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Introductory

SPECIES: Empetrum nigrum | Black Crowberry
ABBREVIATION : EMPNIG SYNONYMS : Empetrum hermaphroditum (Lange) Hagerup SCS PLANT CODE : EMNI COMMON NAMES : black crowberry crowbery curlewberry TAXONOMY : The currently accepted scientific name of black crowberry is Empetrum nigrum L. [2,13,18]. There are two recognized subspecies, both having a circumpolar distribution: Empetrum nigrum ssp. nigrum with unisexual flowers and Empetrum nigrum ssp. hermaphroditum with bisexual flowers [19]. LIFE FORM : Shrub FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY COMPILED BY AND DATE : Robin F. Matthews, March 1992 LAST REVISED BY AND DATE : NO-ENTRY AUTHORSHIP AND CITATION : Matthews, Robin F. 1992. Empetrum nigrum. In: Remainder of Citation

DISTRIBUTION AND OCCURRENCE

SPECIES: Empetrum nigrum | Black Crowberry
GENERAL DISTRIBUTION : Black crowberry is distributed throughout Alaska, across the Yukon Territory and Canada to Labrador, Newfoundland, and Greenland. It occurs south through New England and the Great Lakes states, as well as along the Pacific Coast to northern California. Black crowberry also has a wide distribution throughout Europe [38,42,47]. ECOSYSTEMS : FRES10 White - red - jack pine FRES11 Spruce - fir FRES19 Aspen - birch FRES20 Douglas-fir FRES23 Fir - spruce FRES24 Hemlock - Sitka spruce FRES26 Lodgepole pine FRES44 Alpine STATES : AK CA ME MA MI MN NH NY OR VT WA WI AB BC LB MB NB NF NT NS ON PQ SK YT ADMINISTRATIVE UNITS : ACAD DENA GLBA ISRO LACL MORA NOCA OLYM PIRO WRST YUCH BLM PHYSIOGRAPHIC REGIONS : 1 Northern Pacific Border 2 Cascade Mountains KUCHLER PLANT ASSOCIATIONS : K001 Spruce - cedar - hemlock forest K002 Cedar - hemlock - Douglas-fir forest K003 Silver fir - Douglas-fir forest K004 Fir - hemlock forest K008 Lodgepole pine - subalpine forest K012 Douglas-fir forest K015 Western spruce - fir forest K052 Alpine meadows and barren K093 Great Lakes spruce - fir forest K094 Conifer bog K095 Great Lakes pine forest K096 Northeastern spruce - fir forest K108 Northern hardwoods - spruce forest SAF COVER TYPES : 1 Jack pine 5 Balsam fir 12 Black spruce 13 Black spruce - tamarack 16 Aspen 18 Paper birch 38 Tamarack 107 White spruce 201 White spruce 202 White spruce - paper birch 204 Black spruce 205 Mountain hemlock 206 Engelmann spruce - subalpine fir 218 Lodgepole pine 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 253 Black spruce - white spruce 254 Black spruce - paper birch SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Black crowberry is a dominant or codominant in a variety of different habitats. It may occur as an understory dominant in open conifer woodlands with black spruce (Picea mariana), white spruce (P. glauca), or shore pine (Pinus contorta var. contorta). Black crowberry can dominate shrub-types with dwarf birch (Betula nana), willow (Salix spp.), and ericaceous shrubs in bogs or muskegs and on open, moist tundra [1,8,33,37,46]. Other commonly associated species include: paper birch (Betula papyrifera), quaking aspen (Populus tremuloides), Alaska cedar (Chamaecyparis nootkatensis), bog birch (Betula glandulosa), Labrador tea (Ledum glandulosum and L. groenlandicum), various Vaccinium and Carex species, feathermosses (Hylocomium spp. and Pleurozium spp.), lichens (Cladonia spp. and Cladina spp.), and sphagnum mosses. Published classification schemes listing black crowberry as a major component of plant associations (pas), community types (cts), or vegetation types (vts) are as follows: AREA CLASSIFICATION AUTHORITY AK gen. veg. pas Viereck & Dyrness 1980 Kenai Peninsula, AK vts Reynolds 1990 Canadian Rocky Mtns. old growth cts Achuff 1989 NF peatland pas Pollett 1972 sw YT cts Douglas 1974

VALUE AND USE

SPECIES: Empetrum nigrum | Black Crowberry
WOOD PRODUCTS VALUE : NO-ENTRY IMPORTANCE TO LIVESTOCK AND WILDLIFE : Black crowberry fruits are utilized as fall and winter forage by over 40 species of songbirds, waterfowl, and upland game birds [27,28,44,47]. The berries are especially important to grouse and ptarmigan [10,27,47]. Black crowberry seeds are a major component of the red-backed vole's fall diet [51]. Big game animals that browse black crowberry foliage include reindeer, caribou, and bear [4,17,41]. Bear also eat the berries, so black crowberry utilization by bear increases in summer as fruits become ripe. Occurrence of black crowberry fruits in bear scat samples increased from 5.9 percent in early spring to 12.9 percent by late summer [26]. PALATABILITY : NO-ENTRY NUTRITIONAL VALUE : Black crowberry in barren-ground caribou forage areas consists of 6.27 percent protein and releases energy in the amount of 5.51 kilocalories per gram [31] Digestibility of black crowberry has been classified as low [40]. COVER VALUE : Dense mats of black crowberry probably provide cover for small rodents and mammals. VALUE FOR REHABILITATION OF DISTURBED SITES : Black crowberry has been broadly successful at naturally colonizing borrow pits in the tundra regions of northwestern Canada, and may be of use in managed reclamation projects [21]. Black crowberry has followed cottongrass (Eriophorum spissum) in the colonization of mined peatlands, but only after decades have elapsed [12]. Dense black crowberry mats catch blowing soils in areas of high wind exposure, and its interlocking roots may help stabilize the steep, rocky slopes it often inhabits. Black crowberry could not be established by seed on test plots in simulated pipeline trenches near Fort Norman, Northwest Territories [29]. OTHER USES AND VALUES : Black crowberry fruits are used, but usually mixed with other berries, in pies or jellies. In the winter, Native Americans gather the persistant berries buried beneath the snow [19,47]. MANAGEMENT CONSIDERATIONS : Black crowberry can be grown from stem cuttings and has been used as ground cover in rough, low areas in interior Alaska [47]. Black crowberry showed no signs of recovery 2 years after clearcutting and subsequent burning near Fairbanks, Alaska [9]. Three years after defoliation, black crowberry in barren-ground caribou forage areas had not recovered [31].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Empetrum nigrum | Black Crowberry
GENERAL BOTANICAL CHARACTERISTICS : Black crowberry is a low, creeping evergreen shrub that generally reaches 6 inches (15 cm) in height and often forms dense mats. The leaves are linear to elliptic, and the lower surface is grooved to reduce evapotranspiration in harsh climates. Black crowberry has inconspicuous purple flowers [2,13,47,49]. Young black crowberry plants have a strong primary root, but as the plants age, a shallow root system with many lateral roots develops [5]. RAUNKIAER LIFE FORM : Chamaephyte REGENERATION PROCESSES : Sexual reproduction: Black crowberry is classified as polygamous, dioecious, or monoecious. The dark-blue to black fruit is a drupe containing six to nine nutlets [2,13,18]. Seeds are dispersed by birds and animals [20]. Some seeds may become established under the parent, but seedling mortality is generally high [5]. Black crowberry seeds have been found buried beneath the soil, although only a small percent of the seeds are actually viable [20,32]. Seeds were found in 71 percent of soil cores taken from plots near Great Slave Lake, Northwest Territories [20]. Vegetative reproduction: Sprouting from underground or basal portions is the main form of reproduction of black crowberry [5,20,39]. In addition, adventitious roots form where procumbent branches come in contact with the ground [5]. SITE CHARACTERISTICS : Black crowberry is found from sea level to alpine zones. It occurs in a wide variety of habitats including sphagnum bogs or muskegs, open tundra, rockfields, conifer forests, coastal bluffs, and exposed sea cliffs [3,38,47,49]. Black crowberry is tolerant of a wide range of soil moisture conditions, but is intolerant of prolonged water logging, and on wet sites it is found in better drained areas [5]. Black crowberry is adapted to harsh climates and it often inhabits sites exposed to wind, fog, and salt aerosals. Site characteristics influence black crowberry morphology: on sites with high wind exposure, black crowberry is branched and prostrate; on wet sites it is sparsely branched and has long annual growth increments; on dry sites it has branching shoots and is bushy [5]. Black crowberry is found in sandy to rocky soils, glacial till, and alluvial deposits [8,42]. Soil pH ranges from 2.5 to 7.7 [5]. Black crowberry establishes itself on mineral soils and stagnant surfaces that are nutrient enriched [7] but is also classified as an indicator of nitrogen-poor soils [22]. SUCCESSIONAL STATUS : Black crowberry is a pioneer on sandy blowouts, dry, lichen-covered depressions on eskers [3], and in avalanche areas [30]. However, it is more often associated with late seral or climax communities, particularily white or black spruce types [8,24,45]. Black crowberry is common and abundant in old forests that have had no recent fires [14]. SEASONAL DEVELOPMENT : Flowering occurs in spring in areas of early snowmelt and continues through July. Fruits mature from August to late fall and persist through the winter under snow cover [18,32,42,47].

FIRE ECOLOGY

SPECIES: Empetrum nigrum | Black Crowberry
FIRE ECOLOGY OR ADAPTATIONS : Black crowberry generally occurs in communities with long fire intervals or in communities that lack the dry fuel to sustain a fire [7,24,45]. Low growth form and small stems make black crowberry liable to top-kill by fire. Belowground parts are also very susceptible to fire damage because most of them are located near the soil surface [14,35]. Postfire seedlings may arise from seed banks but are not a regular occurrence [24]. Black crowberry can regenerate vegetatively following fire [5,20,39], but this process is slow. Normal or prefire densities may not be reached for 20 to 30 years [24]. POSTFIRE REGENERATION STRATEGY : survivor species; on-site surviving root crown or caudex off-site colonizer; seed carried by animals or water; postfire yr 1&2

FIRE EFFECTS

SPECIES: Empetrum nigrum | Black Crowberry
IMMEDIATE FIRE EFFECT ON PLANT : Fire top-kills black crowberry; moderate or severe fires also readily kill underground parts close to the soil surface [14,35]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Black crowberry is slow to recover following fire [5,48,50]. In Labrador, black crowberry decreased significantly in frequency and abundance following fire. Preburn frequency was 61 percent, while postburn frequency was 0 percent after 5 years [14]. It also showed little or no recovery in 2- or 7 year-old burns in the Seward Peninsula, Alaska [35]. In the Wickersham Dome Fire near Fairbanks, Alaska, black crowberry in black spruce stands responded differently in lightly and heavily burned areas. In the lightly burned sites, percent cover was 1.4, 1.1, 0.9, and 1.25 in postfire years 1, 2, 3, and 4, respectively. However, in the heavily burned sites, black crowberry cover was 0 percent in the 4 years immediately following the fire [46]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : NO-ENTRY FIRE MANAGEMENT CONSIDERATIONS : NO-ENTRY

REFERENCES

SPECIES: Empetrum nigrum | Black Crowberry
REFERENCES : 1. Achuff, Peter L. 1989. Old-growth forests of the Canadian Rocky Mountain national parks. Natural Areas Journal. 9(1): 12-26. [7442] 2. Anderson, J. P. 1959. Flora of Alaska and adjacent parts of Canada. Ames, IA: Iowa State University Press. 543 p. [9928] 3. Argus, George W. 1966. Botanical investigations in northeastern Saskatchewan: the subarctic Patterson-Hasbala Lakes region. Canadian Field-Naturalist. 80(3): 119-143. [8406] 4. Banfield, A. W. F.; Tener, J. S. 1958. A preliminary study of the Ungava caribou. Journal of Mammalogy. 39(4): 560-573. [12994] 5. Bell, J. N. B.; Tallis, J. H. 1973. Biological flora of the British Isles: Empertrum nigrum L. Journal of Ecology. 61: 289-305. [25083] 6. 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] 7. Damman, A. W. H. 1977. Geographical changes in the vegetation pattern of raised bogs in the Bay of Fundy region of Maine and New Brunswick. Vegetatio. 35(3): 137-151. [10158] 8. Douglas, George W. 1974. Montane zone vegetation of the Alsek River region, southwestern Yukon. Canadian Journal of Botany. 52: 2505-2532. [17283] 9. 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. [7471] 10. Ellison, Laurence. 1966. Seasonal foods and chemical analysis of winter diet of Alaskan spruce grouse. Journal of Wildlife Management. 30(4): 729-735. [9735] 11. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905] 12. Famous, Norman C.; Spencer, M. 1989. Revegetation patterns in mined peatlands in central and eastern North America studied. Restoration and Management Notes. 7(2): 95-96. [10171] 13. 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] 14. Foster, David R. 1985. Vegetation development following fire in Picea mariana (black spruce) - Pleurozium forests of south-eastern Labrador, Canada. Journal of Ecology. 73: 517-534. [7222] 15. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; [and others]. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998] 16. Hatler, David F. 1972. Food habits of black bears in interior Alaska. Canadian Field-Naturalist. 86(1): 17-31. [10389] 17. Henry, G. H. R.; Gunn, A. 1991. Recovery of tundra vegetation after overgrazing by caribou in arctic Canada. Arctic. 44(1): 38-42. [14747] 18. Hitchcock, C. Leo; Cronquist, Arthur. 1964. Vascular plants of the Pacific Northwest. Part 2: Salicaceae to Saxifragaceae. Seattle, WA: University of Washington Press. 597 p. [1166] 19. Hulten, Eric. 1968. Flora of Alaska and neighboring territories. Stanford, CA: Stanford University Press. 1008 p. [13403] 20. Johnson, E. A. 1975. Buried seed populations in the subarctic forest east of Great Slave Lake, Northwest Territories. Canadian Journal of Botany. 53: 2933-2941. [6466] 21. Kershaw, G. Peter; Kershaw, Linda J. 1987. Successful plant colonizers on disturbances in tundra areas of northwestern Canada. Arctic and Alpine Research. 19(4): 451-460. [6115] 22. Klinka, K.; Krajina, V. J.; Ceska, A.; Scagel, A. M. 1989. Indicator plants of coastal British Columbia. Vancouver, BC: University of British Columbia Press. 288 p. [10703] 23. Kuchler, A. W. 1964. Manual to accompany the map of potential vegetation of the conterminous United States. Special Publication No. 36. New York: American Geographical Society. 77 p. [1384] 24. 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. [7653] 25. Lyon, L. Jack; Stickney, Peter F. 1976. Early vegetal succession following large northern Rocky Mountain wildfires. In: Proceedings, Tall Timbers fire ecology conference and Intermountain Fire Research Council fire and land management symposium; 1974 October 8-10; Missoula, MT. No. 14. Tallahassee, FL: Tall Timbers Research Station: 355-373. [1496] 26. MacHutchon, A. Grant. 1989. Spring and summer food habits of black bears in the Pelly River Valley, Yukon. Northwest Science. 63(3): 116-118. [12249] 27. Martell, Arthur M.; Dickinson, Dawn M.; Casselman, Lisa M. 1984. Wildlife of the Mackenzie Delta region. Occasional Publ. No. 15. Edmonton, AB: The University of Alberta, Boreal Institute for Northern Studies. 214 p. [15014] 28. Martin, Alexander C.; Zim, Herbert S.; Nelson, Arnold L. 1951. American wildlife and plants. New York: McGraw-Hill Book Company, Inc. 500 p. [4021] 29. 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: Walker, D. G.; Powter, C. B.; Pole, M. W., compilers. Reclamation, a global perspective: Proceedings of the conference; 1989 August 27-31; Calgary, AB. Rep. No. RRTAC 89-2. Vol. 1. Edmonton, AB: Alberta Land Conservation and Reclamation Council: 81-90. [14363] 30. Meehan, William R. 1974. The forest ecosystem of southeast Alaska: 4. Wildlife habitats. Gen. Tech. Rep. PNW-16. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 32 p. [13479] 31. Miller, Donald R. 1976. Taiga winter range relationships and diet. Canadian Wildlife Service Rep. Series No. 36. Ottawa, ON: Environment Canada, Wildlife Service. 42 p. (Biology of the Kaminuriak population of barren-ground caribou; pt 3). [13007] 32. Morin, Hubert; Payette, Serge. 1988. Buried seed populations in the montane, subalpine, and alpine belts of Mont Jacques-Cartier, Quebec. Canadian Journal of Botany. 66: 101-107. [6376] 33. Pollett, Frederick C. 1972. Classification of peatlands in Newfoundland. In: Proceedings, 4th International Peat Congress. 1: 101-110. [15403] 34. Racine, Charles H. 1979. The 1977 tundra fires in the Seward Peninsula, Alaska: effects and initial revegetation. BLM-Alaska Technical Report 4. U.S. Department of the Interior, Bureau of Land Management, Alaska State Office. 51 p. [8330] 35. Racine, Charles H. 1981. Tundra fire effects on soils and three plant communities along a hill-slope gradient in the Seward Peninsula, Alaska. Arctic. 34(1): 71-84. [7233] 36. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843] 37. Reynolds, Keith M. 1990. Preliminary classification of forest vegetation of the Kenai Penninsula, Alaska. Res. Pap. PNW-RP-424. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station. 67 p. [14581] 38. Roland, A. E.; Smith, E. C. 1969. The flora of Nova Scotia. Halifax, NS: Nova Scotia Museum. 746 p. [13158] 39. Rowe, J. S. 1983. Concepts of fire effects on plant individuals and species. In: Wein, Ross W.; MacLean, David A., eds. SCOPE 18: The role of fire in northern circumpolar ecosystems. Chichester; New York: John Wiley & Sons: 135-154. [2038] 40. Schoen, John W.; Kirchhoff, Matthew D. 1990. Seasonal habitat use by Sitka black-tailed deer on Admiralty Island, Alaska. Journal of Wildlife Management. 54(3): 371-378. [11940] 41. Scotter, George W. 1967. The winter diet of barren-ground caribou in northern Canada. Canadian Field-Naturalist. 81: 33-39. [16672] 42. Soper, James H.; Heimburger, Margaret L. 1982. Shrubs of Ontario. Life Sciences Misc. Publ. Toronto, ON: Royal Ontario Museum. 495 p. [12907] 43. U.S. Department of Agriculture, Soil Conservation Service. 1982. National list of scientific plant names. Vol. 1. List of plant names. SCS-TP-159. Washington, DC. 416 p. [11573] 44. Van Dersal, William R. 1938. Native woody plants of the United States, their erosion-control and wildlife values. Washington, DC: U.S. Department of Agriculture. 362 p. [4240] 45. Viereck, Leslie A. 1982. Effects of fire and firelines on active layer thickness and soil temperatures in interior Alaska. In: Proceedings, 4th Canadian permafrost conference; 1981 March 2-6; Calgary, AB. The Roger J.E. Brown Memorial Volume. Ottawa, ON: National Research Council of Canada: 123-135. [7303] 46. 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. [2431] 47. 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. [6884] 48. Viereck, Leslie A.; Schandelmeier, Linda A. 1980. Effects of fire in Alaska and adjacent Canada--a literature review. BLM-Alaska Tech. Rep. 6. Anchorage, AK: U.S. Department of the Interior, Bureau of Land Mangement, Alaska State Office. 124 p. [7075] 49. Voss, Edward G. 1972. Michigan flora. Part I. Gymnosperms and monocots. Bloomfield Hills, MI: Cranbrook Institute of Science; Ann Arbor, MI: University of Michigan Herbarium. 488 p. [11471] 50. Wein, R. W. 1974. Recovery of vegetation in arctic regions after burning. Rep. 74-6. Ottawa, ON: Canadian Task Force on Northern Oil Development. 41 p. [13001] 51. West, Stephen D. 1982. Dynamics of colonization and abundance in central Alaskan populations of the northern red-backed vole, Clethrionomys rutilus. Journal of Mammalogy. 63(1): 128-143. [7300]

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