A study of postfire recovery in a southwestern Montana rangeland by Nimir and Payne [98] found limited evidence that fire reduces old man's whiskers immediately after burning. Although specific fire dates were not provided, it may be inferred that prescribed burning was conducted sometime in late May or early June on 2 sites. The burns were described as incomplete, "leaving some areas completely unburned, some partially burned, and others completely burned." Site 1 had "the most complete burn," perhaps due to its greater exposure to sun and wind and earlier snowmelt, compared with site 2 where "the fire was much less intense." Weekly estimates of old man's whiskers cover were averaged over a 12-week period following burning. On site 1, average cover was 1.70 sq dm2 / 20 sq dm on the unburned plots and 0.90 dm2 /20 dm2 on the burned plots. On site 2, average cover was 0.29 dm2/20 dm2 on the unburned plots and 0.20 dm2 /20 dm2 on the burned plots. The average cover values on site 1 were described as significantly (p< 0.01) different, while no difference in old man's whiskers cover between burned and unburned plots was detected for site 2. Old man's whiskers productivity, as measured in clipped plots, was described as significantly (no p-value provided) different between burned (32 kg/ha) and unburned (294 kg/ha) plots at site 1, while no information on old man's whiskers productivity at site 2 was provided. Statistical methods were not described for this study. Based on the above information it was concluded that old man's whiskers was "reduced by burning at both sites," and old man's whiskers was included in a list of species "damaged" by fire [98].
Schwecke and Hann [110] described a monitoring program that tracked the effects of prescribed burning on vegetation in a Rocky Mountain Douglas-fir (Pseudotsuga menziesii var. glauca)/rough fescue habitat type in western Montana. Their postfire observations suggest no negative effect of fire. Within sampling plots, individual plants (quantity not provided) were marked for monitoring prior to burning. Following both spring and fall burning, there was 0% mortality among the old man's whiskers plants that were monitored. Burn conditions were described (see [110]). The publication provides no qualitative description of direct fire effects on individual plants and no explanation for the lack of fire-caused mortality.
Several studies indicate that old man's whiskers populations may increase following fire.
Bailey and Anderson [14] reported that old man's whiskers was among several perennial forbs
that "responded quickly" following prescribed burning in rough fescue-porcupine
grass (Hesperostipa spartea) grasslands in central Alberta, but no data or further
description were provided. Bork and others [27] reported that old man's whiskers increased
"prominently" following prescribed burning in shrub-encroached grassland in
northwestern Alberta, but again, no data or further description were provided.
Bork and others [26] examined vegetation response following a December wildfire that
burned in rough fescue-Parry oatgrass (Danthonia parryi) grassland in southwestern
Alberta. No preburn data were collected. Estimated June mean canopy cover of old man's whiskers
in the interior of the burned area was 1.6% and 2.2% in postfire years 1 and 2, respectively.
Similar estimates were 0.1% and 0.3% along the perimeter of the burned area, and adjacent
unburned plots contained no old man's whiskers. Estimated mean 1- and 2-year postfire litter cover
was 11.2% and 55.4% in the interior of the burned area, 39.6% and 76.3% along the perimeter
of the burned area, and 91.7% and 97.9% in the adjacent unburned plots [26]. Although not
discussed by the authors, old man's whiskers cover following the fire may have been related to
litter consumption.
These data strongly suggest that old man's whiskers presence was diminished by both fire treatments.
Another study demonstrated no net effect of fire on old man's whiskers populations. Menke and Muir [92]
studied the effects of fire on a grassland community in the Snake River Canyon, southwestern Idaho.
Mean prefire cover and 1- and 2-year postfire cover of old man's whiskers were not significantly (p<0.05)
different within either the burned or unburned plots, and there was no significant difference (p = 0.70)
in magnitude and direction of year-to-year changes between burned and unburned areas [92].
The following Research Project Summary provides information on prescribed fire use and postfire response of many
plant species, including old man's whiskers:
Understory recovery after low- and high-intensity fires in northern Idaho
ponderosa pine forests
This description provides characteristics that may be relevant to fire ecology, and is not meant for identification. Keys for identification are available (e.g. [25,36,50,51,54,58,60,61,69,73,87,95, 123,126]).
Old man's whiskers is a perennial forb [25,36,50,51,54,58,61,62,70,73,82,87, 126] with erect or ascending stems [60,73,82] 5.9 to 20 inches (15-50 cm) tall [25,36,50,54,58,60,61,73,82,87], often forming clumps 8 to 16 inches (20-40 cm) or more wide [54,61].
Leaves are generally basal [58,60,61], 0.8 to 7.9 inches (2-20 cm) long [25,36,50,54,58,60,61,73,82,126], these being numerous [73] and unequally or interruptedly pinnate to pinnatifid or lyrate [50,54,73,87], with 7-19 leaflets [50,54,73,87], 0.5 to 2 inches (13-50 mm) long [36,50,54], progressively larger toward the apex [54]. There are also typically 1 to 4 small leaves about mid-length on the stem [54,61,73,82].
The stalked flowers [73] are perfect [36,61,62,87,126], 0.3 to 0.4 inch (7-10 mm) long [126] and number from 1 to 9 [36,54,61,73,82,126], but usually 3 [25,73,82]. Styles are long (1 to 2 inches (2.5-5 cm)) and plumose [51,54,123]. The terminal segment of the style is generally persistent in Geum triflorum var. triflorum, while in G.t. var. campanulatum and G.t. var. ciliatum it is generally deciduous [62]. The fruit is a more or less flat [60] achene [36,43,51,54,54,58,60,61,62,70,73,82, 82,87,123,126], 0.1 to 0.5 inches (2.5-12 mm) long [36,54,58,60,61,61].
Old man's whiskers has a thick scaly caudex [36,70]. It also produces short, thick rhizomes [43,50,54,60,73,126].
Old man's whiskers is native to North America and can be found from the Yukon and Northwest Territories, south through British Columbia, Washington and Oregon (mostly east of the Cascades), the Klamath, Cascade and Sierra Nevada Ranges of California, and into northern and central Arizona and northern New Mexico. Its range extends across Canada to Newfoundland. In the United States it is distributed eastward from the Great Basin and Rocky Mountains, across the northern Great Plains and into the prairie region of the Lake States and parts of the Northeast [2,3,6,7,16,21,25,34,36,39,43,48,50,51,54,60,61,62,66,69, 70,73,74,81,87,88,92,95,103,109,120,123,126].
Geum triflorum var. ciliatum occurs in and east of the Cascades from British Columbia to California, east to Alberta and the Rocky Mountain states, probably intergrading with G. t. var. triflorum and G. t. var. campanulatum [4,36,61,62,69,70,82]. According to Great Plains Flora Association [54], a few specimens from the western Great Plains "approach" G. t. var. ciliatum in similarity of diagnostic traits.
Geum triflorum var. campanulatum is apparently a montane ecotype of the Olympic Mountains in Washington, and of Saddle Mountain, Clatsop County, Oregon, transitional to G. t. var. ciliatum [61,62].
Geum triflorum var. canescens is found at higher elevations (8,000 to 9,000 feet (2,400-2,700 m)) in the Sierra Nevada Range and on Hinkey Summit, Santa Rosa Range, northwestern Nevada [69]. Plants Database [121] indicates that G. t. var. canescens also occurs in Washington, Oregon, Idaho, Montana, and Wyoming.
Geum triflorum var. triflorum occurs from Alberta to southeastern Canada and the northeastern U.S., and south in the Rockies, where it more or less straddles the continental divide from Montana to northern New Mexico, and includes some populations as far west as northeastern Arizona [36,50,61,62]. Most Great Plains specimens are G. t. var. triflorum [54].
Plants Database provides state distribution maps of old man's whiskers and its infrataxa.
Fire adaptations: Several studies have documented old man's whiskers population-level responses to fire (see Fire Effects), including postfire increases [6,7,26]. Yet, as of this writing (2006), there is no published information documenting specific plant-level mechanisms by which old man's whiskers responds to fire. Important areas for future inquiry might include, for example, the importance of vegetative (rhizome or caudex) postfire recovery compared with postfire seedling establishment, or the relative influence of fire severity or season of burning on these adaptations.
FIRE REGIMES: As of this writing (2006) there is very little published information that specifically addresses fire regime characteristics that might influence old man's whiskers productivity or distribution. Limited research has suggested that factors such as fire intensity [7] or season [6] may be important (see Fire Effects), but these few studies are too limited in number and scope to provide much clarity or certainty about their effect.
There is also no published information, as of this writing (2006), that indicates the importance of fire return interval for old man's whiskers productivity or distribution. However, it may be inferred from studies of FIRE REGIMES in plant communities and ecosystems where old man's whiskers can be found, that old man's whiskers seems to occur where fire return intervals are relatively short. See the following table, which provides fire return intervals for plant communities and ecosystems where old man's whiskers may occur.Find fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find FIRE REGIMES"
Community or ecosystem Dominant species Fire return interval range (years) grand fir Abies grandis 35-200 [10] bluestem prairie Andropogon gerardii var. gerardii-Schizachyrium scoparium <10 [77,99] Nebraska sandhills prairie Andropogon gerardii var. paucipilus-Schizachyrium scoparium 99] silver sagebrush steppe Artemisia cana 5-45 [59,101,128] sagebrush steppe Artemisia tridentata/Pseudoroegneria spicata 20-70 [99] basin big sagebrush Artemisia tridentata var. tridentata 12-43 [106] mountain big sagebrush Artemisia tridentata var. vaseyana 15-40 [11,32,93] Wyoming big sagebrush Artemisia tridentata var. wyomingensis 10-70 ( x=40) [122,129] desert grasslands Bouteloua eriopoda and/or Pleuraphis mutica 10 to <100 [90,99] plains grasslands Bouteloua spp. 99,128] blue grama-needle-and-thread grass-western wheatgrass Bouteloua gracilis-Hesperostipa comata-Pascopyrum smithii 99,105,128] blue grama-buffalo grass Bouteloua gracilis-Buchloe dactyloides 99,128] grama-galleta steppe Bouteloua gracilis-Pleuraphis jamesii 99] cheatgrass Bromus tectorum 100,127] curlleaf mountain-mahogany* Cercocarpus ledifolius 13-1,000 [13,107] western juniper Juniperus occidentalis 20-70 Rocky Mountain juniper Juniperus scopulorum <35 [99] wheatgrass plains grasslands Pascopyrum smithii <5-47+ [99,101,128] Engelmann spruce-subalpine fir Picea engelmannii-Abies lasiocarpa 35 to >200 blue spruce* Picea pungens 35-200 [10] pinyon-juniper Pinus-Juniperus spp. <35 [99] Rocky Mountain bristlecone pine P. aristata 9-55 [41,42] whitebark pine* Pinus albicaulis 50-200 [1,8] Rocky Mountain lodgepole pine* Pinus contorta var. latifolia 25-340 [17,18,119] Sierra lodgepole pine* Pinus contorta var. murrayana 35-200 [10] Colorado pinyon Pinus edulis 10-400+ [47,53,71,99] Pacific ponderosa pine* Pinus ponderosa var. ponderosa 1-47 [10] interior ponderosa pine* Pinus ponderosa var. scopulorum 2-30 [10,15,83] Arizona pine Pinus ponderosa var. arizonica 2-15 [15,35,112] galleta-threeawn shrubsteppe Pleuraphis jamesii-Aristida purpurea <35 to <100 eastern cottonwood Populus deltoides <35 to 200 [99] quaking aspen-paper birch Populus tremuloides-Betula papyrifera 35-200 [44,124] quaking aspen (west of the Great Plains) Populus tremuloides 7-120 [10,56,91] mountain grasslands Pseudoroegneria spicata 3-40 ( x=10) [9,10] Rocky Mountain Douglas-fir* Pseudotsuga menziesii var. glauca 25-100 [10,11,12] California mixed evergreen Pseudotsuga menziesii var. menziesii-Lithocarpus densiflorus-Arbutus menziesii <35 [10] bur oak Quercus macrocarpa <10 [124] oak savanna Quercus macrocarpa/Andropogon gerardii-Schizachyrium scoparium 2-14 [99,124] little bluestem-grama prairie Schizachyrium scoparium-Bouteloua spp. <35 [99] mountain hemlock* Tsuga mertensiana 35 to >200 [10] *fire return interval varies widely; trends in variation are noted in the species reviewBiogeographic: The published literature describes a variety of site types where old man's whiskers may be found. These include, but may not be limited to: montane, alpine or high valley grasslands, meadows, balds and tundra [2,36,51,69,82]; sometimes rocky and often open mountain slopes, hillsides, and foothills [69,70,82,87]; western grasslands and sagebrush (Artemisia spp.) plains [24,25,28,39,51,60,61,66,75,82,84,85, 109]; Missouri River bottomlands [3]; Great Plains and Midwestern prairie and grasslands [5,6,21,30,37,54,64,95,123]; old fields [16]; woodlands and open forests [7,51,54,60,69,70,82,85,123].
Elevation: Old man's whiskers is found at a range of elevations, from the subalpine and alpine zones of the western mountains [43,61,81,82,86], to Great Lakes prairies [95,123], and old fields in the New Jersey Piedmont [16]. In the mountainous West its elevation range might be quite broad. For instance, Lackschewitz [81] described old man's whiskers habitat in west-central Montana from valley bottoms to upper subalpine, and Hitchcock and Cronquist [61] indicated that it could be found from the lower foothills to subalpine ridges in the Pacific Northwest.
The following table lists published accounts of elevation ranges where old man's whiskers can be found in western North America. These examples should not necessarily be interpreted as elevational limits to old man's whiskers distribution, but simply as a demonstration of the kinds of elevations, particularly upper elevations, where old man's whiskers might occur.
Location Elevation Colorado 7,000 to 11,000 feet (2,100-3,400 m) [58] Intermountain West 5,200 to 11,000 feet (1,600-3,400 m) [36] east-central Arizona 10,400 to 10,900 feet (3170-3320 m) [2] California 4,300 to 10,500 feet (1,300-3,200 m) [60] Uinta Basin (Utah) 7,500 to 10,000 feet (2300-3050m) [51] Greater Yellowstone 6,800 to 10,000 feet (2,070-3050 m) [39] southern Montana above 9,800 feet (>3000 m) [20] Arizona (Geum triflorum var. ciliatum) 6,000 to 9,500 feet (1,800-2,900 m) [70] northern New Mexico 7,000 to 9,000 feet (2,100-2,700 m) [87] northwestern Wyoming 6,700 to 9,000 feet (2,040-2,700 m) [28] Nevada (G. t. var. canescens) 8,000 to 9,000 feet (2,400-2,700 m) northern Nevada (G. t. var. ciliatum) 5,000 to 8,000 feet (1,500-2,400 m) [69] British Columbia 3,480 to 7,434 feet (1,060-2,266 m) [73] southeastern Oregon (G. t. var. ciliatum) 6,000 to 7,000 feet (1,800-3,400 m) [4] northern Idaho to 2,900 feet (880 m) [7]Climate: While to date (2006) there is no comprehensive source for climate parameters describing the distribution of old man's whiskers, the following examples provide some indication of climate for habitats where old man's whiskers occurs. At a south-central Saskatchewan prairie site, mean monthly temperatures range from -0.4 °F (-18 °C) in January to 66 °F (19 °C) in July, and average annual precipitation is 14 inches (360 mm), 30% falling as snow [6]. A northern Idaho ponderosa pine (Pinus ponderosa) habitat receives an average of 22 inches (560 mm) of annual precipitation, mostly falling from October to March [7]. A study site in the upper Blackfoot Valley in western Montana receives an average of about 16 inches (400 mm) of precipitation annually, nearly a third occurring in May and June [24]. A southeastern Oregon montane habitat containing a population of G. t. var. ciliatum has an average frost-free period of 10-50 days and annual precipitation estimated at 12-18 inches (300-460 mm), about 40% occurring in the growing season [4].
Moisture regime: The site characteristic that is perhaps most frequently mentioned in relation to old man's whiskers occurrence is moisture regime. In the more arid regions of western North America, old man's whiskers habitat is usually characterized as moist or wet. This is particularly true where old man's whiskers is found in isolated moist areas surrounded by a wider landscape of dry habitat. Some examples include wet alpine meadows in Nevada [69], more mesic areas of sagebrush-dominated plains and lower foothills in the Pacific Northwest [61], moist streambanks and wet meadows in high valley and montane sites in the Intermountain West [36], and moist slopes in Colorado [58].
In grassland habitats of the northern Rockies, several studies have documented old man's whiskers's affinity for moister areas. A study of grassland phytogeography along the North Fork Flathead River in Glacier National Park, northwestern Montana, indicated that while old man's whiskers was present in most grassland habitats studied, it became increasingly more prevalent towards the mesic end of the moisture gradient in habitats dominated by rough fescue (Festuca altaica) [75]. In an Idaho fescue/bluebunch wheatgrass habitat type in western Montana, old man's whiskers is among several species that were more likely to be prominent in the moister Columbia needlegrass (Achnatherum nelsonii) phase, compared with other phases of the habitat type [96]. A study in southwestern Montana mountain grasslands, comprised of variations of an Idaho fescue-bluebunch wheatgrass habitat type, indicated greater old man's whiskers productivity where moisture availability was greatest. Mean old man's whiskers aboveground biomass productivity was 7 lb/ac (range 2-13) on a southwest exposure (typical Idaho fescue-bluebunch wheatgrass habitat type), compared with 98 lb/ac (range 45-139) on a northeast exposure (Columbia needlegrass phase of the same habitat type). It was suggested that the northeast exposure had greater moisture availability during the growing season [97]. However, old man's whiskers presence is probably reduced on sites where snowpack lasts into summer (Weaver 1970 as cited in [94]).
Conversely, where grassland habitats persist within a larger landscape of moister, often forested habitats, old man's whiskers sites are typically characterized as dry. These areas are most common along the prairie/forest interface in the midwestern and north-eastern U.S. and Canadian prairie provinces. Some examples include isolated dry grasslands in Wood Buffalo National Park, northern Alberta [109], dry prairie sites along the forest-prairie transition zone in north-central Minnesota [30], dry prairies in northern Illinois [95], xeric prairie habitats in Wisconsin [37], and "dry habitats" in Michigan [123]. A similar situation may be found in western North America where dry, grassy isolates occur within a larger, moister forested matrix. For instance, Franklin and Dyrness [48] noted the presence of old man's whiskers on steep, south-facing slopes in the Olympic Mountains, northwestern Washington.
In some areas, old man's whiskers may occur locally within a relatively wide range of moisture regimes. Lackschewitz [81] indicated that old man's whiskers occurs on sites that are mesic (adequate moisture during all or most of the growing season, but rarely if ever flooded) to meso-xeric (moisture abundant in the early growing season but dry later on) sites in west-central Montana. According to Klinkenberg [73] old man's whiskers can be found within dry to mesic grasslands, meadows, rocky slopes and open forests in the steppe, montane and subalpine zones of British Columbia. Lloyd and others [85] indicated that old man's whiskers is found from xeric to subhydric soil moisture regimes within the bunch grass, Rocky Mountain Douglas-fir (Pseudotsuga menziesii var. glauca), and montane spruce (Picea spp.) zones in the Kamloops Forest Region, southern British Columbia.
Soils: Although as of this writing (2006) there is no comprehensive information on old man's whiskers soil affinities, it appears from several sources that old man's whiskers occurs on a wide variety of soils. For example, it is found on heavy clay soils in northern Nevada [69], as well as moist, well-drained soils in the Uinta Basin of northeastern Utah [51].
Limited published information suggests that old man's whiskers is probably not an important forage species for domestic livestock. Kearney and others [70] indicated it may provide forage for domestic sheep. Cattle were not observed grazing old man's whiskers while on summer range on lower slopes of the Elkhorn Range, west-central Montana [52].
A smattering of published information provides little in the way of definitive conclusions about the response of old man's whiskers to domestic livestock grazing. Old man's whiskers was labeled an "increaser" in response to grazing on Montana rangelands by Mueggler and Stewart [96] and Montana State University Extension Service [80]. Daubenmire [38] considered prairie smoke a "decreaser" in response to grazing within the Idaho fescue-common snowberry (Symphoricarpos albus) habitat type in eastern Washington. In a rough fescue/bluebunch wheatgrass habitat type in western Montana, Mueggler and Stewart [96] indicated that prairie smoke increases with "light" grazing, but declines with "heavy use," particularly by domestic sheep.
While old man's whiskers is probably utilized by native ungulates, at least seasonally, it appears not to be a major forage source. Gordon [52] reported elk utilization of small amounts of old man's whiskers as winter forage (1% of diet during January-March) on lower slopes of the Elkhorn Range, west-central Montana. Old man's whiskers grazing by elk was not observed on these sites during spring (April-May). Kamps [65] recorded evidence of mule deer feeding on old man's whiskers in central Montana, mainly in spring, and Singer [114] documented spring utilization by white-tailed deer in the North Fork area of Glacier National Park, Montana. Hansen and others ([57] and references therein) ranked old man's whiskers "food value" as "poor" for elk, mule deer, pronghorn, upland game birds, waterfowl, small nongame birds, and small mammals, and "fair" for white-tailed deer in southwestern Montana.
Palatability/nutritional value: Old man's whiskers does not seem particularly palatable or nutritious for livestock. Hansen and others ([57] and references therein) ranked old man's whiskers "energy value" and "protein value" as "poor." Palatability was considered "poor" for cattle and horses and "fair" for domestic sheep ([57] and references therein). Mueggler and Stewart [96] characterized old man's whiskers as having "very poor" palatability for cattle, domestic sheep, deer, and elk. Gross energy value of old man's whiskers seeds, in average gram-calories per gram, were 4,998 for whole and 5,693 for hulled seeds [72].
Cover value: No information is available on this topic.
Vegetation classifications describing plant communities where old man's whiskers is a dominant species
are:
AB:
prairie Junegrass (Koeleria macrantha)-old man's whiskers [118]
WY:
mountain big sagebrush (Artemisia tridentata ssp. vaseyana)/needle-and-thread grass
(Hesperostipa comata)-old man's whiskers
mountain big sagebrush/Idaho fescue (Festuca idahoensis)-old man's whiskers
mountain big sagebrush/bluebunch wheatgrass (Pseudoroegneria spicata)-prairie
smoke [28]
CA:
stemless goldenweed (Stenotus acaulis)-G. t. var. canescens [86]
Old man's whiskers phenology may vary according to site conditions. A study of variation in plant seasonal development on a southwestern Montana mountain grassland site showed that old man's whiskers phenological development initiated earlier and proceeded more rapidly, on average, on a southwest exposure compared with a northeast exposure [97]. Hitchcock and Cronquist [61] indicated that flowering occurred from April to early August in the Pacific Northwest, depending on elevation.
Some examples of flowering phenology from across old man's whiskers's North American range are:
Location Approximate flowering times south-central Wisconsin [5] mid-May-early June Uintas (northeastern Utah) [51] May-June Pacific Northwest [61] April-early August Intermountain West [36] June-August Great Plains [54] April-June northern New Mexico [87] May-June Arizona [70] May-August northern Illinois [95] May-June western New York to Minnesota and Iowa [50] May-June (Geum triflorum var. triflorum) Beartooth Plateau, southern Montana [20] late June-July southern Wisconsin, northern Illinois [40] May-late JuneCurtis [37] indicated that old man's whiskers was among the earliest plants to flower on xeric prairie sites in Wisconsin.
The following old man's whiskers flowering data were collected over 6 seasons near Woodworth, central North Dakota [33].
Earliest 1st bloom Latest 1st bloom Median date of full flowering Median date when flowering 95% complete Mean # days in flower May 10 1983 June 6, 1979 May 22 June 19 37The following old man's whiskers flowering data were collected over 13 seasons at Swift Current, southwestern Saskatchewan [29].
Mean flowering date Earliest flowering date Latest flowering date Latest date in flower Mean # days in flower May 13 April 24, 1946 May 25, 1954 August 1 62Old man's whiskers regenerates from seeds, rhizomes, and probably by sprouting from the caudex. However, as of this writing (2006) there is very little published information describing old man's whiskers regeneration processes.
Pollination: No information is available on this topic.
Breeding system: Given that the flowers are perfect [36,61,62,87,126], old man's whiskers is also monoecious.
Seed production: Stevens [116] collected 142 seeds from a single "mature" old man's whiskers plant "of average size," "growing where competition was low."
Seed dispersal: Fruits are wind-dispersed [123]. Zimmerman [130] noted that "seed heads do not all ripen together...., yet suddenly and individually fall off, especially during rainstorms."
Seed banking: No information is available on this topic.
Germination: Germination is probably variable, depending on conditions, but not dependent upon stratification. Zimmerman [130] suggested that old man's whiskers seeds do not require stratification, and can germinate soon after dispersal. Baskin and Baskin [19] also provided qualified support for the idea that seeds are nondormant at maturity. Greene and Curtis [55] conducted germination tests on a variety of Wisconsin prairie species. Stratification for 3 months appeared to have little effect on prairie smoke germination, with 64% of stratified seeds germinating compared with 50% for unstratified.
Old man's whiskers seed viability may be variable from year to year. Unstratified seed collected in southern and western Wisconsin in 1937, 1940, and 1941, yielded germination rates of 80%, 14%, and 50%, respectively [55]. Germination testing on old man's whiskers seed collected from an eastern South Dakota prairie resulted in a 90% germination rate, requiring 7-16 days [115]. Zimmerman [130] suggested that old man's whiskers seeds do require prolonged wetting prior to germination [130].
Seedling establishment/growth: As of this writing (2006) there is very little published information describing old man's whiskers seedling establishment and growth. A study of the effects of pocket gopher disturbance on shortgrass prairie forbs suggests that old man's whiskers seedling establishment and growth may be enhanced by small-scale disturbance [88].
Asexual regeneration: Several sources indicate that old man's whiskers has both rhizomes [43,50,54,60,73,126] and a caudex [36,70], suggesting that asexual regeneration can occur. However, there is currently (2006) very little published information describing the importance of asexual regeneration for its survival and spread. Curtis [37] wrote that old man's whiskers was among several species "with a strongly developed rhizome method of vegetative propagation."
Although research describing the relative benefit to old man's whiskers provided by fire-caused disturbance appears ambiguous (see Fire Effects), several studies suggest old man's whiskers presence may be enhanced by other types of disturbance. Bramble-Brodahl [28] classified the sagebrush-grass vegetation of the Gros Ventre District of the Bridger-Teton National Forest. In the mountain big sagebrush/Idaho fescue-old man's whiskers habitat type, old man's whiskers was among species said to increase with disturbance. In the mountain big sagebrush/needle-and-thread grass-old man's whiskers habitat type, old man's whiskers was among species said to increase with disturbance on wetter sites [28]. Redmann and Schwartz [103] studied the dry grassland vegetation of Wood Buffalo National Park, Alberta. Sampling at 4 paired sites indicated that old man's whiskers had greater presence on sites disturbed by "compaction and erosion", compared with undisturbed sites [103]. A study of the effects of pocket gopher disturbance on shortgrass prairie forbs indicated that old man's whiskers benefits from small-scale disturbance. Comparisons of undisturbed habitat and habitat disturbed by pocket gopher burrowing indicated that old man's whiskers seedling establishment and growth were greater on pocket gopher mounds [88].
Since old man's whiskers seems to benefit from disturbance, it is most likely to be found in association with early-successional rather than late-successional communities. Bunting and others [31] provided some support for this idea in a study of the western juniper (Juniperus occidentalis)/mountain big sagebrush mosaic of the South Fork Owyhee River, southwestern Idaho. Old man's whiskers was among several forb species associated primarily with early-successional grassland and sagebrush steppe within this mosaic. These species were not typically found in late-successional juniper (Juniperus spp.) woodland communities [31].
The currently accepted scientific name of old man's whiskers is Geum triflorum Pursh.(Rosaceae)
[25,36,43,50,51,54,60,61,62,63,68,69,70,73,81,82,87,95,126]. The following varieties are recognized:
G. triflorum var. campanulatum (Greene) C.L. Hitchc. [61,62,68]
G. triflorum var. canescens (Greene) Kartesz & Gandhi [67,68,69]
G. triflorum var. ciliatum (Pursh) Fassett [48,54,61,62,68,69,70,82,87,126]
G. triflorum var. triflorum [50,54,61,62,68,73]
For the purposes of this review, the common name "old man's whiskers" is used when discussing
characteristics common to (or assumed to be common to) the species in general. When referring to
infrataxa, the scientific names for the varieties listed above are used.
Geum triflorum, commonly known as prairie smoke,[3] old man's whiskers,[4] torchflower,[4] three-sisters,[5] long-plumed purple avens,[4] lion's beard,[4] or three-flowered avens,[4] is a spring-blooming perennial herbaceous plant[4] of the Rosaceae family. It is a hemiboreal continental climate species that is widespread in colder and drier environments of western North America, although it does occur in isolated populations as far east as New York and Ontario.[6] It is particularly known for the long feathery plumes on the seed heads that have inspired many of the regional common names and aid in wind dispersal of its seeds.
Geum triflorum is a perennial herb with short, spreading rhizomes, which form colonies of stemless rosettes.[3] The roots are fiberous[7] and have a sassafras-like flavor.[8] The leaves grow from a caudex and are 4–30 cm long. They are divided into leaflets with deep divisions that makes the leaves resemble the leaves of a fern. The leaflets are arranged pinnately along a common leaf stem with smaller leaflets mixed in with 7–18 larger ones and single larger leaflet at the end of the leaf. The leaves are covered with extremely small downy hairs.[9][8]
Early in the spring, the leaves often lie flat to the ground and are in poor condition, but they soon become more upright in response to the warmer days and lack of snow cover. In the heat of a dry summer, the leaves also will lie down closer to the earth. The plants resume growth in the fall as other plants are starting to go dormant, developing a mound of deep grey-green leaves.[6] The leaves are evergreen in areas without severe cold or there is protective snow cover, though they often turn purple, orange, or reddish.[10][11]
The flowers of G. triflorum appear from mid-spring to early summer. The flowering stalks stand well above the leaves on red-purple-maroon stems 10–45 cm in height. The flowering stem is almost bare with a few very small leaves called bractlets on the main stem and where the arching flower stalks (pedicels) attach to the main stem.[8] Each flower hangs upside down by itself from a separate pedicle. There are usually three flowers on each flower stalk, but sometimes one, five, or even seven per stalk. The sepals are strongly closed and pink to maroon in color, covered in fine downy hairs, with five narrow pointed bracts radiating outward toward the base of the flower. The flowers contain five 7–13 mm long elliptical petals mostly to entirely hidden under the sepals. They are most often a light yellow to cream, but sometimes have a blush of pink or purple; they have purple veins.[9][12]
When pollination is completed, the flower heads turn upright and the sepals begin to open. The petals may be visible at this stage. The many styles grow longer, eventually becoming 15–70 mm in length. The styles are densely covered in fine hairs making them resemble downy bird feathers or wisps of mauve smoke.[13] The seed heads start out pale pink and fade to tan or grey as the seeds mature in mid-summer.[6] The seeds do not appear to need cold stratification, as germination did not change significantly when tested.[3]
Geum triflorum was named and described by German–American botanist Frederick Traugott Pursh in his book Flora americae septentrionalis using an 1811 collection by naturalist John Bradbury. He placed the species within Carl Linnaeus's Geum, a genus with a name derived from Greek for "taste", with the species name of Geum triflorum for the three flowers usually present on each flower stalk. The species previously had been collected in Idaho on 12 June 1806 by American explorer Meriwether Lewis but was not described by him. That specimen was incorrectly described as a new species named Geum ciliatum by Pursh.[14] German botanist Kurt Sprengel placed it in Sieversia as S. triflora in his update of Systema Vegetabilium published in 1825.[15] This classification was eventually rejected as was the 1906 attempt by American botanist Edward Lee Greene to create a new genus that would reclassify G. triflorum as Erythrocoma triflora and separate out a dozen regional varieties as separate species.[16][17][5]
G. triflorum has three varieties that are accepted by many, but not all, authorities, as of January 2023.[9]
Geum triflorum var. campanulatum was described as a separate species, Erythrocoma campanulata, by Greene[18] and as Geum campanulatum by English-born botanist George Neville Jones, but American botanist Charles Leo Hitchcock argued for its classification as a variety of G. triflorum in 1961, writing, "There has been much diversity of opinion regarding both the generic status of, and significance of the variation in, this complex. In general the several taxa that have been recognized at the specific level are largely sympatric and completely transitional and there seems to be no good reason to recognize more than 3 races for our area...".[19][20] This has become the accepted view as researched by botanist Richard Pankhurst. It differs in having leaflets that are rounder (obovate-cuneiform instead of cuneiform), shorter leaves overall, and flowers that are more open or bell-shaped (campanulate). It was described from a type specimen found in the Olympic Mountains.[18] It is recorded by the USDA Natural Resources Conservation Service PLANTS database (PLANTS) as growing in Washington state and Oregon.[21] This variety is accepted by World Flora Online (WFO) and PLANTS, but not Plants of the World Online (POWO) or Flora of North America (FNA).[1][5]
Geum triflorum var. canescens was similarly described by Greene as Erythrocoma canescens in his 1906 book. Arguing against Erythrocoma, the Swedish-American botanist Per Axel Rydberg classified it as Sieversia canescens in 1913.[22] In 1958, American botanist and plant taxonomist Philip A. Munz accepted it as a species but moved it back to Geum as Geum canescens.[23] In 1990, John T. Kartesz and Kanchi Gandhi published an article on the nomenclature of North American plants giving it its current status as a variety of G. triflorum.[24][25] The type specimen was collected in the northern Sierra Nevada Mountains in 1863 near Ebbett's Pass. It is distinguished by being stouter and sometimes taller than Geum triflorum var. campanulatum and covered in grayish-white hairs on the leaves "canescently soft-villous and sparsely pilose", but with similarly shorter leaves than the species.[18] It is recorded by PLANTS as growing in the Pacific Northwest of the United States including California, Idaho, Montana, Nevada, Oregon, Washington state, and Wyoming with county level distribution records in northern California and Nevada. This variety is accepted by WFO and PLANTS, but not POWO or FNA.[1][5]
Geum triflorum var. ciliatum was first described as species Geum ciliatum by Pursh at the same time as he described G. triflorum.[26] Scottish botanist and plant collector George Don reclassified it as Sieversia ciliata in 1832 in the book A General History of the Dichlamydeous Plants.[27] It was similarly placed by Greene into his proposed Erythrocoma, but he separated it into two species, Erythrocoma ciliata[28] and Erythrocoma grisea.[29] In 1913, Rydberg described Sieversia grisea, but this is now accepted as a synonym for the variety.[30] While there were other published classifications, the one currently accepted was written by Norman Carter Fassett and published in 1928 in the journal Rhodora.[31] The variety is distinguished by having larger leaflets that are cleft (having a deep division) for more than half their lengths and the seed heads having shorter styles, 15–40 mm instead of 15–70 mm.[31] The PLANTS database records it as growing from British Columbia to northern California and from the mountains of New Mexico to Saskatchewan. It is recorded by them as growing in both Colorado and Montana without county level distributions.[32] This variety is accepted by WFO, PLANTS, POWO, and FNA.[1][5]
Geum triflorum f. ornatum was published by Fassett at the same time as G. triflorum var. ciliatum in Rhodora. It is not accepted as a form of G. triflorum by most authorities,[33] including WFO, PLANTS, POWO, and FNA.[1][5]
G. triflorum is widely distributed in North America, but only common in the Upper Midwest and mountainous west. In Canada, it is common from British Columbia to Manitoba. It also grows in parts of the Yukon Territory and Northwest Territory. It is a common prairie species in Minnesota, Wisconsin, and North Dakota. Contiguous with this it is reported in nine of the north-eastern counties in Illinois and eight counties in north-eastern South Dakota. It is recorded in PLANTS as growing in Iowa without a specific location. It is common throughout the prairies of Montana and Wyoming and into the mountains of Idaho, Colorado, Utah, and Nevada. It is present in the adjoining Black Hills in South Dakota, the eastern portions of Washington state, Oregon, and the north east mountainous portion of California. Populations are also found in the mountains of northern New Mexico and Arizona.
In the eastern United States and Ontario populations are much rarer and isolated. It grows in six counties of the lower peninsula of Michigan and just Chippewa County in the upper peninsula. In New York state it is only recorded in PLANTS as growing in Jefferson County and Oswego County, which are adjacent to Lake Ontario.[34]
The New York Flora Atlas only records vouchered specimens from Jefferson County, NY.[35]
G. triflorum grows in open, non-forested habitat with well drained soils. This includes meadows in montane forests, prairies of the upper Midwest, and alvars in Michigan, New York, Ontario, and Manitoba.[36][37] Plants are tolerant of clay soils when slopes provide drainage, but intolerant of being waterlogged, particularly in winter.[4] They require good moisture for new plants to establish and prefer areas that are well supplied with water in the spring, but tolerant of drought in summer and healthier with drier soils in summer.[11] In habitats with more precipitation they will often be found in areas with more freely draining soils and/or on areas raised up above the surrounding landscape. In drier habitats plants will often be found in lower areas that have additional moisture from spring runoff. Prairie smoke plants are tolerant of some shade and prefer some afternoon shade in climates with hot summers.[4]
In the central Rocky Mountains G. triflorum will grow to around timberline at 3,800 meters and as low as 2,000 meters in the foothills.[38] In New York it grows at an altitude of 90 meters. In the midwest it grows near to the level of the great lakes at 90 meters in elevation in Chaumont Barrens Preserve.[37]
The flowers produce both nectar and pollen. They are frequently visited by bumblebees, which are able to force their way into the mostly closed flowers and reach the nectar.[39] They also buzz-pollinate to dislodge pollen from the stamens and gather it to feed their young. As it blooms early in the season and in large amounts in suitable habitats it is suggested that it is one of the plants that is critical to the success of queen bumblebees in establishing their first brood.[40] Smaller bees such as sweat bees from the genus Lasioglossum feed on pollen grains from the opening of the flower.[41] Bumblebees are the only effective cross-pollinators.[10] Various insects become nectar robbers by chewing holes in the top of the flower, near the stem, to reach the nectar.[41]
It is not a major source of forage for most native herbivores including mule deer, elk, pronghorn, and game birds, but is foraged by white tailed deer. It is similarly not generally eaten by cattle, horses, or domestic sheep and has a low amount of protein.[3]
Prairie smoke is planted in gardens as an ornamental for its interesting seed heads and for the persistent foliage in the winter.[11] It is popular for native gardens, rock gardens, and with gardeners that wish to have an informal natural look. Plants are winter hardy in USDA zones 3–7 and do not have any serious disease or pest problems.[4] It is planted in either the spring or fall from cultivated plants or by seed. The size is dependent on the richness of the soil. Very lean soils without organic matter will grow compact plants, and in rich soils, the plants will become large and competitive.[6] It is not tolerant of deep shade and can be crowded out by taller species which grow over it and deprive it of light.[11]
Some Indigenous peoples of the Northwest Plateau have used three-flowered avens to treat tuberculosis.[42]
As recorded by Walter McClintock and later John C. Hellson, people of the Blackfeet Nation made an infusion of the roots to treat sore throats, canker sores in the mouth, and application to wounds[43] and the infusion was also mixed with grease to create a salve used for the treatment of rashes, blisters, sores, and wounds.[44] Infusions of the whole plant were used to treat severe coughs.[45] The roots of G. Triflorum were scraped and mixed with tobacco and then smoked by the Blackfeet to "clear the head".[46] They also drank a tea made of the whole plant to promote general health.[47]
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USFWS Mountain-Prairie
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Flower detail
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Prairie smoke (Geum triflorum) on Olympic National Forest Mount Townsend Trail
Geum triflorum Saskatchewan, flower side view
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Geum triflorum in meadow habitat Yellowstone National Park. 11 June 2017
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Detail of the styles on a seed head
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Immature seed head opened to show the achenes sitting in the persistent hypanthium and bearing plumose tails (modified styles).
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Meadow habitat mid-summer development of seed heads Olympic National Park.
Geum triflorum, commonly known as prairie smoke, old man's whiskers, torchflower, three-sisters, long-plumed purple avens, lion's beard, or three-flowered avens, is a spring-blooming perennial herbaceous plant of the Rosaceae family. It is a hemiboreal continental climate species that is widespread in colder and drier environments of western North America, although it does occur in isolated populations as far east as New York and Ontario. It is particularly known for the long feathery plumes on the seed heads that have inspired many of the regional common names and aid in wind dispersal of its seeds.
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