Scott, Joe H.; Hartwell, Michael G. 1995. Age-class structure of old growth ponderosa pine/Douglas-fir stands and its relationship to fire history. Res. Pap. INT-RP-481. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 25 p. 
16. Atzet, Thomas; White, Diane E.; McCrimmon, Lisa A.; Martinez, Patricia A.; Fong, Paula Reid; Randall, Vince D., tech. coords. 1996. Field guide to the forested plant associations of southwestern Oregon. Technical Paper R6-NR-ECOL-TP-17-96. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. Available online: https://www.fs.fed.us /r6/siskiyou/guide.htm [2004, October 7]. 
17. Bailey, Arthur W. 1970. Barrier effect of the shrub Elaeagnus commutata on grazing cattle and forage production in central Alberta. Journal of Range Management. 23(4): 248-251. 
18. Bailey, Warren Hutchinson. 1963. Revegetation in the 1914-1915 devastated area of Lassen Volcanic National Park. Corvallis, OR: Oregon State University. 195 p. Dissertation. 
19. Baisan, Christopher H.; Swetnam, Thomas W. 1990. Fire history on a desert mountain range: Rincon Mountain Wilderness, Arizona, U.S.A. Canadian Journal of Forest Research. 20: 1559-1569. 
20. Baker, William L. 1989. Classification of the riparian vegetation of the montane and subalpine zones in western Colorado. The Great Basin Naturalist. 49(2): 214-228. 
21. Bakuzis, E. V; Hansen, H. L. 1962. Ecographs of herb species of Minnesota forest communities. Minnesota Forestry Notes. 118: 1-2. 
22. Bare, Janet E. 1979. Wildflowers and weeds of Kansas. Lawrence, KS: The Regents Press of Kansas. 509 p. 
23. Barrett, Stephen W. 1993. Fire regimes on the Clearwater and Nez Perce National Forests north-central Idaho. Final Report: Order No. 43-0276-3-0112. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory. 21 p. Unpublished report on file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. 
24. Barrett, Stephen W.; Arno, Stephen F.; Key, Carl H. 1991. Fire regimes of western larch - lodgepole pine forests in Glacier National Park, Montana. Canadian Journal of Forest Research. 21: 1711-1720. 
25. Bartos, D. L.; Mueggler, W. F. 1981. Early succession in aspen communities following fire in western Wyoming. Journal of Range Management. 34(4): 315-318. 
26. Bartos, Dale L.; Brown, James K.; Booth, Gordon D. 1994. Twelve years biomass response in aspen communities following fire. Journal of Range Management. 47: 79-83. 
27. Bellemare, Jesse; Motzkin, Glenn; Foster, David R. 2002. Legacies of the agricultural past in the forested present: an assessment of historical land-use effects on rich mesic forests. Journal of Biogeography. 29(10/11): 1401-1420. 
28. Bentz, Jerry A.; Woodard, Paul M. 1988. Vegetation characteristics and bighorn sheep use on burned and unburned areas in Alberta. Wildlife Society Bulletin. 16(2): 186-193. 
29. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. 
30. Biondini, M. E.; Steuter, A. A.; Grygiel, C. E. 1989. Seasonal fire effects on the diversity patterns, spatial distribution and community structure of forbs in the northern mixed prairie, USA. Vegetatio. 85: 21-31. 
31. Bork, Edward W.; Adams, Barry W.; Willms, Walter D. 2002. Resilience of foothills rough fescue, Festuca campestris, rangeland to wildfire. The Canadian Field-Naturalist. 116(1): 51-59. 
32. Bormann, F. H.; Buell, M. F. 1964. Old-age stand of hemlock-northern hardwood forest in central Vermont. Bulletin of the Torrey Botanical Club. 91(6): 451-465. 
33. Bradley, Anne F.; Noste, Nonan V.; Fischer, William C. 1992. Fire ecology of forests and woodlands in Utah. Gen. Tech. Rep. INT-287. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 128 p. 
34. Brown, Dalton Milford. 1941. Vegetation of Roan Mountain: a phytosociological and successional study. Ecological Monographs. 11: 61-97. 
35. Brumelis, G.; Carleton, T. J. 1989. The vegetation of post-logged black spruce lowlands in central Canada. II. Understory vegetation. Journal of Applied Ecology. 26: 321-339. 
36. Buell, Murray F.; Martin, William E. 1961. Competition between maple-basswood and fir-spruce communities in Itasca Park, Minnesota. Ecology. 42(2): 428-429. 
37. Burkhardt, Wayne J.; Tisdale, E. W. 1976. Causes of juniper invasion in southwestern Idaho. Ecology. 57: 472-484. 
38. Burrill, L. C. 1992. WEEDS--Catchweed bedstraw (Galium aparine L.). PNW 388. Corvallis, OR: Pacific Northwest Extension Service. 2 p. 
39. Butler, Jack; Goetz, Harold. 1984. Influence of livestock on the composition and structure of green ash communities in the Northern Great Plains. In: Noble, Daniel L.; Winokur, Robert P., eds. Wooded draws: characteristics and values for the Northern Great Plains: Symposium proceedings; 1984 June 12-13; Rapid City, SD. Great Plains Agricultural Council Publication No. 111. Rapid City, SD: South Dakota School of Mines and Technology, Biology Department: 44-49. 
40. Callow, J. Michael; Kantrud, Harold A.; Higgins, Kenneth F. 1992. First flowering dates and flowering periods of prairie plants at Woodworth, North Dakota. Prairie Naturalist. 24(2): 57-64. 
41. Carter, Christy Tucker; Ungar, Irwin A. 2002. Aboveground vegetation, seed bank and soil analysis of a 31-year-old forest restoration on coal mine spoil in southeastern Ohio. The American Midland Naturalist. 147(1): 44-59. 
42. Chapman, Rachel Ross; Crow, Garrett E. 1981. Application of Raunkiaer's life form system to plant species survival after fire. Bulletin of the Torrey Botanical Club. 108(4): 472-478. 
43. Cholewa, Anita F. 1977. Successional relationships of vegetational composition to logging, burning, and grazing in the Douglas-fir/Physocarpus habitat type of northern Idaho. Moscow, ID: University of Idaho. 65 p. [+ appendices]. Thesis. 
44. Cholewa, Anita F.; Johnson, Frederic D. 1983. Secondary succession in the Pseudotsuga menziesii/Physocarpus malvaceus association. Northwest Science. 57(4): 273-282. 
45. Christensen, E. M.; Clausen, J. J. (Jones); Curtis, J. T. 1959. Phytosociology of the lowland forests of northern Wisconsin. The American Midland Naturalist. 62(1): 232-247. 
46. Clambey, Gary K. 1992. Ecological aspects of the Knife River Indian Villages National Historic Site, west-central North Dakota. In: Smith, Daryl D.; Jacobs, Carol A., eds. Recapturing a vanishing heritage: Proceedings, 12th North American prairie conference; 1990 August 5-9; Cedar Falls, IA. Cedar Falls, IA: University of Northern Iowa: 75-78. 
47. Clark, David Lee. 1991. The effect of fire on Yellowstone ecosystem seed banks. Bozeman, MT: Montana State University. 115 p. Thesis. 
48. Clary, Warren P. 1983. Overstory-understory relationships: spruce-fir forests. In: Bartlett, E. T.; Betters, David R., eds. Overstory-understory relationships in western forests. Western Regional Research Publication No. 1. Fort Collins, CO: Colorado State University, Agriculture Experiment Station: 9-12. 
49. Cole, David N. 1988. Disturbance and recovery of trampled montane grassland and forests in Montana. Res. Pap. INT-389. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 37 p. 
50. Collins, Ellen I. 1984. Preliminary classification of Wyoming plant communities. Cheyenne, WY: Wyoming Natural Heritage Program/The Nature Conservancy. 42 p. 
51. Cooper, Charles F. 1961. Pattern in ponderosa pine forests. Ecology. 42(3): 493-499. 
52. Cormack, R. G. H. 1953. A survey of coniferous forest succession in the eastern Rockies. Forestry Chronicle. 29: 218-232. 
53. Corns, I. G. W.; Annas, R. M. 1986. Field guide to forest ecosystems of west-central Alberta. Edmonton, AB: Canadian Forestry Service, Northern Forestry Centre. 251 p. 
54. Corns, Ian G.; La Roi, George H. 1976. A comparison of mature with recently clear-cut and scarified lodgepole pine forests in the Lower Foothills of Alberta. Canadian Journal of Forest Research. 6(1): 20-32. 
55. Costello, David F. 1944. Important species of the major forage types in Colorado and Wyoming. Ecological Monographs. 14: 107-134. 
56. Cowan, Ian McTaggart. 1945. The ecological relationships of the food of the Columbian black-tailed deer, Odocoileus hemionus columbianus (Richardson), in the coast forest region of southern Vancouver Island, British Columbia. Ecological Monographs. 15(2): 110-139. 
57. Cox, Stephen William. 1970. Microsite selection of resident and invading plant species following logging and slash burning on Douglas fir clear-cuts in the Oregon Coast Range. Corvallis, OR: Oregon State University. 49 p. Thesis. 
58. Crane, M. F.; Habeck, James R.; Fischer, William C. 1983. Early postfire revegetation in a western Montana Douglas-fir forest. Res. Pap. INT-319. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 29 p. 
59. Cronquist, Arthur; Holmgren, Arthur H.; Holmgren, Noel H.; [and others]. 1984. Intermountain flora: Vascular plants of the Intermountain West, U.S.A. Vol. 4. Subclass Asteridae, (except Asteraceae). New York: The New York Botanical Garden. 573 p. 
60. Crouch, Glenn L. 1985. Effects of clearcutting a subalpine forest in central Colorado on wildlife habitat. Res. Pap. RM-258. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 12 p. 
61. Daubenmire, Rexford F. 1936. The "big woods" of Minnesota: its structure, and relation to climate, fire, and soils. Ecological Monographs. 6(2): 233-268. 
62. Daubenmire, Rexford. 1953. Notes on the vegetation of forested regions of the far northern Rockies and Alaska. Northwest Science. 27: 125-138. 
63. Davis, Kathleen M. 1980. Fire history of a western larch/Douglas-fir forest type in northwestern Montana. In: Stokes, Marvin A.; Dieterich, John H., technical coordinators. Proceedings of the fire history workshop; 1980 October 20-24; Tucson, AZ. Gen. Tech. Rep. RM-81. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 69-74. 
64. Davis, Kathleen M.; Clayton, Bruce D.; Fischer, William C. 1980. Fire ecology of Lolo National Forest habitat types. INT-79. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 77 p. 
65. De Grandpre, Louis; Gagnon, Daniel; Bergeron, Yves. 1993. Changes in the understory of Canadian southern boreal forest after fire. Journal of Vegetation Science. 4: 803-810. 
66. Del Tredici, Peter. 1977. The buried seeds of Comptonia peregrina, the sweet fern. Bulletin of the Torrey Botanical Club. 104(3): 270-275. 
67. Despain, Don G. 1973. Vegetation of the Big Horn Mountains, Wyoming, in relation to substrate and climate. Ecological Monographs. 43(3): 329-355. 
68. DeVelice, Robert L.; Ludwig, John A.; Moir, William H.; Ronco, Frank, Jr. 1986. A classification of forest habitat types of northern New Mexico and southern Colorado. Gen. Tech. Rep. RM-131. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 59 p. 
69. Diggs, George M., Jr.; Lipscomb, Barney L.; O'Kennon, Robert J. 1999. Illustrated flora of north-central Texas. Sida Botanical Miscellany No. 16. Fort Worth, TX: Botanical Research Institute of Texas. 1626 p. 
70. Doak, Daniel F.; Loso, Michael G. 2003. Effects of grizzly bear digging on alpine plant community structure. Arctic, Antarctic, and Alpine Research. 35(4): 421-428. 
71. Doyle, Kathleen M.; Knight, Dennis H.; Taylor, Dale L.; [and others]. 1998. Seventeen years of forest succession following the Waterfalls Canyon Fire in Grand Teton National Park, Wyoming. International Journal of Wildland Fire. 8(1): 45-55. 
72. Driscoll, K. G.; Arocena, J. M.; Massicotte, H. B. 1999. Post-fire soil nitrogen content and vegetation composition in sub-boreal spruce forests of British Columbia's central interior, Canada. Forest Ecology and Management. 121: 227-237. 
73. Ducey, Mark J.; Moser, W. Keith; Ashton, P. Mark S. 1996. Effect of fire intensity on understory composition and diversity in a Kalmia-dominated oak forest, New England, USA. Vegetatio. 123: 81-90. 
74. Duchesne, Luc C.; Hawkes, Brad C. 2000. Fire in northern ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 35-51. 
75. Duncan, Wilbur H.; Duncan, Marion B. 1987. The Smithsonian guide to seaside plants of the Gulf and Atlantic coasts from Louisiana to Massachusetts, exclusive of lower peninsular Florida. Washington, DC: Smithsonian Institution Press. 409 p. 
76. Dziadyk, Bohdan; Clambey, Gary K. 1983. Floristic composition of plant communities in a western Minnesota tallgrass prairie. In: Kucera, Clair L., ed. Proceedings, 7th North American prairie conference; 1980 August 4-6; Springfield, MO. Columbia, MO: University of Missouri: 45-54. 
77. Edgerton, Paul J. 1987. Influence of ungulates on the development of the shrub understory of an upper slope mixed conifer forest. In: Provenza, Frederick D.; Flinders, Jerran T.; McArthur, E. Durant, compilers. Proceedings--symposium on plant-herbivore interactions; 1985 August 7-9; Snowbird, UT. Gen. Tech. Rep. INT-222. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 162-167. 
78. Eis, S.; Craigdallie, D. 1980. Shore and landscape analysis of the western section of the Capital Regional District of British Columbia. BC-X-208. Victoria, BC: Canadian Forestry Service, Pacific Forest Research Centre. 43 p. 
79. Emmingham, W. H. 1972. Conifer growth and plant distribution under different light environments in the Siskiyou Mountains of southwestern Oregon. Corvallis, OR: Oregon State University. 50 p. Thesis. 
80. Evenden, Angela G. 1989. Ecology and distribution of riparian vegetation in the Trout Creek Mountains of southeastern Oregon. Corvallis, OR: Oregon State University. 156 p. Dissertation. 
81. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. 
82. Finney, Mark A.; Martin, Robert E. 1989. Fire history in a Sequoia sempervirens forest at Salt Point State Park, California. Canadian Journal of Forest Research. 19: 1451-1457. 
83. Flaccus, Edward; Ohmann, Lewis F. 1964. Old-growth northern hardwood forests in northeastern Minnesota. Ecology. 45(3): 448-459. 
84. Flora of North America Association. 2004. Flora of North America: The flora. [Online]. Flora of North America Association (Producer). Available: http://www.fna.org/FNA. 
85. Foote, M. Joan. 1983. Classification, description, and dynamics of plant communities after fire in the taiga of interior Alaska. Res. Pap. PNW-307. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 108 p. 
86. Foote, M. Joan. 1993. Revegetation following the 1950 Porcupine River Fire: 1950-1981. Fairbanks, AK: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station, Institute of Northern Forestry. 71 p. Review draft. 
87. Freedman, June D.; Habeck, James R. 1985. Fire, logging, and white-tailed deer interrelationships in the Swan Valley, northwestern Montana. In: Lotan, James E.; Brown, James K., compilers. Fire's effects on wildlife habitat--symposium proceedings; 1984 March 21; Missoula, MT. Gen. Tech. Rep. INT-186. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 23-35. 
88. Frissell, Sidney S., Jr. 1968. A fire chronology for Itasca State Park, Minnesota. Minnesota Forestry Research Notes No. 196. St. Paul, MN: University of Minnesota. 2 p. 
89. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; Lewis, Mont E.; Smith, Dixie R. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. 
90. Girard, Michele M.; Goetz, Harold; Bjugstad, Ardell J. 1989. Native woodland habitat types of southwestern North Dakota. Res. Pap. RM-281. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 36 p. 
91. Gleason, Henry A.; Cronquist, Arthur. 1991. Manual of vascular plants of northeastern United States and adjacent Canada. 2nd ed. New York: New York Botanical Garden. 910 p. 
92. Great Plains Flora Association. 1986. Flora of the Great Plains. Lawrence, KS: University Press of Kansas. 1392 p. 
93. Green, R. N.; Marshall, P. L.; Klinka, K. 1989. Estimating site index of Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) from ecological variables in southwestern British Columbia. Forest Science. 35(1): 50-63. 
94. Greene, H. C.; Curtis, J. T. 1950. Germination studies of Wisconsin prairie plants. The American Midland Naturalist. 43(1): 186-194. 
95. Greenlee, Jason M.; Langenheim, Jean H. 1990. Historic fire regimes and their relation to vegetation patterns in the Monterey Bay area of California. The American Midland Naturalist. 124(2): 239-253. 
96. Gruell, G. E.; Loope, L. L. 1974. Relationships among aspen, fire, and ungulate browsing in Jackson Hole, Wyoming. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 33 p. In cooperation with: U.S. Department of the Interior, National Park Service, Rocky Mountain Region. 
97. Habeck, James R. 1978. A study of climax western redcedar (Thuja plicata Donn.) forest communities in the Selway-Bitterroot Wilderness, Idaho. Northwest Science. 52(1): 67-76. 
98. Hadley, E. B.; Buccos, R. P. 1967. Plant community composition and net primary production within a native eastern North Dakota prairie. The American Midland Naturalist. 77: 116-127. 
99. Hadley, Elmer B. 1970. Net productivity and burning response of native eastern North Dakota prairie communities. The American Midland Naturalist. 84(1): 121-135. 
100. Hall, James B.; Hansen, Paul L. 1997. A preliminary riparian habitat type classification system for the Bureau of Land Management districts in southern and eastern Idaho. Tech. Bull. No. 97-11. Boise, ID: U.S. Department of the Interior, Bureau of Land Management; Missoula, MT: University of Montana, School of Forestry, Riparian and Wetland Research Program. 381 p. 
101. Halpern, Charles B.; Harmon, Mark E. 1983. Early plant succession on the Muddy River mudflow, Mount St. Helens, Washington. The American Midland Naturalist. 110(1): 97-106. 
102. Halverson, Nancy M., compiler. 1986. Major indicator shrubs and herbs on national forests of western Oregon and southwestern Washington. R6-TM-229. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 180 p. 
103. Hamilton, Evelyn; Peterson, Les. 2003. Response of vegetation to burning in a subalpine forest cutblock in central British Columbia: Otter Creek site. Res. Pap. 23. Victoria, BC: British Columbia Ministry of Forestry, Research Branch. 60 p. 
104. Hansen, Paul L.; Hoffman, George R.; Steinauer, Gerry A. 1984. Upland forest and woodland habitat types of the Missouri Plateau, Great Plains Province. In: Noble, Daniel L.; Winokur, Robert P., eds. Wooded draws: characteristics and values for the Northern Great Plains: Symposium proceedings; 1984 June 12-13; Rapid City, SD. Great Plains Agricultural Council Publ. No. 111. Rapid City, SD: South Dakota School of Mines and Technology, Biology Department: 15-26. 
105. Hansen, Paul L.; Pfister, Robert D.; Boggs, Keith; [and others]. 1995. Classification and management of Montana's riparian and wetland sites. Miscellaneous Publication No. 54. Missoula, MT: The University of Montana, School of Forestry, Montana Forest and Conservation Experiment Station. 646 p. 
106. Harmon, Janice M.; Franklin, Jerry F. 1995. Seed rain and seed bank of third- and fifth-order streams on the western slope of the Cascade Range. Res. Pap. PNW-RP-480. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 27 p. 
107. Harper, Karen A.; Macdonald, S. Ellen. 2002. Structure and composition of edges next to regenerating clear-cuts in mixed-wood boreal forest. Journal of Vegetation Science. 13: 535-546. 
108. Harrington, H. D. 1964. Manual of the plants of Colorado. 2d ed. Chicago: The Swallow Press, Inc. 666 p. 
109. Hatler, David F. 1972. Food habits of black bears in interior Alaska. Canadian Field-Naturalist. 86(1): 17-31. 
110. Hawk, G. M.; Zobel, D. B. 1974. Forest succession on alluvial landforms of the McKenzie River Valley, Oregon. Northwest Science. 48(4): 245-265. 
111. Hawk, Glenn Martin. 1977. Comparative study of temperate Chamaecyparis forests. Corvallis, OR: Oregon State University. 195 p. Dissertation. 
112. Hayward, C. Lynn. 1945. Biotic communities of the southern Wasatch and Uinta Mountains, Utah. The Great Basin Naturalist. 6(1-4): 1-124. 
113. Heinselman, Miron L. 1970. The natural role of fire in northern conifer forests. In: The role of fire in the Intermountain West: Symposium proceedings; 1970 October 27-29; Missoula, MT. Missoula, MT: Intermountain Fire Research Council: 30-41. In cooperation with: University of Montana, School of Forestry. 
114. Henderson, Jan A.; Peter, David H.; Lesher, Robin D.; Shaw, David C. 1989. Forested plant associations of the Olympic National Forest. R6-ECOL-TP 001-88. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 502 p. 
115. Hickman, James C., ed. 1993. The Jepson manual: Higher plants of California. Berkeley, CA: University of California Press. 1400 p. 
116. Higgins, Kenneth F.; Kruse, Arnold D.; Piehl, James L. 1989. Effects of fire in the Northern Great Plains. Ext. Circ. EC-761. Brookings, SD: South Dakota State University, Cooperative Extension Service, South Dakota Cooperative Fish and Wildlife Research Unit. 47 p. 
117. Hitchcock, C. Leo; Cronquist, Arthur. 1973. Flora of the Pacific Northwest. Seattle, WA: University of Washington Press. 730 p. 
118. Hitchcock, C. Leo; Cronquist, Arthur; Ownbey, Marion. 1959. Vascular plants of the Pacific Northwest. Part 4: Ericaceae through Campanulaceae. Seattle, WA: University of Washington Press. 510 p. 
119. Holcroft, Anne C.; Herrero, Stephen. 1991. Black bear, Ursus americanus, food habits in southwestern Alberta. Canadian Field-Naturalist. 105(3): 335-345. 
120. Holloway, Patricia S.; Alexander, Ginny. 1990. Ethnobotany of the Fort Yukon region, Alaska. Economic Botany. 44(2): 214-225. 
121. Holsten, Edward H.; Werner, Richard A.; Develice, Robert L. 1995. Effects of a spruce beetle (Coleoptera: Scolytidae) outbreak and fire on Lutz spruce in Alaska. Environmental Entomology. 24(6): 1539-1547. 
122. Hooven, Edward F. 1973. Response of the Oregon creeping vole to the clearcutting of a Douglas-fir forest. Northwest Science. 47(4): 256-264. 
123. Hopkins, Rick B.; Cassel, J. Frank; Bjugstad, Ardell J. 1986. Relationships between breeding birds and vegetation in four woodland types of the Little Missouri National Grasslands. Res. Pap. RM-270. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 12 p. 
124. Houston, Kent E.; Hartung, Walter J.; Hartung, Carol J. 2001. A field guide for forest indicator plants, sensitive plants, and noxious weeds of the Shoshone National Forest, Wyoming. Gen. Tech. Rep. RMRS-GTR-84. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 184 p. 
125. Huff, Mark Hamilton. 1984. Post-fire succession in the Olympic Mountains, Washington: forest vegetation, fuels, and avifauna. Seattle, WA: University of Washington. 235 p. Dissertation. 
126. Hulett, G. K.; Coupland, R. T.; Dix, R. L. 1966. The vegetation of dune sand areas within the grassland region of Saskatchewan. Canadian Journal of Botany. 44: 1307-1331. 
127. Hulten, Eric. 1968. Flora of Alaska and neighboring territories. Stanford, CA: Stanford University Press. 1008 p. 
128. Hutchinson, Todd F.; Sutherland, Steve. 2000. Fire and understory vegetation: a large-scale study in Ohio and a search for general response patterns in central hardwood forests. In: Yaussy, Daniel A., compiler. Proceedings: workshop on fire, people, and the central hardwoods landscape; 2000 March 12-14; Richmond, KY. Gen. Tech. Rep. NE-274. Newtown Square, PA: U.S. Department of Agriculture, Forest Service, Northeastern Research Station: 64-74. 
129. Jones, Stanley D.; Wipff, Joseph K.; Montgomery, Paul M. 1997. Vascular plants of Texas. Austin, TX: University of Texas Press. 404 p. 
130. Kamps, G. F. 1969. Whitetail deer and mule deer relationships in the Snowy Mountains of central Montana. Bozeman, MT: Montana State University. 59 p. Thesis. 
131. Kartesz, John T.; Meacham, Christopher A. 1999. Synthesis of the North American flora (Windows Version 1.0), [CD-ROM]. Available: North Carolina Botanical Garden. In cooperation with the Nature Conservancy, Natural Resources Conservation Service, and U.S. Fish and Wildlife Service [2001, January 16]. 
132. Kartesz, John Thomas. 1988. A flora of Nevada. Reno, NV: University of Nevada. 1729 p. [In 3 volumes]. Dissertation. 
133. Kilgore, Bruce M. 1971. Response of breeding bird populations to habitat changes in a giant sequoia forest. The American Midland Naturalist. 85(1): 135-152. 
134. Klinka, K.; Carter, R. E. 1980. Ecology and silviculture of the most productive ecosystems for growth of Douglas-fir in southwestern British Columbia. Land Management Report Number 6. Victoria, BC: Province of British Columbia, Ministry of Forests. 12 p. 
135. Klinka, K.; Krestov, P. V.; Chourmouzis, C. 2002. Classification and ecology of the mid-seral Picea mariana forests of British Columbia. Applied Vegetation Science. 5(2): 227-235. 
136. Klinka, K.; Wang, Q.; Carter, R. E. 1990. Relationships among humus forms, forest floor nutrient properties, and understory vegetation. Forest Science. 36(3): 564-581. 
137. Kovalchik, Bernard L.; Hopkins, William E.; Brunsfeld, Steven J. 1988. Major indicator shrubs and herbs in riparian zones on national forests of central Oregon. R6-ECOL-TP-005-88. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 159 p. 
138. Kramer, Neal B.; Johnson, Frederic D. 1987. Mature forest seed banks of three habitat types in central Idaho. Canadian Journal of Botany. 65: 1961-1966. 
139. Krefting, Laurits W.; Ahlgren, Clifford E. 1974. Small mammals and vegetation changes after fire in a mixed conifer-hardwood forest. Ecology. 55: 1391-1398. 
140. Kruse, Arnold D.; Higgins, Kenneth F. 1998. Effects of prescribed fire upon wildlife habitat in northern mixed-grass prairie. In: Alexander, M. E.; Bisgrove, G. F., tech. coords. The art and science of fire management: Proceedings of the 1st Interior West Fire Council annual meeting and workshop; 1988 October 24-27; Kananaskis Village, AB. Information Report NOR-X-309. Edmonton, AB: Forestry Canada, Northwest Region, Northern Forestry Centre: 182-193. 
141. Krzic, M.; Newman, R. F.; Broersma, K. 2003. Plant species diversity and soil quality in harvested and grazed boreal aspen stands of northeastern British Columbia. Forest Ecology and Management. 182: 315-325. 
142. Kucera, Clair L. 1981. Grasslands and fire. In: Mooney, H. A.; Bonnicksen, T. M.; Christensen, N. L.; Lotan, J. E.; Reiners, W. A., technical coordinators. Fire regimes and ecosystem properties: Proceedings of the conference; 1978 December 11-15; Honolulu, HI. Gen. Tech. Rep. WO-26. Washington, DC: U.S. Department of Agriculture, Forest Service: 90-111. 
143. Kuchler, A. W. 1964. United States [Potential natural vegetation of the conterminous United States]. Special Publication No. 36. New York: American Geographical Society. 1:3,168,000; colored. 
144. Kudish, Michael. 1992. Adirondack upland flora: an ecological perspective. Saranac, NY: The Chauncy Press. 320 p. 
145. Kufeld, Roland C. 1973. Foods eaten by the Rocky Mountain elk. Journal of Range Management. 26(2): 106-113. 
146. La Roi, George Henri. 1964. An ecological study of the boreal spruce-fir forests of the North American taiga. Durham, NC: Duke University. 397 p. Dissertation. 
147. Lacey, John; Mosley, John. 2002. 250 plants for range contests in Montana. MONTGUIDE MT198402 AG 6/2002. Range E-2 (Misc.). Bozeman, MT: Montana State University, Extension Service. 4 p. 
148. Lackschewitz, Klaus. 1991. Vascular plants of west-central Montana--identification guidebook. Gen. Tech. Rep. INT-227. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 648 p. 
149. Lamb, Eric G.; Mallik, Azim U.; Mackereth, Robert W. 2003. The early impact of adjacent clearcutting and forest fire on riparian zone vegetation in northwestern Ontario. Forest Ecology and Management. 177: 529-538. 
150. Langenheim, Jean H. 1962. Vegetation and environmental patterns in the Crested Butte Area, Gunnison County, Colorado. Ecological Monographs. 32(3): 249-285. 
151. Larsen, J. A. 1922. Effect of removal of the virgin white pine stand upon the physical factors of site. Ecology. 3(4): 302-305. 
152. Larsen, J. A. 1924. Some factors affecting reproduction after logging in northern Idaho. Journal of Agricultural Research. 28(11): 1149-1157. 
153. Laughlin, Daniel C. 2003. Lack of native propagules in a Pennsylvania, USA, limestone prairie seed bank: futile hopes for a role in ecological restoration. Natural Areas Journal. 23(2): 158-164. 
154. Laven, R. D.; Omi, P. N.; Wyant, J. G.; Pinkerton, A. S. 1980. Interpretation of fire scar data from a ponderosa pine ecosystem in the central Rocky Mountains, Colorado. In: Stokes, Marvin A.; Dieterich, John H., technical coordinators. Proceedings of the fire history workshop; 1980 October 20-24; Tucson, AZ. Gen. Tech. Rep. RM-81. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 46-49. 
155. Lawrence, Donna L.; Romo, J. T. 1995. Tree and shrub communities of wooded draws near the Matador Research Station in southern Saskatchewan. The Canadian Field Naturalist. 108(4): 397-412. 
156. Lawrence, George; Biswell, Harold. 1972. Effect of forest manipulation on deer habitat in giant sequoia. Journal of Wildlife Management. 36(2): 595-605. 
157. Leckie, Sara; Vellend, Mark; Bell, Graham; [and others]. 2000. The seed bank in an old-growth, temperate deciduous forest. Canadian Journal of Botany. 78(2): 181-192. 
158. Lee, Philip. 2004. The impact of burn intensity from wildfires on seed and vegetative banks, and emergent understory in aspen-dominated boreal forests. Canadian Journal of Botany. 82(10): 1468-1480. 
159. Lee, Philip; Sturgess, Kelly. 2002. Assemblages of vascular plants on logs and stumps within 28-year-old aspen-dominated boreal forests. In: Laudenslayer, William F., Jr.; Shea, Patrick J.; Valentine, Bradley E.; Weatherspoon, C. Phillip; Lisle, Thomas E., tech. coords. Proceedings of the symposium on the ecology and management of dead wood in western forests; 1999 November 2-4; Reno, NV. Gen. Tech. Rep. PSW-GTR-181. Albany, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Research Station: 369-380. 
160. Leege, Thomas A.; Godbolt, Grant. 1985. Herbaceous response following prescribed burning and seeding of elk range in Idaho. Northwest Science. 59(2): 134-143. 
161. Lesica, Peter. 2001. Recruitment of Fraxinus pennsylvanica (Oleaceae) in eastern Montana woodlands. Madrono. 48(4): 286-292. 
162. Lloyd, D.; Angove, K.; Hope, G.; Thompson, C. 1990. A guide to site identification and interpretation for the Kamloops Forest Region. Part 1. Land Management Handbook No. 23. Victoria, BC: British Columbia Ministry of Forests, Research Branch. 191 p. 
163. Lonner, Terry. 1975. Elk and logging--an update. Montana Outdoors. 6(4): 38-42. 
164. Losensky, Jack. 1987. A strategy to implement ecosystem maintenance burning on the Lolo National Forest. Missoula, MT: U.S. Department of Agriculture, Forest Service, Lolo National Forest. 89 p. Unpublished report on file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. 
165. Lura, Charles L.; Barker, William T.; Nyren, Paul E. 1988. Range plant communities of the Central Grasslands Research Station in south central North Dakota. Prairie Naturalist. 20(4): 177-192. 
166. Lutz, H. J. 1930. The vegetation of Heart's Content, a virgin forest in northwestern Pennsylvania. Ecology. 11(1): 2-29. 
167. Mace, Richard D. 1986. Analysis of grizzly bear habitat in the Bob Marshall Wilderness, Montana. In: Contreras, Glen P.; Evans, Keith E., compilers. Proceedings--grizzly bear habitat symposium; 1985 April 30 - May 2; Missoula, MT. Gen. Tech. Rep. INT-207. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station: 136-149. 
168. Mahony, Thomas M.; Stuart, John D. 2000. Old-growth forest associations in the northern range of coastal redwood. Madrono. 47(1): 53-60. 
169. Manning, Mary E.; Padgett, Wayne G. 1995. Riparian community type classification for Humboldt and Toiyabe National Forests, Nevada and eastern California. R4-Ecol-95-01. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Region. 306 p. 
170. Marcum, C. Les. 1975. Summer-fall habitat selection and use by a western Montana elk herd. Missoula, MT: University of Montana. 188 p. Dissertation. 
171. Martin, Jon Randall. 1989. Vegetation and environment in old growth forests of northern southeast Alaska. Tempe, AZ: Arizona State University. 221 p. Thesis. 
172. Martin, William C.; Hutchins, Charles R. 1981. A flora of New Mexico. Volume 2. Germany: J. Cramer. 2589 p. 
173. Maryland Department of Natural Resources. 2003. Rare, threatened, and endangered plants of Maryland, [Online]. In: Endangered species--endangered plants. Anapolis, MD: Maryland Department of Natural Resources, Wildlife and Heritage Service, Natural Heritage Program (Producer). Available: http://dnrweb.dnr.state.md.us/download/rteplants.pdf [2005, June 15]. 
174. Massachusetts Division of Fisheries and Wildlife. 2002. Massachusetts list of endangered, threatened and special concern species, [Online]. In: Official state rare species list. Westborough, MA: Natural Heritage and Endangered Species Program (Producer). Available: http://www.mass.gov/dfwele/dfw/nhesp/nhrare.htm [2005, March 18]. 
175. Matlack, Glenn R. 1994. Plant species in a mixed-history forest landscape in eastern North America. Ecology. 75(5): 1491-1502. 
176. McGee, Ann; Feller, M. C. 1993. Seed banks of forested and disturbed soils in southwestern British Columbia. Canadian Journal of Botany. 71: 1574-1583. 
177. McGee, John M. 1977. Effects of prescribed burning on a sagebrush ecosystem in northwestern Wyoming. Final report: Cooperative Agreement No. 16-675-CA. Laramie, WY: University of Wyoming. 134 p. 
178. McKenzie, Donald; Halpern, Charles B.; Nelson, Cara R. 2000. Overstory influences on herb and shrub communities in mature forests of western Washington, U.S.A. Canadian Journal of Forest Research. 30(10): 1655-1666. 
179. Meades, W. J.; Moores, L. 1989. Forest site classification manual: A field guide to the Damman forest types of Newfoundland. Forest Resources Development Agreement FRDA Report 003. St. Johns, NF: Environment Canada. 295 p. 
180. Meidinger, D.; Lewis, T.; Kowall, R. 1986. Biogeoclimatic zones and subzones of the northern portion of the Mackenzie Timber Supply Area, British Columbia. In: Northern Fire Ecology Project: Northern Mackenzie Timber Supply Area. Victoria, BC: Province of British Columbia, Ministry of Forests. 44 p. 
181. Meinecke, E. P. 1929. Quaking aspen: A study in applied forest pathology. Tech. Bull. No. 155. Washington, DC: U.S. Department of Agriculture. 34 p. 
182. Merrill, Evelyn H.; Callahan-Olson, Angela; Raedeke, Kenneth J.; [and others]. 1995. Elk (Cervus elaphus roosevelti) dietary composition and quality in the Mount St. Helens blast zone. Northwest Science. 69(1): 9-18. 
183. Meyer, Marvis I. 1985. Classification of native vegetation at the Woodworth Station, North Dakota. Prairie Naturalist. 17(3): 167-175. 
184. Miller, Richard F.; Rose, Jeffery A. 1995. Historic expansion of Juniperus occidentalis (western juniper) in southeastern Oregon. The Great Basin Naturalist. 55(1): 37-45. 
185. Moffatt, S. F.; McLachlan, S. M. 2004. Understorey indicators of disturbance for riparian forests along an urban-rural gradient in Manitoba. Ecological Indicators. 4: 1-16. 
186. Morrison, Peter H.; Swanson, Frederick J. 1990. Fire history and pattern in a Cascade Range landscape. Gen. Tech. Rep. PNW-GTR-254. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 77 p. 
187. Moss, E. H. 1955. The vegetation of Alberta. Botanical Review. 21(9): 493-567. 
188. Mueggler, Walter F. 1983. Variation in production and seasonal development of mountain grasslands in western Montana. Research Paper INT-316. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 16 p. 
189. Muldavin, Esteban H.; De Velice, Robert L.; Ronco, Frank, Jr. 1996. A classification of forest habitat types: southern Arizona and portions of the Colorado Plateau. RM-GTR-287. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 130. 
190. Munz, Philip A. 1974. A flora of southern California. Berkeley, CA: University of California Press. 1086 p. 
191. Myers, Ronald L. 2000. Fire in tropical and subtropical ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 161-173. 
192. Neiland, Bonita J. 1958. Forest and adjacent burn in the Tillamook Burn area of northwestern Oregon. Ecology. 39(4): 660-671. 
193. Nimlos, Thomas J.; Van Meter, Wayne P.; Daniels, Lewis A. 1968. Rooting patterns of forest understory species as determined by radioiodine absorption. Ecology. 49(6): 1145-1151. 
194. Nuzzo, Victoria. 1978. Propagation and planting of prairie forbs and grasses in southern Wisconsin. In: Glenn-Lewin, David C.; Landers, Roger Q., Jr., eds. Proceedings, 5th Midwest prairie conference; 1976 August 22-24; Ames, IA. Ames, IA: Iowa State University: 182-189. 
195. Outcalt, Kenneth Wayne; White, Edwin H. 1981. Phytosociological changes in understory vegetation following timber harvest in northern Minnesota. Canadian Journal of Forest Research. 11: 175-183. 
196. Pabst, Robert J.; Spies, Thomas A. 2001. Ten years of vegetation succession on a debris-flow deposit in Oregon. Journal of the American Water Resources Association. 37(6): 1693-1708. 
197. Palmer, Michael W.; McAlister, Suzanne D.; Arevalo, Jose Ramon; DeCoster, James K. 2000. Changes in the understory during 14 years following catastrophic windthrow in two Minnesota forests. Journal of Vegetation Science. 11(6): 841-854. 
198. Patterson, Patricia A.; Neiman, Kenneth E.; Tonn, Jonalea. 1985. Field guide to forest plants of northern Idaho. Gen. Tech. Rep. INT-180. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 246 p. 
199. Paulsen, Harold A., Jr. 1969. Forage values on a mountain grassland-aspen range in western Colorado. Journal of Range Management. 22: 102-107. 
200. Paysen, Timothy E.; Ansley, R. James; Brown, James K.; [and others]. 2000. Fire in western shrubland, woodland, and grassland ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-volume 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 121-159. 
201. Pemble, R. H.; Van Amburg, G. L.; Mattson, Lyle. 1981. Intraspecific variation in flowering activity following a spring burn on a northwestern Minnesota prairie. In: Stuckey, Ronald L.; Reese, Karen J., eds. The prairie peninsula--in the "shadow" of Transeau: Proceedings, 6th North American prairie conference; 1978 August 12-17; Columbus, OH. Ohio Biological Survey: Biological Notes No. 15. Columbus, OH: Ohio State University, College of Biological Sciences: 235-240. 
202. Peters, Erin F.; Bunting, Stephen C. 1994. Fire conditions pre- and postoccurrence of annual grasses on the Snake River Plain. 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: 31-36. 
203. Peterson, Kristopher I. 1999. Whitebark pine (Pinus albicaulis) decline and restoration in Glacier National Park. Grand Forks, ND: University of North Dakota. 75 p. Thesis. 
204. Pfister, Robert D.; Kovalchik, Bernard L.; Arno, Stephen F.; Presby, Richard C. 1977. Forest habitat types of Montana. Gen. Tech. Rep. INT-34. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 174 p. 
205. Pojar, Jim; MacKinnon, Andy, eds. 1994. Plants of the Pacific Northwest Coast: Washington, Oregon, British Columbia and Alaska. Redmond, WA: Lone Pine Publishing. 526 p. 
206. Powell, David C. 1994. Effects of the 1980's western spruce budworm outbreak on the Malheur National Forest in northeastern Oregon. Tech. Pub. R6-FI&D-TP-12-94. Portland, OR: U.S. Department of Agriculture, Forest Service, Natural Resources Staff, Forest Insects and Diseases Group. 176 p. 
207. Pratt, David W.; Black, R. Alan; Zamora, B. A. 1984. Buried viable seed in a ponderosa pine community. Canadian Journal of Botany. 62: 44-52. 
208. Qi, Meiqin; Scarratt, John B. 1998. Effect of harvesting method on seed bank dynamics in a boreal mixedwood forest in northwestern Ontario. Canadian Journal of Botany. 76: 872-883. 
209. Quinn, Ronald D. 1990. The status of walnut forests and woodlands (Juglans californica) in southern California. In: Schoenherr, Allan A., ed. Endangered plant communities of southern California: Proceedings, 15th annual symposium; 1989 October 28; Fullerton, CA. Special Publication No. 3. Claremont, CA: Southern California Botanists: 42-54. 
210. Quintilio, D.; Alexander, M. E.; Ponto, R. L. 1991. Spring fires in a semimature trembling aspen stand in central Alberta. Information Report NOR-X-323. Edmonton, AB: Forestry Canada, Northwest Region, Northern Forestry Centre. 30 p. 
211. Radford, Albert E.; Ahles, Harry E.; Bell, C. Ritchie. 1968. Manual of the vascular flora of the Carolinas. Chapel Hill, NC: The University of North Carolina Press. 1183 p. 
212. Rafaill, Barbara L. 1988. Soil characteristics and vegetational features of abandoned and artificially revegetated surface mines in the Cumberland Mountains. Carbondale, IL: Southern Illinois University. 192 p. Dissertation. 
213. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. 
214. Redmann, Robert E.; Schwarz, Arthur G. 1986. Dry grassland plant communities in Wood Buffalo National Park, Alberta. Canadian Field-Naturalist. 100(4): 526-532. 
215. Reed, Catherine C. 1995. Species richness of insects on prairie flowers in southeastern Minnesota. In: Hartnett, David C., ed. Prairie biodiversity: Proceedings, 14th North American prairie conference; 1994 July 12-16; Manhattan, KS. Manhattan, KS: Kansas State University: 103-115. 
216. Rees, Daniel C.; Juday, Glenn Patrick. 2002. Plant species diversity on logged versus burned sites in central Alaska. Forest Ecology and Management. 155: 291-302. 
217. Reich, Peter B.; Bakken, Peter; Carlson, Daren; Frelich, Lee E.; Friedman, Steve K.; Grigal, David F. 2001. Influence of logging, fire, and forest type on biodiversity and productivity in southern boreal forests. Ecology. 82(10): 2731-2748. 
218. Revel, Richard D. 1993. Canada's rough fescue grasslands. Restoration & Management Notes. 11(2): 117-124. 
219. Riegel, Gregg M.; Miller, Richard F.; Krueger, William C. 1992. Competition for resources between understory vegetation and overstory Pinus ponderosa in northeastern Oregon. Ecological Applications. 2(1): 71-85. 
220. Riegel, Gregg M.; Miller, Richard F.; Krueger, William C. 1995. The effects of aboveground and belowground competition in a Pinus ponderosa forest. Forest Science. 41(4): 864-889. 
221. Ripple, William J. 1994. Historic spatial patterns of old forests in western Oregon. Journal of Forestry. 92(11): 45-49. 
222. Roberts, M. R.; Gilliam, F. S. 1995. Disturbance effects on herbaceous layer vegetation and soil nutrients in Populus forests of northern lower Michigan. Journal of Vegetation Science. 6(6): 903-912. 
223. Roberts, Mark R.; Zhu, Lixiang. 2002. Early response of the herbaecous layer to harvesting in a mixed coniferous-deciduous forest in New Brunswick, Canada. Forest Ecology and Management. 155: 17-31. 
224. Rogers, Lynn L.; Allen, Arthur W. 1987. Habitat suitability index models: Black bear--upper Great Lakes region. Biol. Rep. 82 (10.144). Washington DC: U.S. Department of the Interior, Fish and Wildlife Service. 54 p. 
225. Roper, Laren Alden. 1970. Some aspects of the synecology of Cornus nuttallii in northern Idaho. Moscow, ID: University of Idaho. 81 p. Thesis. 
226. Ross, S. Rachel. 1978. The effects of prescribed burning on ground cover vegetation of white pine and mixed hardwood forests in southeastern New Hampshire. Durham, NH: University of New Hamshire. 151 p. Thesis. 
227. Rowe, J. S. 1969. Lightning fires in Saskatchewan grassland. Canadian Field-Naturalist. 83: 317-324. 
228. Rowe, J. S.; Bergsteinsson, J. L.; Padbury, G. A.; Hermesh, R. 1974. Fire studies in the Mackenzie Valley. ALUR 73-74-61. Ottawa: Canadian Department of Indian and Northern Development. 123 p. 
229. Sapsis, David B. 1990. Ecological effects of spring and fall prescribed burning on basin big sagebrush/Idaho fescue--bluebunch wheatgrass communities. Corvallis, OR: Oregon State University. 105 p. Thesis. 
230. Saunders, Jack K., Jr. 1955. Food habits and range use of the Rocky Mountain goat in the Crazy Mountains, Montana. Journal of Wildlife Management. 19(4): 429-437. 
231. Sawyer, John O.; Thornburgh, Dale A. 1977. Montane and subalpine vegetation of the Klamath Mountains. In: Barbour, Michael G.; Major, Jack, eds. Terrestrial vegetation of California. New York: John Wiley & Sons: 699-732. 
232. Schallenberger, Allen Dee. 1966. Food habits, range use and interspecific relationships of bighorn sheep in the Sun River area, west-central Montana. Bozeman, MT: Montana State University. 44 p. Thesis. 
233. Scheller, Robert M.; Mladenoff, David J. 2002. Understory species patterns and diversity in old-growth and managed northern hardwood forests. Ecological Applications. 12(5): 1329-1343. 
234. Seip, Dale R.; Bunnell, Fred L. 1985. Species composition and herbage production of mountain rangelands in northern British Columbia. Canadian Journal of Botany. 63: 2077-2080. 
235. Seklecki, Mariette T.; Grissino-Mayer, Henri D.; Swetnam, Thomas W. 1996. Fire history and the possible role of Apache-set fires in the Chiricahua Mountains of southeastern Arizona. In: Ffolliott, Peter F.; DeBano, Leonard F.; Baker, Malchus, B., Jr.; [and others], tech. coords. Effects of fire on Madrean Province ecosystems: a symposium proceedings; 1996 March 11-15; Tucson, AZ. Gen. Tech. Rep. RM-GTR-289. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 238-246. 
236. Seymour, Frank Conkling. 1982. The flora of New England. 2d ed. Phytologia Memoirs 5. Plainfield, NJ: Harold N. Moldenke and Alma L. Moldenke. 611 p. 
237. Shanks, Royal E.; Goodwin, Richard H. 1943. Notes on the flora of Monroe County, New York. Proceedings of the Rochester Academy of Science. 8(5-6): 299-331. 
238. Shaw, Charles Hugh. 1916. The vegetation of the Selkirks. Botanical Gazette. 61: 477-494. 
239. Shear, Ted; Young, Mike; Kellison, Robert. 1997. An old-growth definition for red river bottom forests in the eastern United States. Gen. Tech. Rep. SRS-10. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station. 9 p. 
240. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. 
241. Shirley, Hardy L. 1932. Light intensity in relation to plant growth in a virgin Norway pine forest. Journal of Agricultural Research. 44: 227-244. 
242. Siccama, T. G.; Bormann, F. H.; Likens, G. E. 1970. The Hubbard Brook Ecosystem Study: productivity, nutrients and phytosociology of the herbaceous layer. Ecological Monographs. 40(4): 389-402. 
243. Simmerman, Dennis G.; Arno, Stephen F.; Harrington, Michael G.; Graham, Russell T. 1991. A comparison of dry and moist fuel underburns in ponderosa pine shelterwood units in Idaho. In: Andrews, Patricia L.; Potts, Donald F., eds. Proceedings, 11th annual conference on fire and forest meteorology; 1991 April 16-19; Missoula, MT. SAF Publication 91-04. Bethesda, MD: Society of American Foresters: 387-397. 
244. Smith, D. W.; James, T. D. W. 1978. Changes in the shrub and herb layers of vegetation after prescribed burning in Populus tremuloides woodland in southern Ontario. Canadian Journal of Botany. 56: 1792-1797. 
245. Smith, Jane K.; Laven, Richard D.; Omi, Philip N. 1985. Vegetation changes in aspen stands resulting from prescribed burning in recreation areas of the Front Range of Colorado. Final Report. Contract Nos. RM-80-112-GR and RM-81-162-GR (EC-367): Eisenhower Consortium for Western Environmental Forestry Research. 53 p. On file with: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. 
246. Smith, Jane Kapler; Fischer, William C. 1997. Fire ecology of the forest habitat types of northern Idaho. Gen. Tech. Rep. INT-GTR-363. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 142 p. 
247. Spies, Thomas A. 1991. Plant species diversity and occurrence in young, mature, and old-growth Douglas-fir stands in western Oregon and Washington. In: Ruggiero, Leonard F.; Aubry, Keith B.; Carey, Andrew B.; Huff, Mark H., technical coordinators. Wildlife and vegetation of unmanaged Douglas-fir forests. Gen. Tech. Rep. PNW-GTR-285. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station: 111-121. 
248. Staniforth, Richard J.; Scott, Peter A. 1991. Dynamics of weed populations in a northern subarctic community. Canadian Journal of Botany. 69: 814-821. 
249. Steele, Robert; Geier-Hayes, Kathleen. 1995. Major Douglas-fir habitat types of central Idaho: a summary of succession and management. Gen. Tech. Rep. INT-GTR-331. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 23 p. 
250. Steele, Robert; Pfister, Robert D.; Ryker, Russell A.; Kittams, Jay A. 1981. Forest habitat types of central Idaho. Gen. Tech. Rep. INT-114. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 138 p. 
251. Stelfox, John G. 1976. Range ecology of Rocky Mountain bighorn sheep in Canadian national parks. Report Series Number 39. Ottawa, ON: Canadian Wildlife Service. 50 p. 
252. Stevens, O. A. 1932. The number and weight of seeds produced by weeds. American Journal of Botany. 19: 784-794. 
253. Stickney, Peter F. 1986. First decade plant succession following the Sundance Forest Fire, northern Idaho. Gen. Tech. Rep. INT-197. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 26 p. 
254. Stickney, Peter F. 1989. FEIS postfire regeneration workshop--April 12: Seral origin of species comprising secondary plant succession in Northern Rocky Mountain forests. 10 p. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. 
255. Stickney, Peter F.; Campbell, Robert B., Jr. 2000. Data base for early postfire succession in northern Rocky Mountain forests. Gen. Tech. Rep. RMRS-GTR-61-CD, [CD-ROM]. Fort Collins, CO: U.S. Department of Agriculture, Forest Service (Producer). Available: Rocky Mountain Research Station. 
256. Strausbaugh, P. D.; Core, Earl L. 1977. Flora of West Virginia. 2nd ed. Morgantown, WV: Seneca Books, Inc. 1079 p. 
257. Stringer, P. W. 1973. An ecological study of grasslands in Banff, Jasper, and Waterton Lakes national parks. Canadian Journal of Botany. 51: 383-411. 
258. Stuart, John D. 1987. Fire history of an old-growth forest of Sequoia sempervirens (Taxodiaceae) forest in Humboldt Redwoods State Park, California. Madrono. 34(2): 128-141. 
259. Stuever, Mary C.; Hayden, John S. 1996. Plant associations (habitat types) of the forests and woodlands of Arizona and New Mexico. Final report: Contract R3-95-27. Placitas, NM: Seldom Seen Expeditions, Inc. 520 p. 
260. Swan, Frederick R., Jr. 1970. Post-fire response of four plant communities in south-central New York State. Ecology. 51(6): 1074-1082. 
261. Swan, Frederick Robbins, Jr. 1966. The effects of fire on plant communities of south-central New York State. Ithaca, NY: Cornell University. 169 p. Dissertation. 
262. Tande, Gerald F. 1979. Fire history and vegetation pattern of coniferous forests in Jasper National Park, Alberta. Canadian Journal of Botany. 57: 1912-1931. 
263. Tester, John R. 1996. Effects of fire frequency on plant species in oak savanna in east-central Minnesota. Bulletin of the Torrey Botanical Club. 123(4): 304-308. 
264. Thompson, Ralph L. 2001. Botanical survey of Myrtle Island Research Natural Area, Oregon. Gen. Tech. Rep. PNW-GTR-507. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 27 p. 
265. Thysell, David R.; Carey, Andrew B. 2000. Effects of forest management on understory and overstory vegetation: a retrospective study. Gen. Tech. Rep. PNW-GTR-488. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 41 p. 
266. Topik, Christopher; Halverson, Nancy M.; Brockway, Dale G. 1986. Plant association and management guide for the western hemlock zone: Gifford Pinchot National Forest. R6-ECOL-230A. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 132 p. 
267. Turner, Nancy Chapman; Bell, Marcus A. M. 1973. The ethnobotany of the southern Kwakiutl Indians of British Columbia. Economic Botany. 27: 257-310. 
268. Tyser, Robin W. 1992. Vegetation associated with two alien plant species in a fescue grassland in Glacier National Park, Montana. The Great Basin Naturalist. 52(2): 189-193. 
269. U.S. Department of Agriculture, Natural Resources Conservation Service. 2005. PLANTS database (2005), [Online]. Available: https://plants.usda.gov /. 
270. Umbanhowar, Charles E., Jr. 1995. Revegetation of earthen mounds along a topographic-productivity gradient in a northern mixed prairie. Journal of Vegetation Science. 6(5): 637-646. 
271. Viereck, Leslie A. 1970. Forest succession and soil development adjacent to the Chena River in interior Alaska. Arctic and Alpine Research. 2(1): 1-26. 
272. Vincent, Dwain W. 1992. The sagebrush/grasslands of the upper Rio Puerco area, New Mexico. Rangelands. 14(5): 268-271. 
273. Vincent, Gilles; Bergeron, Yves; Meilleur, Alain. 1986. Plant community pattern analysis: a cartographic approach applied in the Lac des Deux-Montagnes area (Quebec). Canadian Journal of Botany. 64: 326-335. 
274. Vogl, Richard J. 1971. Fire and the northern Wisconsin pine barrens. In: Proceedings, annual Tall Timbers Fire ecology conference; 1970 August 20-21; New Brunsick, Canada. No. 10. Tallahassee, FL: Tall Timbers Research Station: 175-209. 
275. Voss, Edward G. 1996. Michigan flora. Part III: Dicots (Pyrolaceae--Compositae). Cranbrook Institute of Science Bulletin 61/University of Michigan Herbarium. Ann Arbor, MI: The Regents of the University of Michigan. 622 p. 
276. Vujnovic, K.; Bentz, J. 2001. Preliminary classification of native wheat grass (Agropyron spp.) community types in Alberta. Edmonton, AB: Alberta Environment, Natural Heritage Centre. 362 p. Unpublished report on file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. 
277. Wade, Dale D.; Brock, Brent L.; Brose, Patrick H.; [and others]. 2000. Fire in eastern ecosystems. In: Brown, James K.; Smith, Jane Kapler, eds. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech. Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station: 53-96. 
278. Wali, M. K.; Killingbeck, K. T.; Bares, R. H.; Shubert, L. E. 1980. Vegetation-environment relationships of woodland and shrub communities, and soil algae in western North Dakota. ND REAP Project No. 7-01-1, No. 79-16. Grand Forks, ND: University of North Dakota, Department of Biology, Project of the North Dakota Regional Environmental Assessment Program (REAP). 159 p. 
279. Weaver, T.; Collins, D. 1977. Possible effects of weather modification (increased snowpack) on Festuca idahoensis meadows. Journal of Range Management. 30(6): 451-456. 
280. Welsh, Stanley L.; Atwood, N. Duane; Goodrich, Sherel; Higgins, Larry C., eds. 1987. A Utah flora. The Great Basin Naturalist Memoir No. 9. Provo, UT: Brigham Young University. 894 p. 
281. Whisenant, Steven G. 1990. Postfire population dynamics of Bromus japonicus. The American Midland Naturalist. 123: 301-308. 
282. White, Alan S. 1983. The effects of thirteen years of annual prescribed burning on a Quercus ellipsoidalis community in Minnesota. Ecology. 64(5): 1081-1085. 
283. White, Alan S. 1986. Prescribed burning for oak savanna restoration in central Minnesota. Res. Pap. NC-266. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station. 12 p. 
284. Wilhelm, Gerould S. 1991. Implicatons of changes in floristic composition of the Morton Arboretum's East Woods. In: Burger, George V.; Ebinger, John E.; Wilhelm, Gerould S., eds. Proceedings of the oak woods management workshop; 1988 October 21-22; Peoria, IL. Charleston, IL: Eastern Illinois University: 31-54. 
285. Willms, W. D.; Smoliak, S.; Dormaar, J. F. 1985. Effects of stocking rate on a rough fescue grassland vegetation. Journal of Range Management. 38(3): 220-225. 
286. Wilm, Harold Gridley. 1932. The relation of successional development to the silviculture of forest burn communities in southern New York. Ithaca, NY: Cornell University. 74 p. [+ appendices]. Thesis. 
287. Wofford, B. Eugene. 1989. Guide to the vascular plants of the Blue Ridge. Athens, GA: The University of Georgia Press. 384 p. 
288. Wolfe-Bellin, Kelly S.; Moloney, Kirk A. 2001. Successional vegetation dynamics on pocket gopher mounds in an Iowa tallgrass prairie. In: Bernstein, Neil P.; Ostrander, Laura J., eds. Seeds for the future; roots of the past: Proceedings of the 17th North American prairie conference; 2000 July 16-20; Mason City, IA. Mason City, IA: North Iowa Community College: 155-163. 
289. Wright, Henry A.; Bailey, Arthur W. 1982. Fire ecology: United States and southern Canada. New York: John Wiley & Sons. 501 p. 
290. Wunderlin, Richard P. 1998. Guide to the vascular plants of Florida. Gainesville, FL: University Press of Florida. 806 p. 
291. Young, James A.; Evans, Raymond A. 1981. Demography and fire history of a western juniper stand. Journal of Range Management. 34(6): 501-505. 
292. Youngblood, Andrew P.; Padgett, Wayne G.; Winward, Alma H. 1985. Riparian community type classification of eastern Idaho - western Wyoming. R4-Ecol-85-01. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Region. 78 p. 
293. Zimmerman, G. T.; Neuenschwander, L. F. 1984. Livestock grazing influences on community structure, fire intensity, and fire frequency within the Douglas-fir/ninebark habitat type. Journal of Range Management. 37(2): 104-110. 
294. Zimmerman, James H. 1972. Propagation of spring prairie plants. In: Zimmerman, James H., ed. Proceedings of the second Midwest prairie conference; 1970 September 18-20; Madison, WI. Madison, WI: University of Wisconsin Arboretum: 153-161.