Fragrant bedstraw: In old-growth deciduous forests of southwestern Quebec, researchers calculated a maximum of 25 fragrant bedstraw seeds/m² from soil samples collected in May. Sugar maple, striped maple, American beech, and white ash between 150 and 450 years old dominated the sites [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 fragrant 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. Fragrant bedstraw occurred in 2 edge plots and 3 forested plots, but no seed germinated from soil collected from any of the 3 sites [153].

On intermittently flooded (2- to 10-year flood intervals, typically) gravel bars, 36 fragrant bedstraw seeds/m² emerged from soil collected on 3rd order streams where coverage of fragrant bedstraw was 1%; on 5th order streams, 36.5 seeds/m² emerged from soil collected on sites with 1.4% fragrant bedstraw coverage. On annually flooded gravel bars, collected soil had 0.5 to 3.5 seeds/m² where coverage of fragrant bedstraw was 1% or less. Streams flowed in Oregon's Cascade Range [106].

The seed banks from undisturbed and disturbed communities in southwestern British Columbia reveal an increased density of fragrant bedstraw seed with increased disturbance levels. Characteristic species in undisturbed and slightly disturbed sites included Douglas-fir, western hemlock, western redcedar, maple, and red alder. Highly disturbed sites occurred within maintained right of ways. Fragrant bedstraw plants were present in each study site. The distribution of fragrant bedstraw along a disturbance gradient and within the soil profile is provided below [176].

Ahlgren [2] compared seedling emergence from intact blocks of soil collected in late summer from severely burned and unburned sites. The author described the fire as recent, but time since fire was unclear. In the 270-year-old red pine stands of northeastern Minnesota, fragrant bedstraw occurred at 40% frequency on burned sites and 93% frequency on unburned sites. Based on greenhouse experiments, the author calculated that 10,890,000 seedlings/ha could germinate from burned soils and 218,000 seedlings/ha could emerge from unburned soils.

In north-central Idaho's western hemlock-western redcedar forests, northern bedstraw did not occur in the earliest seral community (burned 3 years prior), but was present in all others described as immature shrub to near climax communities [225]. Similarly, in northern lower Michigan, studies in mature 2nd growth (55-82 years old) and disturbed (≤15 years old) stands revealed an association between fragrant bedstraw and disturbed mesic sites. Quaking aspen, sugar maple, and American beech dominated the mesic sites [222]. In hybrid white spruce × Engelmann spruce forests of central British Columbia, fragrant bedstraw occurred in all forests 14 to 140 years old [72]. Habeck [97] reports fragrant bedstraw in climax (315- to 600-year old stands) western redcedar forests in Idaho's Selway-Bitterroot Wilderness.

While the above studies suggest a tolerance of early-, mid-, and late seral conditions, the following studies indicate that preferences within a community type or area exist as well. In Douglas-fir forests, fragrant bedstraw frequency of occurrence was significantly (p< 0.01) greater in mature (80-195 years) forests than in old-growth (≥195 years) or young (< 80 years) forests in Oregon's Cascade Mountains. In western Washington's Douglas-fir forests, however, northern bedstraw frequency of occurrence was almost equal in mature and young forests, but was significantly lower (p< 0.01) in old-growth forests [247]. In rich mesic forests of western Massachusetts, researchers found fragrant bedstraw frequency was significantly (p≤0.05) lower in more open sites [27]. In central Idaho's Douglas-fir/ninebark habitat type, fragrant bedstraw is considered a major late seral species that decreases following logging and wildfire disturbances [249].

Light intensity relationships: Much of the following information addresses fragrant bedstraw's light intensity preference as a result of logging practices. While light intensity is indeed altered through logging operations, mechanical soil disturbances also occur and may influence findings. In general, fragrant bedstraw favors diffuse light over full sun or full shade conditions.

In mixed conifer forests of southeastern Oregon's Siskiyou Mountains, the percent cover of fragrant bedstraw was highest in sites receiving 25% to 60% full light. The range of full sunlight received and corresponding fragrant bedstraw coverage were as follows [79]:

Researchers compared old-growth, even-aged, and uneven-aged hardwood (sugar maple, basswood, yellow birch, and eastern hemlock) forests in northern Wisconsin and Michigan. Fragrant bedstraw coverage, frequency, and constancy were greater in uneven-aged forests where photosynthetically active radiation levels were significantly (α = 0.05) greater than in either other type [233]. Likewise, in southern boreal forests of northeastern Minnesota, researchers established that on average, fragrant bedstraw occurred with similar density on postfire and postlogging sites aged 25 to 100 years. This finding suggests that canopy release was the most important factor in fragrant bedstraw occurrence within this time frame [217].

In boreal mixed woods of Thunder Bay, Ontario, fragrant bedstraw abundance increased in riparian areas adjacent to upland burned sites as compared to riparian areas next to undisturbed woodlands. Sites burned in the 1999 Nipigon Fire, and the study was published in 2003. The authors suggest increased light availability as the reason for increases in riparian sites dominated by red-osier dogwood and thinleaf alder [149]. In black spruce forests of northeastern Ontario and western Quebec, fragrant bedstraw reached a high of 1.9% coverage on nutrient-rich logged sites where the stand age averaged 35.5 years and a high of 0.9% cover on nutrient-rich unlogged sites [35]. Increases in fragrant bedstraw density were significant (p<0.01) following thinning treatments in giant sequoia groves of Tulare County, California [156]. Frequency following the thinning reportedly decreased, however [133].

While fragrant bedstraw increases following canopy release predominate, Freedman and Habeck [87] found coverage and presence of fragrant bedstraw to be lower in treated (logged, burned, logged and burned) versus untreated Douglas-fir-, ponderosa pine-, and western larch-dominated forests in Swan Valley, Montana.

Other studies suggest that season of canopy removal and method of removal may affect the response of fragrant bedstraw. In northern Minnesota, researchers monitored understory vegetation changes for 2 years following winter and spring logging and 2 methods of harvest, full-tree logging (trees skidded intact) and tree-length logging (trees limbed and topped on site). All trees greater than 1 inch (2.5 cm) dbh were cut. On the tree-length logged site, piles were burned in July. Fire conditions included a high build-up index of 26, relative humidity of 45%, average temperature of 88 °F (31 °C), and an initial wind speed of 19 km/h with gusts of up to 48 km/h. Density of fragrant bedstraw was significantly lower (p=0.05) on untreated and tree-length burned sites in the 2nd posttreatment year. The density of fragrant bedstraw for each of the treatments is given below [195]:

Major disturbance events: Fragrant bedstraw is often present in very early seral communities resulting from extreme weather events or volcanic activity. After the eruption of Mount St. Helens in Washington, snow and ice rapidly melted from the volcano sides. As water flowed, it collected rocks, debris, and organic materials. Massive amounts of material were deposited along and in the Muddy River. Following these events, researchers recorded fragrant bedstraw on stump bases with deposits of organic material, on root wads of uprooted trees, in moist depressions and sinks of the mudflow channel, and on sites with original soils covered with a layer of mudflow material. The frequency of fragrant bedstraw on the Muddy River was 6% [101].

A debris flow along a 2nd order stream in Douglas-fir, western hemlock, and red alder communities in Oregon's central Coast Range occurred in the winter of 1989 and 1990. Researchers monitored vegetation changes for 10 years following the event. Fragrant bedstraw percent constancy (or percent occurrence in all plots) was greatest the 2nd recovery year. The percent constancy of fragrant bedstraw during the succession of this area is provided below [196].

Bailey [18] revisited sites affected by the 1914-15 eruptions in northeastern California's Lassen Volcanic National park in 1963. Fragrant bedstraw occurred at the edge of aspen stands considered by the researcher to be "far from climax."

SEASONAL DEVELOPMENT:
From areas reporting seasonal development of both northern and fragrant bedstraw, it appears that fragrant bedstraw development is slightly later than northern bedstraw's. The states or regions indicating flower or fruit set timing for bedstraw provide broad ranging dates to incorporate year-to-year variation in climate and wide regional distributions.

Northern bedstraw:

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

Fragrant bedstraw:

FIRE ECOLOGY

• FIRE ECOLOGY OR ADAPTATIONS
• POSTFIRE REGENERATION STRATEGY

Fire regimes: A diversity of communities provide bedstraw habitat, and since fire regimes are dictated by the overstory community, bedstraw experiences a wide range of fire regimes. Davis and others [64], in a review, classify the spruce/fragrant bedstraw and subalpine fir/fragrant bedstraw habitat types as having "infrequent, severe fires with long-lasting effects." The same habitat types described on the Lolo National Forest, Montana, occupy moist environments that burn infrequently. The estimated fire return interval for these sites is 24 to 140 years [164]. Both bedstraw species occupy northern spruce-fir forests that are typically maintained under moist conditions, and the fire return interval in these forests ranges from 35 to more than 200 years [74]. Much shorter fire return intervals are reportedly tolerated by bedstraw as well. Northern bedstraw is typical in fescue-oatgrass mountain grasslands that are characterized by a fire return interval of less than 35 years [200]. Fragrant bedstraw occupies southern California walnut woodlands of southern California that burn annually due to an increased presence of annual grasses [209].

The following table provides fire return intervals for plant communities and ecosystems where bedstraw is important. For further information, see the FEIS review of the dominant species listed below. This list may not be inclusive for all plant communities in which bedstraw occurs. Find further fire regime information for the plant communities in which these species may occur by entering the species' names in the FEIS home page under "Find Fire Regimes".

FIRE EFFECTS

• IMMEDIATE FIRE EFFECT ON PLANT
• DISCUSSION AND QUALIFICATION OF FIRE EFFECT
• PLANT RESPONSE TO FIRE
• DISCUSSION AND QUALIFICATION OF PLANT RESPONSE
• FIRE MANAGEMENT CONSIDERATIONS

Northern and fragrant bedstraw: The postfire response is not always the same for northern and fragrant bedstraw in burned areas where both occur together.

Fire effects related to seasonality/severity: In the early 1960s, 17 wildfires burned in south-central New York. All but 1 of the fires burned in the spring, and sampling occurred 10 to 26 months following fire. Northern and fragrant bedstraw frequencies were significantly higher (p=0.01) on burned sites. Northern bedstraw averaged 2% frequency on unburned sites and 33% on burned sites within goldenrod (Solidago spp.)-poverty oatgrass habitats. Fragrant bedstraw averaged 2% frequency on unburned sites and 29% on burned sites in hardwood and mixed oak forests [260].

The postfire responses for northern and fragrant bedstraw were opposite in quaking aspen boreal forests of northeastern Alberta. Following a lightning-ignited spring wildfire, Lee [158] compared the immediate postfire seed banks and 2nd year postfire vegetation of unburned, "lightly" burned, and severely burned sites. Severely burned sites had all downed wood (≥ 7.9 inches (20 cm)) and the top 2.4 to 4 inches (6-10 cm) of organic material oxidized. Light burns partially oxidized small and mid-sized downed wood and just the top 0.8 inch (2 cm) of organic matter. Seed density estimates came from seedling and vegetative emergence techniques. Fragrant bedstraw rhizomes likely did not survive the fire while seed did, and the reverse was true for northern bedstraw. These findings may reflect different rhizome and seed heat tolerances for the 2 species or may indicate the occupation of different microsites where fire effects were different. Data are summarized below [158]:

Repeated fires: Sites in east-central Minnesota's oak savannas burned at frequencies of 0 to 19 years in a 20-year period. The coverage of fragrant bedstraw decreased with increased fire frequency; however, the r value for this relationship was -0.52 (significant at p < 0.10). Northern bedstraw coverage was not significantly changed by fire [263].

Northern bedstraw: Northern bedstraw recovers quickly following fire. Severe or growing season fires may result in decreased northern bedstraw coverage and/or frequency, but typically decreases are short lived.

Northern bedstraw is often mentioned as an important species in postburn communities of Canada. In the Selkirk Mountains of British Columbia, Shaw [238] lists northern bedstraw as a prominent herb in the early postfire reforestation of western hemlock, lodgepole pine, and quaking aspen forests. Seip and Bunnell [234] describe northern bedstraw in mountain grasslands resulting from stand-replacing fires in subalpine spruce forests of northern British Columbia. In coniferous forests of Alberta's eastern Rockies, northern bedstraw is most frequent in recently burned areas (10 to 20 years since fire) [52].

In interior Alaska's white spruce forests, Foote [85] visited sites burned between 6 months and 200 or more years ago. Northern bedstraw frequency was greatest but coverage was lowest on sites burned 6 months prior in a surface fire that scorched stems and killed some trees [85]. One of the fires that was included in the previous postfire recovery chronosequence was the 1950 Porcupine River fire. Foote [86] investigated the postfire vegetation recovery 1, 4, 7, 10, 23, and 30 years following the fire. Northern bedstraw frequency and coverage were greatest in the 10th postfire year [86].

The following studies highlight fire effects that are likely a result of fire severity or seasonality. These fire characteristics are difficult to consider singly; studies listed in this section highlight severity or seasonality.

Fire effects related mainly to severity: Generally, northern bedstraw increases following low-severity fires. The postfire response of northern bedstraw to high-severity fires is less predictable. Researchers burned ponderosa pine-dominated forests in the fall on the Coeur d'Alene, Idaho, Indian Reservation. Different fire severity levels resulted. High-severity fires consumed 80% of the duff layer, and low-severity fires removed 40%. Comparisons between unburned and burned sites revealed northern bedstraw frequency and coverage were greatest on sites burned in low-severity fires and lowest on sites burned in high-severity fires. However, treatment differences were not significant (p<0.1) [9]. See the Research Project Summary Understory recovery after low- and high-intensity fires in northern Idaho ponderosa pine forests for an extended report on this study.

In mixed oak forests of Eastford, Connecticut, researchers burned 2 sites in April. The 1st site burned in 1984, and the 2nd site burned in 1985. Fires burned under similar conditions; fuel moistures were between 18% and 28%, fire spread was slow (1m/min), and flame lengths were 11.8 inches (30 cm) or less. Within each site, portions burned more severely than others resulting in high mortality of the overstory. On the severely burned portion of site 1, 70% of the density and 60% of the basal area were removed. On the severely burned part of site 2, 95% of the density and basal area were removed. Northern bedstraw occurred only on burned sites. The density and frequency of northern bedstraw 7 to 8 years following these fires are shown below [73].

Prefire vegetation was compared to lightly (1%-20% of litter and duff consumed and 0-few trees killed), moderately (21%-80% of litter and duff consumed and <90% of trees killed), and heavily (81%-100% of litter and duff consumed and >90% of trees killed) burned vegetation following a late August prescription fire in quaking aspen-dominated communities of northwestern Wyoming. Northern bedstraw produced less biomass before the fire than 3 years following the fire on lightly and heavily burned sites. Northern bedstraw productivity was less 12 years following the fire than before the fire [25,26]. See the Research Project Summary Vegetation recovery following a mixed-severity fire in aspen groves of western Wyoming for an extended report on this fire study.

Fall prescription fires burned quaking aspen communities of Colorado's Front Range. Fire severity was greater on plots with an understory of common juniper than on plots with an herbaceous understory. Northern bedstraw densities were significantly (p=0.05) greater 1 year following fire. Increases were greater on less severely burned plots. The differences for pre- and postburn northern bedstraw coverage and density are given below [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.

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.

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]:

On the Namekagon River barrens of northern Wisconsin where jack pine and bur oak codominate, spring prescription fires burned. Northern bedstraw was common on both burned and unburned sites. On unburned sites, northern bedstraw averaged 89% frequency. On burned sites, frequency averaged 76%. An increased frequency of grasses following the fire may explain the slightly lower northern bedstraw frequency on burned sites [274].

McGee [177] compared early spring and late summer prescription fires in northwestern Wyoming's mountain big sagebrush communities. Two years after the fires, northern bedstraw coverage and frequency were greatest the 2nd postfire season on sites burned in the late summer. The coverage and frequency on unburned and spring burned sites were very similar [177].

In a fescue-oatgrass community of southern Alberta, researchers compared burned and unburned vegetation following a mid-December wildfire in 1997. The Granum wildfire burned when temperatures averaged 55 °F (13 °C), relative humidity was 17%, and winds were 19 to 25 mi/hr (30-40 km/hr) with gusts of 43.5 mi/hr (70 km/hr). Prefire fuel loads were unavailable, but nearby unburned sites had 900 kg/ha litter loadings. Postfire growing season precipitation was 46% greater than the long-term average. Northern bedstraw coverage was similar on interior burned plots and unburned plots 2 years following the fire. However, coverage was almost double on perimeter burn sites (those on the blackened side of fire line) when compared to unburned sites the 1st postfire year [31].

An early-spring prescription fire (May 2, 1972) stimulated northern bedstraw flowering on burned undisturbed mesic, highly-disturbed mesic, and on highly-disturbed wet to mesic prairie sites of northeastern Minnesota. Disturbances on the sites included grazing, sod production, and hay production but were discontinued approximately 15 years prior to the study. The fire occurred during periods of high humidity, virtually no wind, and wet to damp soils [201].

Repeated fires: The following studies report mixed postfire responses of northern bedstraw following multiple fires. Some report a tolerance of annual fires while others suggest that multiple fires followed by multiple years of rest are favored by northern bedstraw. Higgins and others [116] in a review suggest that northern bedstraw does not change or slightly decreases following periodic spring fires in the Northern Great Plains. In north-central South Dakota, northern bedstraw density was significantly greater (p<0.05) on northern mixed prairie plots burned annually for 3 consecutive years in the fall (October 5-17) than on unburned control plots [30].

In oak woodlands of east-central Minnesota's Cedar Creek Natural History Area, White [283] compared several burning schedules. All fires burned in the spring. However, particular overstory densities and soil series of the different sites were significantly (p≤0.05) correlated with northern bedstraw and could not be reliably related to burned sites [283].

Fragrant bedstraw: The following information suggests that fragrant bedstraw is not as fire tolerant as northern bedstraw. Fewer studies report increases in fragrant bedstraw following low-severity and dormant season fires than were reported for northern bedstraw. In north-central Idaho western hemlock-western redcedar habitats, fragrant bedstraw was absent 3 years following fire. Fire timing or severity are unknown. Steele and Geier-Hayes [249] consider fragrant bedstraw a major late-seral species in central Idaho's Douglas-fir/ninebark habitat type that decreases following logging and wildfires.

As time since fire increases however, the presence of fragrant bedstraw can decrease as well. In 1955 and 1956, Neiland [192] compared northwestern Oregon's mature (~300 years) western hemlock and Douglas-fir stands to sites that burned severely in 1933, 1939, and 1945. Fragrant bedstraw was absent from unburned forests but averaged 3% frequency on burned sites. In forests codominated by balsam fir, black spruce, and paper birch around Lake Duparquet, Quebec, fragrant bedstraw coverage was 1.9% on sites burned 26 years ago. On other sites that burned between 46 and 230 years ago, coverage of bedstraw varied from 0.1% to 0.5% [65].

The following studies highlight fire effects that are likely a result of fire severity or seasonality. These fire characteristics are difficult to consider singly; studies listed in this section highlight severity or seasonality.

Fire effects related mainly to severity: Fragrant bedstraw can survive low- and high-severity fires, but typically unburned frequencies or coverages are greater than those of burned sites. Following a "holocaustic" fire that killed all above ground vegetation, consumed all litter, and left bare mineral soil in the Pack River Valley of northern Idaho, fragrant bedstraw occurred on 5 of 18 sites and averaged 2% frequency [253]. Following severe fires in 270-year-old red pine stands of northeastern Minnesota, fragrant bedstraw occurred at 40% frequency on burned sites and 93% frequency in unburned stands [2]. See Seed banking for more information on this study.

In the Priest River Experimental Forest of northern Idaho, researchers compared the postfire regeneration following dry and moist prescription fires. Douglas-fir, western redcedar, and grand fir mixed forests were harvested and burned. The moist season burn occurred on June 1, 1989, when air temperatures were 69 °F to 76°F (21-24 °C), relative humidity was 43% to 50%, and winds were 1 to 8 mph. The dry season fire burned on September 13 and 14, 1989, when air temperatures were 54 °F to 77 °F (12-25 °C), relative humidity was 39% to 66%, and winds were 1 to 5 mph. Coverage of fragrant bedstraw decreased on both the moist and dry burn sites. There was no statistical analysis of the data. However, decreases were greater on dry burn sites. Coverage increased on unburned sites [243]:

Two forest sites within the Engelmann spruce-subalpine fir zone of central British Columbia were clearcut in the winter. One site burned in a low-severity prescribed fire the following fall. The coverage of fragrant bedstraw on the burned sites had not regained prefire levels by 11 years postfire. On logged unburned sites, increased fragrant bedstraw coverage lasted for 5 years following the disturbance [103]. See the Research Project Summary Revegetation in a subalpine forest after logging and fire in central British Columbia for an extended report on this study. Cox [57] compared the recovery of fragrant bedstraw in clearcut and clearcut and burned Douglas-fir forests of Oregon's Coast Range. The slash burn produced a moderately severe fire (litter, duff, and woody debris consumed, but mineral soil color unchanged). No prefire data were available. Differences between burned and unburned plots 1 and 2 years following fire were negligible [57].

While decreases in fragrant bedstraw coverage and frequency following fire predominate, the frequency of fragrant bedstraw increased following low-severity, spring prescription fires in quaking aspen woodlands of southern Ontario. The frequency of fragrant bedstraw on unburned sites was 4%. The frequency 4 months postfire was 21% and a little over 1 year postfire was 12.5% [244]. Following a mid-July crown fire near Missoula, Montana, fragrant bedstraw frequency had doubled from the 1st to the 2nd postfire year [58].

Fire effects related mainly to seasonality: Many of the following studies suggest that spring and fall fires may increase the frequency of fragrant bedstraw, while summer fires may decrease its frequency. Following spring fires in American beech-sugar maple and black oak-red maple forests in south-central New York, burned and unburned sites were compared. Fragrant bedstraw frequency was significantly higher (p=0.01) on burned sites; frequency on unburned sites was 2.4% and on burned sites was 28.6% [261].

In mixed conifer-hardwood forests of northeastern Minnesota, researchers assessed vegetation recovery in burned areas. Two sites dominated by black spruce, jack pine, and paper birch burned, one in late April and the other in mid-July. The late April fire occurred during high winds, leaving small unburned patches. Fragrant bedstraw frequency of occurrence on burned sites was over double that of unburned sites 3 years following the spring fire. The data collected on burned and unburned sites are summarized below [139]:

In white pine forests of Strafford County, New Hampshire, fall (1976) and spring (1977) prescription fires burned. The fires produced flame lengths of 3 to 24 inches (7.6-61 cm) and scorched trees at heights of 2 to 8 feet (0.6-2.4 m). Fragrant bedstraw was not on control plots and was not present on plots before the fire. However, it did occur following the fall and spring fires on white pine-dominated forests and following the spring fires in white pine mixed forests. Fragrant bedstraw plants on the burned plots resulted from seed germination. Plants on the spring-burned plots matured by late July and produced seed by the end of the growing season. The survival and/or development of plants on fall burned plots is unknown [42,226].

Fragrant bedstraw frequency decreased, but coverage was unchanged following a prescription fire in beetle-damaged white spruce forests of southern Alaska's Chuguch National Forest. The fire top-killed all overstory and understory vegetation in June of 1984. Prefire (1980) coverage of fragrant bedstraw was 2%, and frequency of occurrence was 24%. Seven years following the fire coverage remained 2% and the frequency of occurrence was 12% [121].

A prescription head fire within the Grand fir-Oregon boxwood (Paxistima myrsinites) habitat type of north-central Idaho also decreased the frequency of fragrant bedstraw. The fire burned mid-May of 1975 when temperatures were 82 °F (28 °C), relative humidity was 25%, and winds were negligible. Decreases in frequency were greater for sites that were grass seeded than unseeded sites following the fire. Statistical significance of the results was not addressed. Fragrant bedstraw frequency of occurrence is provided below [160]:

Repeated fires: The only study reporting fragrant bedstraw recovery following multiple fires indicates a tolerance of annual fires for up to 3 years. In southern Ohio hardwood forests, prescription fires burned some sites once and burned other sites for 3 consecutive years in March and April. Flame lengths were less than 20 inches (50 cm), and fire severity was low. The frequency of fragrant bedstraw increased by more than 10% on burned plots [128].

Hamilton's Research Papers (Hamilton 2006a, Hamilton 2006b) and the following Research Project Summaries provide further information on prescribed fire use and postfire response of many plant species including bedstraw:

• 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

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]:

MANAGEMENT CONSIDERATIONS

• IMPORTANCE TO LIVESTOCK AND WILDLIFE
• VALUE FOR REHABILITATION OF DISTURBED SITES
• OTHER USES
• OTHER MANAGEMENT CONSIDERATIONS

Northern bedstraw -
Livestock: Studies report conflicting responses of northern bedstraw to grazing. Several studies indicate an increased presence of northern bedstraw on sites grazed by livestock. The biomass of northern bedstraw was greater on grazed than ungrazed fescue grasslands of central Alberta. A decrease in grass yields was thought to facilitate northern bedstraw increases [17]. In aspen stands of Colorado and Wyoming, northern bedstraw is constant on moderately grazed ranges, and its removal from grazed vegetation may indicate mismanagement [55]. In rough fescue grasslands of southwestern Alberta, researchers tracked changes in the percent composition of northern bedstraw under different stocking rates and over a 32-year period. Northern bedstraw increased with length of grazing time but was relatively unaffected by stocking rates. Complete study results are shown below [285]:

Others report decreases in northern bedstraw with livestock grazing, or increased utilization of northern bedstraw with increased lengths of grazing time. On Douglas-fir/ninebark habitat types near Moscow, Idaho, the production and frequency of northern bedstraw was greater on ungrazed than cattle grazed sites. Ungrazed sites were not closed to native ungulate grazing, and stocking rates in the area averaged 1 animal/13 ha for 20 years [293]. On aspen ranges within the Black Mesa Experimental Forest of western Colorado, the utilization of northern bedstraw after 21 cattle-grazing days was 1%, after 38 days was 2%, and after 57 days was 6%. No utilization occurred after 78 days of use, but this was because most forbs on the site had senesced [199].

Native ungulates: The amount of northern bedstraw in elk, deer, mountain goat, and bighorn sheep diets is typically low, but season and/or stocking levels can increase utilization rates. Northern bedstraw made up a trace of winter mule deer diets in the Snowy Mountains of central Montana. After monitoring 96 feeding sites and analyzing 21 rumen samples, Kamps [130] found northern bedstraw constituted less than 0.5% of January diets and 1% of February diets. In a review of Rocky Mountain elk forage habits, Kufeld [145] considers northern bedstraw a least valuable forage plant. Least valuable forage is eaten by elk, but either makes up a small portion of the diet or is consumed in a much smaller proportion than is available. In June-collected elk feces from the Mount Saint Helens blast zone in southwestern Washington, just 0.1% of the total density was northern bedstraw [182].

Bentz and Woodard [28] consider northern bedstraw a secondary forage species for bighorn sheep in subalpine forests of southwestern Alberta. In the Sun River area of west-central Montana, 3 of 803 observed plant feeding instances by bighorn sheep were on northern bedstraw [232]. The stomach contents of 27 mountain goats from the Crazy Mountains of Montana contained 0.9% northern bedstraw by volume (0.5% by weight) in the summer and just a trace of the volume (0.1% by weight) in the fall. These findings came from 5 stomachs collected in the summer and 18 collected in the fall [230].

In several Canadian National Parks, Stelfox [251] compared bighorn sheep diets on winter ranges from 1968 to 1970 where the frequency of northern bedstraw ranged from 80% to 100%. In Waterton Lakes National Park, northern bedstraw did not comprise any portion of sheep diets. Zero utilization was likely because ungulate stocking rates were low. In Banff, northern bedstraw made up 9.1% of bighorn sheep's diet composition and was utilized at 20% frequency. Ungulate stocking rates were moderate in Banff. In Jasper National Park, ungulate stocking rates were high, and northern bedstraw comprised 1.1% to 2.2% of sheep diets but was utilized at frequencies of 12% to 86%. Utilization was greatest in the spring in Jasper and in the summer in Banff, but some utilization occurred year round in both parks [251].

Omnivores: Researchers recorded high levels of northern bedstraw usage by black bears in interior Alaska. Young stems and leaves were present in the spring diet. From 23 stomach contents, northern bedstraw occurred at 17% frequency and constituted 10.2% mean volume. From 16 intestines, the frequency of northern bedstraw was 33% and mean volume was 15%. No scat samples contained northern bedstraw [109]. Usage of northern bedstraw was less by black bears in the Rocky Mountains of southwestern Alberta, but research relied on scat samples alone. From scat collected in the summer of 1984 (n=22), northern bedstraw frequency of occurrence was 5%. Frequency was 8% in scat collected in the fall (n=13) of the same year [119].

Birds: Northern bedstraw may be important to breeding and ground foraging birds. Bird surveys in the Little Missouri National Grasslands of western North Dakota revealed heavy usage of ash woodlands where northern bedstraw is a prominent understory herb. Researchers conducted surveys from mid-May through mid-July in 1979, 1980, and 1981. The highest density of ground foragers and 531 nesting pairs/40 ha were in ash woodlands. Three bird species were exclusive to ash woodlands, and 6 species occurred with their highest densities in ash woodlands [123].

Insects: Findings from a single insect study indicate that northern bedstraw may be important to certain insect species. In a southeastern Minnesota pioneer cemetery site, a single collection of insects on northern bedstraw plants yielded 6 total insect species, 3 of which were unique to northern bedstraw. The insect species were not identified [215].

Fragrant bedstraw -
Native ungulates: While fragrant bedstraw has relatively low grazing value it is a valuable indicator of productive elk, deer, and moose habitat. On Vancouver Island, black-tailed deer ate the new growth of fragrant bedstraw in the spring and summer. Utilization was low given the abundance of the plant [56]. Throughout a 3-year-long study of habitat selection by elk in western Montana, the subalpine/fragrant bedstraw habitat type was "strongly selected for." Selection described the use of a vegetation type that exceeded the availability of the type. Elk used this habitat predominantly for feeding, although fragrant bedstraw was not a utilized food source [170]. Lonner [163] highlights moist sites within the same subalpine/fragrant bedstraw habitat type as very important elk summer range. In south-central Montana, the spruce/fragrant bedstraw habitat characterizes good year-round moose habitat and elk and deer winter range. The subalpine fir/fragrant bedstraw habitat in the same area receives moderate to heavy deer and elk use. Moose use valley bottom sites [204].

Other native mammals: Fragrant bedstraw may be an important rodent food source. In the Cascade foothills near Blue River, Oregon, 125-year-old Douglas-fir forests were logged and logged and burned. Fragrant bedstraw was an important herb in the 2nd postfire and postlogging years. The creeping vole increased in density on treated sites. The author considered increased vole densities and herbaceous understory vegetation to be related, as the vole feeds on the leaves and stems of shrubs and herbs [122].

Omnivores: Fragrant bedstraw identifies important grizzly bear habitat and is an important black bear food source. In the Bob Marshall Wilderness, Montana, the spruce/fragrant bedstraw habitat is ranked as the 2nd (out of 10) most important habitats for grizzly bears during the herbaceous foraging season (den emergence to July 31) and 3rd most important during the fruit foraging season (from August 1 to den entry). However, fragrant bedstraw was not listed as important grizzly bear food [167]. In a review, Rogers and Allen [224] list fragrant bedstraw as 1 of several herbaceous species commonly found in the early spring black bear diets in northeastern Minnesota and Massachusetts.

Palatability/nutritional value: Few studies address the palatability and nutritional content of bedstraw. The lack of fragrant bedstraw's inclusion in nutritional studies may be due to its low palatability [137]. Some have even suggested that bedstraw may be poisonous [147]. Below are some specific findings regarding nutritional value of northern and fragrant bedstraw in various environments.

Northern bedstraw: Paulsen [199] found northern bedstraw produces 17 pounds of forage/acre in aspen communities of western Colorado. Herbage production of northern bedstraw taken from western Montana's mountain grasslands ranged from 19 to 72 kg/ha (dried). Production was greater on southwest exposures than on northeast exposures [188]. Northern bedstraw on burned sites may have increased forage value. Following a spring fire in an eastern North Dakota tallgrass prairie, northern bedstraw herbage production was much greater on burned sites even though frequencies were the same on burned and unburned sites. The complete results of this study are summarized in the Fire Management Considerations section [99].

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

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

VALUE FOR REHABILITATION OF DISTURBED SITES:
Bedstraw may be valuable in the revegetation of abandoned mining sites. Northern bedstraw, although not directly seeded onto a coal mine spoil, was present at 20% to 75% frequency on a 31-year-old coal mine restoration site in southeastern Ohio [41]. Fragrant bedstraw made up 1.1% of the vegetative cover on a 15- to 20-year-old abandoned coal surface mine in Campbell County, Tennessee [212].

Northern bedstraw is successfully transplanted using a sod relocation method. Northern bedstraw survived when sod taken from undisturbed rough fescue grasslands in Alberta was transplanted to a new site [218]. During prairie restoration efforts in northern Wisconsin, researchers found direct seeding of northern bedstraw to be fairly successful but rated the transplanting success of seedlings in a sod form and as 1-year-old transplants as excellent. Individual seedlings showed poor survival in the field, but sod transplants survived even when there was no precipitation for the 2 weeks following transplanting [194].

OTHER USES:
There were multiple distinct uses of the 2 bedstraw species by native people.

Northern bedstraw: The Gwich'in Athabaskan people of Fort Yukon, Alaska, used a poultice of northern bedstraw green shoots to treat general aches and pains. The same shoots in tea treated cold symptoms [120].

Fragrant bedstraw: Ditidaht, indigenous people of the Pacific Northwest Coast, used fragrant bedstraw as a rinse to enhance the thickness and luster of their hair. Fragrant bedstraw flowers when dried were used as a perfume [205]. This species is also used to flavor wines [22]. In a review, Turner and Bell [267] report that the Kwakiutl people of British Columbia rubbed fragrant bedstraw on the skin to treat chest pains. Hellebore (Helleborus spp.) roots were often applied following this preparation.

OTHER MANAGEMENT CONSIDERATIONS:
Northern bedstraw: A study of grassland sites dominated by native and nonnative species suggests that northern bedstraw may decrease in coverage on sites invaded by nonnative forbs. Tyser [268] found northern bedstraw coverage was 1.8% in timothy (Phleum pratense)-dominated sites, 0.6% in native fescue-dominated sites, and 0.2% on sites invaded by spotted knapweed (Centaurea maculosa). Likely the differences in coverage relate to changes in species dominance as all sites had homogeneous topography, slopes, aspects, and substrates.

Fragrant bedstraw: Several studies suggest that fragrant bedstraw can indicate environmental conditions and productive sites in several Pacific Northwest forests. In western Oregon and southwestern Washington, fragrant bedstraw is indicative of moist, well-drained sites in low to mid-elevation forests [102]. The presence of fragrant bedstraw in riparian zones of central Oregon suggests high productivity sites for conifers [137]. Fragrant bedstraw is also 1 of several understory species indicating productive Douglas-fir habitat in southwestern British Columbia [134].

An extensive study of trampling in montane grasslands and forests of western Montana, suggests fragrant has high resiliency. The trampling treatments were completed by 130- to 190-pound people wearing lug-soled boots. Seventy-five to 100 trampling passes per year reduced fragrant bedstraw's frequency of occurrence by 50% and coverage increased less than 10% from the end of the 1st trampling season (August) to the beginning of the 2nd trampling season (June). Long-term resilience was high however; fragrant bedstraw increased by more than 30% after given 3 years recovery time. Fragrant bedstraw recovered in 10 months from less than 41 trampling passes without losing more than 20% of pretrampling coverage. If trampled for 3 seasons and given a longer recovery time (3 years), fragrant bedstraw tolerates high trampling levels (≥1,200 passes). Whether or not these findings of human trampling can be related to large herbivore trampling is unknown [49].

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