Galium triflorum

This native perennial plant is 1-3' tall, branching occasionally. The slender stems have a tendency to sprawl; they are light green to reddish green, angular, and strongly ridged. Usually, there are a few short hairs that provide the stems with a slightly rough texture. At intervals along the stems, there are whorls of 6 sessile leaves. These leaves are up to 2½" long and ½" across; they are oblanceolate in shape, while their margins are smooth and ciliate. Each leaf abruptly tapers to a short pointed tip (cuspidate). Occasionally, 1-3 stalks of flowers develop from the middle to upper whorled leaves; these stalks are often longer than the leaves, and they can be axillary or terminal. Each of these stalks may produce 1-3 flowers on slender pedicels. Each flower spans about 1/6" across, consisting of 4 greenish white to white petals, 4 stamens, and a bristly 2-celled ovary. Each petal tapers to a narrow tip. The blooming period occurs during the summer for about 2 months. Each flower is replaced by a bristly 2-celled carpel; each globoid cell contains a single small seed. The cells of the carpel eventually separate. The root system produces rhizomes. This plant can spread by reseeding itself, or by forming vegetative offshoots from the rhizomes.

The common name refers to the vanilla scent of the dried foliage. Sweet-Scented Bedstraw is typically found in moist woodlands. It can be distinguished from other Galium spp. (Bedstraws) by considering the following set of features

(key to state/province abbreviations)

Northern bedstraw:
UNITED STATES

AK AZ CA CO CT DE ID IL IN IA KY ME MD MA MI MN MO MT NE NV NH NJ NM NY ND OH OR PA RI SD TN TX UT VT VA WA WV WI WY DC PR VI

CANADA

AB BC MB NF NT ON SK

Sweetscented bedstraw:
UNITED STATES

AL AK AZ AR CA CO CT DE FL GA ID IL IN IA KS KY LA ME MD MA MI MN MS MO MT NE NV NH NJ NM NY NC ND OH OK OR PA RI SC SD TN TX UT VT VA WA WV WI WY DC PR VI

CANADA

AB BC MB NB NF ON SK

Bedstraw is widely distributed throughout North America and Canada [127,269]. Sweetscented 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.

Sweet-Scented Bedstraw occurs in most areas of Illinois, where it is occasional to locally common (see Distribution Map). Habitats include moist to mesic deciduous woodlands, rocky bluffs, areas along woodland paths, thickets, moist meadows, partially shaded seeps and springs, partially shaded riverbanks, and bogs. In badly degraded woodlands, this plant is replaced by Galium aparine (Cleavers).

More info on this topic.

This species can be found in the following regions of the western United States (according to the Bureau of Land Management classification of Physiographic Regions of the western United States):

BLM PHYSIOGRAPHIC REGIONS [29]:

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

Galium jalapense Schltdl.:
Mexico (Mesoamerica)

Note: This information is based on publications available through Tropicos and may not represent the entire distribution. Tropicos does not categorize distributions as native or non-native.

Galium triflorum var. asprelliforme Fernald:
Canada (North America)
United States (North America)

Note: This information is based on publications available through Tropicos and may not represent the entire distribution. Tropicos does not categorize distributions as native or non-native.

Galium triflorum Michx.:
Canada (North America)
Greenland (North America)
Japan (Asia)
North Korea (Asia)
Russian Federation (Asia)
South Korea (Asia)
United States (North America)
Mexico (Mesoamerica)
China (Asia)

Note: This information is based on publications available through Tropicos and may not represent the entire distribution. Tropicos does not categorize distributions as native or non-native.

Canada

Origin: Unknown/Undetermined

Regularity: Regularly occurring

Currently: Unknown/Undetermined

Confidence: Confident

United States

Origin: Native

Regularity: Regularly occurring

Currently: Present

Confidence: Confident

Type of Residency: Year-round

More info for the terms: forb, rhizome, schizocarp

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 sweetscented 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.

Sweetscented bedstraw: Sweetscented 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].

More info for the terms: mesic, taiga

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].

Sweetscented bedstraw: In the Intermountain West, moist woods and riparian areas are typical sweetscented bedstraw habitat [59,280]. In more southwestern regions, sweetscented bedstraw is restricted to mesic, shady sites [172,190]. In the Great Plains states, sweetscented bedstraw rarely occupies moist prairie sites [92]. Voss [275] describes sweetscented 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, sweetscented bedstraw is common to deciduous forests, fields, brush thickets, and roadsides [75,211].

Elevation: Several western regions report elevational ranges for northern and sweetscented 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]

Sweetscented 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/sweetscented bedstraw HT) 854-2,151 m central and eastern Montana (subalpine fir/sweetscented 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 5.0±1.3% (s x) at snow levels of 24 inches (60 cm), 9.0±1.3% at 48 inches (120 cm), and 11.2±3.2% at 95 inches (240 cm). Flowering was delayed on sites with 95 inches (240 cm) of snow accumulation [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.

Sweetscented bedstraw: Soils described in sweetscented bedstraw habitats on Vancouver Island, British Columbia, are acidic and nitrogen rich [136]. In the subalpine fir/sweetscented bedstraw habitat type of central and eastern Montana, soils range from neutral to strongly acidic [105]. Sweetscented bedstraw habitat in the Adirondacks has "higher" pH soils [144].

More info for the terms: association, climax, constancy, fern, forb, frequency, hardwood, mesic, presence, shrub, swamp, taiga

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

Northern and sweetscented 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 sweetscented bedstraw has greater constancy than northern bedstraw.
In the subalpine fir/sweetscented bedstraw habitat type, sweetscented 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: sweetscented bedstraw [3,68]

ID: sweetscented bedstraw [100,250,292]

MT: sweetscented bedstraw [50,105,204,250]

NM: sweetscented bedstraw [68]

WY: northern bedstraw [50]

        sweetscented 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].

Sweetscented bedstraw-

Northwest: Sweetscented bedstraw is
a common understory species in numerous coniferous, deciduous, and mixed forests
of the northwest.

Coniferous forests:
In southeastern Alaska, sweetscented bedstraw inhabits several Sitka spruce
(Picea sitchensis), western hemlock, and mixed conifer habitat types [171].
Sweetscented bedstraw is characteristic of several productive Douglas-fir-dominated
habitats of southwestern British Columbia [93]. Sweetscented 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, sweetscented bedstraw is common to several western hemlock forests.
In the Gifford Pinchot National Forest, sweetscented 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, sweetscented 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, sweetscented bedstraw occurs in several
community types characterized by the presence of Port-Orford-cedar (Chamaecyparis lawsoniana)
and western hemlock or fir [111]. Constancy of sweetscented 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 sweetscented bedstraw in Idaho.
In the Selway-Bitterroot Wilderness, sweetscented bedstraw persists in 315- to
600-year-old western redcedar stands [97]. In east-central Idaho, the presence of
sweetscented bedstraw identifies the Engelmann spruce/sweetscented bedstraw habitat
type. Other habitat types where sweetscented 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 sweetscented bedstraw as an important understory species.
From Montana to northwestern Wyoming, the Engelmann spruce/sweetscented 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]. Sweetscented bedstraw is common in western larch
(Larix occidentalis)- and whitebark pine-dominated forests of the northern Rockies
[48]. Constancy of sweetscented bedstraw is between 95% and 100% in the spruce/sweetscented
bedstraw, subalpine fir/sweetscented 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 sweetscented bedstraw habitats include aspen,
poplar, alder, and dogwood (Cornus spp.) in the Northwest. In the taiga of interior
Alaska, sweetscented bedstraw is typical of mature balsam poplar/devil's club
stands [85]. In Alberta, researchers found sweetscented bedstraw associated with
and growing on decaying logs and stumps in 28-year-old, aspen-dominated boreal
forests [159].

Sweetscented 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
sweetscented bedstraw in ponderosa pine-California black oak (Q. kelloggii) and western
hemlock-tanoak (Lithocarpus densiflora) vegetation of southwestern Oregon. Sweetscented
bedstraw occurs in riparian vegetation of the Trout Creek Mountains as well [80].

Riparian vegetation typical of Montana and southern Idaho includes sweetscented bedstraw
[100].
Sweetscented 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/sweetscented bedstraw is a major community type at elevations below 6,595 feet
(2,010 m) [292].

Southwest:
Sweetscented bedstraw is a typical understory species in several southwestern coniferous,
deciduous, and mixed forest types.

Coniferous forests:
Sweetscented 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, sweetscented 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]. Sweetscented 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, sweetscented 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, sweetscented bedstraw typifies the
white fir/sweetscented bedstraw riparian forest habitat type [3,68]. Hayward [112]
describes sweetscented 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, sweetscented 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
sweetscented bedstraw habitat. Southern California walnut (J. californica) and
coast live oak (Q. agrifolia) make up the overstory and wild oat (Avena fatua)
and sweetscented bedstraw the understory [209]. In the Humboldt-Toiyabe National Forest,
sweetscented bedstraw occupies several communities dominated by quaking aspen, red-osier
dogwood, and/or willow (Salix spp.) [169]. Kartesz [132] reports sweetscented bedstraw
in Nevada's California red fir forests.

In western Colorado, sweetscented 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 sweetscented bedstraw
communities [20]. Sweetscented bedstraw is a principal understory species in the blue
spruce/red-osier dogwood habitat type of southern Colorado and northern New Mexico [3,68].
Sweetscented 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:
Sweetscented 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, sweetscented bedstraw occurs in green ash-box elder
forests [46]. Sweetscented 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, sweetscented bedstraw frequency is 19% [83].

In cedar swamps of northern Wisconsin, sweetscented 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. Sweetscented 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 sweetscented bedstraw in the
understory.

Deciduous and mixed forests:
Sweetscented 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
sweetscented bedstraw in northwestern Pennsylvania's hemlock-beech vegetation. In New
York's Adirondack uplands, sweetscented bedstraw is found in red maple, striped maple
(Acer pensylvanicum), fir, yellow birch, and beech (Fagus spp.) mixed forests
[144]. Sweetscented 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 sweetscented 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, sweetscented
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, sweetscented
bedstraw is important in swamp white oak (Q. bicolor) communities [273].

Southeast:
Southeastern sweetscented bedstraw habitats include hardwood and river bottom forests.
Deciduous and mixed forests:
Sweetscented 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]. Sweetscented
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].

More info on this topic.

This species is known to occur in association with the following Rangeland Cover Types (as classified by the Society for Range Management, SRM):

More info for the terms: cover, hardwood

SRM (RANGELAND) COVER TYPES [240]:

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

ALASKAN RANGELANDS

901 Alder

904 Black spruce-lichen

906 Broadleaf forest

908 Fescue

920 White spruce-paper birch

921 Willow

More info on this topic.

This species is known to occur in association with the following cover types (as classified by the Society of American Foresters):

More info for the terms: cover, swamp

SAF COVER TYPES [81]:

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

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This species is known to occur in association with the following plant community types (as classified by Küchler 1964):

More info for the terms: bog, shrub

KUCHLER [143] PLANT ASSOCIATIONS:

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

More info on this topic.

This species is known to occur in the following ecosystem types (as named by the U.S. Forest Service in their Forest and Range Ecosystem [FRES] Type classification):

ECOSYSTEMS [89]:

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

Sweet-Scented Bedstraw occurs in most areas of Illinois, where it is occasional to locally common (see Distribution Map). Habitats include moist to mesic deciduous woodlands, rocky bluffs, areas along woodland paths, thickets, moist meadows, partially shaded seeps and springs, partially shaded riverbanks, and bogs. In badly degraded woodlands, this plant is replaced by Galium aparine (Cleavers).

The flowers are occasionally visited by small bees or flies; these insects usually suck nectar. The caterpillars of several moths feed on the foliage of Galium spp. (Bedstraws), including Lobocleta ossularia (Drab Brown Wave), Pleuroprucha insularia (Common Tan Wave), and Scopula limboundata (Large Lace Border). Deer leave the foliage alone. The bristly carpels of Sweet-Scented Bedstraw can cling to the fur of animals and the clothing of humans; by this means, the seeds are distributed to new locations.

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More info for the terms: association, climax, constancy, cover, density, eruption, frequency, hardwood, mesic, presence, severity, shrub, stringer, succession, taiga, tree, wildfire

Northern and sweetscented 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. Sweetscented 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. Sweetscented 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. Sweetscented 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 sweetscented 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 sweetscented 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) 3.82 2 year-old-mound (n = 40) 7.32 undisturbed quadrat
(n = 25) 3.12* * 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].

Sweetscented bedstraw: Like northern bedstraw, sweetscented 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 sweetscented bedstraw.

After reviewing successional change and disturbance dynamics studies within western forests, McKenzie and others [178] classify sweetscented bedstraw as a "release herb," one that responds positively to canopy removal or other disturbance. Sweetscented bedstraw successfully colonized sites that were substantially disturbed in northwestern Connecticut. White pine forests were clearcut and then bulldozed to expose the mineral soil. Sweetscented 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. Sweetscented bedstraw coverage decreased with increasing disturbance intensity. Undisturbed and low- and high-intensity rural sites had significantly (p<0.0001) more sweetscented bedstraw coverage than urban or suburban sites [185].

A study of alluvial deposits along the McKenzie River in Oregon revealed the highest cover of sweetscented bedstraw in the earliest seral community. On low floodplain areas dominated by red alder, sweetscented bedstraw had 6% canopy cover. On high floodplains, grand fir replaced red alder after 30 to 70 years, and here sweetscented bedstraw had 3% cover. In later seral stages dominated by Douglas-fir and western hemlock, sweetscented 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 sweetscented 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, sweetscented bedstraw occurred in all forests 14 to 140 years old [72]. Habeck [97] reports sweetscented 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, sweetscented 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 sweetscented bedstraw frequency was significantly (p≤0.05) lower in more open sites [27]. In central Idaho's Douglas-fir/ninebark habitat type, sweetscented 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 sweetscented 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, sweetscented 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 sweetscented bedstraw was highest in sites receiving 25% to 60% full light. The range of full sunlight received and corresponding sweetscented 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. Sweetscented 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, sweetscented 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 sweetscented bedstraw occurrence within this time frame [217].

In boreal mixed woods of Thunder Bay, Ontario, sweetscented 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, sweetscented 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 sweetscented 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 sweetscented bedstraw increases following canopy release predominate, Freedman and Habeck [87] found coverage and presence of sweetscented 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 sweetscented 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 sweetscented bedstraw was significantly lower (p=0.05) on untreated and tree-length burned sites in the 2nd posttreatment year. The density of sweetscented bedstraw for each of the treatments is given below [195]:

Post-treatment year

1st

2nd 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/m²) 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: Sweetscented 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 sweetscented 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 sweetscented 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. Sweetscented bedstraw percent constancy (or percent occurrence in all plots) was greatest the 2nd recovery year. The percent constancy of sweetscented bedstraw during the succession of this area is provided below [196].

Year 1990 1991 1992 1993 1996 1999 sweetscented 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. Sweetscented bedstraw occurred at the edge of aspen stands considered by the researcher to be "far from climax."

More info for the terms: fire management, presence

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/m²) Calories/g
(ash-free) Calories/m² % total (calories/m²) 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.

More info for the terms: basal area, cover, crown fire, density, duff, fire frequency, fire severity, fire tolerant, fire use, frequency, fuel, hardwood, litter, mesic, mixed-severity fire, natural, prescribed fire, presence, restoration, rhizome, series, severity, surface fire, wildfire

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 sweetscented bedstraw:
The postfire response is not always the same for northern and sweetscented
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 sweetscented 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. Sweetscented 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 sweetscented 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.
Sweetscented 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

(seeds/m²)2nd postfire year coverage (%)
Fire typeUBLSUBLSNorthern bedstraw1.9000.642.63.7Sweetscented bedstraw24.530.111.3

0.66

0.030.01
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 sweetscented 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 (%)SiteDominantsintact overstoryno overstoryintact overstoryno
overstory1eastern white pine
black oak

white oak040,6000502northern red oak
white oak

sweet birch5,60018,4002575
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.

FirePrefireLight Moderate Heavy Light Moderate Heavy Postfire yearNA

3

12

Production (kg/ha)33423150131415
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.
CommunityHerbaceous understory
(low
severity)Juniper understory
(high
severity)Burn statusprefire (1980)postfire (1982)prefire (1980)postfire (1982)Density

(number of stems/0.1m)1.23.11.42.7Coverage (%)0.70.50.50.9
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.
SiteBurned
(October 1972)Reburned
(May 1978)Unburned Frequency (%)20208Cover (%)83.21.2Prominence value*35.814.33.4
* 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 statusUnburned (n=125)Burned (n=125)Time since fire03 monthsCover (%)<0.054Frequency (%)722

Burn statusUnburned (n=23)Burned (n=28)Year197019711972197019711972Cover (%)1<0.05<0.05332Frequency (%)172222293229

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].

Sweetscented bedstraw:
The following information suggests that sweetscented bedstraw is not as fire tolerant
as northern bedstraw. Fewer studies report increases in sweetscented bedstraw following
low-severity and dormant season fires than were reported for northern bedstraw.
In north-central Idaho western hemlock-western redcedar habitats, sweetscented
bedstraw was absent 3 years following fire. Fire timing or severity are unknown.
Steele and Geier-Hayes [249] consider sweetscented 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 sweetscented 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. Sweetscented 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, sweetscented 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:
Sweetscented 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, sweetscented 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, sweetscented 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 76°F (21-24 °C),
relative humidity was 43% to 50%, and winds were 1 to 8 mph. The dry season fire burned
on September 13 and 14, 1989, when air temperatures were 54 °F to 77 °F (12-25 °C),
relative humidity was 39% to 66%, and winds were 1 to 5 mph. Coverage of sweetscented
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 typePrefire cover (%)Postfire cover (%)Unburned1.43.2Moist burn0.90.7Dry burn1.60.8
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 sweetscented bedstraw on the
burned sites had not regained prefire levels by 11 years postfire. On logged
unburned sites, increased sweetscented 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 sweetscented 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 sweetscented bedstraw coverage and frequency following
fire predominate, the frequency of sweetscented bedstraw increased following
low-severity, spring prescription fires in quaking aspen woodlands of southern
Ontario. The frequency of sweetscented 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, sweetscented 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 sweetscented 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. Sweetscented 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. Sweetscented 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]:
FireUnburnedSpring fireSummer fireYears postfire0035142511Occurrence (%)33737033

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). Sweetscented
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. Sweetscented 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].

Sweetscented 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 sweetscented 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 sweetscented 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. Sweetscented bedstraw frequency of occurrence is provided below [160]:
Time since firePrefire3 months1year2 yearsBurned & seeded

(# of occurrences/10 plots)6002Burned
(# of occurrences/10 plots)3211
Repeated fires:
The only study reporting sweetscented 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 sweetscented 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:

• 
  Effects of surface fires in a mixed red and eastern white pine stand in Michigan
• 
  Vegetation response to restoration treatments in ponderosa pine-Douglas-fir forests of western Montana

More info for the term: frequency

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 sweetscented bedstraw as a residual colonizer, coming from an on-site ground source.

More info for the term: rhizome

Powell [206] indicates that northern bedstraw has better chances of surviving fire than sweetscented bedstraw. Sweetscented 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 sweetscented bedstraw a nonsurvivor because of its delicate rhizome; this classification was tentative as the researchers observed few postfire responses. However, Edgerton [77] described sweetscented bedstraw as a "surviving forb" following clearcutting and broadcast burning of a mixed conifer forest in the Umatilla National Forest of Oregon.

More info for the terms: fire severity, severity

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.

More info for the terms: ground residual colonizer, initial off-site colonizer, rhizome, secondary colonizer

POSTFIRE REGENERATION STRATEGY [254]:
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)

More info for the terms: fire regime, fire severity, presence, severity

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 sweetscented 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/sweetscented bedstraw and subalpine fir/sweetscented 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]. Sweetscented 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. If you are interested in plant communities or ecosystems that are not listed below, see the complete FEIS fire regime table.

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 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. 200,289] blue grama-needle-and-thread grass-western wheatgrass Bouteloua gracilis-Hesperostipa comata-Pascopyrum smithii 200,227,289] cheatgrass Bromus tectorum 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 277] western juniper Juniperus occidentalis 20-70 Rocky Mountain juniper Juniperus scopulorum 200] western larch Larix occidentalis 25-350 [13,24,63] yellow-poplar Liriodendron tulipifera 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. 200] whitebark pine* Pinus albicaulis 50-200 [1,10] jack pine Pinus banksiana 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 277] eastern cottonwood Populus deltoides 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. 12] oak-hickory Quercus-Carya spp. < 35 northeastern oak-pine Quercus-Pinus spp. 10 to 277] oak-gum-cypress Quercus-Nyssa-spp.-Taxodium distichum 35 to > 200 [191] southeastern oak-pine Quercus-Pinus spp. 277] coast live oak Quercus agrifolia 2-75 [95] white oak-black oak-northern red oak Quercus alba-Q. velutina-Q. rubra 277] canyon live oak Quercus chrysolepis 12] northern pin oak Quercus ellipsoidalis 277] Oregon white oak Quercus garryana 12] California black oak Quercus kelloggii 5-30 [200] bur oak Quercus macrocarpa 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 to277] little bluestem-grama prairie Schizachyrium scoparium-Bouteloua spp. 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. 74,277] *fire return interval varies widely; trends in variation are noted in the species review
**mean

More info for the terms: constancy, density, frequency, mesic, natural, perfect, rhizome, xeric

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: Sweetscented 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 sweetscented bedstraw seeds/m² although sweetscented 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 sweetscented bedstraw in collected seed rain was 0.2% in clearcut areas and 0.01% in partially-cut areas, although sweetscented 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 sweetscented 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. Sweetscented bedstraw's migration rates were 0.87±0.55 (s x) m/year and 0.91±0.55 m/year based on the furthest 1/2 peak density and furthest individual calculations, respectively [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 50±80 (s) northern bedstraw seeds/m² were in spring seed banks and 133±103 seeds/m² were in fall seed banks. The coverage of northern bedstraw in the study area was 5% and constancy was 96% [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"/m².

Sweetscented bedstraw: In old-growth deciduous forests of southwestern Quebec, researchers calculated a maximum of 25 sweetscented bedstraw seeds/m² from soil samples collected in May. Sugar maple, striped maple, American beech, and white ash between 150 and 450 years old dominated the sites [157]. Soil samples 4 inches (10 cm) deep from a total area of 2.3 m² were collected from early May to late August in Douglas-fir-grand fir forests of central Idaho and contained 23 total viable sweetscented bedstraw seeds. The maximum seed density was 126/m². 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. Sweetscented 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 sweetscented bedstraw seeds/m² emerged from soil collected on 3rd order streams where coverage of sweetscented bedstraw was 1%; on 5th order streams, 36.5 seeds/m² emerged from soil collected on sites with 1.4% sweetscented bedstraw coverage. On annually flooded gravel bars, collected soil had 0.5 to 3.5 seeds/m² where coverage of sweetscented 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 sweetscented 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. Sweetscented bedstraw plants were present in each study site. The distribution of sweetscented 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

144

Mean seeds± SE/m² 20±6 (litter) 7±2
(0-5cm) 7±2
(5-10cm) 60±20 (litter) 2±1
(0-5cm) 80±33
(0-5cm) 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, sweetscented 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 sweetscented 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].

Site

Drier

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 Site

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 sweetscented bedstraw's rhizomes to be more fire sensitive than those of northern bedstraw.

More info on this topic.

More info for the term: geophyte

RAUNKIAER [213] LIFE FORM:
Geophyte

More info for the term: forb

Forb

More info on this topic.

From areas reporting seasonal development of both northern and sweetscented bedstraw, it appears that sweetscented 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.

Sweetscented 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 sweetscented bedstraw sets fruit from late June to late August.

The following is a representative barcode sequence, the centroid of all available sequences for this species.

Barcode of Life Data Systems (BOLDS) Stats
Public Records: 1
Specimens with Barcodes: 7
Species With Barcodes: 1

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

Canada

Rounded National Status Rank: NNR - Unranked

United States

Rounded National Status Rank: N5 - Secure

Rounded Global Status Rank: G5 - Secure

More info for the terms: forbs, frequency, nonnative species, presence

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.

Sweetscented bedstraw:
Several studies suggest that sweetscented bedstraw can indicate environmental conditions and
productive sites in several Pacific Northwest forests. In western Oregon and southwestern
Washington, sweetscented bedstraw is indicative of moist, well-drained sites in low to
mid-elevation forests [102]. The presence of sweetscented bedstraw in riparian zones of central
Oregon suggests high productivity sites for conifers [137]. Sweetscented 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 sweetscented 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 sweetscented 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; sweetscented
bedstraw increased by more than 30% after given 3 years recovery time. Sweetscented
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), sweetscented 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].

The preference is light shade to partial sun, moist conditions, and a fertile loamy soil.