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SPECIES: Galium boreale, G. triflorum


northern bedstraw fragrant bedstraw
2004 Dr. Virginia Kline
University of Wisconsin Arboretum
2001 Thayne Tuason
Central Washington Native Plants

Gucker, Corey L. 2005. Galium boreale, G. triflorum. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: [].


Galium boreale L. [131]
    =Galium boreale subsp. septentrionale Roemer & Schultes [115]


northern bedstraw

fragrant bedstraw
sweetscented bedstraw

The genus name of bedstraw is Galium L. (Rubiaceae) [91,92,117,129,131,275]. This review provides information on the following bedstraw species [129,131]:

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 [115]. 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.



Northern bedstraw is listed as endangered in both Maryland and Massachusetts [173,174,269].


SPECIES: Galium boreale, G. triflorum
Bedstraw is widely distributed throughout North America and Canada [127,269]. Fragrant bedstraw occurs in every U.S. state except Hawaii. Northern bedstraw is also absent from Hawaii and several southeastern states [269].

Distributional maps of bedstraw and the 2 individual species are accessible through the Plants database.

FRES10 White-red-jack pine
FRES11 Spruce-fir
FRES14 Oak-pine
FRES15 Oak-hickory
FRES16 Oak-gum-cypress
FRES17 Elm-ash-cottonwood
FRES18 Maple-beech-birch
FRES19 Aspen-birch
FRES20 Douglas-fir
FRES21 Ponderosa pine
FRES22 Western white pine
FRES23 Fir-spruce
FRES24 Hemlock-Sitka spruce
FRES25 Larch
FRES26 Lodgepole pine
FRES27 Redwood
FRES28 Western hardwoods
FRES29 Sagebrush
FRES35 Pinyon-juniper
FRES36 Mountain grasslands
FRES37 Mountain meadows
FRES38 Plains grasslands
FRES39 Prairie
FRES41 Wet grasslands
FRES42 Annual grasslands

STATES/PROVINCES: (key to state/province abbreviations)

Northern bedstraw:


Fragrant bedstraw:


1 Northern Pacific Border
2 Cascade Mountains
3 Southern Pacific Border
4 Sierra Mountains
5 Columbia Plateau
6 Upper Basin and Range
7 Lower Basin and Range
8 Northern Rocky Mountains
9 Middle Rocky Mountains
10 Wyoming Basin
11 Southern Rocky Mountains
12 Colorado Plateau
13 Rocky Mountain Piedmont
14 Great Plains
15 Black Hills Uplift
16 Upper Missouri Basin and Broken Lands

K001 Spruce-cedar-hemlock forest
K002 Cedar-hemlock-Douglas-fir forest
K004 Fir-hemlock forest
K005 Mixed conifer forest
K006 Redwood forest
K007 Red fir forest
K008 Lodgepole pine-subalpine forest
K010 Ponderosa shrub forest
K011 Western ponderosa forest
K012 Douglas-fir forest
K013 Cedar-hemlock-pine forest
K014 Grand fir-Douglas-fir forest
K015 Western spruce-fir forest
K016 Eastern ponderosa forest
K018 Pine-Douglas-fir forest
K019 Arizona pine forest
K020 Spruce-fir-Douglas-fir forest
K021 Southwestern spruce-fir forest
K024 Juniper steppe woodland
K025 Alder-ash forest
K030 California oakwoods
K038 Great Basin sagebrush
K047 Fescue-oatgrass
K050 Fescue-wheatgrass
K051 Wheatgrass-bluegrass
K055 Sagebrush steppe
K056 Wheatgrass-needlegrass shrubsteppe
K063 Foothills prairie
K064 Grama-needlegrass-wheatgrass
K066 Wheatgrass-needlegrass
K067 Wheatgrass-bluestem-needlegrass
K069 Bluestem-grama prairie
K074 Bluestem prairie
K082 Mosaic of K074 and K100
K083 Cedar glades
K093 Great Lakes spruce-fir forest
K094 Conifer bog
K095 Great Lakes pine forest
K096 Northeastern spruce-fir forest
K097 Southeastern spruce-fir forest
K098 Northern floodplain forest
K099 Maple-basswood forest
K100 Oak-hickory forest
K101 Elm-ash forest
K102 Beech-maple forest
K103 Mixed mesophytic forest
K104 Appalachian oak forest
K106 Northern hardwoods
K107 Northern hardwoods-fir forest
K108 Northern hardwoods-spruce forest
K109 Transition between K104 and K106
K110 Northeastern oak-pine forest
K111 Oak-hickory-pine
K112 Southern mixed forest

1 Jack pine
5 Balsam fir
12 Black spruce
13 Black spruce-tamarack
14 Northern pin oak
15 Red pine
16 Aspen
17 Pin cherry
18 Paper birch
19 Gray birch-red maple
20 White pine-northern red oak-red maple
21 Eastern white pine
22 White pine-hemlock
23 Eastern hemlock
24 Hemlock-yellow birch
25 Sugar maple-beech-yellow birch
26 Sugar maple-basswood
27 Sugar maple
28 Black cherry-maple
30 Red spruce-yellow birch
31 Red spruce-sugar maple-beech
32 Red spruce
33 Red spruce-balsam fir
34 Red spruce-Fraser fir
35 Paper birch-red spruce-balsam fir
37 Northern white-cedar
38 Tamarack
39 Black ash-American elm-red maple
42 Bur oak
44 Chestnut oak
46 Eastern redcedar
51 White pine-chestnut oak
52 White oak-black oak-northern red oak
53 White oak
55 Northern red oak
58 Yellow-poplar-eastern hemlock
59 Yellow-poplar-white oak-northern red oak
60 Beech-sugar maple
61 River birch-sycamore
62 Silver maple-American elm
63 Cottonwood
65 Pin oak-sweetgum
91 Swamp chestnut oak-cherrybark oak
92 Sweetgum-willow oak
93 Sugarberry-American elm-green ash
94 Sycamore-sweetgum-American elm
96 Overcup oak-water hickory
97 Atlantic white-cedar
107 White spruce
108 Red maple
109 Hawthorn
110 Black oak
201 White spruce
202 White spruce-paper birch
203 Balsam poplar
204 Black spruce
205 Mountain hemlock
206 Engelmann spruce-subalpine fir
207 Red fir
208 Whitebark pine
210 Interior Douglas-fir
211 White fir
213 Grand fir
216 Blue spruce
217 Aspen
218 Lodgepole pine
219 Limber pine
220 Rocky Mountain juniper
221 Red alder
222 Black cottonwood-willow
223 Sitka spruce
224 Western hemlock
225 Western hemlock-Sitka spruce
227 Western redcedar-western hemlock
228 Western redcedar
229 Pacific Douglas-fir
230 Douglas-fir-western hemlock
231 Port-Orford-cedar
232 Redwood
234 Douglas-fir-tanoak-Pacific madrone
235 Cottonwood-willow
236 Bur oak
237 Interior ponderosa pine
239 Pinyon-juniper
243 Sierra Nevada mixed conifer
244 Pacific ponderosa pine-Douglas-fir
245 Pacific ponderosa pine
246 California black oak
251 White spruce-aspen
252 Paper birch
253 Black spruce-white spruce
254 Black spruce-paper birch
255 California coast live oak
256 California mixed subalpine

101 Bluebunch wheatgrass
102 Idaho fescue
109 Ponderosa pine shrubland
110 Ponderosa pine-grassland
202 Coast live oak woodland
203 Riparian woodland
214 Coastal prairie
215 Valley grassland
216 Montane meadows
303 Bluebunch wheatgrass-western wheatgrass
304 Idaho fescue-bluebunch wheatgrass
308 Idaho fescue-tufted hairgrass
309 Idaho fescue-western wheatgrass
311 Rough fescue-bluebunch wheatgrass
312 Rough fescue-Idaho fescue
313 Tufted hairgrass-sedge
314 Big sagebrush-bluebunch wheatgrass
315 Big sagebrush-Idaho fescue
316 Big sagebrush-rough fescue
323 Shrubby cinquefoil-rough fescue
401 Basin big sagebrush
402 Mountain big sagebrush
403 Wyoming big sagebrush
409 Tall forb
411 Aspen woodland
412 Juniper-pinyon woodland
418 Bigtooth maple
421 Chokecherry-serviceberry-rose
422 Riparian
504 Juniper-pinyon pine woodland
601 Bluestem prairie
602 Bluestem-prairie sandreed
604 Bluestem-grama prairie
606 Wheatgrass-bluestem-needlegrass
607 Wheatgrass-needlegrass
608 Wheatgrass-grama-needlegrass
609 Wheatgrass-grama
610 Wheatgrass
612 Sagebrush-grass
613 Fescue grassland
805 Riparian
809 Mixed hardwood and pine
815 Upland hardwood hammocks
901 Alder
904 Black spruce-lichen
906 Broadleaf forest
908 Fescue
920 White spruce-paper birch
921 Willow

Fragrant and northern bedstraw occur together in several vegetation types of Canada and the northwestern United States.

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 [146]. Bedstraw occurs in nutrient-rich white spruce-black spruce-highbush cranberry (Viburnum edule) associations in British Columbia [135].

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) [198]. 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 [204].

In western North Dakota, the 2 bedstraw species are present with almost equal frequencies in Rocky Mountain juniper (Juniperus scopulorum) communities [278]. 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 [3].

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 [105], quaking aspen-paper birch (Betula papyrifera) communities of western North Dakota [278], narrowleaf cottonwood/Saskatoon serviceberry (P. angustifolia/Amelanchier alnifolia) communities of Colorado's White River National Forest [3], and bur oak (Quercus macrocarpa) communities of western North Dakota [278].

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.) [280].

Classifications: Bedstraw is recognized as a dominant species in many vegetation classifications including:

CO: fragrant bedstraw [3,68]
ID: fragrant bedstraw [100,250,292]
MT: fragrant bedstraw [50,105,204,250]
NM: fragrant bedstraw [68]
WY: northern bedstraw [50]
        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 [162] and white spruce/twinberry honeysuckle (Lonicera involucrata) vegetation types of British Columbia [180]. 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 [203]. 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 [204]. Engelmann spruce/subalpine fir and ponderosa pine communities are common northern bedstraw habitat in the Big Horn Mountains of Wyoming [67].

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 [187]. Coverage of northern bedstraw decreases with stand age in quaking aspen/highbush cranberry/twinflower (Linnaea borealis) communities in the taiga of interior Alaska [85]. 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 [228].

Northern bedstraw is typical in balsam poplar (Populus balsamifera) forests of Alberta [187]. Spruce/red-osier dogwood (Cornus sericea ssp. sericea) riparian forests in Montana also provide northern bedstraw habitat [105]. In eastern Montana, northern bedstraw occupies green ash (Fraxinus pennsylvanica) woodlands [161].

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 [257]. 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) [214] and rough fescue/shrubby cinquefoil (F. altaica-Dasiphora floribunda) grasslands, northern bedstraw is also prominent [187]. 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 [276].

On steep south-facing slopes in Alaska's Yukon lowlands, northern bedstraw associates with fringed sagebrush/purple pinegrass (Artemisia frigida/Calamagrostis purpurascens) vegetation [62]. Shrubby cinquefoil/tufted hairgrass (Deschampsia cespitosa) riparian habitat types in Montana also provide northern bedstraw habitat [105]. In western Wyoming, mountain big sagebrush (A. tridentata ssp. vaseyana) is a northern bedstraw associate [177]. In montane riparian sites throughout Wyoming, the aster (Aster spp.)-northern bedstraw community type is recognized [50].

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 [259]. Welsh and others [280] 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 [150].

Shrub and grassland communities: In Nevada, northern bedstraw occupies sagebrush and pinyon-juniper (Pinus-Juniperus spp.) vegetation [132]. 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 [150].

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) [155]. In the Great Sand Hills of Saskatchewan, northern bedstraw occupies creeping juniper habitat [126].

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 [104]. 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 [90].

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 [98]. 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) [183]. In south-central North Dakota, northern bedstraw occupies the blue grama (Bouteloua gracilis)-sun sedge (Carex inops ssp. heliophila)- little bluestem vegetation type [165]. 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 [76].

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 [73]. In New York, oak (Quercus spp.), aspen, maple (Acer spp.), and beech (Fagus spp.) forests are described as northern bedstraw habitat [286]. 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 [237].

Fragrant bedstraw-
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 [171]. Fragrant bedstraw is characteristic of several productive Douglas-fir-dominated habitats of southwestern British Columbia [93]. 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 [162].

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 [266]. 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 [114].

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 [111]. 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 [16].

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 [97]. 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 [250]. 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 [292].

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) [50]. Fragrant bedstraw is common in western larch (Larix occidentalis)- and whitebark pine-dominated forests of the northern Rockies [48]. 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 [204].

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 [85]. In Alberta, researchers found fragrant bedstraw associated with and growing on decaying logs and stumps in 28-year-old, aspen-dominated boreal forests [159].

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 [264]. Atzet and others [16] 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 [80].

Riparian vegetation typical of Montana and southern Idaho includes fragrant bedstraw [100]. 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 [105]. 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) [292].

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 [168]. 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 [231]. Fragrant bedstraw is also typical of giant sequoia (Sequoiadendron giganteum)-mixed conifer forests with white fir and incense-cedar (Calocedrus decurrens) [133]. 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 [259].

In northern New Mexico and southern Colorado, fragrant bedstraw typifies the white fir/fragrant bedstraw riparian forest habitat type [3,68]. Hayward [112] 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 [190]. 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 [209]. In the Humboldt-Toiyabe National Forest, fragrant bedstraw occupies several communities dominated by quaking aspen, red-osier dogwood, and/or willow (Salix spp.) [169]. Kartesz [132] 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 [20]. 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 [189].

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 [46]. 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% [83].

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 [45].

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 [128]. Lutz [166] 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 [144]. 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 [32]. Ross [226] 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 [179]. In the Lac des Deuz-Montagnes area of Quebec, fragrant bedstraw is important in swamp white oak (Q. bicolor) communities [273].

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 [34]. 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) [239].


SPECIES: Galium boreale, G. triflorum

northern bedstraw fragrant bedstraw
2005 Dan Tenaglia,
This description provides characteristics that may be relevant to fire ecology, and is not meant for identification. Keys for identification are available [59,91,92,115,117,118,127,132,172,236,275,280,287].

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 [92]. If hairs are present, they are short and without hooks [38,59,118]. Stevens [252] 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 [92]. 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 [92]. Seeds are coated with dense hooked hairs and are typically 1.5 to 2.2 mm in diameter [59,92,118].


Bedstraw produces both rhizomes and seeds allowing for vegetative and sexual reproduction.

Breeding system: Bedstraw has perfect flowers [108].

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 [215]. 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 [252] 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 [252].

Archibold [6] 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 [6].

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/m although fragrant bedstraw occupied no coverage 3.3 feet (1 m) from trapping site [106].

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 [208].

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 0.870.55 (s x) m/year and 0.910.55 m/year based on the furthest 1/2 peak density and furthest individual calculations, respectively [175].

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 5080 (s) northern bedstraw seeds/m were in spring seed banks and 133103 seeds/m were in fall seed banks. The coverage of northern bedstraw in the study area was 5% and constancy was 96% [207].

Following the 1988 Yellowstone fires, Clark [47] 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 [47]. From soil collected in central Saskatchewan's native porcupine grass and Montana wheatgrass prairies, Archibold [7] reported 42 northern bedstraw "root sprouts"/m.

Fragrant bedstraw: In old-growth deciduous forests of southwestern Quebec, researchers calculated a maximum of 25 fragrant bedstraw seeds/m from soil samples collected in May. Sugar maple, striped maple, American beech, and white ash between 150 and 450 years old dominated the sites [157]. Soil samples 4 inches (10 cm) deep from a total area of 2.3 m 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/m. Most seed (87%) came from the top 2 inches (5 cm) of soil [138].

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 [153].

On intermittently flooded (2- to 10-year flood intervals, typically) gravel bars, 36 fragrant bedstraw seeds/m emerged from soil collected on 3rd order streams where coverage of fragrant bedstraw was 1%; on 5th order streams, 36.5 seeds/m 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/m where coverage of fragrant bedstraw was 1% or less. Streams flowed in Oregon's Cascade Range [106].

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 [176].

Site condition Undisturbed (n=18)

Low disturbance (n=11)

High disturbance (n=18)

Total number of germinants 60 68


Mean seeds SE/m 206 (litter) 72
6020 (litter) 21
Constancy (%) 33 22 11 28 6 17

Ahlgren [2] 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 [94]. 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 [194].

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 [193] 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 [193].



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

More mesic

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 [70]. Stickney and Campbell [255] consider fragrant bedstraw's rhizomes to be more fire sensitive than those of northern bedstraw.

Bedstraw occupies a diversity of moist sites [59]. Woodlands, prairies, meadows, riparian areas, and swamps are all potential bedstraw habitats [92,172,275].

Northern bedstraw: Northern bedstraw often occupies stony slopes and meadows of Alaska and Canada [127] and meadows and damp slopes in the Southwest [172]. 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 [188].

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 [92]. Voss [275] 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.

Northern bedstraw:
State, province, or region Elevational range
Alberta 500-1,750 m [53]
California 15-2,000 m [115]
Colorado 1,520-3,050 m [108]
Intermountain West up to 2,700 m [59]
New Mexico 1,830-3,050 m [172]
Utah 1,650-3,100 m [280]

Fragrant bedstraw:
State, province, or region Elevational range
Adirondacks 290-900 m [144]
Alberta 500-1,500 m [53]
California 10-3,000 m [115]
southern California below 2,440 m [190]
Colorado 1,980-2,740 m [108]
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 [105]
New Mexico 2,130-2,740 m [172]
Utah 1,220-2,500 m [280]

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 [85]. 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 [158]. In northeastern Oregon's bedstraw habitats, winters are cold and wet, and summers are hot and dry [219]. 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 [278].

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 5.01.3% (s x) at snow levels of 24 inches (60 cm), 9.01.3% at 48 inches (120 cm), and 11.23.2% at 95 inches (240 cm). Flowering was delayed on sites with 95 inches (240 cm) of snow accumulation [279].

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 [78]. 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 [214]. In southwestern North Dakota woodlands, soil pH ranged from 6.8 to 8.4 on sites where northern bedstraw occurred [90]. Strausbaugh and Core [256] 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 [136]. In the subalpine fir/fragrant bedstraw habitat type of central and eastern Montana, soils range from neutral to strongly acidic [105]. Fragrant bedstraw habitat in the Adirondacks has "higher" pH soils [144].

Northern and fragrant bedstraw: Rarely did both bedstraw species occur together on study sites in different stages of succession or following a disturbance. In quaking aspen-dominated boreal forests of central Alberta, researchers compared the composition of forests on the edge of a clearcut and interior forests. Fragrant bedstraw cover was greater in 1-year-old edge forests but less on 5- and 16-year-old forests than in interior forests. Northern bedstraw cover was greater in 1-, 5-, and 16-year-old edge forests than in interior forests [107]. These findings are similar to those of Bakuzis and Hansen [21] who established the edaphic and climatic conditions preferred by herbs in Minnesota forests. Fragrant bedstraw favored increased moisture and decreased light conditions, while northern bedstraw tolerated lower moisture levels and higher light conditions.

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 [197].

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 [207]. In bunchberry (Cornus canadensis)-dominated sites of central Alaska, northern bedstraw was a principal species in both early and late seral communities [216]. 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% [85]. Stringer [257] 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.) [248].

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 [271].

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 [241].

Northern bedstraw persists in recently clearcut (6- to 12-year-old-stands) and mature lodgepole pine forests in the Lower Foothills of Alberta [54]. Likewise, Crouch [60] 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 [288].

1 year-old-mound (n = 40) 5.85 undisturbed quadrat
(n = 49)
2 year-old-mound (n = 40) 7.32 undisturbed quadrat
(n = 25)
* Difference significant (p<0.05)

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 [270].

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 [141]:

Treatment Uncut Harvested Uncut/grazed Harvested/grazed
Cover (%) 0.7 1.3 0.9 2.5

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 [238] 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 [234]. 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 [52].

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 [178] 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 [66].

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 [185].

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 [110].

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 [225]. 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 [222]. In hybrid white spruce Engelmann spruce forests of central British Columbia, fragrant bedstraw occurred in all forests 14 to 140 years old [72]. Habeck [97] 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 [247]. In rich mesic forests of western Massachusetts, researchers found fragrant bedstraw frequency was significantly (p≤0.05) lower in more open sites [27]. 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 [249].

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 [79]:

Percentage of full light 0-3.5 3.5-6 6-11 11-25 25-60 60-100+
Cover (%) 1 5 5 1 25 5

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 [233]. 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 [217].

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 [149]. 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 [35]. Increases in fragrant bedstraw density were significant (p<0.01) following thinning treatments in giant sequoia groves of Tulare County, California [156]. Frequency following the thinning reportedly decreased, however [133].

While fragrant bedstraw increases following canopy release predominate, Freedman and Habeck [87] 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 [195]:

Post-treatment year


Treatment Control Full-tree logging (winter) Full-tree logging (spring) Control Full-tree logging (winter) Full-tree logging (spring) Tree-length logging (winter) & burning
Density (stems/m) 0.68 1.52 0.73 0.09* 1.19 0.63 0.10*
* Values are significantly (p=0.05) lower than other presented values.

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% [101].

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 [196].

Year 1990 1991 1992 1993 1996 1999
fragrant bedstraw
constancy %
13 20 10 5 8 6

Bailey [18] 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.

Northern bedstraw:

State, region Flowering dates
Blue Ridge Province May-August [287]
Great Plains states June-September [92]
northern Idaho June-August [198]
southeastern Illinois flowering begins late May-late June, typically lasts 30 days [40]
western Montana 1st bloom: early May-mid-July, end of blooming: late July-mid-August [188].
North Dakota late May-early September [284]
Utah's Wasatch Mountains mid-June-early August [4]
West Virginia May-August [256]
Wisconsin spring [294]

Fruiting dates for northern bedstraw are between mid-July and mid-August in New England [236]. In subarctic northern Manitoba, Staniforth and Scott [248] report that northern bedstraw had immature fruits as of mid-September. From data collected over 10 years in western Montana's mountain grasslands, Mueggler [188] indicated fruit dissemination occurred from early August through early September, and plants were dry by mid-September.

Fragrant bedstraw:
State, region Flowering dates
Atlantic and Gulf coast states May-August [75]
Blue Ridge Province July-August [287]
southern California May-July [190]
Carolinas April-May [211]
Florida spring-summer [290]
northern Idaho June-August [198]
southeastern Illinois early June-late August [284]
Kansas June-July [22]
north-central Texas June-July [69]
West Virginia May-September [256]
Wyoming June-August [124]

Seymour [236] suggests that fragrant bedstraw sets fruit from late June to late August.


SPECIES: Galium boreale, G. triflorum
Fire adaptations: Fire severity likely dictates which bedstraw recolonization strategy prevails. When burned in low-severity fires, bedstraw likely sprouts from rhizomes. However, high-severity fires may damage fragrant bedstraw's more delicate rhizomes [255], and recolonization of the area would likely be from on- or off-site seed sources. Northern bedstraw's more robust rhizomes are capable of withstanding more severe fires, but on- or off-site seed sources may still colonize disturbed sites.

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 [64], 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 [164]. 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 [74]. 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 [200]. Fragrant bedstraw occupies southern California walnut woodlands of southern California that burn annually due to an increased presence of annual grasses [209].

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 [12]
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 [277]
bluestem prairie Andropogon gerardii var. gerardii-Schizachyrium scoparium < 10 [142,200]
sagebrush steppe Artemisia tridentata/Pseudoroegneria spicata 20-70 [200]
basin big sagebrush Artemisia tridentata var. tridentata 12-43 [229]
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 [277]
western juniper Juniperus occidentalis 20-70
Rocky Mountain juniper Juniperus scopulorum < 35 [200]
western larch Larix occidentalis 25-350 [13,24,63]
yellow-poplar Liriodendron tulipifera < 35 [277]
Great Lakes spruce-fir Picea-Abies spp. 35 to > 200
northeastern spruce-fir Picea-Abies spp. 35-200 [74]
southeastern spruce-fir Picea-Abies spp. 35 to > 200 [277]
Engelmann spruce-subalpine fir Picea engelmannii-Abies lasiocarpa 35 to > 200 [12]
black spruce Picea mariana 35-200
conifer bog* Picea mariana-Larix laricina 35-200 [74]
blue spruce* Picea pungens 35-200 [12]
red spruce* Picea rubens 35-200 [74]
pinyon-juniper Pinus-Juniperus spp. < 35 [200]
whitebark pine* Pinus albicaulis 50-200 [1,10]
jack pine Pinus banksiana <35 to 200 [74]
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 [12]
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 [277]
eastern cottonwood Populus deltoides < 35 to 200 [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 [277]
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 [12]
oak-hickory Quercus-Carya spp. < 35
northeastern oak-pine Quercus-Pinus spp. 10 to < 35 [277]
oak-gum-cypress Quercus-Nyssa-spp.-Taxodium distichum 35 to > 200 [191]
southeastern oak-pine Quercus-Pinus spp. < 10 [277]
coast live oak Quercus agrifolia 2-75 [95]
white oak-black oak-northern red oak Quercus alba-Q. velutina-Q. rubra < 35 [277]
canyon live oak Quercus chrysolepis <35 to 200 [12]
northern pin oak Quercus ellipsoidalis < 35 [277]
Oregon white oak Quercus garryana < 35 [12]
California black oak Quercus kelloggii 5-30 [200]
bur oak Quercus macrocarpa < 10 [277]
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 [277]
little bluestem-grama prairie Schizachyrium scoparium-Bouteloua spp. < 35 [200]
redwood Sequoia sempervirens 5-200 [12,82,258]
western redcedar-western hemlock Thuja plicata-Tsuga heterophylla > 200 [12]
eastern hemlock-yellow birch Tsuga canadensis-Betula alleghaniensis > 200 [277]
western hemlock-Sitka spruce Tsuga heterophylla-Picea sitchensis > 200
mountain hemlock* Tsuga mertensiana 35 to > 200 [12]
elm-ash-cottonwood Ulmus-Fraxinus-Populus spp. < 35 to 200 [74,277]
*fire return interval varies widely; trends in variation are noted in the species review

Rhizomatous herb, rhizome in soil
Ground residual colonizer (on-site, initial community)
Initial off-site colonizer (off-site, initial community)
Secondary colonizer (on-site or off-site seed sources)


SPECIES: Galium boreale, G. triflorum
Bedstraw can be killed by fire [33,246], but underground structures often survive low-severity fires. Likely fire severity and/or seasonality dictate the survival of bedstraw.

Powell [206] indicates that northern bedstraw has better chances of surviving fire than fragrant bedstraw. Fragrant bedstraw has less than 35% chance of 50% population survival [137,206] while northern bedstraw's chances are 35% to 64% for 50% of the species population to survive fires with average flame lengths of 12 inches (30.5 cm) [206]. Stickney and Campbell [255] consider fragrant bedstraw a nonsurvivor because of its delicate rhizome; this classification was tentative as the researchers observed few postfire responses. However, Edgerton [77] described fragrant bedstraw as a "surviving forb" following clearcutting and broadcast burning of a mixed conifer forest in the Umatilla National Forest of Oregon.

Bedstraw regenerates from rhizomes or seeds [33,137,206] and likely regains prefire frequency or coverage 5 to 10 years after fire [206]. Stickney [253] classifies fragrant bedstraw as a residual colonizer, coming from an on-site ground source.

Severe fires can cause large decreases in bedstraw; however, bedstraw can remain unchanged or increase following low-severity understory fires or cool season spring and fall fires [33,140,206,246].

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 [260].

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 [158] 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 [158]:

Postfire characteristic Mean seed density
2nd postfire year coverage (%)
Fire type UB L S UB L S
Northern bedstraw 1.9 0 0 0.64 2.6 3.7
Fragrant bedstraw 24.5 30.1 11.3


0.03 0.01
UB-unburned, L-light, S-severe

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 [263].

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 [238] lists northern bedstraw as a prominent herb in the early postfire reforestation of western hemlock, lodgepole pine, and quaking aspen forests. Seip and Bunnell [234] 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) [52].

In interior Alaska's white spruce forests, Foote [85] 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 [85]. One of the fires that was included in the previous postfire recovery chronosequence was the 1950 Porcupine River fire. Foote [86] 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 [86].

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) [9]. 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 [73].

Density (stems/ha) Frequency (%)
Site Dominants intact overstory no overstory intact overstory no
1 eastern white pine
black oak
white oak
0 40,600 0 50
2 northern red oak
white oak
sweet birch
5,600 18,400 25 75

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.

Fire Prefire Light Moderate Heavy Light Moderate Heavy
Postfire year NA



Production (kg/ha) 33 42 31 50 13 14 15

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 [245]. 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.

Community Herbaceous understory
(low severity)
Juniper understory
(high severity)
Burn status prefire (1980) postfire (1982) prefire (1980) postfire (1982)
(number of stems/0.1m)
1.2 3.1 1.4 2.7
Coverage (%) 0.7 0.5 0.5 0.9

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 [8]. 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 [210]. 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.

Site Burned
(October 1972)
(May 1978)
Frequency (%) 20 20 8
Cover (%) 8 3.2 1.2
Prominence value* 35.8 14.3 3.4
* Prominence value= % cover *(√frequency)

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 [5]:

Burn status Unburned (n=125) Burned (n=125)
Time since fire 0 3 months
Cover (%) <0.05 4
Frequency (%) 7 22
Burn status Unburned (n=23) Burned (n=28)
Year 1970 1971 1972 1970 1971 1972
Cover (%) 1 <0.05 <0.05 3 3 2
Frequency (%) 17 22 22 29 32 29

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 [274].

McGee [177] 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 [177].

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 [31].

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 [201].

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 [116] 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 [30].

In oak woodlands of east-central Minnesota's Cedar Creek Natural History Area, White [283] 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 [283].

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 [249] 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 [192] 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% [65].

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 [253]. 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 [2]. 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 76F (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 [243]:

Fire type Prefire cover (%) Postfire cover (%)
Unburned 1.4 3.2
Moist burn 0.9 0.7
Dry burn 1.6 0.8

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 [103]. 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 [57] 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 [57].

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% [244]. Following a mid-July crown fire near Missoula, Montana, fragrant bedstraw frequency had doubled from the 1st to the 2nd postfire year [58].

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% [261].

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 [139]:

Fire Unburned Spring fire Summer fire
Years postfire 0 0 3 5 14 2 5 11
Occurrence (%) 3 3 7 3 7 0 3 3

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% [121].

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 [160]:

Time since fire Prefire 3 months 1year 2 years
Burned & seeded
(# of occurrences/10 plots)
6 0 0 2
(# of occurrences/10 plots)
3 2 1 1

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 [128].

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:

Fire management decisions are likely unaffected by the presence of bedstraw in the understory. Bedstraw recovers quickly, remains unchanged, or increases following fire. Special consideration of bedstraw when developing a fire management plan is unnecessary in most cases.

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 [99]:

Site condition Herbage weight
(dry g/m)
Calories/m % total (calories/m)
Unburned 2.8 4,690 12,991 0.85
Burned 8.8 4,654 40,723 2.07

To read more about the use of bedstraw by animals see Importance To Livestock And Wildlife.


SPECIES: Galium boreale, G. triflorum
Bedstraw is not particularly palatable to livestock and native ungulates, but often makes up a small portion of their diets. Usage may increase with stocking rates or with length of grazing time. While not necessarily an important food source for herbivores, bedstraw is an early season food source for black bears. The presence of bedstraw indicates important elk, deer, and moose habitats.

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 [17]. In aspen stands of Colorado and Wyoming, northern bedstraw is constant on moderately grazed ranges, and its removal from grazed vegetation may indicate mismanagement [55]. 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 [285]:

Sampling times 1st 6 years of grazing
Last 6 years of grazing
Stocking rates
1.2 1.6 2.4 4.8 1.2 1.6 2.4 3.2 (mean)
Percent composition 1.1 1.7 1.4 1.9 4.3 4.2 3.7 3.9

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 [293]. 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 [199].

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 [130] 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 [145] 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 [182].

Bentz and Woodard [28] 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 [232]. 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 [230].

In several Canadian National Parks, Stelfox [251] 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 [251].

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 [109]. 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 [119].

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 [123].

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 [215].

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 [56]. 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 [170]. Lonner [163] 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 [204].

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 [122].

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 [167]. In a review, Rogers and Allen [224] 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 [137]. Some have even suggested that bedstraw may be poisonous [147]. Below are some specific findings regarding nutritional value of northern and fragrant bedstraw in various environments.

Northern bedstraw: Paulsen [199] 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 [188]. 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 [99].

Fragrant bedstraw: Fragrant bedstraw collected in July from Hubbard Brook, New Hampshire's hardwood and boreal forests had the following nutritional composition [242]:

Element K N Mg Ca S P Mn Fe Zn Na Cu
Content 2.1% 2.8% 0.2% 1.7% 0.3% 0.2% 318 ppm 109 ppm 294 ppm 20 ppm 12 ppm

Cover value: In south-central Montana, the subalpine fir/fragrant bedstraw habitat provides important big game cover [204].

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 [41]. 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 [212].

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 [218]. 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 [194].

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 [120].

Fragrant bedstraw: 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 [205]. This species is also used to flavor wines [22]. In a review, Turner and Bell [267] 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.

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 [268] 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 [102]. The presence of fragrant bedstraw in riparian zones of central Oregon suggests high productivity sites for conifers [137]. Fragrant bedstraw is also 1 of several understory species indicating productive Douglas-fir habitat in southwestern British Columbia [134].

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 [49].

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