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SPECIES:  Chimaphila umbellata


Introductory

SPECIES: Chimaphila umbellata
AUTHORSHIP AND CITATION : Matthews, Robin F. 1994. Chimaphila umbellata. 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/shrub/chiumb/all.html []. Revisions : On 15 December 2014, the common name of this species was changed from: prince's-pine to: pipsissewa. ABBREVIATION : CHIUMB SYNONYMS : Pyrola umbellata L. [41,82] SCS PLANT CODE : CHUM COMMON NAMES : pipsissewa prince's-pine prince's pine waxflower wintergreen TAXONOMY : The currently accepted scientific name of pipsissewa is Chimaphila umbellata (L.) Barton [17,39,57,81,82]. Pipsissewa comprises a circumboreal complex in which several geographical varieties have been recognized [70]. The following varieties and subspecies are accepted: Chimaphila umbellata var. umbellata (Eurasia) [70] Chimaphila umbellata var. occidentalis (Rydb.) Blake (western North America) [17,38,39,57,82] Chimaphila umbellata var. cisatlantica Blake (eastern North America) [17,24,63,70] Chimaphila umbellata var. acuta (Rydb.) Blake (Arizona and New Mexico) [35,44] Chimaphila umbellata subsp. domingensis (S.F. Blake) Dorr (Dominican Republic) [84] Chimaphila umbellata subsp. mexicana (DC.) Hulten (Mexico) [41] LIFE FORM : Shrub, Forb FEDERAL LEGAL STATUS : See OTHER STATUS OTHER STATUS : Chimaphila umbellata var. cisatlantica is listed as threatened in Ohio by the Natural Heritage Program [85]. Chimaphila species are considered vulnerable in New York and may become rare, threatened, or endangered in the future if collection and/or development continues. They are protected under the 1974 New York State Wildflower Law [50].

DISTRIBUTION AND OCCURRENCE

SPECIES: Chimaphila umbellata
GENERAL DISTRIBUTION : Pipsissewa is a circumboreal species that is widely distributed in the northern hemisphere.  It is found from Newfoundland to Alaska south to California and Mexico, and east to New Mexico, Colorado, and South Dakota.  It is also found in the eastern United States from Maine south in the mountains to Georgia and west to Minnesota [25,44,63,67,81,82]. ECOSYSTEMS :    FRES10  White - red - jack pine    FRES11  Spruce - fir    FRES13  Loblolly - shortleaf pine    FRES15  Oak - hickory    FRES17  Elm - ash - cottonwood    FRES18  Maple - beech - birch    FRES19  Aspen - birch    FRES20  Douglas-fir    FRES21  Ponderosa pine    FRES22  Western white pine    FRES23  Fir - spruce    FRES24  Hemlock - Sitka spruce    FRES25  Larch    FRES26  Lodgepole pine    FRES27  Redwood    FRES28  Western hardwoods STATES :      AK  AZ  CA  CO  CT  DE  GA  ID  IL  IN      IA  KY  ME  MD  MA  MI  MN  MT  NV  NH      NJ  NM  NY  NC  OH  OR  PA  RI  SC  SD      TN  UT  VT  VA  WA  WV  WI  WY  AB  BC      MB  NB  NF  NT  NS  ON  PE  PQ  SK  YT      MEXICO BLM PHYSIOGRAPHIC REGIONS :     1  Northern Pacific Border     2  Cascade Mountains     3  Southern Pacific Border     4  Sierra Mountains     5  Columbia Plateau     6  Upper Basin and Range     7  Lower Basin and Range     8  Northern Rocky Mountains     9  Middle Rocky Mountains    10  Wyoming Basin    11  Southern Rocky Mountains    12  Colorado Plateau    13  Rocky Mountain Piedmont    15  Black Hills Uplift    16  Upper Missouri Basin and Broken Lands KUCHLER PLANT ASSOCIATIONS :    K001  Spruce - cedar - hemlock forest    K002  Cedar - hemlock - Douglas-fir forest    K003  Silver fir - Douglas-fir forest    K004  Fir - hemlock forest    K005  Mixed conifer forest    K006  Redwood forest    K007  Red fir forest    K008  Lodgepole pine - subalpine forest    K010  Ponderosa shrub forest    K011  Western ponderosa forest    K012  Douglas-fir forest    K013  Cedar - hemlock - pine forest    K014  Grand fir - Douglas-fir forest    K015  Western spruce - fir forest    K016  Eastern ponderosa forest    K017  Black Hills pine forest    K018  Pine - Douglas-fir forest    K019  Arizona pine forest    K020  Spruce - fir - Douglas-fir forest    K021  Southwestern spruce - fir forest    K026  Oregon oakwoods    K029  California mixed evergreen forest    K030  California oakwoods    K093  Great Lakes spruce - fir forest    K094  Conifer bog    K095  Great Lakes pine forest    K096  Northeastern spruce - fir forest    K097  Southeastern spruce - fir forest    K098  Northern floodplain forest    K099  Maple - basswood forest    K100  Oak - hickory forest    K102  Beech - maple forest    K103  Mixed mesophytic forest    K104  Appalachian oak forest    K106  Northern hardwoods    K107  Northern hardwoods - fir forest    K108  Northern hardwoods - spruce forest    K109  Transition between K104 and K106    K110  Northeastern oak - pine forest SAF COVER TYPES :      1  Jack pine      5  Balsam fir     12  Black spruce     13  Black spruce - tamarack     14  Northern pin oak     15  Red pine     16  Aspen     17  Pin cherry     18  Paper birch     19  Gray birch - red maple     20  White pine - northern red oak - red maple     21  Eastern white pine     22  White pine - hemlock     23  Eastern hemlock     24  Hemlock - yellow birch     25  Sugar maple - beech - yellow birch     26  Sugar maple - basswood     27  Sugar maple     30  Red spruce - yellow birch     31  Red spruce - sugar maple - beech     32  Red spruce     33  Red spruce - balsam fir     34  Red spruce - Fraser fir     35  Paper birch - red spruce - balsam fir     37  Northern white-cedar     38  Tamarack     42  Bur oak     45  Pitch pine     51  White pine - chestnut oak     52  White oak - black oak - northern red oak     53  White oak     55  Northern red oak     58  Yellow-poplar - eastern hemlock     59  Yellow-poplar - white oak - northern red oak     60  Beech - sugar maple     63  Cottonwood    107  White spruce    108  Red maple    110  Black oak    201  White spruce    202  White spruce - paper birch    203  Balsam poplar    204  Black spruce    205  Mountain hemlock    206  Engelmann spruce - subalpine fir    207  Red fir    208  Whitebark pine    210  Interior Douglas-fir    211  White fir    212  Western larch    213  Grand fir    215  Western white pine    217  Aspen    218  Lodgepole pine    222  Black cottonwood - willow    223  Sitka spruce    224  Western hemlock    225  Western hemlock - Sitka spruce    226  Coastal true fir - hemlock    227  Western redcedar - western hemlock    228  Western redcedar    229  Pacific Douglas-fir    230  Douglas-fir - western hemlock    231  Port-Orford-cedar    232  Redwood    233  Oregon white oak    234  Douglas-fir - tanoak - Pacific madrone    235  Cottonwood - willow    236  Bur oak    237  Interior ponderosa pine    243  Sierra Nevada mixed conifer    244  Pacific ponderosa pine - Douglas-fir    245  Pacific ponderosa pine    246  California black oak    249  Canyon live oak    251  White spruce - aspen    252  Paper birch    253  Black spruce - white spruce    254  Black spruce -  paper birch    256  California mixed subalpine SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Pipsissewa is a common understory species in many habitat types throughout its range but often does not reach dominance.  It is found in coniferous and mixed forests with numerous tree species.  In addition to those already mentioned, pipsissewa may occur with sugar pine (Pinus lambertiana), giant sequoia (Sequoia gigantea), and Brewer spruce (Picea breweriana) [6,65,79].  Several publications that list pipsissewa as a dominant understory species in the western United States follow. Description and classification of the forests of the upper Illinois   River drainage of southwestern Oregon [6] Preliminary plant associations of the Siskiyou Mountain Province [8] Preliminary plant associations of the southern Oregon Cascade Mountain   Province [7] Terrestrial vegetation of California [66] Natural vegetation of Oregon and Washington [21] Plant association and management guide: Willamette National Forest [37] Plant associations of south Chiloquin and Klamath Ranger   Districts--Winema National Forest [40] Vegetation and fire history of a ponderosa pine-white fir forest in    Crater Lake National Park [53] Associated species are well described for the Northwest and include baldhip rose (Rosa gymnocarpa), Nootka rose (R. nutkana), Greene mountain-ash (Sorbus scopulina), common snowberry (Symphoricarpos albus), Saskatoon serviceberry (Amelanchier alnifolia), Pacific yew (Taxus brevifolia), huckleberry (Vaccinium spp.), russet buffaloberry (Shepherdia canadensis), spiraea (Spiraea spp.), menziesia (Menziesia ferruginea), creeping juniper (Juniperus horizontalis), common juniper (J. communis), pachistima (Pachistima myrsinites), snowbrush ceanothus (Ceanothus velutinus), Utah honeysuckle (Lonicera utahensis), currant (Ribes spp.), raspberry (Rubus spp.), twinflower (Linnaea borealis), bunchberry (Cornus canadensis), beargrass (Xerophyllum tenax), Oregon-grape (Mahonia repens), queencup beadlily (Clintonia uniflora), violet (Viola spp.), strawberry (Fragaria spp.), sweet-scented bedstraw (Galium trifolium), pyrola (Pyrola spp.), oneleaf foamflower (Tiarella unifoliata), western rattlesnake plantain (Goodyera oblongifolia), pinegrass (Calamagrostis rubescens), and elk sedge (Carex geyeri) [1,7,14,28,37].

MANAGEMENT CONSIDERATIONS

SPECIES: Chimaphila umbellata
IMPORTANCE TO LIVESTOCK AND WILDLIFE : Pipsissewa is of minor importance in winter, spring, and fall diets of Roosevelt elk in the Pacific Northwest [43].  It is a component of white-tailed deer winter diets in the Swan Valley, Montana [56]. Mature stands of white fir (Abies concolor)-giant chinquapin (Chrysolepsis chrysophylla)/pachistima (Pachistima myrsinites)-prince's pine and Shasta red fir (Abies magnifica var. shastensis)-white fir-giant chinquapin/pipsissewa-long stolon sedge (Carex inops) plant associations in the Winema National Forest are critical elk calving and deer fawning habitat.  They are also important for feeding and nesting sites for birds and are suitable habitats for spotted owls, goshawks, and pileated woodpeckers [40].  White fir-Brewer spruce/pipsissewa plant associations in the southern Oregon Cascade Mountain Province are also excellent wildlife habitat [8]. PALATABILITY : NO-ENTRY NUTRITIONAL VALUE : NO-ENTRY COVER VALUE : NO-ENTRY VALUE FOR REHABILITATION OF DISTURBED SITES : Pipsissewa is sensitive to trampling and has a low potential for recovery.  It may, however, recover from very low (less than 40 passes per year) or low (75-100 passes per year) trampling intensities [13]. OTHER USES AND VALUES : Historically, pipsissewa roots and leaves were boiled and the infusion was ingested as a treatment for tuberculosis and long-lasting colds.  The leaves were also used as an astringent.  Pipsissewa can also be used as an ingredient in root beer [34,40].   OTHER MANAGEMENT CONSIDERATIONS : In a study of lodgepole pine stands in spruce (Picea engelmannii and P. glauca)/queencup beadlily (Clintonia uniflora) habitat types in Glacier National Park, pipsissewa displayed a significant (p<.10) decrease in frequency in response to a mountain pine beetle infestation and was more numerous on sites never infested than sites infested 80 years earlier. It was negatively correlated to overstory removal and increased light intensity, as shown by a steady decline in cover and frequency following the epidemic [3]. Pipsissewa is a major constituent of old-growth forests in the Swan Valley, Montana.  It often persists only on sheltered, unburned microsites.  It is present (39% frequency) in untreated old-growth and mature stands but is absent from burned clearcuts and plantation sites (20-30 years old).  Where standing trees remain to provide cover, frequency may be as high as 67 percent in stands that have been select cut without burning [22].  At other locations pipsissewa has essentially disappeared from stands or has had a major decrease in frequency or cover following stand removal with or without subsequent burning [4,5,14,74]. In the Vancouver Forest Region of British Columbia, pipsissewa is an indicator species in several variants of biogeoclimatic units for which guidelines for site diagnosis, tree species selection, and slash burning have been developed [26].  It is used as an indicator of good forest sites in the Winema and Fremont National Forests, Oregon.  When associated with twinflower, it is an indicator of the best fir (Abies spp.) sites [32,40].  The presence of pipsissewa is used to predict natural regeneration success under partially cut stands on the Dead Indian Plateau in southwest Oregon [55]. Pipsissewa is not a serious competitor to conifer seedlings [7].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Chimaphila umbellata
GENERAL BOTANICAL CHARACTERISTICS : Pipsissewa is a native evergreen low shrub or perennial rhizomatous herb.  The woody stems are usually 4 to 12 inches (10-30 cm) tall and the leathery, whorled leaves are sharply serrate.  The fertile stems are generally erect and may have 2 to 15 flowers.  Fruits are depressed, globose capsules which often persist through the winter [25,39,57,70,72,82]. Across its range, pipsissea was show wide variation in size; in leaf blade length, number of teeth, and prominence of lower surface venation; in sepal shape; and in stigma and capsule size [31]. RAUNKIAER LIFE FORM :    Chamaephyte    Geophyte REGENERATION PROCESSES : Pipsissewa reproduces both sexually and vegetatively.  Flowers have been observed being pollinated by bumblebees and staphylinid beetles [11]. Pipsissewa develops numerous, minute seeds [25,57,72].  Their dispersal mechanism has not been documented.  A New Brunswick study of boreal herb reproductive biology found that pipsissewa flowered for an average of 30 days.  Fruit set was low for flowers opening at the beginning of the flowering period.  Eighty-three percent of buds opened and 76.5 percent survived the flowering period.  Forty-seven percent of flower buds eventually developed fruit, and 45.6 percent actually matured fruit [36]. Pipsissewa produces long rhizomes that normally grow at a fast rate. Genets are generally long-lived [83].  Reports differ concerning the depth of pipsissewa rhizomes.  In a study of the Douglas-fir forest zone in southern interior British Columbia, McLean [52] listed prince's pine with species that have rhizomes growing from 2 to 5 inches (5-13 cm) below the mineral soil surface.  Most of those species are able to regenerate from those depths, but he stated that only pipsissewa rhizomes near the soil surface are able to produce new shoots.  Stickney [78] reported that in the northern Rocky Mountains, pipsissewa rhizomes are confined to the duff near or above the mineral soil surface. SITE CHARACTERISTICS : Pipsissewa is found in a wide variety of soils and soil moisture regimes.  It most commonly occurs in mixed woods and coniferous forests [17,24,35,38,44] on dry, well-drained, rocky or sandy soils [7,18,24,26].  In coastal regions of British Columbia, pipsissewa is an indicator of dry to very dry, nutrient-poor soils in montane boreal, temperate, and cool mesothermal climates.  Its occurrence decreases with increasing elevation and precipitation, and increases with continentality [47].  In Ontario, pipsissewa most often occurs on sandy or rocky soil on well-drained sites, on gravel terraces, and in jack pine (Pinus banksiana) barrens [70].  In red pine (P. resinosa)-white pine (P. strobus) forests of Voyageurs National Park, Minnesota, pipsissewa is found on dry, shallow, well-drained, nutrient poor to medium loamy sand to sandy loam soils [51].  In the nearby Boundary Waters Canoe area, pipsissewa is found on shallow, sandy soils to deep soils with a high clay content [59]. Pipsissewa also occurs in moist or imperfectly-drained situations throughout its range.  It is found on moist sites in oak ecosystems of Michigan [2], on moist sites in the Black Hills of South Dakota [72], and in lodgepole pine (P. contorta) forests in Alberta [9].  In the Adirondack Mountains of New York, pipsissewa occurs on well- to imperfectly-drained sites, most often under pines (Pinus spp.) on outwash soils, but also on tills in mixed woods [50]. Pipsissewa occurs in the following elevational ranges:                          feet              meters                _________________________________________________                UT     6,930-9,570     2,100-2,900 [82]                AZ     6,000-9,570     1,800-2,900 [31]                CA     1,000-9,570       300-2,900 [38,57]                CO     8,000-11,500    2,400-3,500 [35] SUCCESSIONAL STATUS : Pipsissewa has been classified as moderately shade tolerant to tolerant throughout its range [26,29,42,47,50,60].  Its highest frequency or cover is probably reached at intermediate light levels, such as in relatively open conifer stands in the Siskiyou Mountains of Oregon [15]. In the western Cascades, pipsissewa is significantly more frequent (p<.05) under a Douglas-fir (Pseudotsuga menziesii) canopy than under a western hemlock (Tsuga heterophylla) canopy (71% vs. 4% frequency). Average cover under western hemlock is less than 1.0 percent compared to 12.5 percent under Douglas-fir.  The difference may be due to less direct radiation in the western hemlock stands [73]. Pipsissewa is present throughout succession and occurs in stands of all ages [28,29,59,64,71].  It is found in relatively young stands [1,20,45], but is probably more frequent in mid-successional stages and mature forests [3,5,21,29].  Pipsissewa is a common understory component in many old-growth and climax forests of the Pacific Northwest [20,27,30,42]. SEASONAL DEVELOPMENT : Pipsissewa flowers from June to August throughout its range [17,24,25,31,57].

FIRE ECOLOGY

SPECIES: Chimaphila umbellata
FIRE ECOLOGY OR ADAPTATIONS : Pipsissewa is a fire-sensitive species that is very susceptible to damage and often shows a strong decline following fire [33,52,71,78]. Survival probably depends to a great extent on damage to rhizomes, so it depends on depth of rhizomes, fire severity, and consumption of duff [68,78].  Loss of the long-lived evergreen leaves may also reduce survival.  Postfire vegetative recovery depends primarily on the survival of scattered individuals in undisturbed microsites [33]. POSTFIRE REGENERATION STRATEGY :    Rhizomatous low woody plant, rhizome in organic mantle    Rhizomatous herb, rhizome in soil

FIRE EFFECTS

SPECIES: Chimaphila umbellata
IMMEDIATE FIRE EFFECT ON PLANT : Pipsissewa has a moderate to high probability of being killed by fire [40,76].  Low-severity fires that do not consume the organic mantle may only top-kill it. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Postfire response of pipsissewa is variable and is probably most dependent on fire severity and the uniformity of the burn.  Some studies have reported pipsissewa surviving fire.  In mixed western hemlock-Douglas-fir-western redcedar (Thuja plicata) stands in North Cascades National Park, Washington, pipsissewa was considered a residual species following a July wildfire.  Its frequency in postfire years 1, 2, and 3 was 65.3, 52.1, and 52.1 percent, respectively [54]. Pipsissewa appeared to survive on moderately burned sites following the Waterfalls Canyon Fire in Grand Teton National Park in July, 1974, but was eliminated from severely burned sites.  The prefire vegetation was spruce-fir with lodgepole pine (Pinus contorta) and whitebark pine (P. albicaulis).  Pipsissewa had the following percent frequency and cover as measured in 1975 [10]:                          Frequency     Cover ______________________________________________ Unburned sites              52           5 Sites burned in 1932         2         trace Moderately burned sites     17           1 Severely burned sites        0           0  In the northern Rocky Mountains, slow recovery after fire has been reported.  Pipsissewa was eliminated from initial postfire communities by a severe wildfire in western larch (Larix occidentalis)-Douglas-fir stands on the Flathead National Forest, Montana [75].  In western larch-fir (Abies grandis and A. lasiocarpa) stands on the Flathead and Lolo National Forests, Montana, pipsissewa had not recovered by postfire year 9 following logging and broadcast burning [74].  Pipsissewa was also absent 10 months after a late-summer wildfire in lodgepole pine stands in the Chamberlain Basin, Idaho.  It was found on adjacent unburned sites and was present on burned sites 5 years after the fire, but had less biomass production than on unburned sites [61]. Variable responses to fire have been reported for pipsissewa in Minnesota.  It survived the Little Sioux Wildfire in May, 1971, in mixed conifer-hardwood stands in northeastern Minnesota.  Number of individuals (on seventy 0.605 sq m plots) and aboveground average dry weight per individual pipsissewa were measured at the end of each growing season for the first 5 postfire years [58]:                        1971     1972      1973     1974     1975 ________________________________________________________________ No. of individuals      15     no data     57       30        7 Ave. dry wt. (g)       .07     no data    .33      .29      .46 Pipsissewa responded more slowly after wildfires in second-growth mixed conifer-hardwood forests in northeastern Minnesota.  It was not present in postfire years 3, 5, or 14 after the April Heartlake Fire. It was not present on the Kelley Creek Burn, resulting from a July fire, at postfire year 2 but had a frequency of 3 percent in postfire years 5 and 11 [48]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : The Research Project Summary Vegetation response to restoration treatments in ponderosa pine-Douglas-fir forests of western Montana provides
information on prescribed fire and postfire response of plant community species, including pipsissewa, that was not available when this species review was written. FIRE MANAGEMENT CONSIDERATIONS : Pipsissewa is a component in many subzones in which guidelines for prescribed burning and tree species selection have been developed in the Vancouver Forest District, British Columbia [46].

REFERENCES

SPECIES: Chimaphila umbellata
REFERENCES :  1.  Antos, J. A.; Habeck, J. R. 1981. Successional development in Abies        grandis (Dougl.) Forbes forests in the Swan Valley, western Montana.        Northwest Science. 55(1): 26-39.  [12445]  2.  Archambault, Louis; Barnes, Burton V.; Witter, John A. 1989. Ecological        species groups of oak ecosystems of southeastern Michigan. Forest        Science. 35(4): 1058-1074.  [9768]  3.  Armour, Charles David. 1982. Fuel and vegetation succession in response        to mountain pine beetle epidemics in northwestern Montana. Moscow, ID:        University of Idaho. 47 p. Thesis.  [16488]  4.  Arno, Stephen F.; Simmerman, Dennis G. 1982. Succession after cutting        and fire treatments on forest habitat types in western Montana. In:        Baumgartner, David M., compiler. Site preparation and fuels management        on steep terrain: Proceedings of a symposium; 1982 February 15-17;        Spokane, WA. Pullman, WA: Washington State University, Cooperative        Extension: 113-117.  [18537]  5.  Arno, Stephen F.; Simmerman, Dennis G.; Keane, Robert E. 1985. Forest        succession on four habitat types in western Montana. Gen. Tech. Rep.        INT-177. Ogden, UT: U.S. Department of Agriculture, Forest Service,        Intermountain Forest and Range Experiment Station. 74 p.  [349]  6.  Atzet, Thomas. 1979. Description and classification of the forests of        the upper Illinois River drainage of southwestern Oregon. Corvallis, OR:        Oregon State University. 211 p. Dissertation.  [6452]  7.  Atzet, Thomas; McCrimmon, Lisa A. 1990. Preliminary plant associations        of the southern Oregon Cascade Mountain Province. Grants Pass, OR: U.S.        Department of Agriculture, Forest Service, Siskiyou National Forest. 330        p.  [12977]  8.  Atzet, Thomas; Wheeler, David L. 1984. Preliminary plant associations of        the Siskiyou Mountain Province. Portland, OR: U.S. Department of        Agriculture, Forest Service, Pacific Northwest Region. 278 p.  [9351]  9.  Baranyay, J. A.; Safranyik, L. 1970. Effect of dwarf mistletoe on growth        and mortality of lodgepole pine in Alberta. Publ. No. 1285. Ottawa:        Canadian Forestry Service, Department of Fisheries and Forestry. 19 p.        [8286] 10.  Barmore, William J., Jr.; Taylor, Dale; Hayden, Peter. 1976. Ecological        effects and biotic succession following the 1974 Waterfalls Canyon Fire        in Grand Teton National Park. Research Progress Report 1974-1975.        Unpublished report on file at: U.S. Department of Agriculture, Forest        Service, Intermountain Fire Sciences Laboratory, Missoula, MT. 99 p.        [16109] 11.  Barrett, Spencer C.; Helenurm, Kaius. 1987. The reproductive biology of        boreal forest herbs. I. Breeding systems and pollination. Canadian        Journal of Botany. 65: 2036-2046.  [6624] 12.  Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals,        reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's        associations for the eleven western states. Tech. Note 301. Denver, CO:        U.S. Department of the Interior, Bureau of Land Management. 169 p.        [434] 13.  Cole, David N. 1988. Disturbance and recovery of trampled montane        grassland and forests in Montana. Res. Pap. INT-389. Ogden, UT: U.S.        Department of Agriculture, Forest Service, Intermountain Research        Station. 37 p.  [3622] 14.  Edgerton, Paul J. 1987. Influence of ungulates on the development of the        shrub understory of an upper slope mixed conifer forest. In: Provenza,        Frederick D.; Flinders, Jerran T.; McArthur, E. Durant, compilers.        Proceedings--symposium on plant-herbivore interactions; 1985 August 7-9;        Snowbird, UT. Gen. Tech. Rep. INT-222. Ogden, UT: U.S. Department of        Agriculture, Forest Service, Intermountain Research Station: 162-167.        [7411] 15.  Emmingham, W. H. 1972. Conifer growth and plant distribution under        different light environments in the Siskiyou Mountains of southwestern        Oregon. Corvallis, OR: Oregon State University. 50 p. Thesis.  [9651] 16.  Eyre, F. H., ed. 1980. Forest cover types of the United States and        Canada. Washington, DC: Society of American Foresters. 148 p.  [905] 17.  Fernald, Merritt Lyndon. 1950. Gray's manual of botany. [Corrections        supplied by R. C. Rollins]. Portland, OR: Dioscorides Press. 1632 p.        (Dudley, Theodore R., gen. ed.; Biosystematics, Floristic & Phylogeny        Series; vol. 2).  [14935] 18.  Filip, Stanley M.; Little, Elbert L., Jr. 1971. Trees and shrubs of the        Bartlett Experimental Forest, Carroll County, New Hampshire. Res. Pap.        NE-211. 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