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Potentilla hippiana



INTRODUCTORY


Photo by Jennifer Stewart, www.plantsofmagnolia.net
AUTHORSHIP AND CITATION:
Meyer, Rachelle. 2009. Potentilla hippiana. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.usda.gov/database/feis/plants/forb/pothip/all.html [].

FEIS ABBREVIATION:
POTHIP

NRCS PLANT CODE [69]:
POHI

COMMON NAMES:
woolly cinquefoil
wooly cinquefoil
horse cinquefoil
Hipp's cinquefoil
Hipp cinquefoil
cinquefoil

TAXONOMY:
The scientific name of woolly cinquefoil is Potentilla hippiana Lehm. (Rosacea) [12,17,18,21,23,44,75].

Two varieties of woolly cinquefoil are recognized, P. hippiana var. argyrea (Rydb.) Boivin and P. hippiana var. hippiana [27]. The scientific name is used in this review when information applies to a specific variety.

Woolly cinquefoil hybridizes with the following Potentilla species: beautiful cinquefoil (P. pulcherrima) [12,74,75], graceful cinquefoil (P. gracilis var. brunnescens) [75], varileaf cinquefoil (P. diversifolia), elegant cinquefoil (P. concinna), and possibly bearded cinquefoil (P. crinita) [12].

SYNONYMS:
For Potentilla hippiana var. argyrea:
Potentilla argyrea Rydb [21,27]

For Potentilla hippiana var. hippiana:
Potentilla hippiana var. diffusa Gray ex Lehm
Potentilla leucophylla Torr. non Pallas [27]
Potentilla leneophylla Torr. & Jones ex Eat. [27,28]
Potentilla propinqua (Rydb.) Rydb. [21,27]

Although some systematists consider branched cinquefoil (P. effusa) a subspecies of woolly cinquefoil [12,21,23], this review considers it a separate species (as in [27,60,74]). Therefore, its biology and ecology are not addressed in this review.

LIFE FORM:
Forb

FEDERAL LEGAL STATUS:
None

OTHER STATUS:
Information on state- and province-level protection status of plants in the United States and Canada is available at NatureServe.

DISTRIBUTION AND OCCURRENCE

SPECIES: Potentilla hippiana
GENERAL DISTRIBUTION:
Woolly cinquefoil occurs throughout much of Canada and the western United States. It is native in most of its range, but considered nonnative in Nova Scotia [57,63], Quebec, Ontario, [63], and Michigan [72]. The distributions of P. hippiana var. hippiana and P. hippiana var. argyrea overlap. Potentilla hippiana var. hippiana occurs in Alaska [26,27] and British Columbia east to Nova Scotia [12,21,27,63] and south through Idaho, Montana, and the Dakotas to New Mexico, Arizona [12,21,27,44,75], and southern Nevada [12,28]. It also occurs at low levels in Nebraska [27,35,63] and possibly Minnesota [27,60]. Potentilla hippiana var. argyrea occurs only in North Dakota, Alberta, Saskatchewan, and Manitoba [27,63]. Plants Database provides a distributional map of woolly cinquefoil and its varieties.

HABITAT TYPES AND PLANT COMMUNITIES:
Woolly cinquefoil occurs in foothill plain, montane and subalpine meadow, sagebrush (Artemisia spp.), forest, and alpine communities. Outside its native range, it has been found in "pastures" in Ontario, in "fields" Nova Scotia [63], and on a mine site in Michigan [72].

Woolly cinquefoil occurs in several grassland communities including Great Plains prairies [7,17,18,21,24,46,55,60,71] and subalpine meadows [11,16,40,50]. It has been reported it Great Plains grasslands of Montana [17,21], North Dakota [21], South Dakota [21,24], Wyoming [7,18,21], Colorado [21,46,55,60,71], and New Mexico [21]. Woolly cinquefoil commonly occurs in montane [2,16,55,73] and subalpine meadows [11,16,40,50] of Colorado [50,55,73], Utah [75], Arizona [2,29,40], and New Mexico [11,16,44]. It occupies both dry grasslands [16,61] and wet meadows [2,11,16,20,53]. Prairie Junegrass (Koeleria macrantha) [7,16,24,73], blue grama (Bouteloua gracilis) [7,14,24,71], fescues (Festuca spp.)—especially Arizona fescue (F. arizonica) [2,9,14,16,53], Muhlenbergia species—especially mountain muhly (M. montana) [7,14,16,73], oatgrasses (Danthonia spp.) [11,16,73], and/or bluegrasses (Poa spp.) [11,16,20,45,53,73] often dominate or codominate grasslands with woolly cinquefoil.

Woolly cinquefoil may occupy a few shrubland cover types. It occurs in sagebrush communities in the Intermountain region [12], west-central Idaho [1], and Colorado. Woolly cinquefoil was a consistent component of the Wyoming big sagebrush/bluebunch wheatgrass (Artemisia tridentata subsp. wyomingensis/Pseudoroegneria spicata subsp. spicata) community on the Arapaho-Roosevelt National Forest in north-central Colorado [73]. Sagebrush meadow was included in a list of intermountain habitat types with woolly cinquefoil [12]. Woolly cinquefoil also occurs in juniper (Juniperus spp.) scablands in west-central Idaho [1]. It occurred on sites in 5 age classes of riparian habitat in Colorado. Ragweed sagebrush (A. franseriodes), common juniper (J. communis), and Fendler's meadow-rue (Thalictrum fendleri) were the only species that occurred in 2 or more of these 5 age classes at coverages of 5.0% or greater. Species that occurred in all 5 age classes, with coverage of at least 4% in 1 of the age classes, include common juniper, Woods' rose (Rosa woodsii), Saskatoon serviceberry (Amelanchier alnifolia), fringed brome (Bromus ciliatus), and small-leaf pussytoes (Antennaria parvifolia) [5].

Woolly cinquefoil occurs from lower montane to subalpine woodlands and forests. These communities may be dominated by ponderosa pine (Pinus ponderosa), Douglas-fir (Pseudotsuga menziesii), quaking aspen (Populus tremuloides), spruce (Picea spp.), fir (Abies spp.), and/or Rocky Mountain bristlecone pine (Pinus aristata). Woolly cinquefoil was included in a list of species occurring in pinyon (Pinus spp.)-juniper woodlands on the Gila National Forest in western New Mexico [25]. It is widespread in open ponderosa pine forests from west-central Idaho [1] southeast through Colorado [31,42,58] and south to New Mexico [11,38,41] and southern Nevada [6,28]. In upper montane forests of Colorado, woolly cinquefoil occurs in ponderosa pine-Douglas-fir communities [32,42]. On the east slope of the Front Range in Colorado, it was documented in an upper montane forest comprised primarily of quaking aspen, Douglas-fir, and ponderosa pine [42]. Woolly cinquefoil was present in mixed-conifer communities with ponderosa pine, Douglas-fir, white fir (A. concolor), and blue spruce (Picea pungens) in southwestern Colorado [58]. It occurred in a mixed-conifer forest in Arizona comprised of those species, Engelmann spruce (Picea engelmannii), subalpine fir (A. lasiocarpa), southwestern white pine (Pinus strobiformis), and quaking aspen [52]. It has been reported in quaking aspen stands in Colorado [31] and quaking aspen-meadow mosaics in Arizona [9] and New Mexico [11]. Woolly cinquefoil occurred in a Douglas-fir-limber pine-Rocky Mountain bristlecone pine woodland in New Mexico [51] and Rocky Mountain bristlecone pine/fescue communities in Colorado [31].

Woolly cinquefoil has been reported in alpine communities in the Intermountain region [12], Utah [64,75], Wyoming [64], and Colorado [54]. In Colorado it occurred in an alpine community with American bistort (Polygonum bistortoides), alpine clover (Trifolium dasyphyllum), Parry's lousewort (Pedicularis parryi), and sticky polemonium (Polemonium viscosum) [54].

Several species cooccur with woolly cinquefoil across several habitats and stand structures. Prairie Junegrass has been reported with woolly cinquefoil in grasslands [7,16,24,73] shrublands [73], and coniferous forests [42,58]. Sedges (Carex spp.) [2,7,16,20,53] such as sun sedge (C. inops subsp. heliophila) [14,32,42,73] and elk sedge (C. geyeri) [31,58] cooccur with woolly cinquefoil in grasslands, shrublands, and forests. Western yarrow (Achillea millefolium) [11,41,42,52,53,58], silvery lupine (Lupinus argenteus) [9,42,52], common dandelion (Taraxacum officinale) [20,42,53], and fleabanes (Erigeron spp.) [14,16,41,42,51,53] such as beautiful fleabane (E. formosissimus) [11,51,58] have been found in grasslands and forests with woolly cinquefoil.

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Potentilla hippiana
Photo by Al Schneider, www.swcoloradowildflowers.com

GENERAL BOTANICAL CHARACTERISTICS:
This description provides characteristics that may be relevant to fire ecology and is not meant for identification. Keys for identification are available (e.g., [12,17,18,21,23,44,75]).

Woolly cinquefoil is a perennial forb from 4 to 20 inches (10-50 cm) tall [1,12,21,35,44,75]. It has a stout, branched caudex [12,21,64,75] over a thick taproot [12]. Its basal leaves range from 1 to 7.5 inches long (2.5-19 cm) or more [75]. They are odd-pinnately divided, typically with 7 to 13 toothed leaflets [1,12,21,23,26,35,44,75] from 0.8 to 2.0 inches (2-5 cm) long [1,75]. The perfect flowers [1,12,75] occur in a branching cyme [1,26,64] and range from 5 to 9.5 mm wide [1,75]. Leaf pubescence is variable, ranging from almost hairless [44], to hirsute [1], sericeous [29,44], or tomentose [1,44,75]. In Potentilla hippiana var. hippiana, the upper surface of the leaf may be less hairy than the underside [29,44]. Woolly cinquefoil produces numerous achenes [12,75] from 1.4 to 1.9 mm long [21,75].

RAUNKIAER [56] LIFE FORM:
Chamaephyte
Hemicryptophyte

SEASONAL DEVELOPMENT:
Woolly cinquefoil typically flowers from June to August [6,12,21,29,44], although it has been reported flowering as late as September in Arizona [29]. In an artificial heat experiment in Gunnison County, Colorado, woolly cinquefoil flowering was significantly (P<0.001) associated with snowmelt date, with earlier melting leading to earlier flowering. The period from budding to fruit dehiscence was significantly (P=0.024) longer in warmed plots than in controls [19]. In the same area, the date of first woolly cinquefoil flowering was not significantly related to peak flowering date or maximum number of plants flowering [50]. For effects of warming on woolly cinquefoil growth and abundance, see Growth [15]. For effects of warming on woolly cinquefoil pathogens and herbivores, see Palatability and/or nutritional value [59].

REGENERATION PROCESSES:
As of 2009, only limited information on woolly cinquefoil pollinators, seed production, seedling establishment, and vegetative regeneration was available. In an alpine community of Colorado, most woolly cinquefoil flowers were pollinated by flies, often syrphids, and less than 10% were pollinated by bees [54].

Woolly cinquefoil has potential for high seed production. Floras have noted that woolly cinquefoil produces several [75] to numerous achenes [12,75]. A review of rangeland species states that cinquefoils (Potentilla spp.) generally have numerous flowers that produce "innumerable" seeds that apparently have "good viability" [68].

 

Western New Mexico University Department of Natural Sciences,
Dale A. Zimmerman Herbarium

Vegetative spread of woolly cinquefoil was reported in an early-seral grassland on compacted granite-gravel soil in the Front Range [71]. The mechanism of vegetative reproduction was not discussed.

As of 2009, no information was available on woolly cinquefoil germination rates, seed dispersal or seed banking.

Growth: Woolly cinquefoil may grow faster on certain disturbed sites and in warm areas or years. Woolly cinquefoil leaf number (P=0.004) and leaf length (P=0.04) were significantly greater on soil mounds created by Botta's pocket gophers than on undisturbed sites [45]. In the Gunnison area of Colorado, woolly cinquefoil biomass was significantly (P<0.10) greater on an artificially warmed plot than on a control plot in 1997. Woolly cinquefoil was marginally more abundant (P=0.13) and larger (P=0.18) on the warmed plot in 1997 [15].

SITE CHARACTERISTICS:
Sites with woolly cinquefoil are often open [32,37,48,53] (see Successional Status) and have coarse-textured soil. Woolly cinquefoil has a wide elevation range. Topographic characteristics of sites with woolly cinquefoil are variable, although it is often reported on south-facing slopes. Due to their exposure and soil characteristics, many sites with woolly cinquefoil are dry. However, woolly cinquefoil may occur in wet meadows and riparian areas.

Soil: Woolly cinquefoil typically occurs in well-drained loam or sand with high content of coarse fragments and variable depth. Limited information suggests that woolly cinquefoil occurs on moderately acidic soils. Soil pH of north-central Colorado sites with woolly cinquefoil ranged from 6.4 in a Parry's oatgrass/sun sedge community to 7.0 in a Wyoming big sagebrush/bluebunch wheatgrass community [76]. Parent materials on sites with woolly cinquefoil are highly variable and may be derived from sedimentary [37,76], metamorphic [76], or igneous rock [74,76]. In Colorado, woolly cinquefoil has been reported on parent materials with glacial [44], alluvial, and colluvial origins [76].  
Photo by Doug Waylett

Texture: Woolly cinquefoil typically occurs in coarse-textured soils [35] and occasionally occurs in clay [11,42]. It occupies sand and loamy sand in western New Mexico [41] and Colorado [42,73]. Woolly cinquefoil has been documented in sandy loam on sites in northern Alberta [62], Wyoming [7], Colorado [31,42], and New Mexico [11,41]. In north-central Colorado, woolly cinquefoil occurred on sites with sand, "coarse-loam", and loam [73]. Woolly cinquefoil occupied sites with loam in woodlands on the Gunnison National Forest [31]. It was present on sites in New Mexico with sandy clay loam [11]. In a subalpine clearing with woolly cinquefoil on the east slope of the Front Range, soil texture was sandy clay loam at depths from 6 to 12 inches (15-30 cm) [42]. On a site in northern Alberta, woolly cinquefoil occurred in a dry grassland with loam at the surface, clay loam at shallow depths, and clay from 11 to 25.5 inches (28-65 cm) deep [62]. Woolly cinquefoil occurred in wet meadows of New Mexico that occurred on clay [11].

Descriptions of soil on sites with woolly cinquefoil suggest that coarse-fragment content is often high. From Arizona and New Mexico [38] to Montana [35], sites with woolly cinquefoil have been described as "stony" [35,38], "rocky" [31], "granite-gravel" [35], and "gravelly and cobbly" [73]. Percentage of coarse fragments had only been reported on sites in north-central Colorado as of 2009. A Parry's oatgrass/sun sedge community with 3% coverage of woolly cinquefoil occurred on a site with less than 20% coarse fragment content. A mountain big sagebrush/bluebunch wheatgrass community with trace coverage of woolly cinquefoil occurred on a site with 35% to 55% coarse fragment content [73]. The high coarse fragment content and coarse texture of the soil likely contributes to the "well-drained" [31] and "excessively drained" [38] nature of some sites with woolly cinquefoil. A grassland site on the Front Range with several Potentilla species probably experienced rapid drainage. There was little organic matter on this site [35] (see Soil moisture for more information). Although these rocky sites suggest little organic matter and poor soil nutrient availability, along the Animas River in Colorado woolly cinquefoil occurred in mature riparian vegetation that was associated with fine organic sediments and higher nitrate levels than those found in younger communities, where woolly cinquefoil did not occur [5].

Depth: Woolly cinquefoil has been reported on sites with shallow to deep soil. In Colorado it has been reported on "shallow" soil [31], moderately shallow soils up to 16 inches (40 cm) deep [73], "moderately deep" soils [31], and soils more than 24 inches (60 cm) deep [73]. Woolly cinquefoil was present in ponderosa pine/kinnikinnick (Arctostaphylos uva-ursi) communities in Arizona and New Mexico that occurred on "shallow soils" [38]. Based on the maximum depth of soil samples, woolly cinquefoil occurred in dry grasslands of northern Alberta that had soil depths of at least 41.5 inches (105 cm) in some areas [62].

Elevation: Woolly cinquefoil occurs from about 4,300 feet (1,220 m) to as high as 12,000 feet (3,660 m) [54,60]. Individuals were collected at 4,300 feet (1,300 m) in Teton and 4,500 feet (1,370 m) in Teton and Powell counties in Montana [70]. Woolly cinquefoil has been reported as high as 12,000 feet in the Gunnison region of Colorado [54]. The following table includes elevations occupied by woolly cinquefoil in several states. Woolly cinquefoil is likely to occur outside of these elevational ranges, especially in areas where little information is available such as Wyoming and Montana.

Reported elevations of sites with woolly cinquefoil
State Elevation range (feet)
Arizona 7,000-11,500 [29]
Colorado 5,300 [55] -12,000 [54]
Montana 4,300-7,500 [70]
Nevada 8,000-9,000 [27,28]
New Mexico 7,000 [44] -11,500 [16]
Utah 7,380-11,320 [75]
Wyoming 7,100-7,940 [7]

Topography: Woolly cinquefoil occurs in valleys [16,38,42] and on gently sloping uplands [7,73], hillsides [16,38,42,73], and ridgetops [38,42,51].

Woolly cinquefoil has been observed in flat areas, moderate slopes, and steep slopes. It occupied a sedge meadow in north-central New Mexico with a 1.8% slope [11] and woodlands of the Gunnison National Forest with slopes less than 9%. Woolly cinquefoil has been recorded on moderate slopes from 10% to 25% in woodlands of this area [31] and in montane forests with ponderosa pine and Douglas-fir on the east slope of the Front Range [42]. It has been recorded in a Wyoming big sagebrush/bluebunch wheatgrass community that occurred on 35% to 55% slopes in north-central Colorado [73], a Rocky Mountain bristlecone pine/Arizona fescue community on 51% slopes, and a limber pine/matted saxifrage (Saxifraga bronchialis subsp. austromontana) community on 53% slopes on the Gunnison National Forest [31].

Communities with woolly cinquefoil occur on all aspects but may be most common on south-facing slopes. Woolly cinquefoil has been reported on southern aspects in grasslands of New Mexico [16] and northern Alberta [62], Wyoming big sagebrush/bluebunch wheatgrass shrublands of north-central Colorado [73], and montane and subalpine forest communities [31] such as blue spruce/Arizona fescue in northern New Mexico and Arizona [38] and Douglas-fir-limber pine-Rocky Mountain bristlecone pine in New Mexico [51]. Woolly cinquefoil has been reported on an east-facing slope in a quaking aspen stand in Colorado [31], on western aspects in blue spruce/Arizona fescue communities in northern New Mexico and Arizona [38], and on northern aspects in montane ponderosa pine- and Douglas-fir-dominated forests in the Front Range [42].

Moisture: On a moisture gradient, woolly cinquefoil occurs in communities ranging from those generally considered dry [34]—ponderosa pine/Idaho fescue, blue spruce/Arizona fescue [31], and dry grasslands [62]—to wet communities such as wet meadows [2,11,16,20,53] and sedge meadows [11]. In northwestern Montana, woolly cinquefoil was categorized as occurring in a "mesoxeric" moisture regime, intermediate between mesic and xeric [34].

Precipitation: Annual precipitation in areas with woolly cinquefoil ranges from 14 inches (350 mm) [41] to over 30 inches (760 mm) [40,52]. In Arizona and New Mexico, much of this falls from July to September [41,52], and the dry season occurs during spring [38,52]. Along the Front Range, the highest average monthly rainfall from 1952 to 1964 was in May [32], and fall and winter are the driest seasons [32,42]. The following table gives annual precipitation for several communities grouped by location.

Annual precipitation for sites with woolly cinquefoil
Location Habitat Annual precipitation (inches)
Front Range, Colorado
opening in a subalpine lodgepole pine (Pinus contorta) stand 25.7
upper montane dry grass complex 17
montane forests of ponderosa pine or ponderosa pine/Douglas-fir 20 [42]
Douglas-fir-ponderosa pine woodland ≈21.5 [32]
Southwestern Colorado ponderosa pine forest 20-24 [58]
Northern Arizona Engelmann spruce-Rocky Mountain bristlecone pine krummholz and adjacent meadows 39 (3-year mean at weather station nearest to the study site) [40]
Eastern Arizona mixed-conifer forest 30 [52]
Northern Arizona and New Mexico white fir/Arizona fescue 26-28
ponderosa pine/Arizona fescue 20-25 [38]
Western New Mexico ponderosa pine, narrowleaf cottonwood (Populus angustifolia), and Arizona alder (Alnus oblongifolia) riparian community 14 [41]

Soil moisture: Soil moisture on many sites with woolly cinquefoil is often low, although there are exceptions. Low montane habitats of the Front Range generally have low soil moisture, and soil moisture content is close to wilting point in winter and fall [42]. In Douglas-fir-ponderosa pine woodland on the Front Range, soil moisture content was just under 10% from July to September [32]. Average percent soil moisture content from June to August on sites in north-central New Mexico with woolly cinquefoil generally ranged from 9.6% in a quaking aspen-meadow mosaic to 21.9% in a wet meadow. Woolly cinquefoil also occurred in a sedge meadow with 65.6% soil moisture content [11]. High evaporation rates have been reported on sites with woolly cinquefoil in Arizona [38,40] and New Mexico [38,51]. High winds [40,42], high insolation [51,62], and/or soil factors [51] (see Soil) may explain the dry conditions on many sites with woolly cinquefoil.

Temperature: Sites with woolly cinquefoil experience a wide range of temperatures throughout the year. On the Front Range, the coldest reported temperatures on sites with woolly cinquefoil were -10 °F (-23 °C) in January in a Douglas-fir-ponderosa pine woodland [32] and -20 °F (-29 °C) in an upper-montane dry grassland subject to cold air drainage [42]. The mean annual winter temperature in watersheds with woolly cinquefoil in western New Mexico was 30 °F (-1 °C) [41]. The hottest temperature reported on a site with woolly cinquefoil was 96 °F (36 °C) in a low montane ponderosa pine woodland on the Front Range [42]. Mean summer temperatures in study areas with woolly cinquefoil include 47.9 °F (9 °C) near timberline in the San Francisco Mountains of northern Arizona [40] and 64 °F (18 °C) in 2 watersheds in western New Mexico [41]. In north-central New Mexico, June to August weekly minimum temperatures ranged from 31.6 °F (-0.2 °C) in sedge meadows to 41 °F (4.9 °C) in blue spruce-dominated forest [11]. On the Front Range, annual mean temperatures of communities with woolly cinquefoil ranged from 34 °F (1 °C) in a clearing in a lodgepole pine forest to 47 °F (8 °C) in a lower-montane ponderosa pine woodland [42].

On the Front Range, the longest frost-free period in communities with woolly cinquefoil ranged from 75 days in an upper-montane dry grass complex to 137 days in a lower-montane ponderosa pine forest [42]. Lengths of the frost-free period on other sites with woolly cinquefoil are within this range. Woolly cinquefoil was common in a ponderosa pine forest in southwestern Colorado with a frost-free period of about 90 to 110 days [58]. Near timberline in the San Francisco Mountains of northern Arizona, frost-free period from 1917 to 1919 ranged from 101 and 113 days [40].

For effects of temperature on phenology [19], see Seasonal Development. Effects of temperature on plant size and abundance [15] are discussed in Growth, and effects on herbivore and pathogen damage are found in Palatability and/or nutritional value.

SUCCESSIONAL STATUS:
Woolly cinquefoil is most often reported in open, mature, and climax communities but may occur in early-seral vegetation and communities that generally have closed canopies.

Communities with woolly cinquefoil are often described as open [35], although woolly cinquefoil can occur in rather dense forests [31,42]. Grasslands [46,62,71] and woodlands [31,42,51] with woolly cinquefoil in northern Alberta [62], west-central Montana [35], on the east slope of the Front Range [42,46,71], southwestern Colorado [31], and New Mexico [51] are considered open. In northern Arizona and New Mexico, woolly cinquefoil occurred in a blue spruce/Arizona fescue habitat type that was often adjacent to meadows [38]. On the east slope of the Front Range, woolly cinquefoil occurred in a stand with very dense patches as well as comparatively open areas of quaking aspen, Douglas-fir, and ponderosa pine [42]. On the Gunnison National Forest, woolly cinquefoil occurred at 0.2% coverage in a stand with 35% and 40% canopy coverage of Engelmann spruce and quaking aspen, respectively [31].

Woolly cinquefoil habitat, while open, commonly lacks evidence of recent disturbance. In a mosaic of riparian vegetation of different ages in southwestern Colorado, it occurred in communities that were over 61 years old [5]. In northern Arizona, woolly cinquefoil was numerous on a site with interspersed quaking aspen and meadow communities that had not been grazed in about 25 years and had not been cultivated in over 60 years [9,45]. Some of the grasslands where woolly cinquefoil often occurs may be climax communities [3,16,48,61]. Based on the soils present, grasslands with woolly cinquefoil in northern Alberta were concluded to have persisted for "a long period of time". Species in this area that are more common outside of the boreal zone, including woolly cinquefoil, were generally late-successional herbs [61]. Woolly cinquefoil occurred in a soil-climax, dry-grass community on the east slope of the Front Range [42]. Woolly cinquefoil occurred in a Douglas-fir-limber pine-Rocky Mountain bristlecone pine community in New Mexico that may have originated from conifer encroachment into montane grasslands [51]. Subalpine and montane meadows remain stable or succeed to shrubland or woodland due to a complex interaction of aridity, grazing, and fire [3,4,8,33,47,48,49,76]. A discussion of the role of these factors in meadow persistence or conversion is available in the FEIS review of Thurber fescue.

Woolly cinquefoil is occasionally documented on recently-disturbed sites and in early-seral vegetation. In northern Arizona, woolly cinquefoil had significantly (P<0.05) greater occurrence than expected on sites disturbed by Botta's pocket gophers [45]. It occurred in a dry grassland on the east slope of the Front Range that had "recently" been cleared of brush and grazed [42] (see Grazing). According to a 1917 classification of foothills vegetation of the Front Range, woolly cinquefoil was a component of a "primitive grassland community" on the Front Range, including a subtype on compacted granite-gravel. Cover on a few granite-gravel sites was comprised entirely of woolly cinquefoil [71].

FIRE EFFECTS AND MANAGEMENT

SPECIES: Potentilla hippiana
As of 2009, there were no data on woolly cinquefoil's immediate or long-term response to fire. There were no studies comparing abundance of woolly cinquefoil either before and after fire or on burned and unburned sites. There were no anecdotal reports of woolly cinquefoil in burned areas. The following is speculation based on woolly cinquefoil morphology, the disturbance histories on sites with woolly cinquefoil, and responses of other cinquefoils (Potentilla spp.) and/or species with similar morphology.

FIRE EFFECTS:
Immediate fire effect on plant: Due to a lack of information as of 2009, the immediate effect of fire on woolly cinquefoil is uncertain. A report of vegetative spread [71] and the presence of a caudex [12,21,64,75] suggest that top-kill may be more likely than complete mortality, although there are no reports of woolly cinquefoil sprouting following damage.

Postfire regeneration strategy [65]:
Caudex or an herbaceous root crown, growing points in soil

Possible:
Ground residual colonizer (on site, initial community)
Initial off-site colonizer (off site, initial community)
Secondary colonizer (on- or off-site seed sources)

Fire adaptations and plant response to fire:

Fire adaptations: As of 2009, sprouting of woolly cinquefoil following fire and the location of woolly cinquefoil's perennating buds in relation to the soil surface had not been reported. Some species with a caudex over a thick taproot, such as sulfur cinquefoil (Potentilla recta) and arrowleaf balsamroot (Balsamorhiza sagittata), may sprout after fire. Other species with a caudex over a thick taproot, such as gooseberryleaf globemallow (Sphaeralcea grossulariifolia), may increase or show no change in response to fire. Soil may protect most or all of the caudex from the heat of low- to moderate-severity fire, and nutrients stored in the taproot may facilitate production of new aboveground tissue following fire.

Woolly cinquefoil's abundant seed production suggests potential for establishment from on- or off-site seed sources. However, the longevity of woolly cinquefoil seeds on burned or unburned sites and the ability of those seeds to germinate in recently burned areas are unknown. Greater woolly cinquefoil abundance has been observed on sites disturbed by Botta's pocket gophers [45] and on sites subject to season-long grazing [14] than on undisturbed or less disturbed sites. However, the ability of woolly cinquefoil to establish in burned areas is unknown. Small sulfur cinquefoil individuals were observed 1 year after fire in an arid grassland of northwestern Montana. It is not known if these plants established from seed dispersed from off site-sources, seed that was present in the seed bank and survived the fire, or from caudex sprouts of previously established individuals [39]. Seeds of sticky cinquefoil (Potentilla glandulosa) in soil collected from unburned sites in the Yellowstone ecosystem germinated after soil was heated to 212 °F (100 °C) in the laboratory [10].

Plant response to fire: There is potential for woolly cinquefoil to recover quickly following fire through caudex sprouting and germination of numerous seeds. This would be most likely following a single, low-severity fire. High fire severity would result in greater soil heating and a greater chance of mortality, while frequent fires may exhaust nutrient reserves in the taproot. Widespread fires may delay recovery due to long distances from available seed sources. Woolly cinquefoil was observed on a southwestern New Mexico site a year after a prescribed fire burned patchily and at low severity in a ponderosa pine forest. It is not known if it occurred in areas that had been burned or if it occurred only in unburned patches (Kleinman 2009 personal communication [30]). Either way, it would have potential to spread from those patches. Sulfur cinquefoil had more positive short-term responses to fall fires in western Montana, which were more severe than spring fires [39]. Other factors that may impact woolly cinquefoil's response include site characteristics and postfire weather. For instance, woolly cinquefoil's general trend of occurring in open areas (see Successional Status) suggests that fire exclusion may be more detrimental to woolly cinquefoil in communities where fire historically maintained an open canopy. Small sulfur cinquefoil plants declined on both burned and control plots in the year after burning treatments, likely due to receiving 51% of the average precipitation [39].

The disturbance histories of sites with woolly cinquefoil are varied, with woolly cinquefoil occurring on recently-disturbed sites as well as long-undisturbed sites (see Successional Status). Woolly cinquefoil has been documented in habitats with no known disturbance in at least 70 years [5], habitats that were heavily disturbed 60 or more years ago [5,9,45], habitats that were minimally disturbed recently [2,45], and heavily- and recently-grazed habitats [14]. Several sites with woolly cinquefoil, even those historically maintained by fire, have experienced little or no recent disturbance. For instance, dry grasslands occupied by woolly cinquefoil in northern Alberta had not burned in about 70 years. It is possible that continued fire exclusion on this site would ultimately result in conversion to another habitat type and that woolly cinquefoil would decline with other grassland species [61].

FUELS AND FIRE REGIMES:
Fuels: Fuels on sites with woolly cinquefoil are likely to vary greatly with cover type and site characteristics. On the Front Range, woolly cinquefoil occurred in sparsely vegetated communities [71] and in a stand with dense patches of Douglas-fir, ponderosa pine, and quaking aspen [42]. Estimates of woolly cinquefoil biomass on 2 transects in an Arizona meadow where it had the 3rd highest importance value were 212.31 kg/ha and 268.5 kg/ha. Its estimated biomass on transects in a meadow where it was a minor species were 0 and 122.36 kg/ha [2].

Fire regimes: Communities with woolly cinquefoil have varying fire regimes. Many habitats with woolly cinquefoil are maintained by frequent, low-severity fire. Great Plains grasslands—such as those occupied by woolly cinquefoil in Montana [17,21], North Dakota [21], South Dakota [21,24], Wyoming [7,18,21], Colorado [21,46,55,60,71], and New Mexico [21], typically have fire-return intervals of less than 20 years. Ponderosa pine woodlands, which may also be occupied by woolly cinquefoil [11,28,38,41]—generally experience low-severity fires every 10 to 20 years. Prescribed burning was suggested in a ponderosa pine/Arizona fescue community with woolly cinquefoil in northern New Mexico to maintain the open, "parklike" structure of the stand [38]. However, several shrublands and forests occupied by woolly cinquefoil have fire-return intervals of 100 years or more. Woolly cinquefoil has been documented in southwestern moist, mixed-conifer-quaking aspen forests [52] with fire-return intervals of over 100 years for fire of any severity. See the Fire Regime Table for further information on fire regimes of vegetation communities in which woolly cinquefoil may occur. Find further 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".

Fire frequency of many montane meadows has been reduced due to fire exclusion and grazing, which reduce fuel loads [3,66]. Milchunas [47] reviews the interaction of fire, weather, and grazing in ponderosa pine forests and adjacent montane meadows. A discussion of the role of fire in meadow persistence or conversion to shrubland, woodland, or forest is available in the FEIS review of Thurber fescue.

FIRE MANAGEMENT CONSIDERATIONS:
More research is needed to determine recommendations regarding woolly cinquefoil and fire. Woolly cinquefoil has potential for rapid recovery and occurs in habitats maintained by frequent fire, suggesting that any negative impacts of a single low-severity fire are likely short term. Fire exclusion could result in shading [61] and ultimately type conversion of some grassland habitats [4,33,47,61]. Prescribed burning has been recommended for white fir/Arizona fescue and ponderosa pine/Arizona fescue communities with woolly cinquefoil in northern Arizona and New Mexico [38] and dry grasslands with woolly cinquefoil in northern Alberta [61]. However, woolly cinquefoil occurrence in fire-excluded grasslands [61] and cover types with fairly long fire-return intervals [31,42,52] suggests that woolly cinquefoil tolerates long periods without fire in some areas.

MANAGEMENT CONSIDERATIONS

SPECIES: Potentilla hippiana
IMPORTANCE TO WILDLIFE AND LIVESTOCK:
Woolly cinquefoil generally has low forage and cover value for wildlife species. Ungulates, small mammals, birds, and insects eat it to some extent. Although woolly cinquefoil provides little cover, it occurs in habitats that are important for wildlife.

Palatability and/or nutritional value: Woolly cinquefoil generally has low palatability and forage value. The flowers are apparently the most palatable part of the plant. The tannin content of the leaves likely contributes to their low palatability. According to a 1937 US Department of Agriculture publication, woolly cinquefoil provides no or poor forage for cattle and poor to fair forage for domestic sheep. Horses occasionally eat the flowers. Elk use woolly cinquefoil lightly in spring [68], summer, and fall [67,68]. Woolly cinquefoil's forage value for deer may be little to none [67], or deer may "frequently eat considerable quantities" [68]. According to a review, cinquefoils (Potentilla spp.) occur in the bighorn sheep diet to some extent and comprise from 0.5% to 5% of the diet of some birds and small mammals such as ruffed grouse, rosy finch, cottontails (Sylvilagus spp.), and Richardson's ground squirrel. Cinquefoils comprised from 5% to 10% of the diet of the least chipmunk in the western prairie and "mountain-desert" regions [43].

Caterpillars and a grasshopper were observed feeding on woolly cinquefoil in an artificial heat experiment in Colorado. Woolly cinquefoil was also damaged by 3 different pathogens. Caterpillar rasping damage was significantly (P=0.05) less in heated plots than in control plots. Warming explained >10% and melting date explained <1% of the variance in number of herbivores and pathogens on woolly cinquefoil [59].

Cover value: Although woolly cinquefoil does not provide cover for wildlife species, it occurs in meadows and forests important to wildlife. It was a common forb in meadows used by greater sage-grouse in North Park, Colorado [20]. Woolly cinquefoil occurs at low frequency and density in wet meadows and moist openings that provide high forage value and habitat heterogeneity in the Southwest [53]. It occurred in white fir/Arizona fescue stands that provide good habitat for wildlife when stands form a mosaic of different successional stages [38].

VALUE FOR REHABILITATION OF DISTURBED SITES:
No information is available on this topic.

OTHER USES:
According to Kartesz [27,28], Native Americans used woolly cinquefoil to treat burns, as a dermatological treatment, and to expedite childbirth.

OTHER MANAGEMENT CONSIDERATIONS:
As noted in a guide to growing and propagating wild flowers, cinquefoils (Potentilla spp.) "walk that delicate—some would say imaginary—line between wildflower and weed" [13]. The weediness of woolly cinquefoil, potential need for control, or methods of control had not been addressed in the literature as of 2009.

Grazing: Livestock grazing history on sites with woolly cinquefoil is variable and includes sites unlikely to have been grazed, historically grazed, and recently grazed. Woolly cinquefoil occurs in riparian communities in southwestern Colorado that have "probably not" been grazed by livestock [5]. Bison were the main grazers in an area of northern Alberta where woolly cinquefoil occurred in dry grasslands [62]. Sites in Arizona with abundant woolly cinquefoil had no recent [9,45] or minimal [2] livestock grazing. Woolly cinquefoil occurred on sites in the San Francisco Peaks area that had not been been grazed in over 25 years and had not been cultivated in over 60 years [9,45]. A meadow with 11 to 12% cover of woolly cinquefoil in Arizona was either unused or grazed infrequently at light intensity [2]. Woolly cinquefoil occurred in a grassland on the east slope of the Front Range that had "recently" been cleared of brush and grazed during summer [42].

Density of woolly cinquefoil on plots grazed at different times suggests increased density with increased use. Woolly cinquefoil density was about 1 plant/foot² in plots near Colorado Springs, Colorado, that were either grazed in September and October, alternately grazed in spring and fall, or ungrazed. Density was about 2 plants/foot² in plots grazed in May and June or throughout the season (May-October). Heavy, season-long grazing had occurred on this site for 23 years before grazing treatments were implemented [14]. A flora of west-central Montana groups woolly cinquefoil with cinquefoil species that often occur in "overgrazed" areas [35].

APPENDIX: FIRE REGIME TABLE

SPECIES: Potentilla hippiana
The following table provides fire regime information that may be relevant to woolly cinquefoil habitats. Find further 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".

Fire regime information on vegetation communities in which woolly cinquefoil may occur. This information is taken from the LANDFIRE Rapid Assessment Vegetation Models [37], which were developed by local experts using available literature, local data, and/or expert opinion. This table summarizes fire regime characteristics for each plant community listed. The PDF file linked from each plant community name describes the model and synthesizes the knowledge available on vegetation composition, structure, and dynamics in that community. Cells are blank where information is not available in the Rapid Assessment Vegetation Model.
Southwest Great Basin Northern and Central Rockies Northern Great Plains
Southwest
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Southwest Grassland
Shortgrass prairie Replacement 87% 12 2 35
Mixed 13% 80    
Shortgrass prairie with shrubs Replacement 80% 15 2 35
Mixed 20% 60    
Shortgrass prairie with trees Replacement 80% 15 2 35
Mixed 20% 60    
Montane and subalpine grasslands Replacement 55% 18 10 100
Surface or low 45% 22    
Montane and subalpine grasslands with shrubs or trees Replacement 30% 70 10 100
Surface or low 70% 30    
Southwest Shrubland
Mountain sagebrush (cool sage) Replacement 75% 100    
Mixed 25% 300    
Mountain-mahogany shrubland Replacement 73% 75    
Mixed 27% 200    
Southwest Woodland
Pinyon-juniper (mixed fire regime) Replacement 29% 430    
Mixed 65% 192    
Surface or low 6% >1,000    
Pinyon-juniper (rare replacement fire regime) Replacement 76% 526    
Mixed 20% >1,000    
Surface or low 4% >1,000    
Ponderosa pine/grassland (Southwest) Replacement 3% 300    
Surface or low 97% 10    
Bristlecone-limber pine (Southwest) Replacement 67% 500    
Surface or low 33% >1,000    
Southwest Forested
Riparian forest with conifers Replacement 100% 435 300 550
Ponderosa pine-Douglas-fir (southern Rockies) Replacement 15% 460    
Mixed 43% 160    
Surface or low 43% 160    
Southwest mixed conifer (warm, dry with aspen) Replacement 7% 300    
Mixed 13% 150 80 200
Surface or low 80% 25 2 70
Southwest mixed conifer (cool, moist with aspen) Replacement 29% 200 80 200
Mixed 35% 165 35  
Surface or low 36% 160 10  
Aspen with spruce-fir Replacement 38% 75 40 90
Mixed 38% 75 40  
Surface or low 23% 125 30 250
Stable aspen without conifers Replacement 81% 150 50 300
Surface or low 19% 650 600 >1,000
Spruce-fir Replacement 96% 210 150  
Mixed 4% >1,000 35 >1,000
Great Basin
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Great Basin Grassland
Mountain meadow (mesic to dry) Replacement 66% 31 15 45
Mixed 34% 59 30 90
Great Basin Shrubland
Basin big sagebrush Replacement 80% 50 10 100
Mixed 20% 200 50 300
Wyoming sagebrush steppe Replacement 89% 92 30 120
Mixed 11% 714 120  
Mountain big sagebrush Replacement 100% 48 15 100
Mountain big sagebrush with conifers Replacement 100% 49 15 100
Mountain sagebrush (cool sage) Replacement 75% 100    
Mixed 25% 300    
Mountain shrubland with trees Replacement 22% 105 100 200
Mixed 78% 29 25 100
Great Basin Woodland
Ponderosa pine Replacement 5% 200    
Mixed 17% 60    
Surface or low 78% 13    
Great Basin Forested
Interior ponderosa pine Replacement 5% 161   800
Mixed 10% 80 50 80
Surface or low 86% 9 8 10
Ponderosa pine-Douglas-fir Replacement 10% 250   >1,000
Mixed 51% 50 50 130
Surface or low 39% 65 15  
Great Basin Douglas-fir (dry) Replacement 12% 90   600
Mixed 14% 76 45  
Surface or low 75% 14 10 50
Aspen with conifer (low to midelevation) Replacement 53% 61 20  
Mixed 24% 137 10  
Surface or low 23% 143 10  
Douglas-fir (warm mesic interior) Replacement 28% 170 80 400
Mixed 72% 65 50 250
Aspen with conifer (high elevation) Replacement 47% 76 40  
Mixed 18% 196 10  
Surface or low 35% 100 10  
Spruce-fir-pine (subalpine) Replacement 98% 217 75 300
Mixed 2% >1,000    
Aspen with spruce-fir Replacement 38% 75 40 90
Mixed 38% 75 40  
Surface or low 23% 125 30 250
Stable aspen without conifers Replacement 81% 150 50 300
Surface or low 19% 650 600 >1,000
Northern and Central Rockies
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Northern and Central Rockies Grassland
Northern prairie grassland Replacement 55% 22 2 40
Mixed 45% 27 10 50
Mountain grassland Replacement 60% 20 10  
Mixed 40% 30    
Northern and Central Rockies Shrubland
Wyoming big sagebrush Replacement 63% 145 80 240
Mixed 37% 250    
Basin big sagebrush Replacement 60% 100 10 150
Mixed 40% 150    
Mountain shrub, nonsagebrush Replacement 80% 100 20 150
Mixed 20% 400    
Mountain big sagebrush steppe and shrubland Replacement 100% 70 30 200
Northern and Central Rockies Forested
Ponderosa pine (Northern Great Plains) Replacement 5% 300    
Mixed 20% 75    
Surface or low 75% 20 10 40
Ponderosa pine (Northern and Central Rockies) Replacement 4% 300 100 >1,000
Mixed 19% 60 50 200
Surface or low 77% 15 3 30
Ponderosa pine (Black Hills, low elevation) Replacement 7% 300 200 400
Mixed 21% 100 50 400
Surface or low 71% 30 5 50
Ponderosa pine (Black Hills, high elevation) Replacement 12% 300    
Mixed 18% 200    
Surface or low 71% 50    
Ponderosa pine-Douglas-fir Replacement 10% 250   >1,000
Mixed 51% 50 50 130
Surface or low 39% 65 15  
Douglas-fir (xeric interior) Replacement 12% 165 100 300
Mixed 19% 100 30 100
Surface or low 69% 28 15 40
Douglas-fir (warm mesic interior) Replacement 28% 170 80 400
Mixed 72% 65 50 250
Douglas-fir (cold) Replacement 31% 145 75 250
Mixed 69% 65 35 150
Lower subalpine (Wyoming and Central Rockies) Replacement 100% 175 30 300
Upper subalpine spruce-fir (Central Rockies) Replacement 100% 300 100 600
Northern Great Plains
Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics
Percent of fires Mean interval
(years)
Minimum interval
(years)
Maximum interval
(years)
Northern Plains Grassland
Nebraska Sandhills prairie Replacement 58% 11 2 20
Mixed 32% 20    
Surface or low 10% 67    
Northern mixed-grass prairie Replacement 67% 15 8 25
Mixed 33% 30 15 35
*Fire Severities
Replacement: Any fire that causes greater than 75% top removal of a vegetation-fuel type, resulting in general replacement of existing vegetation; may or may not cause a lethal effect on the plants.
Mixed: Any fire burning more than 5% of an area that does not qualify as a replacement, surface, or low-severity fire; includes mosaic and other fires that are intermediate in effects.
Surface or low: Any fire that causes less than 25% upper layer replacement and/or removal in a vegetation-fuel class but burns 5% or more of the area [22,36].

Potentilla hippiana: REFERENCES


1. Aitken, Marti; Parks, Katherine Gray. 2004. Guide to the common Potentilla species of the Blue Mountains ecoregion. Gen. Tech. Rep. PNW-GTR-603. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 50 p. [49296]
2. Aldon, Earl F.; Barstad, Janet F. 1987. Escudilla Mountain Research Natural Area: a study of an undisturbed montane grassland in Arizona. Natural Areas Journal. 7(3): 107-117. [40109]
3. Allen, Craig Daniel. 1989. Changes in the landscape of the Jemez Mountains, New Mexico. Berkeley, CA: University of California. 346 p. Dissertation. [42116]
4. Arno, Stephen F.; Gruell, George E. 1986. Douglas-fir encroachment into mountain grasslands in southwestern Montana. Journal of Range Management. 39(3): 272-276. [343]
5. Baker, William L.; Walford, Gillian M. 1995. Multiple stable states and models of riparian vegetation succession on the Animas River, Colorado. Annals of the Association of American Geographers. 85(2): 320-338. [74816]
6. Beatley, Janice C. 1976. Vascular plants of the Nevada Test Site and central-southern Nevada: ecologic and geographic distributions. [Washington, DC]: U.S. Energy Research and Development Administration, Office of Technical Information, Technical Information Center. 308 p. Available from U.S. Department of Commerce, National Technical Information Service, Springfield, VA. TID-26881/DAS. [63152]
7. Boutton, Thomas W.; Harrison, A. Tyrone; Smith, Bruce N. 1980. Distribution of biomass of species differing in photosynthetic pathway along an altitudinal transect in southeastern Wyoming grassland. Oecologia. 45(3): 287-298. [74817]
8. Butler, David R. 1986. Conifer invasion of subalpine meadows, central Lemhi Mountains, Idaho. Northwest Science. 60(3): 166-173. [571]
9. Cantor, Lisa F.; Whitham, Thomas G. 1989. Importance of belowground herbivory: pocket gophers may limit aspen to rock outcrop refugia. Ecology. 70(4): 962-970. [7935]
10. Clark, David Lee. 1991. The effect of fire on Yellowstone ecosystem seed banks. Bozeman, MT: Montana State University. 115 p. Thesis. [36504]
11. Coop, Jonathan D.; Givnish, Thomas J. 2007. Gradient analysis of reversed treelines and grasslands of the Valles Caldera, New Mexico. Journal of Vegetation Science. 18: 43-54. [67321]
12. Cronquist, Arthur; Holmgren, Noel H.; Holmgren, Patricia K. 1997. Intermountain flora: Vascular plants of the Intermountain West, U.S.A. Vol. 3, Part A: Subclass Rosidae (except Fabales). New York: The New York Botanical Garden. 446 p. [28652]
13. Cullina, William. 2000. The New England Wild Flower Society guide to growing and propagating wildflowers of the United States and Canada. A Francis Tenenbaum Book. New York: Houghton Mifflin Company. 322 p. [75528]
14. Currie, Pat O. 1976. Recovery of ponderosa pine-bunchgrass ranges through grazing and herbicide or fertilizer treatments. Journal of Range Management. 29(6): 444-448. [15824]
15. de Valpine, Perry; Harte, John. 2001. Plant responses to experimental warming in a montane meadow. Ecology. 82(3): 637-648. [74818]
16. Dick-Peddie, William A. 1993. New Mexico vegetation: past, present, and future. Albuquerque, NM: University of New Mexico Press. 244 p. [21097]
17. Dorn, Robert D. 1984. Vascular plants of Montana. Cheyenne, WY: Mountain West Publishing. 276 p. [819]
18. Dorn, Robert D. 1988. Vascular plants of Wyoming. Cheyenne, WY: Mountain West Publishing. 340 p. [6129]
19. Dunne, Jennifer A.; Harte, John; Taylor, Kevin J. 2003. Subalpine meadow flowering phenology responses to climate change: integrating experimental and gradient methods. Ecological Monographs. 73(1): 69-86. [73236]
20. Gill, R. Bruce. 1965. Effects of sagebrush control on distribution and abundance of sage grouse. Colorado Federal Aid in Wildlife Restoration Project No. W-37-R-17. Job completion report--Research project segment: April 1, 1963 to December 6, 1965. [Denver, CO]: Colorado Game, Fish, and Parks Department. 185 p. [36876]
21. Great Plains Flora Association. 1986. Flora of the Great Plains. Lawrence, KS: University Press of Kansas. 1392 p. [1603]
22. Hann, Wendel; Havlina, Doug; Shlisky, Ayn; [and others]. 2008. Interagency fire regime condition class guidebook. Version 1.3, [Online]. In: Interagency fire regime condition class website. U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior; The Nature Conservancy; Systems for Environmental Management (Producer). 119 p. Available: http://frames.nbii.gov/frcc/documents/FRCC_Guidebook_2008.07.10.pdf [2008, September 03]. [70966]
23. Harrington, H. D. 1964. Manual of the plants of Colorado. 2nd ed. Chicago, IL: The Swallow Press, Inc. 666 p. [6851]
24. Harvey, LeRoy Harris. 1908. Floral succession in the prairie-grass formation of southeastern South Dakota, serotinal and autumnal floral aspects. Botanical Gazette. 46(4): 277-298. [55844]
25. Hill, Alison; Pieper, Rex D.; Southward, G. Morris. 1992. Habitat-type classification of the pinyon-juniper woodlands in western New Mexico. Bulletin 766. Las Cruces, NM: New Mexico State University, College of Agriculture and Home Economics, Agricultural Experiment Station. 80 p. [37374]
26. Hulten, Eric. 1968. Flora of Alaska and neighboring territories. Stanford, CA: Stanford University Press. 1008 p. [13403]
27. Kartesz, John T. 1999. A synonymized checklist and atlas with biological attributes for the vascular flora of the United States, Canada, and Greenland. 1st ed. In: Kartesz, John T.; Meacham, Christopher A. Synthesis of the North American flora (Windows Version 1.0), [CD-ROM]. Chapel Hill, NC: North Carolina Botanical Garden (Producer). In cooperation with: The Nature Conservancy; U.S. Department of Agriculture, Natural Resources Conservation Service; U.S. Department of the Interior, Fish and Wildlife Service. [36715]
28. Kartesz, John Thomas. 1988. A flora of Nevada. Reno, NV: University of Nevada. 1729 p. [In 2 volumes]. Dissertation. [42426]
29. Kearney, Thomas H.; Peebles, Robert H.; Howell, John Thomas; McClintock, Elizabeth. 1960. Arizona flora. 2nd ed. Berkeley, CA: University of California Press. 1085 p. [6563]
30. Kleinman, Russ. 2009. [Email to Rachelle Meyer]. August 8. Regarding Potentilla hippiana information. Silver City, NM: Western New Mexico University, Dale A. Zimmerman Herberium. On file with: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT; FEIS files. [75495]
31. Komarkova, Vera; Alexander, Robert R.; Johnston, Barry C. 1988. Forest vegetation of the Gunnison and parts of the Uncompahgre National Forests: a preliminary habitat type classification. Gen. Tech. Rep. RM-163. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 65 p. [5798]
32. Kooiman, Marianne; Linhart, Yan B. 1986. Structure and change in herbaceous communities of four ecosystems in the Front Range, Colorado, U.S.A. Arctic and Alpine Research. 18(1): 97-110. [4076]
33. Koterba, Wayne D.; Habeck, James R. 1971. Grasslands of the North Fork Valley, Glacier National Park, Montana. Canadian Journal of Botany. 49: 1627-1636. [6401]
34. Lackschewitz, Klaus. 1986. Plants of west-central Montana--identification and ecology: annotated checklist. Gen. Tech. Rep. INT-217. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 128 p. [2955]
35. Lackschewitz, Klaus. 1991. Vascular plants of west-central Montana--identification guidebook. Gen. Tech. Rep. INT-227. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 648 p. [13798]
36. LANDFIRE Rapid Assessment. 2005. Reference condition modeling manual (Version 2.1), [Online]. In: LANDFIRE. Cooperative Agreement 04-CA-11132543-189. Boulder, CO: The Nature Conservancy; U.S. Department of Agriculture, Forest Service; U.S. Department of the Interior (Producers). 72 p. Available: https://www.landfire.gov /downloadfile.php?file=RA_Modeling_Manual_v2_1.pdf [2007, May 24]. [66741]
37. LANDFIRE Rapid Assessment. 2007. Rapid assessment reference condition models, [Online]. In: LANDFIRE. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Lab; U.S. Geological Survey; The Nature Conservancy (Producers). Available: https://www.landfire.gov /models_EW.php [2008, April 18] [66533]
38. Larson, Milo; Moir, W. H. 1987. Forest and woodland habitat types (plant associations) of northern New Mexico and northern Arizona. 2nd ed. Albuquerque, NM: U.S. Department of Agriculture, Forest Service, Southwestern Region. 90 p. [8947]
39. Lesica, Peter; Martin, Brian. 2003. Effects of prescribed fire and season of burn on recruitment of the invasive exotic plant, Potentilla recta, in a semi-arid grassland. Restoration Ecology. 11(4): 516-523. [45123]
40. Little, Elbert L., Jr. 1941. Alpine flora of San Francisco Mountain, Arizona. Madrono. 6: 65-96. [14668]
41. Lucas, Richard W.; Baker, Terrell T.; Wood, M. Karl; Allison, Christopher D.; Vanleeuwen, Dawn M. 2004. Riparian vegetation response to different intensities and seasons of grazing. Journal of Range Management. 57(5): 466-474. [50449]
42. Marr, John W. 1961. Ecosystems of the east slope of the Front Range in Colorado. University of Colorado Studies, Series in Biology: No. 8. Boulder, CO: University of Colorado Press. 134 p. [5724]
43. Martin, Alexander C.; Zim, Herbert S.; Nelson, Arnold L. 1951. American wildlife and plants. New York: McGraw-Hill Book Company, Inc. 500 p. [4021]
44. Martin, William C.; Hutchins, Charles R. 1981. A flora of New Mexico. Volume 2. Germany: J. Cramer. 2589 p. [37176]
45. Martinsen, Gregory D.; Cushman, J. Hall; Whitham, Thomas G. 1990. Impact of pocket gopher disturbance on plant species diversity in a shortgrass prairie community. Oecologia. 83: 132-138. [11828]
46. McGinnies, William J.; Shantz, Homer L.; McGinnies, William G. 1991. Changes in vegetation and land use in eastern Colorado: A photographic study, 1904-1986. ARS-85. Washington, DC: U.S. Department of Agriculture, Agricultural Research Service. 165 p. [18825]
47. Milchunas, Daniel G., ed. 2006. Responses of plant communities to grazing in the southwestern United States. Gen. Tech. Rep. RMRS-GTR-169. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 126 p. [63342]
48. Miller, Eric A.; Halpern, Charles B. 1998. Effects of environment and grazing disturbance on tree establishment in meadows of the central Cascade Range, Oregon, USA. Journal of Vegetation Science. 9(2): 265-282. [60189]
49. Miller, Philip Clement. 1964. Factors influencing the vegetation pattern of the White River Plateau in northwestern Colorado. Boulder, CO: University of Colorado. 232 p. Dissertation. [73694]
50. Miller-Rushing, Abraham J.; Inouye, David W.; Primack, Richard B. 2008. How well do first flowering dates measure plant responses to climate change? The effects of population size and sampling frequency. Journal of Ecology. 96(6): 1289-1296. [73018]
51. Moir, W. H. 1993. Alpine tundra and coniferous forest. In: Dick-Peddie, William A., ed. New Mexico vegetation: Past, present, and future. Albuquerque, NM: University of New Mexico Press: 47-84. [21099]
52. Patton, David R. 1976. Timber harvesting increases deer and elk use of a mixed conifer forest. Res. Note RM-329. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 3 p. [14448]
53. Patton, David R.; Judd, B. Ira. 1970. The role of wet meadows as wildlife habitat in the Southwest. Journal of Range Management. 23(4): 272-275. [74819]
54. Pleasants, John M. 1980. Competition for bumblebee pollinators in Rocky Mountain plant communities. Ecology. 61(6): 1446-1459. [74820]
55. Ramaley, Francis. 1919. Xerophytic grasslands at different altitudes in Colorado. Bulletin of the Torrey Botanical Club. 46(2): 37-52. [63275]
56. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843]
57. Roland, A. E.; Smith, E. C. 1969. The flora of Nova Scotia. Halifax, NS: Nova Scotia Museum. 746 p. [13158]
58. Romme, William H.; Jamieson, David W.; Redders, Jeffery S.; Bigsby, Ginger; Lindsey, J. Page; Kendall, Deborah; Cowen, Robert; Kreykes, Thomas; Spencer, Albert W.; Ortega, Joseph C. 1992. Old-growth forests of the San Juan National Forest in southwestern Colorado. In: Kaufmann, Merrill R.; Moir, W. H.; Bassett, Richard L., tech. coords. Old-growth forests in the Southwest and Rocky Mountain regions: Proceedings of a workshop; 1992 March 9-13; Portal, AZ. Gen. Tech. Rep. RM-213. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 154-165. [19050]
59. Roy, Bitty A.; Gusewell, Sabine; Harte, John. 2004. Response of plant pathogens and herbivores to a warming experiment. Ecology. 85(9): 2570-2581. [75311]
60. Rydberg, Per Axel. 1906. Flora of Colorado. Bulletin 100. Fort Collins, CO: Colorado Agricultural College, Agricultural Experiment Station. 448 p. [63874]
61. Schwarz, A. G.; Wein, Ross W. 1997. Threatened dry grasslands in the continental boreal forests of Wood Buffalo National Park. Canadian Journal of Botany. 75(8): 1363-1370. [28005]
62. Schwarz, Arthur G.; Thorpe, Jeffrey P.; Redmann, Robert E. 1988. Isolated grasslands in the boreal forest region of western Canada. In: Davis, Arnold; Stanford, Geoffrey, eds. The prairie: roots of our culture; foundation of our economy: Proceedings, 10th North American prairie conference; 1986 June 22-26; Denton, TX. Dallas, TX: Native Prairie Association of Texas: 01.09. [25577]
63. Scoggan, H. J. 1978. The flora of Canada. Part 3: Dicotyledoneae (Saururaceae to Violaceae). National Museum of Natural Sciences: Publications in Botany, No. 7(3). Ottawa: National Museums of Canada. 568 p. [75493]
64. Scott, Richard W. 1995. The alpine flora of the Rocky Mountains. Volume I: The Middle Rockies. Salt Lake City, UT: University of Utah Press. 901 p. [74878]
65. Stickney, Peter F. 1989. Seral origin of species comprising secondary plant succession in Northern Rocky Mountain forests. FEIS workshop: Postfire regeneration. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. 10 p. [20090]
66. Stuever, Mary C.; Hayden, John S. 1996. Plant associations (habitat types) of the forests and woodlands of Arizona and New Mexico. Final report: Contract R3-95-27. Placitas, NM: Seldom Seen Expeditions, Inc. 520 p. [28868]
67. Thill, Ronald E.; Ffolliott, Peter F.; Patton, David R. 1983. Deer and elk forage production in Arizona mixed conifer forests. Res. Pap. RM-248. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 13 p. [14381]
68. U.S. Department of Agriculture, Forest Service. 1937. Range plant handbook. Washington, DC. 532 p. [2387]
69. U.S. Department of Agriculture, Natural Resources Conservation Service. 2009. PLANTS Database, [Online]. Available: https://plants.usda.gov /. [34262]
70. University of Montana. 2009. Montana vascular plant database, [Online]. In: University of Montana Herbarium. Missoula, MT: University of Montana (Producer). Available: http://herbarium.dbs.umt.edu/database/Default.aspx [2009, July 20]. [75045]
71. Vestal, Arthur G. 1917. Foothills vegetation in the Colorado Front Range. Botanical Gazette. 64(5): 353-385. [64489]
72. Voss, Edward G. 1985. Michigan flora. Part II. Dicots (Saururaceae--Cornaceae). Bulletin 59. Bloomfield Hills, MI: Cranbrook Institute of Science; Ann Arbor, MI: University of Michigan Herbarium. 724 p. [11472]
73. Wasser, C. H.; Hess, Karl. 1982. The habitat types of Region II--U.S. Forest Service: a synthesis. Final report: Cooperative Agreement No. 16-845-CA. Lakewood, CO: U.S. Department of Agriculture, Forest Service, Region 2. 140 p. [5594]
74. Weber, William A.; Wittmann, Ronald C. 1996. Colorado flora: eastern slope. 2nd ed. Niwot, CO: University Press of Colorado. 524 p. [27572]
75. Welsh, Stanley L.; Atwood, N. Duane; Goodrich, Sherel; Higgins, Larry C., eds. 1987. A Utah flora. The Great Basin Naturalist Memoir No. 9. Provo, UT: Brigham Young University. 894 p. [2944]
76. Zier, James L.; Baker, William L. 2006. A century of vegetation change in the San Juan Mountains, Colorado: an analysis using repeat photography. Forest Ecology and Management. 228(1-3): 251-262. [62625]

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