Red fruit bearberry is a frequent shrub associate in boreal and subalpine spruce (Picea spp.) woodlands and forests. It is not usually a dominant understory species in spruce ecosystems [2,6,15,87,95]. Peinado and others  describe red fruit bearberry as "characteristic" of boreal spruce forests. Studies in white spruce (P. glauca) stands across boreal North America found red fruit bearberry had mean cover of <2%. It was not among the 15 most common species in the dwarf shrub-herb stratum . It also occurs in boreal mixed spruce-tamarack (Larix laricina) associations . Red fruit bearberry abundance in northern ecosystems may increase to the west. It is a common associate in white spruce taiga of Alaska and western Canada  and is sometimes "abundant" in black spruce (P. mariana) taiga . Red fruit bearberry was dominant in a subarctic black spruce/littletree willow-bog Labrador tea (Salix arbusculoides-Ledum groenlandicum)-red fruit bearberry/splendid feather moss (Hylocomium splendens) community near Fort Worth, Northwest Territories . In a vegetation survey along the Alaska Highway in Yukon, however, red fruit bearberry was "characteristic" but not dominant in wet spruce forests .
Red fruit bearberry grows in Rocky Mountain lodgepole pine (Pinus contorta var. latifolia) forests of Banff and Jasper National Parks, Alberta , in Engelmann spruce (Picea engelmannii) forests of British Columbia [35,85], and near white spruce swamps in Wyoming .
Red fruit bearberry occurs in and may dominate the understory of balsam poplar (Populus balsamifera subsp. balsamifera) floodplains in northern Alaska (Hettinger and Janz 1974, cited in ).
Red fruit bearberry occurs in and may dominate tundra dwarf heath and other tundra shrub communities of Alaska and northwestern Canada . Dwarf shrub communities typically occupy only a few hundred square meters and are surrounded by other tundra vegetation types, usually spruce woodlands and forests . Red fruit bearberry grows in willow/water sedge (Salix myrtillifolia and/or S. planifolia/Carex aquatilis) shrublands along the Alaskan Highway in Yukon  and in grayleaf willow (S. glauca) communities of the coastal plain of central arctic Alaska  and southwestern Yukon . Red fruit bearberry is codominant in low shrub bog birch-red fruit bearberry/snow lichen (Cetraria nivalis) tundra in Mackenzie Valley, Northwest Territories .
Red fruit bearberry is commonly found in mountain-avens (Dryas spp.) communities of alpine and arctic Alaska . In arctic regions, red fruit bearberry occurs or dominates in arctic mountain-avens (Dryas integrifolia) cushion-plant tundra [42,94]. Alaska willow (S. alaxensis) and/or russet buffaloberry (Shepherdia canadensis) are common codominants or associates .
The following vegetation classifications describe plant communities in which red fruit bearberry is a dominant species.Alaska
Red fruit bearberry is a prostrate, dwarf shrub , about 1 to 6 inches (3-15 cm) tall [1,87]. Leaves are considered deciduous [14,74] or, more accurately, marcescent, meaning they tend to wither and whiten with age but persist on the stem [51,74]. Leaves are 0.8 to 2.4 inches (2-6 cm) long . The inflorescence is a glandular, hairy raceme . Flowers are arranged in small clusters of 1 to 6 ; they are about 0.25 inch (6 mm) long . The fruit is a drupe . Red fruit bearberry forms trailing mats  via spread from stolons  or rhizomes [34,71]. Lateral stem habit is not well distinguished for red fruit bearberry. It is possible that lateral stems of red fruit bearberry grow both above ground (stolons) and below ground (rhizomes), depending upon topography, soil movement, and/or genetic make-up of individual plants. Further studies are needed on stem habit of red fruit bearberry.
Red fruit bearberry is described as "long lived" .RAUNKIAER  LIFE FORM:
Seed dispersal: Bears, other mammals, and birds eat red fruit bearberry fruits (see Importance to Wildlife and Livestock); presumably, these frugivorous animals disperse the seeds in their droppings.
Seed banking: Although information specific to red fruit bearberry was lacking as of 2008, bearberries and manzanitas (Arctostaphylos spp.) generally maintain a soil seed bank . In Alaska, red fruit bearberry seedlings established from planted seed 1 or 2 years after sowing , suggesting that red fruit bearberry can establish from soil-stored seed.
Germination: Red fruit bearberry embryos are dormant. Scarification of the seedcoat and/or overwinter cold stratification are required to break dormancy [13,90]. In the laboratory, stratification for 60 to 90 days at 36 to 41 °F (2-5 °C) broke dormancy . Seeds can germinate at low temperatures (41-77 °F (2-25 °C)) [13,90]. Light enhanced germination in an Alaskan field experiment . Fire scarification typically breaks dormancy of bearberries and manzanitas ; however, no information was available on the effects of fire on red fruit bearberry seeds as of 2008.
Seedling establishment/growth: Red fruit bearberry established from seed in an area where a debris pile had been removed from an arctic coastal plain in Alaska. Four years after removal, red fruit bearberry had trace coverage and 1% frequency .
Vegetative regeneration: Red fruit bearberry sprouts [8,24,71] from stolons  or rhizomes  following top-kill. Sprout origin is not well described in much of the literature describing red fruit bearberry sprouting. Further studies are needed on red fruit bearberry's ability to regenerate vegetatively.SITE CHARACTERISTICS:
Soils: Red fruit bearberry prefers moist to wet soils [30,59,76], (review by ), although drainage may vary from good to poor , (review by ). On sites in northeastern British Columbia and southeastern Yukon, red fruit bearberry was most common on wet sites with a black spruce overstory, while prickly rose and kinnikinnick (Arctostaphylos uva-ursi) were more common on dry to moist sites with a Rocky Mountain lodgepole pine-black spruce overstory . On lowlands on the western shore of the Hudson Bay, red fruit bearberry had greatest coverage (10%) on a poorly drained site with permafrost 35 inches (90 cm) beneath the soil surface .
Red fruit bearberry typically grows in loams, sands, or gravels [30,90] of variable pH [30,59,75]. A permafrost layer underlies the soil on many red fruit bearberry sites . On the Tetlin National Wildlife Refuge in southeastern Alaska, red fruit bearberry grows in a 6- to 18-inch (15-45 cm)- deep organic soil layer that overlies alluvial mineral soil. Permafrost occurs about 12 inches (30 cm) below the organic soil layer . In Yukon, red fruit bearberry is typical on low-lying terrain with underlying permafrost. Soil texture is variable, with the organic layer at a depth of 16 inches (40 cm) or more . Red fruit bearberry-sheathed cottonsedge-northern Labrador tea communities in Yukon generally occur on sandy clay with stones, with an organic horizon up to 4 inches (10 cm) thick. The permafrost layer is 15 to 22 inches (39-55 cm) deep. Soil pH ranges from 3.4 to 3.9. White spruce/red fruit bearberry communities in Yukon often occur on sands, silts, and/or gravels with organic layers 2 to 18 inches (4-45 cm) deep, over permafrost. Soil pH ranges from 5.7 to 7.0 and slopes are moderate . White spruce/mountain alder-red fruit bearberry floodplain communities in the Mackenzie River Delta region have a soil pH range of 8.0 to 8.2 . In Ontario, red fruit bearberry occurs on sandy and gravelly beach ridges, calcareous till, and clay riverbanks . In the Swamp Lake Botanic Area of Wyoming, red fruit bearberry occurs on moist, calcareous sites bordering white spruce swamps .
In an analysis of 95 sites across northwestern and north-central Canada, red fruit bearberry occurred most often on calcareous, fine-textured loams in low arctic and high subarctic tundra shrublands in northwestern Canada. Moisture regime was borderline between wet-mesic and mesic .
Topography and elevation: Red fruit bearberry often occurs on gentle terrain [30,49,59,75], although slopes in alpine locations may be steep. Red fruit bearberry typically grows near treeline . In a floristic study of arctic and alpine plants of North America, Raup  usually found red fruit bearberry at timberline. Because the elevation of treeline increases with decreasing latitude, red fruit bearberry occurs at the highest elevations in its southern distribution, while it occurs at low elevations in Alaska . In boreal locations, global climate change is apparently resulting in upward elevational and latitudinal expansions of red fruit bearberry and other treeline species [79,86].
|Elevations reported for red fruit bearberry, from south to north|
|Swamp Lake Botanic Area, Wyoming||6,600 feet (2,000 m) |
|British Columbia||overall elevational range not available|
|Mt Robson||5, 450 (1,660 m) |
|Kootenay National Park||4,890 feet (1,490 m) |
|Yukon||around 2,000 feet (600 m) elevation |
|Mackenzie Valley||at treeline, 2,000 feet (600 m) |
|Doll Creek area||2,030 feet (620 m) |
Climate: Red fruit bearberry occurs in areas with cold temperate , boreal , and arctic  climates. It is rated moderately tolerant  to intolerant  of drought, with very high winter hardiness .SUCCESSIONAL STATUS:
Red fruit bearberry pioneers on gravel floodplains (review by ) and other bare substrates in primary succession. It colonizes bare mineral soil in tundra and taiga sites in Alaska . Red fruit bearberry colonized bare, unstable dunelands along the Meade River near Barrow, but was not dominant on stable dunes . However, eightpetal mountain-avens and red fruit bearberry formed a prostrate shrub community on stable dunes along the Atigun River of Alaska's North Slope .
Flooding disturbance may favor red fruit bearberry. On the Mackenzie Delta, red fruit bearberry cover apparently increased following spring ice breakup in 1982. The plant community, a white spruce/mountain alder-red fruit bearberry association, experiences flooding approximately once a decade. Red fruit bearberry had 10% cover and 100% frequency 4 years after flooding; its preflood abundance was not described . In a russet buffaloberry-diamondleaf willow-prickly rose (Salix pulchra-Rosa acicularis) floodplain community in the central Alaskan Range, red fruit bearberry occurred on open sites in the mostly dense shrubfield. The area, on the Riley Creek floodplain, was in primary and secondary succession. The lower floodplain was scoured by an ice jam in December 1995, and several centimeters of sand were deposited during flooding and spring ice breakup in 1996. Areas left bare by heavy ice scouring, and therefore forced back to primary succession, were colonized by alpine snow lichen (Stereocaulon alpinum). The area with red fruit bearberry also supported kinnikinnick and common juniper (Juniperus communis) . Cooccurrence of common juniper, which does not sprout, suggests that the floodplain area with red fruit bearberry was only lightly scoured and in early secondary succession. On upland sites that were not scoured or flooded recently, red fruit bearberry occurred in the understory of an open balsam poplar-white spruce stand that apparently established after a fire 71 years prior to the study .
As a species associated with treeline, red fruit bearberry often grows on the edges of receding glaciers in early primary succession. In a study of succession after glacial scouring in the northern Rocky Mountains, Heusser  found forbs generally colonized the scoured area, followed by low shrubs. Red fruit bearberry and eightpetal mountain-avens often dominated the low shrub stage. Shrubs and tree seedlings established on recessional moraines 10 to 60 years after glacial diminution; typically, these species include willows, bog birch, and spruces. The final stage was mature spruce forest, which developed around 100 years after spruce seedling establishment. On Mt Robson in British Columbia, red fruit bearberry followed the forbs northern sweetvetch (Hedsarium boreal subsp. mackenzie) and eightpetal mountain-avens after recession of Robson Glacier, codominating the low shrub stage with willows. At 12,972 feet (3,954 m), Mt Robson is the highest mountain in the Canadian Rocky Mountains, and Robson Glacier is the largest glacier on the mountain [82,83].
|Mean cover and frequency of red fruit bearberry on different-aged recessional moraines on Mt Robson, British Columbia [82,83]|
|50-year-old moraine||73-year-old moraine||160-year-old moraine|
Red fruit bearberry persists on Mt Robson in mature (135-year-old) Engelmann spruce communities [35,85]. In a chronosequence study of primary succession on recessional moraines of the Robson Glacier, Blundon and Dale  found red fruit bearberry dominated the 3rd and last phase of successional development. A timeline for this development was not given.
|Primary succession on the Robinson Glacier, British Columbia |
|Phase 1||rock willow-grayleaf willow/Drummond mountain avens (Salix vestita-S. glauca/Dryas drummondi)/northern sweetvetch|
|Phase 2||willow/Drummond mountain avens/northern sweetvetch|
|Phase 3||Engelmann spruce/red fruit bearberry-northern sweetvetch/brachythecium moss-reindeer lichens (Brachythecium spp.-Cladonia spp.)|
Red fruit bearberry also occurs on disturbed sites in secondary succession. Canopy closure takes about 100 years on northern spruce ecosystems [41,52,53]. As of 2008, the only successional studies conducted in spruce forests with red fruit bearberry were short-term studies spanning no more than 10 years [19,34,93], so the ability of red fruit bearberry to persist in later stages of succession is unknown.
A year after blading, red fruit bearberry established from surviving "rootstalks" on a bare bulldozer track in the Mackenzie Delta . In a study of succession 48 years following construction of the CANOL Pipeline in the Northwest Territories, red fruit bearberry grew in vehicle tracks, although it was at least twice as frequent on undisturbed plots compared to plots in vehicle tracks. Red fruit bearberry was not found on plots in borrow pits . Red fruit bearberry dominated bulldozer lines 2 and 3 years following bulldozer blading in the Mackenzie Delta, recolonizing from intact rhizomes below the bulldozer line. On bulldozer lines in a littletree willow (Salix arbusculoides) community, red fruit bearberry cover was greater on bulldozed edges than on bulldozed line centers or control plots 2 years after blading. Where construction ran through a nearby white spruce/mountain alder community, red fruit bearberry approximated predisturbance cover 3 years after blading .
|Changes in mean red fruit bearberry cover (SD) on bulldozer lines following bulldozing in winter 1969 in the Mackenzie Delta, Northwest Territories |
|Littletree willow community|
|July 1970||July 1971|
|Control||12.0 (1.6)||11.1 (1.9)|
|Bulldozed, on line edge||20.0 (2.6)||18.1 (2.2)|
|Bulldozed, in line center||0.3 (0.1)||0.9 (0.7)|
|White spruce/mountain alder community|
|July 1970||July 1971||July 1972|
|Control||18.3 (2.0)||19.8 (2.0)||16.0 (1.6)|
|Bulldozed||0.9 (0.6)||7.8 (1.2)||17.1 (1.3)|
Few studies had been conducted on red fruit bearberry's response to logging as of 2008. In the short term, logging treatments in an open white spruce/thinleaf alder/splendid feather moss (Alnus incana subsp. tenuifolia/Hylocomium splendens) community slightly reduced red fruit bearberry abundance on Willow Island near Fairbanks, Alaska :
|Mean cover and frequency of red fruit bearberry before and after logging treatments in a white spruce forest on Willow Island, Alaska |
|Posttreatment year||Cover (%)||Frequency (%)|
|Shelterwood (9-m spacing)|
Red fruit bearberry showed no change in cover 10 years after tree harvest or fire compared to an untreated control in a white spruce-paper birch-quaking aspen forest near Fairbanks, Alaska. Red fruit bearberry had 5% cover on clearcut, shelterwood, burned, and undisturbed control plots at postlogging year 10 .SEASONAL DEVELOPMENT:
|Phenological development of red fruit bearberry in various locations|
fruits July-September 
fruits ripe July-August 
|Sheep Mountain, Yukon||flowered from 11 to 20 May over 3 years |
Fire regimes: The boreal spruce, tundra, and alpine communities in which red fruit bearberry grows generally experience long intervals between fires. Red fruit bearberry is most common in early postfire succession within these communities (see Plant Response to Fire).
Northern conifer ecosystems: Fires in spruce forests with red fruit bearberry generally occur more than 100 years apart, although shorter fire-return intervals also occur. Boreal black spruce forests have fire-return intervals ranging from about 50 to 100 years [41,53]. A black spruce-jack pine forest in northern boreal Quebec, for example, experienced fire at intervals of 47 to 270 years . Northern spruce forests experience a combination of crown and ground fires [41,53]. Fires in spruce taiga may last for months, burning thousands to millions of acres .
White spruce forests of Kluane National Park, Yukon, have 150- to 400-year fire-return intervals. Mean fire-return interval increases with increasing latitude, from 133 years in the southern end of the Park, to 164 years in the middle of the Park, and 234 years in the northern end of the Park . Driscoll and others  reported a return interval of 140+ years for fires in hybrid spruce (Picea glauca × P. engelmannii) forests of central British Columbia.
Western conifer ecosystems: As of 2008, studies on red fruit bearberry's postfire occurrence in coniferous forests in western North America were lacking, so it is difficult to project what role fire plays in red fruit bearberry abundance in those forests. Red fruit bearberry grows in white spruce forests of Wyoming and Engelmann spruce forests of British Columbia, which have relatively long fire-free intervals. Arno  estimated fire-return intervals for fir-spruce (Abies-Picea spp.) forests of the northern Rocky Mountains at 150+ years. Intervals of 100 to 250 years between fires are characteristic of Rocky Mountain lodgepole pine in the northern Rockies [4,5]; however, mean fire intervals may be as short as 20 to 50 years in small stands .
As of 2008, no information was available on fire regimes in shrub tundra and alpine ecosystems with red fruit bearberry. Tundra fires are usually small and stand replacing. In Alaska and western Canada, tundra fires typically burn only 2.5 to 25 acres (1-10 ha). Some tundra fires, however, have burned up to 247,000 acres (100,000 ha) . Alpine ecosystems with red fruit bearberry probably burn infrequently due to sparse fuels.
The following table provides fire regime information that may be relevant to red fruit bearberry. 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 red fruit bearberry occurs. For each community, fire regime characteristics are taken from the LANDFIRE Rapid Assessment Vegetation Models . These vegetation models were developed by local experts using available literature, local data, and/or expert opinion as documented in the PDF file linked from the name of each Potential Natural Vegetation Group listed below. Cells are blank where information is not available in the Rapid Assessment Vegetation Model.|
|Northern and Central Rockies|
|Vegetation Community (Potential Natural Vegetation Group)||Fire severity*||Fire regime characteristics|
|Percent of fires||Mean interval
|Northern and Central Rockies Forested|
|Persistent lodgepole pine||Replacement||89%||450||300||600|
|Whitebark pine-lodgepole pine (upper subalpine, Northern and Central Rockies)||Replacement||38%||360|
|Vegetation Community (Potential Natural Vegetation Group)||Fire severity*||Fire regime characteristics|
|Percent of fires||Mean interval
|Northeast spruce-fir forest||Replacement||100%||265||150||300|
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 [27,47].
fruit bearberry's sprouting response is best documented in spruce ecosystems,
where limited studies suggest that it may gain cover slowly
after fire. Origin of postfire sprouts (rhizomes and/or stolons) had not been determined as of 2008.
A white birch-black spruce-tundra ecotone on the Mackenzie Delta was burned under
prescription on 3 July 1973. Red fruit bearberry sprouts were observed on study plots on 1 August
1973, following observations of earlier-sprouting cloudberry
(Rubus chamaemorus) and bluejoint grass (Calamagrostis
canadensis) plants . Red fruit bearberry also sprouted after the
stand-replacing Porcupine Wildfire in northeastern Alaska, gaining
prefire coverage slowly. Its cover on 23-year-old burn plots was less than half
that of that on control plots in adjacent, unburned black spruce forest :
|Red fruit bearberry abundance following the 1950 Porcupine Fire in northeastern Alaska |
|Year||Cover (%)||Frequency (%)|
|1951 (postfire year 1)||0.4||18|
|1954 (postfire year 4)||0.9||9.0|
|1957 (postfire year 7)||1.9||12.3|
|1961 (postfire year 11)||2.2||12.8|
|1973 (postfire year 23)||2.6||5.5|
|1981 (postfire year 31)||4.3||29.3|
In spruce forests of the Mackenzie Valley, red fruit bearberry had low cover in both young and old burns .
|Mean cover and frequency of red fruit bearberry on different-aged burns in the Northwest Territories |
|Habitat type||Burn age (years)||Cover (%)||Frequency (%)|
|black spruce/mosses||4||trace||no data|
|white spruce/splendid feather moss||30||2||no data|
|black spruce/bog Labrador tea/splendid feather moss||old growth; stand age not determined||0.5||30|
Red fruit bearberry recovered more quickly from fire on a black spruce-tundra ecotone site than on a tundra site .
|Mean cover (%) of red fruit bearberry before and after a 8-18 August 1968 wildfire near Inuvik, Northwest Territories |
|Year||Forest-tundra ecotone (n=9)||Tundra
|1973 (unburned control)||0.6 (0.6)*||4.5 (1.6)|
|1973 (postfire year 5)||trace||1.9 (0.7)|
|1990 (postfire year 22)||2.0 (2.0)||1.9 (0.7)|
Prefire brush clearing resulted in slow red fruit bearberry recovery compared to sites that were undisturbed before fire. On a subarctic black spruce woodland in the Northwest Territories, red fruit bearberry frequency was inventoried 8 years before a wildfire and in postfire year 3. Postfire red fruit bearberry frequency was least on right-of-way sites that were cleared 10 years before the wildfire .
|Mean frequency (%) of red fruit bearberry before and after a 6 June 1995 wildfire near Tulita, Northwest Territories |
|Burned right-of-way||Burned forest|
|1987 (prefire year 8)||94.7||90.0|
|1997 (postfire year 3)||44.7||73.7|
Prescribed burning may have promoted red fruit bearberry seedling establishment in the Kenai Mountains of Alaska. Burning was conducted on 13 May 1983 in a Lutz's spruce/Barclay's willow (Picea × lutzii/Salix barclayi) forest to promote growth of moose browse. Red fruit bearberry was not present on transects before burning and had 1% cover in postfire year 15. The author found that postfire cover of browse species was generally greatest on sites where palatable shrubs were present before fire, while sites dominated by bluejoint grass had lowest cover of browse species after fire .
A wildfire on the Seward Peninsula of Alaska apparently had little effect on red fruit bearberry. The 1977 wildfire burned through previously-sampled plots in tussock-shrub tundra dominated by sheathed cottonsedge, Bigelow's sedge (Carex bigelowii), bog birch, and bog blueberry (Vaccinium uliginosum). Red fruit bearberry frequency was about 10% higher in postfire year 24 than before the fire, but its cover remained low and little changed .
|Mean cover and frequency of red fruit bearberry after a 1977 summer wildfire on a tussock-shrub tundra hillslope in the central Steward Peninsula, Alaska |
|Year||Cover (%)||Frequency (%)|
|1973 (prefire; averaged across the hillslope)||1||20|
|1978 (postfire year 1)||1||20|
|1980 (postfire year 3)||1||20|
|2001 (postfire year 24)||1||30|
|1978 (postfire year 1)||trace||10|
|1980 (postfire year 3)||trace||30|
|2001 (postfire year 24)||trace||30|
|1978 (postfire year 1)||trace||20|
|1980 (postfire year 3)||1||40|
|2001 (postfire year 24)||1||40|
Similarly, red fruit bearberry's frequency increased with time since fire, while its cover decreased, on study sites in interior Alaska :
|Mean cover and frequency of red fruit bearberry on burned paper birch-black spruce forests in the Kobuk River valley of interior Alaska |
|Years since fire||Cover (%)||Frequency (%)|
DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
No additional information is available on this topic.
FIRE MANAGEMENT CONSIDERATIONS:
Sparse data make it impossible to recommend fire management practices for red fruit bearberry. Since red fruit bearberry is a low-growing, early-successional plant (see Successional Status), it is likely that red fruit bearberry declines with canopy closure after fire in coniferous forests and tundra shrublands. Further studies are needed on the fire ecology of red fruit bearberry.
Palatability/nutritional value: Hardy  rated red fruit bearberry browse as moderately palatable and moderately tolerant of browsing pressure.
Cover value: Red fruit bearberry is an "especially important" cover and forage species for meadow voles in Fort Churchill, Manitoba .
Woodland caribou in the Mackenzie and Selwyn mountains of Yukon and the Northwest Territories use snowfields within arctic mountain-avens-red fruit bearberry tundra cushion plant communities as relief habitat to escape insect harassment .VALUE FOR REHABILITATION OF DISTURBED SITES:
Red fruit bearberry plants can be obtained from stem cuttings. Propagation from seed may be a "lengthy and difficult process" . In an experimental restoration in the Northwest Territories, red fruit bearberry failed to establish from locally-collected seed broadcast in a trench [21,55]. However, shrub establishment was monitored for only 1 year after sowing, so it is possible that the seed was still dormant and established in later years . Red fruit bearberry-dominated sites along the Alaskan Highway in Yukon are rated "relatively insensitive" to "highly sensitive" to disturbance, depending on soil type. Red fruit bearberry vegetation types on sand or gravel are rated relatively insensitive, while those on peat or other organic soils are rated highly sensitive .OTHER USES:
1. Anderson, J. P. 1959. Flora of Alaska and adjacent parts of Canada. Ames, IA: Iowa State University Press. 543 p. 
2. Arii, Ken; Turkington, Roy. 2002. Do nutrient availability and competition limit plant growth of herbaceous species in the boreal forest understory? Arctic, Antarctic, and Alpine Research. 34(3): 251-261. 
3. Arno, Stephen F. 1980. Forest fire history in the Northern Rockies. Journal of Forestry. 78(8): 460-465. 
4. Barrett, Stephen W.; Arno, Stephen F. 1990. Fire history of the Lamar River drainage, Yellowstone National Park. In: Boyce, Mark S.; Plumb, Glenn E., eds. National Park Service Research Center, 14th annual report. Laramie, WY: University of Wyoming, National Park Service Research Center: 131-133. 
5. Barrett, Stephen W.; Arno, Stephen F.; Menakis, James P. 1997. Fire episodes in the Inland Northwest (1540-1940) based on fire history data. Gen. Tech. Rep. INT-GTR-370. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 17 p. 
6. Birks, H. J. B. 1980. Modern pollen assemblages and vegetational history of the moraines of the Klutlan Glacier and its surroundings, Yukon Territory, Canada. Quaternary Research. 14(1): 101-129. 
7. Bliss, L. C. 1988. Arctic tundra and polar desert biome. In: Barbour, Michael G.; Billings, William Dwight, eds. North American terrestrial vegetation. Cambridge, New York: Cambridge University Press: 1-32. 
8. Bliss, L. C.; Wein, R. W. 1972. Plant community responses to disturbances in the western Canadian Arctic. Canadian Journal of Botany. 50: 1097-1109. 
9. Blundon, D. J.; Dale, M. R. T. 1990. Dinitrogen fixation (Acetylene reduction) in primary succession near Mount Robson, British Columbia, Canada. Arctic and Alpine Research. 22(3): 255-263. 
10. Bockheim, J. G.; O'Brien, J. D.; Munroe, J. S.; Hinkel, K. M. 2003. Factors affecting the distribution of Populus balsamifera on the North Slope of Alaska. Arctic, Antarctic, and Alpine Research. 35(3): 331-340. 
11. Boucher, Tina V. 2003. Vegetation response to prescribed fire in the Kenai Mountains, Alaska. Res. Pap. PNW-RP-554. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 59 p. 
12. Dale, M. R. T.; John, E. A.; Blundon, D. J. 1991. Contact sampling for the detection of interspecific association: a comparison in two vegetation types. Journal of Ecology. 79(3): 781-792. 
13. Densmore, Roseann Van Essen. 1979. Aspects of the seed ecology of woody plants of the Alaskan taiga and tundra. Durham, NC: Duke University. 285 p. Dissertation. 
14. Dorn, Robert D. 1988. Vascular plants of Wyoming. Cheyenne, WY: Mountain West Publishing. 340 p. 
15. Douglas, George W. 1974. Montane zone vegetation of the Alsek River region, southwestern Yukon. Canadian Journal of Botany. 52: 2505-2532. 
16. Doutt, J. Kenneth. 1967. Polar bear dens on the Twin Islands, James Bay, Canada. Journal of Mammalogy. 48(3): 468-471. 
17. Driscoll, K. G.; Arocena, J. M.; Massicotte, H. B. 1999. Post-fire soil nitrogen content and vegetation composition in sub-boreal spruce forests of British Columbia's central interior, Canada. Forest Ecology and Management. 121: 227-237. 
18. Dyrness, C. T. 1980. White spruce. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 81. 
19. Dyrness, C. T.; Viereck, L. A.; Foote, M. J.; Zasada, J. C. 1988. The effect on vegetation and soil temperature of logging flood-plain white spruce. Res. Pap. PNW-RP-392. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 45 p. 
20. Ebersole, James J. 1987. Short-term vegetation recovery at an Alaskan arctic coastal plain site. Arctic and Alpine Research. 19(4): 442-450. 
21. Evans, Kevin E.; Kershaw, G. Peter. 1989. Productivity of agronomic and native plants under various fertilizer and seed application rates on a simulated transport corridor, Fort Norman, Northwest Territories. In: Walker, D. G.; Powter, C. B.; Pole, M. W., compilers. Proceedings of the conference: Reclamation, a global perspective; 1989 August 27-31; Calgary, AB. Edmonton, AB: Alberta Land Conservation and Reclamation Council: 279-287. 
22. Felix, Nancy A.; Raynolds, Martha K. 1989. The effects of winter seismic trails on tundra vegetation in northeastern Alaska, U.S.A. Arctic and Alpine Research. 21(2): 188-202. 
23. Felix, Nancy A.; Raynolds, Martha K.; Jorgenson, Janet C.; DuBois, Kristen E. 1992. Resistance and resilience of tundra plant communities to disturbance by winter seismic vehicles. Arctic and Alpine Research. 24(1): 69-77. 
24. Foote, M. Joan. 1993. Revegetation following the 1950 Porcupine River Fire: 1950-1981. Fairbanks, AK: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station, Institute of Northern Forestry. 71 p. Review draft. 
25. Foster, J. Bristol. 1961. Life history of the Phenacomys vole. Journal of Mammalogy. 42(2): 181-198. 
26. Haag, Richard W. 1974. Nutrient limitations to plant production in two tundra communities. Canadian Journal of Botany. 52(1): 103-116. 
27. Hann, Wendel; Havlina, Doug; Shlisky, Ayn; [and others]. 2005. Interagency fire regime condition class guidebook. Version 1.2, [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). Variously paginated [+ appendices]. Available: http://www.frcc.gov/docs/18.104.22.168/Complete_Guidebook_V1.2.pdf [2007, May 23]. 
28. Hansen, R. M. 1975. Foods of the hoary marmot on Kenai Peninsula, Alaska. The American Midland Naturalist. 94(2): 348-353. 
29. Hanson, Herbert C. 1951. Characteristics of some grassland, marsh, and other plant communities in western Alaska. Ecological Monographs. 21(4): 317-378. 
30. Hardy BBT Limited. 1989. Manual of plant species suitability for reclamation in Alberta. 2d ed. Edmonton, AB: Alberta Land Conservation and Reclamation Council. Report No. RRTAC 89-4. 436 p. 
31. Harper, Karen A.; Kershaw, G. Peter. 1996. Natural revegetation on borrow pits and vehicle tracks in shrub tundra, 48 years following construction of the CANOL No. 1 Pipeline, N.W.T., Canada. Arctic and Alpine Research. 28(2): 163-171. 
32. Hawkes, Brad C. 1983. Fire history and ecology of forest ecosystems in Kluane National Park. In: Wein, Ross Wallace; Riewe, Roderick R.; Methven, Ian R., eds. Resources and dynamics of the boreal zone: Proceedings of a conference; 1982 August; Thunder Bay, ON. Ottawa, ON: Association of Canadian Universities for Northern Studies: 266-280. 
33. Hechtel, John L. 1985. Activity and food habits of barren-ground grizzly bears in arctic Alaska. Missoula, MT: University of Montana. 74 p. Thesis. 
34. Hernandez, Helios. 1973. Natural plant recolonization of surficial disturbances, Tuktoyaktuk Peninsula region, Northwest Territories. Canadian Journal of Botany. 51: 2177-2196. 
35. Heusser, Calvin J. 1956. Postglacial environments in the Canadian Rocky Mountains. Ecological Monographs. 26(4): 263-302. 
36. Hoefs, Manfred. 1979. Flowering plant phenology at Sheep Mountain, southwest Yukon Territory. Canadian Field Naturalist. 93(2): 183-187. 
37. Hogg, Edward H. 1993. An arctic-alpine flora at low elevation in Marble Canyon, Kootenay National Park, British Columbia. Canadian Field-Naturalist. 107: 282-292. 
38. Holloway, Patricia S.; Alexander, Ginny. 1990. Ethnobotany of the Fort Yukon region, Alaska. Economic Botany. 44(2): 214-225. 
39. Houston, Kent E.; Hartung, Walter J.; Hartung, Carol J. 2001. A field guide for forest indicator plants, sensitive plants, and noxious weeds of the Shoshone National Forest, Wyoming. Gen. Tech. Rep. RMRS-GTR-84. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 184 p. 
40. Hulten, Eric. 1968. Flora of Alaska and neighboring territories. Stanford, CA: Stanford University Press. 1008 p. 
41. Hungerford, Roger D.; Frandsen, William H.; Ryan, Kevin C. 1995. Ignition and burning characteristics of organic soils. In: Cerulean, Susan I.; Engstrom, R. Todd, eds. Fire in wetlands: a management perspective: Proceedings, 19th Tall Timbers fire ecology conference; 1993 November 3-6; Tallahassee, FL. No. 19. Tallahassee, FL: Tall Timbers Research Station: 78-91. 
42. Ion, Peter G.; Kershaw, G. Peter. 1989. The selection of snowpatches as relief habitat by woodland caribou (Rangifer tarandus caribou), Macmillan Pass, Selwyn/Makenzie Mountains, N.W.T., Canada. Arctic and Alpine Research. 21(2): 203-211. 
43. Kari, Priscilla Russell. 1987. Tanaina plantlore. Dena'ina K'et'una: An ethnobotany of the Dena'ina Indians of southcentral Alaska. 2nd ed. [Revised]. Anchorage, AK: U.S. Department of the Interior, National Park Service, Alaska Region. 205 p. 
44. 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. 
45. La Roi, George H.; Hnatiuk, Roger J. 1980. The Pinus contorta forests of Banff and Jasper National Parks: a study in comparative synecology and syntaxonomy. Ecological Monographs. 50(1): 1-29. 
46. La Roi, George Henri. 1964. An ecological study of the boreal spruce-fir forests of the North American taiga. Durham, NC: Duke University. 397 p. Dissertation. 
47. 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: http://www.landfire.gov/downloadfile.php?file=RA_Modeling_Manual_v2_1.pdf [2007, May 24]. 
48. 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: http://www.landfire.gov/models_EW.php [2008, April 18] 
49. Landhausser, Simon M.; Wein, Ross W. 1993. Postfire vegetation recovery and tree establishment at the Arctic treeline: climate-change--vegetation response hypotheses. Journal of Ecology. 81: 665-672. 
50. Larsen, James A. 1980. Boreal communities and ecosystems: the broad view. In: Larsen, James A., ed. The boreal ecosystem. New York: Academic Press: 128-236. 
51. Larsen, James A. 1982. Ecology of the northern lowland bogs and conifer forests. New York: Academic Press. 307 p. 
52. Le Goff, Heloise; Sirois, Luc. 2004. Black spruce and jack pine dynamics simulated under varying fire cycles in the northern boreal forest of Quebec, Canada. Canadian Journal of Forest Research. 34(12): 2399-2409. 
53. Lutz, H. J. 1956. Ecological effects of forest fires in the interior of Alaska. Tech. Bull. No. 1133. Washington, DC: U.S. Department of Agriculture, Forest Service. 121 p. 
54. Mann, Daniel H.; Plug, Lawrence J. 1999. Vegetation and soil development at an upland taiga site, Alaska. Ecoscience. 6(2): 272-285. 
55. Maslen, Lynn; Kershaw, G. Peter. 1989. First year results of revegetation trials using selected native plant species on a simulated pipeline trench, Fort Norman, N.W.T., Canada. In: In: Proceedings of the conference: reclamation, a global perspective; 1989 August 27-31; Calgary, AB. Edmonton, AB: Alberta Land Conservation and Reclamation Council, Reclamation Research Technical Advisory Committee: 81-90. 
56. Meyer, Susan E. [In press]. Arctostaphylos Adans.: manzanita. In: Bonner, Franklin T.; Nisley, Rebecca G.; Karrfait, R. P., coords tech. coord. Woody plant seed manual. Agric. Handbook 727. Washington, DC: U.S. Department of Agriculture, Forest Service (Producer). Available: http://www.nsl.fs.fed.us/wpsm/Arctostaphylos.pdf [2008, July 22]. 
57. Miller, Donald R. 1976. Biology of the Kaminuriak population of barren-ground caribou. Part 3. Taiga winter range relationships and diet. Canadian Wildlife Service Rep. Series No. 36. Ottawa, ON: Environment Canada, Wildlife Service. 42 p. 
58. Nowak, Stephanie; Kershaw, G. Peter; Kershaw, Linda J. 2002. Plant diversity and cover after wildfire on anthropogenically disturbed and undisturbed sites in subarctic upland Picea mariana forest. Arctic. 55(3): 269-280. 
59. Orloci, Laszlo; Stanek, Walter. 1979. Vegetation survey of the Alaska highway, Yukon Territory: types and gradients. Vegetatio. 41(1): 1-56. 
60. Oswald, E. T.; Brown, B. N. 1990. Vegetation establishment during 5 years following wildfire in northern British Columbia and southern Yukon Territory. Information Report BC-X-320. Victoria, BC: Forestry Canada, Pacific and Yukon Region, Pacific Forestry Centre. 46 p. 
61. Pearce, C. M.; McLennan, D.; Cordes, L. D. 1988. The evolution and maintenance of white spruce woodlands on the Mackenzie Delta, N. W. T., Canada. Holarctic Ecology. 11(4): 248-258. 
62. Peinado, M.; Aguirre, J. L.; Delgadillo, J. 1997. Phytosociological, bioclimatic and biogeographical classification of woody climax communities of western North America. Journal of Vegetation Science. 8: 505-528. 
63. Peterson, K. M.; Billings, W. D. 1980. Tundra vegetational patterns and succession in relation to microtopography near Atkasook, Alaska. Arctic and Alpine Research. 12(4): 473-482. 
64. Ping, C. L.; Michaelson, G. J.; Packee, E. C.; Stiles, C. A.; Swanson, D. K.; Yoshikawa, K. 2005. Soil catena sequences and fire ecology in the boreal forest of Alaska. Soil Science Society of America Journal. 69(6): 1761-1772. 
65. Racine, Charles; Jandt, Randi; Meyers, Cynthia; Dennis, John. 2004. Tundra fire and vegetation change along a hillslope on the Seward Peninsula, Alaska, U.S.A. Arctic, Antarctic, and Alpine Research. 36(1): 1-10. 
66. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. 
67. Raup, Hugh M. 1947. Some natural floristic areas in boreal America. Ecological Monographs. 17(2): 221-234. 
68. Ritchie, J. C. 1982. The modern and late-quaternary vegetation of the Doll Creek Area, North Yukon, Canada. New Phytologist. 90(3): 563-603. 
69. Rouse, W. R.; Kershaw, K. A. 1973. Studies on lichen-dominated systems. VI. Interrelations of vegetation and soil moisture in the Hudson Bay Lowlands. Canadian Journal of Botany. 51(7): 1309-1317. 
70. Rowe, J. S.; Bergsteinsson, J. L.; Padbury, G. A.; Hermesh, R. 1974. Fire studies in the Mackenzie Valley. ALUR 73-74-61. Ottawa: Canadian Department of Indian and Northern Development. 123 p. 
71. Shaver, G. R.; Chapin, F. S., III. 1986. Effect of fertilizer on production and biomass of Tussock Tundra, Alaska, U.S.A. Arctic and Alpine Research. 18(3): 261-268. 
72. Skogland, Terje. 1980. Comparative summer feeding strategies of arctic and alpine rangifer. The Journal of Animal Ecology. 49(1): 81-98. 
73. Smith, Donald A.; Foster, J. Bristol. 1957. Notes on the small mammals of Churchill, Manitoba. Journal of Mammalogy. 38(1): 98-115. 
74. Soper, James H.; Heimburger, Margaret L. 1982. Shrubs of Ontario. Life Sciences Miscellaneous Publications. Toronto, ON: Royal Ontario Museum. 495 p. 
75. Stanek, W.; Alexander, K.; Simmons, C. S. 1981. Reconnaissance of vegetation and soils along the Dempster Highway, Yukon Territory: I. Vegetation types. BC-X-217. Victoria, BC: Environment Canada, Canadian Forestry Service, Pacific Forest Research Centre. 32 p. 
76. Stanek, Walter. 1980. Vegetation types and environmental factors associated with Foothills Gas Pipeline route, Yukon Territory. BC-X-205. Victoria, BC: Environment Canada, Canadian Forestry Service, Pacific Forest Research Centre. 48 p. 
77. Stelmock, Jim J.; Dean, Frederick C. 1986. Brown bear activity and habitat use, Denali National Park: 1980. In: Proceedings, 6th international conference on bear research and management; 1983 February; Grand Canyon, AZ. [Place of publication unknown]: International Association of Bear Research and Management: 155-167. 
78. 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. 
79. Suarez, Frank; Binkely, Dan; Kaye, Margot W. 1999. Expansion of forest stands into tundra in the Noatak National Preserve, northwest Alaska. Ecoscience. 6(3): 465-470. 
80. Swanson, David K. 1996. Susceptibility of permafrost soils to deep thaw after forest fires in interior Alaska, U.S.A., and some ecologic implications. Arctic and Alpine Research. 28(2): 217-227. 
81. Timoney, Kevin P.; La Roi, George H.; Zoltai, Stephen C.; Robinson, Anne L. 1993. Vegetation communities and plant distributions and their relationships with parent materials in the forest-tundra of northwestern Canada. Ecography. 16: 174-188. 
82. Tisdale, E. W.; Fosberg, M. A.; Poulton, C. E. 1964. Vegetation and soil development of a recently glaciated area near Mt. Robson, British Columbia. Bulletin of the Ecological Society of America. 45(3): 107-108. 
83. Tisdale, E. W.; Fosberg, M. A.; Poulton, C. E. 1966. Vegetation and soil development of a recently glaciated area near Mount Robson, British Columbia. Ecology. 47(4): 517-523. 
84. U.S. Department of Agriculture, Natural Resources Conservation Service. 2008. PLANTS Database, [Online]. Available: http://plants.usda.gov/. 
85. Viereck, L. A.; Dyrness, C. T.; Batten, A. R.; Wenzlick, K. J. 1992. The Alaska vegetation classification. Gen. Tech. Rep. PNW-GTR-286. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 278 p. 
86. Viereck, Leslie A. 1979. Characteristics of treeline plant communities in Alaska. Holarctic Ecology. 2: 228-238. 
87. Viereck, Leslie A.; Little, Elbert L., Jr. 1972. Alaska trees and shrubs. Agric. Handb. 410. Washington, DC: U.S. Department of Agriculture, Forest Service. 265 p. 
88. Walker, D. A.; Short, S. K.; Andrews, J. T.; Webber, P. J. 1981. Late holocene pollen and present-day vegetation, Prudhoe Bay and Atigun River, Alaskan north slope. Arctic and Alpine Research. 13(2): 153-172. 
89. Walker, Marilyn D.; Walker, Donald A.; Auerbach, Nancy A. 1994. Plant communities of a tussock tundra landscape in the Brooks Range Foothills, Alaska. Journal of Vegetation Science. 5(6): 843-866. 
90. Watson, L. E.; Parker, R. W.; Polster, D. F. 1980. Manual of plant species suitability for reclamation in Alberta. Vol. 2: Forbs, shrubs and trees. RRTAC 80-5. Edmonton, AB: Land Conservation and Reclamation Council. 537 p. 
91. Weber, Michael G. 1974. Nutrient budget changes following fire in arctic plant communities. In: Wein, R. W., ed. Recovery of vegetation in arctic regions after burning. Supplementary report. Ottawa, ON: Information Canada, Environmental Social Committee, Northern Pipelines Task Force on Northern Oil Development. 43-63. 
92. Wein, Ross W.; Bliss, L. C. 1973. Experimental crude oil spills on arctic plant communities. Journal of Applied Ecology. 10(3): 671-682. 
93. Werner, Richard A. 2002. Effect of ecosystem disturbance on diversity of bark and woodboring beetles (Coleoptera: Scolytidae, Buprestidae, Cerambycidae) in white spruce (Picea glauca (Moench) Voss) ecosystems of Alaska. Res. Pap. PNW-RP-546. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 15 p. 
94. Winterberger, Kenneth C. 1994. SRM 907: Dryas. In: Shiflet, Thomas N., ed. Rangeland cover types of the United States. Denver, CO: Society for Range Management: 130-131. 
95. Zoltai, S. C.; Pettapiece, W. W. 1973. Studies of vegetation, landform and permafrost in the Mackenzie Valley: Terrain, vegetation and permafrost relationships in the northern part of the Mackenzie Valley. Report No. 73-4. Task Force on Northern Oil Development, Environmental-Social Committee, Northern Pipelines. 105 p.