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SPECIES:  Aralia nudicaulis

Introductory

SPECIES: Aralia nudicaulis
AUTHORSHIP AND CITATION : Pavek, Diane S. 1993. Aralia nudicaulis. 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/aranud/all.html [].
ABBREVIATION : ARANUD SYNONYMS : NO-ENTRY SCS PLANT CODE : ARNU2 COMMON NAMES : wild sarsaparilla aralia salsepareille TAXONOMY : The currently accepted scientific name of wild sarsaparilla is Aralia nudicaulis L. [39,57]. It is a member of the ginseng family (Araliaceae). There are no subspecies, varieties, or forms. LIFE FORM : Forb FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY

DISTRIBUTION AND OCCURRENCE

SPECIES: Aralia nudicaulis
GENERAL DISTRIBUTION : Wild sarsaparilla is distributed from Newfoundland south to North Carolina and west through the Great Plains [39,48,110,116,120]. Its range continues in Canada to British Columbia and in the United States through the Intermountain West to northeastern Washington [34,54,57]. ECOSYSTEMS : FRES10 White - red - jack pine FRES11 Spruce - fir FRES14 Oak - 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 FRES25 Larch STATES : AR CO CT GA ID IL IN IA KY ME MD MA MI MN MO MT NE NH NJ NY NC OH PA RI SD TN VT VA WA WV WI WY AB BC MD NB NF NS ON PQ SK BLM PHYSIOGRAPHIC REGIONS : 8 Northern Rocky Mountains 9 Middle Rocky Mountains 11 Southern Rocky Mountains 13 Rocky Mountain Piedmont 14 Great Plains 15 Black Hills Uplift 16 Upper Missouri Basin and Broken Lands KUCHLER PLANT ASSOCIATIONS : K012 Douglas-fir forest K013 Cedar - hemlock - pine forest K014 Grand fir - Douglas-fir forest K015 Western spruce - fir forest K018 Pine - Douglas-fir forest 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 K101 Elm - ash 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 K111 Oak - hickory - pine forest SAF COVER TYPES : 1 Jack pine 5 Balsam fir 12 Black spruce 13 Black spruce - tamarack 15 Red pine 16 Aspen 17 Pin cherry 18 Paper birch 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 28 Black cherry - 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 39 Black ash - American elm - red maple 42 Bur oak 44 Chestnut oak 46 Eastern redcedar 52 White oak - black oak - northern red oak 58 Yellow-poplar - eastern hemlock 60 Beech - sugar maple 107 White spruce 108 Red maple 110 Black oak 201 White spruce 202 White spruce - paper birch 203 Balsam poplar 204 Black spruce 206 Engelmann spruce - subalpine fir 210 Interior Douglas-fir 212 Western larch 213 Grand fir 215 Western white pine 217 Aspen 218 Lodgepole pine 227 Western redcedar - western hemlock 228 Western redcedar 236 Bur oak 237 Interior ponderosa pine 251 White spruce - aspen 252 Paper birch 253 Black spruce - white spruce 254 Black spruce - paper birch SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Wild sarsaparilla is a widespread, dominant understory species throughout the boreal coniferous and mixed-wood forests [8,13,26,27, 67,77,88]. It is a major understory species of the chestnut oak (Quercus prinus) type and sugar maple-beech (Acer saccharum-Fagus grandiflora) and the bottomland cottonwood or cottonwood-birch associations of the Great Plains [16,33,46,70,87,102]. Wild sarsaparilla is an indicator species in several phases of western redcedar (Thuja plicata), western hemlock (Tsuga heterophylla), and subalpine fir (Abies lasiocarpa) habitat type series [24,31,41,92,93,111]. Numerous classifications use wild sarsaparilla as a dominant or indicator understory species, including the following: (1) Classification of the forest vegetation of Wyoming [5] (2) Field guide habitat classification system for Upper Peninsula of Michigan and Northeast Wisconsin [21] (3) Native woodland habitat types of southwestern North Dakota [45] (4) Classification of quaking aspen stands in the Black Hills and Bear Lodge Mountains [109] (5) Classification and gradient analysis of forest vegetation of Cape Enrage, Bic Park, Quebec [125].

MANAGEMENT CONSIDERATIONS

SPECIES: Aralia nudicaulis
IMPORTANCE TO LIVESTOCK AND WILDLIFE : Wild sarsaparilla is browsed by livestock and wildlife [71]. Grizzly and black bear consume the fruits [73,104,105,106]. Wild sarsaparilla is frequently browsed in summer by caribou, and it is a preferred spring food of moose [12,28,36]. In Minnesota, use of wild sarsaparilla by ruffed grouse occurs throughout the year [86]. Wild sarsaparilla seeds have been found in the stomachs of thrushes [36]. White-tailed deer in northeastern Minnesota used wild sarsaparilla 18 percent of the time during feeding in June [63]. Wild sarsaparilla was so heavily browsed by white-tailed deer on Fire Island National Seashore, New York, during periods of overpopulation that it was only found in deer exclosures or within dense thickets [95]. PALATABILITY : NO-ENTRY NUTRITIONAL VALUE : The current year's growth of wild sarsaparilla in Manitoba had 11.1 percent crude protein, 34.1 percent acid detergent fiber, and 63.4 percent dry matter digestibility [107]. It had only 1 percent dry matter digestibility on Isle Royale, Michigan [12]. Leaf dry matter contained 1.2 percent nitrogen [89]. COVER VALUE : NO-ENTRY VALUE FOR REHABILITATION OF DISTURBED SITES : NO-ENTRY OTHER USES AND VALUES : The Nuxalk of east-central British Columbia collected wild sarsaparilla rhizomes for food from May to September [82]. The rhizomes have been used to make beverages such as root beer [48,75,116]. OTHER MANAGEMENT CONSIDERATIONS : After 6 years, wild sarsaparilla was infrequent to numerous on various logged areas (370 acres [150 ha] logged in alternate strips, a 32-acre [80 ha] clearcut, and an 81-acre [200 ha] clearcut) [37]. Winter and spring clearcuts had significantly (p<0.05) less wild sarsaparilla biomass 2 years after logging compared to an uncut control [100]. Three years after clearcutting, wild sarsaparilla had higher frequency (22.5 percent) on sites prepared by V-blade than on sites prepared by toothed brush rake (7.5 percent) or disking (2.5 percent) [65]. Wild sarsaparilla decreased on clearcuts and shelterwood cuts in areas open 0.25 mile (0.4 km) or more in virgin western white pine (Pinus monticola)-western redcedar in northern Idaho, possibly due to increased air and soil temperatures and evaporation [79,80]. Data on wild sarsaparilla abundance have been used in multiple regression models for predicting forest site quality for regeneration of spruce-fir, lodgepole pine (Pinus contorta), and white spruce (Picea glauca) [78,121,124]. Two years after a clearcut was mulched with various straws, wild sarsaparilla was less frequent in mulch treatments (8.3 percent) than in controls (10 percent) [66]. In ecological classifications that are useful for biomass and productivity estimates, wild sarsaparilla had average rank abundances between 0.2 and 1.3 in northwestern Michigan [58]. Wild sarsaparilla was included in a suitability index for summer food for black bear in the upper Great Lakes Region [104]. Herbivory can reduce the number of wild sarsaparilla flowering stems and fruit production. Clipped shoots produced significantly (p<0.005) fewer fruits and smaller leaves compared to intact shoots. One year later, chance of flowering and leaf production were significantly (p<0.05) less [36]. Wild sarsaparilla was indirectly affected by aerial spraying for spruce budworm in New Brunswick. Wild sarsaparilla had lower fruit set in sprayed areas compared with unsprayed [10]. Fruit collection, seed extraction, and nursery methods have been discussed in detail [17]. Wild sarsaparilla top-growth increased but stem density did not after shoot removal and rhizome transplantation into uniform soils during summer [42]. Wild sarsaparilla was the most common forb in an area impacted by metal aerosols from a zinc smelter in Pennsylvania. Wild sarsaparilla was generally chlorotic [68]. In a boreal forest in Ontario, wild sarsaparilla leaves were significantly (p<0.005) less able to neutralize simulated acid raindrops at pH 3.8 or 5.6 [43].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Aralia nudicaulis
GENERAL BOTANICAL CHARACTERISTICS : Wild sarsaparilla is a native, dioecious, perennial forb [36,39]. It has long rhizomes that are between 1.2 and 4.7 inches (3-12 cm) deep in the mineral soil, with an average depth of 2.4 inches (6 cm) [42]. Wild sarsaparilla is acaulescent; vegetative shoots may be up to 27.6 inches (70 cm) tall with two to three compound leaves [34,36,54]. Reproductive shoots are shorter with two to seven umbels [39]. There is an average of five seeds per fruit [10,48]. RAUNKIAER LIFE FORM : Hemicryptophyte Geophyte REGENERATION PROCESSES : Wild sarsaparilla forms extensive colonies by vegetative reproduction [11,116]. Its rhizomes branch and can produce ramets up to 39 inches (1 m) from the original plant [23]. An exclusion experiment showed that wild sarsaparilla was dependent on insects such as bumblebees, solitary bees, and syrphids for pollination. Germination rates with or without stratification are low (34 percent) [17]. Seedling establishment probably occurs rarely and is dependent on major disturbances [36]. Wild sarsaparilla seedbanks under canopies of stands aged 3 to 75 years since disturbance yielded 28 seeds that had 18 percent germination [3]. Wild sarsaparilla seed rain and seedbank were measured following canopy removal of mature northern hardwood forest in north-central New Hampshire. One year after harvest, 1.3 wild sarsaparilla seeds germinated from soil samples; no fruits were collected in seedtraps during 2 years of sampling [62]. Seeds are disseminated by animals such as birds and black bears [73,91]. While 61 percent of the flowers in a wild sarsaparilla population in New Brunswick set fruit, 95 percent of the fruits were consumed [10]. Germination rates for wild sarsaparilla seeds taken from black bear scat (62-93 percent) were significantly (p<0.01) higher than for uneaten seeds (27-28 percent) [105]. SITE CHARACTERISTICS : Wild sarsaparilla has a relatively wide ecological amplitude [112]. It is common in moist or dry woodlands, thickets, riparian areas, and prairie or bog edges [22,29,39,76,84,110]. It may occur on sand plains or dunes, rocky ridges, and canyon sides [55,126]. Wild sarsaparilla is found in continental climates that are subhumid to humid and may be influenced by maritime air masses [7,47,93,100,101]. Winters are long and cool to cold; summers are short and warm [4,9]. Moderate amounts of precipitation occur throughout its range, varying from 16.1 to 65 inches (409-1,650 mm) [7,64]. Wild sarsaparilla is found on all aspects and slopes [24]. Textures of the soils on which it occurs range from fine loamy clay to coarse loam [4,42,67,70]. Soils are moderate to rich in nutrients [18,35,112]. Sites are poorly drained to well drained [35]. Wild sarsaparilla occurs at low to mid-elevations (2,500 to 4,700 feet [760-1,433 m]) in Montana and Idaho and up to 8,000 feet (2,438 m) in Colorado [24,53,81,102]. In the northeastern United States, it is found at elevations ranging from 680 to 2,995 feet (198-913 m) [40]. Species commonly associated with wild sarsaparilla are broadleaf arnica (Arnica latifolia), bigleaf aster (Aster macrophyllus), bunchberry (Cornus canadensis), queencup beadlily (Clintonia uniflora), and Canada mayflower (Maianthemum canadense) [53,61,99,101]. Often, oakfern (Gymnocarpium dryopteris) and bracken fern (Pteridium aquilinum) are found with wild sarsaparilla [9,24]. SUCCESSIONAL STATUS : Facultative Seral Species Wild sarsaparilla is shade tolerant [75]. In mixed-hardwood stands in New Brunswick aged 7 to 37 years, wild sarsaparilla frequency increased with stand age [85]. Wild sarsaparilla was distributed at three plants per 269 square feet (3 plants/25 sq m) in a 20-year-old eastern white pine (Pinus strobus) stand in Vermont [60]. In a spruce-fir stand in Maine where trees averaged 60 or more years old, wild sarsaparilla occurred with 21 percent frequency on plots [32]. Wild sarsaparilla is a characteristic forb of a wide range of climax forests [25,49]. It occurred in plots with differing amounts of canopy closure in late successional stands of western hemlock-western redcedar stands in northern Idaho [92]. The frequency of wild sarsaparilla varied across western hemlock-western redcedar stands aged less than 50 years to more than 400 years old [50,51]. In old-growth (200-560 years) eastern hemlock (Tsuga canadensis) in northwestern Pennsylvania, wild sarsaparilla frequency averaged 2 percent in plots [59]. In older seral to climax stages of sugar maple-basswood stands aged 100 to 165 years, wild sarsaparilla had 1 to 3 percent cover measured over 7 years [69]. However, in an old-growth eastern hemlock-northern hardwoods forest in Michigan, frequency of wild sarsaparilla was higher in treefall gaps (4.4 percent) than under closed canopy (2.2 percent) [91]. On sand dunes at Lake Michigan, Michigan, wild sarsaparilla was only present under black oak (Quercus velutina) on older, stabilized dunes, aged 600 to 1,100 years [98]. SEASONAL DEVELOPMENT : Wild sarsaparilla emerges from leaf litter by mid-April or May, and leaves expand before the canopy closes [23,30]. It flowers from May to July throughout its range [76,110,120]. Fruits mature in about 32 days [56]. Leaves begin to drop by mid-September. Wild sarsaparilla is dormant during the winter [30].

FIRE ECOLOGY

SPECIES: Aralia nudicaulis
FIRE ECOLOGY OR ADAPTATIONS : Mineral soil protects wild sarsaparilla rhizomes from most fires [42]. Wild sarsaparilla occurs in communities with diverse fire regimes. In Saskatchewan, it is a dominant understory component in aspen-birch forests that experience frequent fires [33]. Wild sarsaparilla occurs in the understory of mixed woods in New Brunswick where fires occur every 9 to 45 years [42]. It is common in aspen-Douglas-fir (Pseudotsuga menziesii) stands in the Black Hills, South Dakota, that have fire return intervals of 80 to 90 years [102]. Wild sarsaparilla is codominant with queencup beadlily in the moist lower elevation habitat types of subalpine fir series where fires are infrequent and severe [31,41]. Wild sarsaparilla is present in woodland communities, such as black oak in Indiana, that are present due to fire suppression since the time of settlement [15]. FIRE REGIMES : Find fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes". POSTFIRE REGENERATION STRATEGY : Rhizomatous herb, rhizome in soil Geophyte, growing points deep in soil Ground residual colonizer (on-site, initial community)

FIRE EFFECTS

SPECIES: Aralia nudicaulis
IMMEDIATE FIRE EFFECT ON PLANT : Fire top-kills wild sarsaparilla; vegetative and reproductive buds are destroyed. Surviving rhizomes sprout and vigorously grow following fall or spring fires. New rhizomes are produced. Flowers are not initiated during the first growing season following fire [19]. Very few wild sarsaparilla come in as seed immediately following fire [1]. Following a light- to moderate-severity lightning fire in April, wild sarsaparilla plants sprouted from surviving rhizomes; no seeds germinated [6]. After a July prescribed fire, surviving wild sarsaparilla sprouted within 2 weeks and was common on all sites. It did decrease in cover, however, and in community importance [114]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Wild sarsaparilla has been classified as a decreaser. It initially decreases in frequency and biomass following fire [72,113]. The amount of decrease does not depend on the type of fire (headfire or backfire) but does depend on fire intensity and time of burning [113]. Wild sarsaparilla generally decreases in frequency by about 50 percent 1 year following fire. Within 4 years, wild sarsaparilla can recover, with an increase in frequency ranging from 50 to 90 percent of preburn levels [1,97]. Two years after fire in northern Ontario, there was a three-fold increase in frequency of wild sarsaparilla [119]. In the red pine (Pinus resinosa) and eastern white pine forests of the Lake States, wild sarsaparilla does well following fire and is prevalent on fresh burns [123]. After various silvicultural methods combined with prescribed burning, the density of wild sarsaparilla usually decreased compared to logging without burning or to control treatments [1,52,94,114]. Wild sarsaparilla had less biomass (1.31 stems/sq m) compared to the control (3.77 stems/sq m) when a winter clearcut in northern Minnesota was followed by summer burning [100]. One year after a wildfire in northern Idaho, wild sarsaparilla sprouted in one out of 21 sites. It entered communities only at postfire years 9 and 10 in low densities of 0.2-2 sq m/0.01 ha [117]. Following two successive annual, low-severity fires in Ontario, wild sarsaparilla decreased in density from preburn levels. Surviving plants decreased following the second fire from 172 stems per hectare to 21 stems per hectare. One year after the fires, wild sarsaparilla began to increase [90]. In northern Michigan, wild sarsaparilla had peak frequency at postfire year 31 [108]. Thirty-three years after fire in northeastern Minnesota, it occurred on all plots with an average frequency of 57 percent [96]. Following a light-severity surface fire in Ontario, wild sarsaparilla only occurred on two sites aged 25 and 50 years and had importance values of 3.6 and 2.0, respectively [115]. The average cover of wild sarsaparilla was measured over 80 years on burned-over jack pine-black spruce (Pinus banksiana-Picea mariana) in northeastern Minnesota. Wild sarsaparilla remained at 1 to 2 percent cover until postfire year 15; it increased to 8 to 12 percent cover from postfire years 15 to 80 [2]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : The Research Project Summary Understory recovery after burning and reburning quaking aspen stands in central Alberta provides information on prescribed fire and postfire response of plant community species including wild sarsaparilla. FIRE MANAGEMENT CONSIDERATIONS : Summer moisture content of wild sarsaparilla averaged 253 percent in northeastern Minnesota. These data were used to assess flammability and test the herbaceous fuel moisture algorithm of the National Fire Danger Rating System [83]. Wild sarsaparilla does not contribute much volume to fine fuels since it decomposes relatively rapidly; leaf organic matter had decreased by 37.4 percent after 4 months [89]. Ground vegetation that included wild sarsaparilla was used to develop regression equations for predicting changes in forest floor moisture [20].

REFERENCES

SPECIES: Aralia nudicaulis
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Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 10 p. [304] 6. Archibold, O. W. 1979. Buried viable propagules as a factor in postfire regeneration in northern Saskatchewan. Canadian Journal of Botany. 57: 54-58. [5934] 7. Arno, Stephen F. 1979. Forest regions of Montana. Res. Pap. INT-218. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 39 p. [340] 8. Arnott, J. T. 1989. Managing for aspen--a shared responsibility. Forestry Chronicle. Feb: 16-22. [6350] 9. Barclay-Estrup, P. 1987. A new shrub for Ontario: mountain bilberry, Vaccinium membranaceum, in Pukaskwa National Park. Canadian Field-Naturalist. 101(4): 526-531. [6233] 10. 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] 11. Bell, A. D.; Tomlinson, P. B. 1980. Adaptive architecture in rhizomatous plants. Botanical Journal of the Linnean Society. 80: 125-160. [11822] 12. Belovsky, G. E.; Jordan, P. A. 1978. The time energy budget of a moose. Theoretical Population Biology. 14: 76-104. [10100] 13. Bergeron, Jean-Francois; Saucier, Jean-Pierre; Robitaille, Andre; Robert, Denis. 1992. Quebec forest ecological classification program. Forestry Chronicle. 68(1): 53-63. [18571] 14. 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] 15. Betz, Robert F. 1978. The prairies of Indiana. In: Glenn-Lewin, David C.; Landers, Roger Q., Jr., eds. Proceedings, 5th Midwest prairie conference; 1976 August 22-24; Ames, IA. Ames, IA: Iowa State University: 25-31. [3292] 16. Bird, Ralph D. 1930. Biotic communities of the aspen parkland of central Canada. Ecology. 11(2): 356-442. [15277] 17. Blum, Barton M. 1974. Aralia L. aralia. In: Schopmeyer, C. S., technical coordinator. Seeds of woody plants in the United States. Agric. Handb. 450. Washington, DC: U.S. Department of Agriculture, Forest Service: 220-222. [7459] 18. Brumelis, G.; Carleton, T. J. 1989. The vegetation of post-logged black spruce lowlands in central Canada. II. Understory vegetation. Journal of Applied Ecology. 26: 321-339. [7864] 19. Chapman, Rachel Ross; Crow, Garrett E. 1981. Application of Raunkiaer's life form system to plant species survival after fire. Torrey Botanical Club. 108(4): 472-478. [7432] 20. Chrosciewicz, Z. 1989. Prediction of forest-floor moisture content under diverse jack pine canopy conditions. Canadian Journal of Forestry. 19: 1483-1487. [9734] 21. Coffman, Michael S.; Alyanak, Edward; Resovsky, Richard. 1980. 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Managing for aspen--a shared responsibility: Proceedings of the Joint TechnicalSession of the Forest Ecology, Silviculture and Tree Improvement Forest Management, and Forest Economics and Policy Working Groups; September 1988; Prince Albert, SK. In: The Forestry Chronicle. February: 16-22. [6919] 27. Crandall, Dorothy L. 1958. Ground vegetation patterns of the spruce-fir area of the Great Smoky Mountains National Park. Ecological Monographs. 28(4): 337-360. [11226] 28. Cringan, Alexander Thom. 1957. History, food habits and range requirements of the woodland caribou of continental North America. Transactions, North American Wildlife Conference. 22: 485-501. [15651] 29. Cronan, Christopher S.; DesMeules, Marc R. 1985. A comparison of vegetative cover and tree community structure in three forested Adirondack watersheds. Canadian Journal of Forest Research. 15: 881-889. [7296] 30. Daubenmire, Rexford F. 1936. The "big woods" of Minnesota: its structure, and relation to climate, fire, and soils. Ecological Monographs. 6(2): 233-268. [2697] 31. Davis, Kathleen M.; Clayton, Bruce D.; Fischer, William C. 1980. Fire ecology of Lolo National Forest habitat types. INT-79. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 77 p. [5296] 32. Davis, Ronald B. 1966. Spruce-fir forests of the coast of Maine. Ecological Monographs. 36(2): 79-94. [8228] 33. Dix, R. L.; Swan, J. M. A. 1971. The roles of disturbance and succession in upland forest at Candle Lake, Saskatchewan. Canadian Journal of Botany. 49: 657-676. [12808] 34. Dorn, R. D. 1977. Manual of the vascular plants of Wyoming. New York: Garland Publ. 2 vols. [21082] 35. Drew, T. J. 1988. Managing white spruce in Alberta's mixedwood forest: the dilemma. In: Samoil, J. K., ed. Management and utilization of northern mixedwoods: Proceedings of a symposium; 1988 April 11-14; Edmonton, AB. Inf. Rep. NOR-X-296. Edmonton, AB: Canadian Forestry Service, Northern Forestrty Centre: 35-40. [13045] 36. Edwards, Joan. 1985. Effects of herbivory by moose on flower and fruit production of Aralia nudicaulis. Journal of Ecology. 73: 861-868. [13626] 37. Eis, S. 1981. Effect of vegetative competition on regeneration of white spruce. Canadian Journal of Forest Research. 11: 1-8. [10104] 38. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905] 39. 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] 40. Filip, Stanley M.; Little, Elbert L., Jr. 1971. Trees and shrubs of the Bartlett Experimental Forest, Carroll County, New Hampshire. Res. Pap. NE-211. Upper Darby, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station. 20 p. [13635] 41. Fischer, William C.; Clayton, Bruce D. 1983. Fire ecology of Montana forest habitat types east of the Continental Divide. Gen. Tech. Rep. INT-141. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 83 p. [923] 42. Flinn, Marguerite Adele. 1980. Heat penetration and early postfire regeneration of some understory species in the Acadian forest. Halifax, NB: University of New Brunswick. 87 p. Thesis. [9876] 43. Gaber, B. A.;Hutchinson, T. C. 1988. The neutralization of acid rain by the leaves of four boreal forest species. Canadian Journal of Botany. 66(9): 1877-1882. [8872] 44. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; [and others]. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. 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