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Introductory

SPECIES: Salix glauca | Grayleaf Willow
ABBREVIATION : SALGLA SYNONYMS : Salix pseudolapponum Seemann SCS PLANT CODE : SAGL SAGLA SAGLV COMMON NAMES : grayleaf willow gray willow gray-leaved willow glaucous willow white willow TAXONOMY : The currently accepted scientific name of grayleaf willow is Salix glauca L. [3,18]. Because it exhibits considerable geographic variation across its extensive range, it has been divided into numerous varieties or subspecies. Argus [3] recognizes three varieties: var. villosa (Hooker) Anderson var. acutifolia (Hooker) Schneider var. glauca Alternately, Hulten [18] recognizes four subspecies: ssp. acutifolia (Hook.) Hult. ssp. callicarpaea (Trautv.) Bocher ssp. desertorum (Richards.) Anderss. ssp. glabrescens (Anderss.) Hult. LIFE FORM : Tree, Shrub FEDERAL LEGAL STATUS : No special status OTHER STATUS : Grayleaf willow is ranked as a critically endangered plant in Idaho and Washington under The Nature Conservancy's Natural Heritage ranking system [43]. COMPILED BY AND DATE : Ronald Uchytil, February 1992 LAST REVISED BY AND DATE : NO-ENTRY AUTHORSHIP AND CITATION : Uchytil, Ronald J. 1992. Salix glauca. In: Remainder of Citation

DISTRIBUTION AND OCCURRENCE

SPECIES: Salix glauca | Grayleaf Willow
GENERAL DISTRIBUTION : Grayleaf willow grows throughout most of Alaska except for the Alleutian Islands and along the southeastern coast [35]. It grows through much of northern Canada from Newfoundland northwest to the northern Yukon Territory, and south to southern British Columbia and Alberta. In the contiguous United States, it grows in alpine and subalpine habitats in Montana, Wyoming, eastern Idaho, Colorado, Utah, and northern New Mexico [10]. ECOSYSTEMS : FRES11 Spruce - fir FRES23 Fir - spruce FRES44 Alpine STATES : AK CO ID MT NM UT WY AB BC LB MB NB NT ON PQ SK YT ADMINISTRATIVE UNITS : CEBR DENA GLAC GRTE LACL ROMO WRST YELL YUCH BLM PHYSIOGRAPHIC REGIONS : 8 Northern Rocky Mountains 9 Middle Rocky Mountains 10 Wyoming Basin 11 Southern Rocky Mountains KUCHLER PLANT ASSOCIATIONS : K015 Western spruce - fir forest K052 Alpine meadows and barren SAF COVER TYPES : 12 Black spruce 107 White spruce 201 White spruce 204 Black spruce 206 Engelmann spruce - subalpine fir 251 White spruce - aspen 253 Black spruce - white spruce SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : In Alaska and northwestern Canada, grayleaf willow dominates or codominates numerous seral willow (Salix spp.) and mixed-shrub floodplain communities. Riparian community associates include Alaska willow (S. alaxensis), littletree willow (S. arbusculoides), Richardson willow (S. lanata), diamondleaf willow (S. planifolia), and green alder (Alnus crispa) [34]. It also codominates in some mixed-shrub tundra communities with birches (Betula spp.), alders (Alnus spp.), and other willows [34]. In the Rocky Mountain States, grayleaf willow/tufted hairgrass (Deschampsia cespitosa) communities occupy well-drained, open alpine and upper subalpine habitats [20,27]. Grayleaf willow occurs as scattered individuals in many boreal forests and woodlands. It is seldom an understory dominant, except in early seral stages. Douglas [11], however, described a 130- to 160-year-old white spruce (Picea glauca)/grayleaf willow community in southwestern Yukon Territory. Classifications listing grayleaf willow as a dominant in community types (cts) and habitat types (hts) are presented below: Area Classification Authority AK general veg. cts Viereck & Dyrness 1980 sw YT montane veg. cts Douglas 1974 CO: Gunnison & general veg. hts Komarkova 1986 Uncompahgre NF UT, se ID riparian cts Padgett & others 1989 Forest Service R-2 forest, shrub, grass Wasser & Hess 1982 & forb hts

VALUE AND USE

SPECIES: Salix glauca | Grayleaf Willow
WOOD PRODUCTS VALUE : NO-ENTRY IMPORTANCE TO LIVESTOCK AND WILDLIFE : Grayleaf willow is a moderately important moose browse in some areas primarily because of its abundance [11,29]; in other areas it is poorly utilized [14,25]. Caribou use is probably moderate at best, and primarily in the summer [7]. In some areas, grayleaf willow makes up a large part of the winter diet of snowshoe hares [31]. Willows are generally a preferred food and building material of beaver [1]. Willow shoots, catkins, leaves, and buds are eaten by numerous small mammals and birds [16]. PALATABILITY : In interior Alaska, grayleaf willow is less palatable to moose than Alaska willow, littletree willow, diamondleaf willow, or sandbar willow (S. interior). Moose browse grayleaf willow lightly in comparison with the others in areas where they grow together [25]. Grayleaf is more palatable to moose than aspen (Populus tremuloides), balsam poplar (Populus balsamifera), or paper birch (Betula papyrifera) [38]. NUTRITIONAL VALUE : Grayleaf willow is a relatively high-quality food for wintering ungulates. Winter stem crude protein content is about 6.4 percent. Grayleaf willow is also a good source of calcium and phosphorus, and its digestibility is relatively high [29,30]. COVER VALUE : In thickets grayleaf willow may provide cover for small animals, but its small stature limits its value as cover for large mammals. VALUE FOR REHABILITATION OF DISTURBED SITES : Densmore and Zasada [8] reported that under laboratory conditions grayleaf willow stem cuttings taken in the fall or spring rarely produce roots and, therefore, do not recommend planting grayleaf willow stem cuttings for rehabilitation purposes. However, grayleaf willow stem cuttings were successfully used to revegetate unstable sand dunes in northern Alberta [42]. Seeding disturbed sites with this species may be a useful establishment measure. Grayleaf willow has been observed naturally invading barrow pits and mine tailings in arctic regions [19,35]. The seeds may be stored for up to 3 years, and require cold stratification before sowing [40,41]. OTHER USES AND VALUES : All willows produce salacin, which is closely related chemically to aspirin. Native Americans used various preparations from willow to treat tooth ache, stomach ache, diarrhea, dysentery, and dandruff [26]. Native Americans also used flexible willow stems for making baskets, bows, arrows, scoops, snares, and fish and muskrat traps [17,21]. MANAGEMENT CONSIDERATIONS : Grayleaf willow is tolerant of heavy browsing [42].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Salix glauca | Grayleaf Willow
GENERAL BOTANICAL CHARACTERISTICS : Grayleaf willow commonly grows as an erect shrub 3 to 4 feet (0.9-1.2 m) tall. On exposed tundra sites it grows as a low, semiprostrate shrub, and on favorable sites it sometimes grows up to 20 feet (6 m) in height and 5 inches (12 cm) in diameter [35]. The bark is gray and smooth but may become rough and furrowed on larger individuals. Male and female flowers occur on separate plants in 3/4- to 2-inch-long (2-5 cm) catkins that persist over the summer. The fruit is a 1/32- to 1/16-inch-long (0.8-1.6 mm) two-valved capsule [35]. Two growth forms occur in the Rocky Mountains. In somewhat sheltered locations in subalpine environments, plants are upright and taller, while semiprostrate plants that are often difficult to distinguish from arctic willow (S. arctica) grow in more exposed, alpine situations [10]. RAUNKIAER LIFE FORM : Phanerophyte REGENERATION PROCESSES : Grayleaf willow's primary mode of reproduction is sexual. It produces an abundance of small, lightweight seeds. Like most willows, it probably begins seed production at an early age (between 2 and 10 years) [16]. Seeds are not shed as they ripen but remain on the plant throughout the summer and are dispersed in the fall. Each seed has a cottony down which aids in dispersal by wind and water. Unlike willow seeds dispersed in summer, grayleaf willow seeds overwinter under snow and germinate in the spring soon after snowmelt [9,41]. This cold stratification promotes good germination; seeds germinate over a wide range of temperatures (from 41 to 77 degrees F [5-25 C]) [9]. Spring germination is advantageous in arctic and alpine environments; the growing season of grayleaf willow seedlings is 3 to 6 weeks longer than that of summer-disersing willows [9]. Exposed mineral soils are required for good germination and seedling establishment [16]. Forest litter generally inhibits germination and establishment. Vegetative Reproduction: Grayleaf willow sprouts from the root crown or stembase if aboveground stems are broken or destroyed by cutting or fire [16]. SITE CHARACTERISTICS : In Alaska and northern Canada, grayleaf willow grows on both uplands and lowlands. In arctic tundra it often grows along river and streambanks, on sandy and gravelly floodplains, and on old benches [3,35]. In boreal environments, it grows as scattered shrubs in white and black spruce (Picea mariana) woodlands, in black spruce muskegs, and on river floodplains [3,35]. In the Rocky Mountains grayleaf willow is restricted to open, alpine and subalpine habitats that commonly have rocky, well-drained soils [10,27]. SUCCESSIONAL STATUS : Grayleaf willow is an early seral species. It pioneers freshly deposited river alluvium, glacial outwash, and disturbed areas with exposed mineral soil, such as road cuts and mine tailings [35]. It is also common in spruce woodlands following fire, especially in stands about 20 to 30 years old [14,23]. It has been found in 160-year-old open spruce woodlands [11], but it is usually displaced in densely forested stands because of its shade intolerance. SEASONAL DEVELOPMENT : Grayleaf willow catkins appear with the leaves. In Alaska and the Yukon, flowering generally occurs in June, the fruits ripen in July and August, and the seeds are dispersed in late August and September [9,35].

FIRE ECOLOGY

SPECIES: Salix glauca | Grayleaf Willow
FIRE ECOLOGY OR ADAPTATIONS : Grayleaf willow is a fire-adapted species. Most plants sprout from the root crown following top-kill by fire. Viereck and Schandelmeier [36] reported that even old, decadent willows sprouted prolifically immediately after fire. The sprouting ability of willows is apparently more vigorous and prolific than that of birches or alders [36]. Grayleaf willow's abundant, wind-dispersed seeds are important in colonizing burned areas. Seeds are dispersed in the fall, overwinter under snow, and germinate in the spring. Thus seedling establishment cannot begin until postfire year 2. POSTFIRE REGENERATION STRATEGY : survivor species; on-site surviving root crown or caudex off-site colonizer; seed carried by wind; postfire years 1 and 2 off-site colonizer; seed carried by animals or water; postfire yr 1&2

FIRE EFFECTS

SPECIES: Salix glauca | Grayleaf Willow
IMMEDIATE FIRE EFFECT ON PLANT : Grayleaf willows that occur in white and black spruce forests can be killed by severe fires that completely remove soil organic layers and char the roots [39]. Less severe fires only top-kill plants. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Grayleaf willow is one of the most abundant willows following fire in white spruce forests of interior Alaska [14]. There are commonly several thousand stems per acre by 10 years after fire. However, its abundance is short-lived, and it is often absent by year 40 as dense white spruce sapling stands develop [14]. Grayleaf willow is also common on mesic black spruce sites after fire. One ll-year-old burn near the Tanana River had about 4,700 grayleaf, Alaska, and diamondleaf willow stems per acre (11,500/ha), and lesser amounts of spruce and poplar [38]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : Fire severity can affect the mode of willow postfire recovery. Following light fires most willows recover quickly, sending up new shoots from undamaged root crowns. Few if any seedlings establish following this type of burn because organic soil layers are only partially consumed, which prevents seedling establishment. Following severe fires, however, the primary mode of recovery is seedling establishment. Severe fires that burn deep into organic soils kill willows but expose mineral soils, which provide excellent seedbeds. FIRE MANAGEMENT CONSIDERATIONS : NO-ENTRY

REFERENCES

SPECIES: Salix glauca | Grayleaf Willow
REFERENCES : 1. Allen, Arthur W. 1983. Habitat suitability index models: beaver. FWS/OBS-82/10.30 (Revised). Washingtion, DC: U.S. Department of the Interior, Fish and Wildlife Service. 20 p. [11716] 2. Viereck, Leslie A. 1966. Plant succession and soil development on gravel outwash of the Muldrow Glacier, Alaska. Ecological Monographs. 36(3): 181-199. [12484] 3. Argus, George W. 1973. The genus Salix in Alaska and the Yukon. Publications in Botany, No. 2. Ottowa, ON: National Museums of Canada, National Museum of Natural Sciences. 279 p. [6167] 4. 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] 5. Black, R. A.; Bliss, L. C. 1978. Recovery sequence of Picea mariana - Vaccinium uliginosum forests after burning near Inuvik, Northwest Territories, Canada. Canadian Journal of Botany. 56: 2020-2030. [7448] 6. Brayshaw, T. Christopher. 1976. Catkin bearing plants of British Columbia. Occas. Pap. No. 18. Victoria, BC: The British Columbia Provincial Museum. 176 p. [6170] 7. Cody, W. J. 1965. Plants of the Mackenzie River Delta and Reindeer Grazing Preserve. Ottawa, ON: Canada Department of Agriculture, Research Branch, Plant Research Institute. 56 p. [13122] 8. Densmore, R.; Zasada, J. C. 1978. Rooting potential of Alaskan willow cuttings. Canadian Journal of Forest Research. 8: 477-479. [5428] 9. Densmore, Roseann; Zasada, John. 1983. Seed dispersal and dormancy patterns in northern willows: ecological and evolutionary significance. Canadian Journal of Botany. 61: 3207-3216. [5027] 10. Dorn, Robert D. 1977. Willows of the Rocky Mountain States. Rhodora. 79: 390-429. [6000] 11. Douglas, George W. 1974. Montane zone vegetation of the Alsek River region, southwestern Yukon. Canadian Journal of Botany. 52: 2505-2532. [17283] 12. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905] 13. Flook, Donald R. 1964. Range relationships of some ungulates native to Banff and Jasper National Parks, Alberta. In: Crisp, D. J., ed. Grazing in terrestrial and marine environments. [Place of publication unknown]: [Publisher unknown]. 119-128. [15688] 14. Foote, M. Joan. 1983. Classification, description, and dynamics of plant communities after fire in the taiga of interior Alaska. Res. Pap. PNW-307. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Forest and Range Experiment Station. 108 p. [7080] 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. Haeussler, S.; Coates, D. 1986. Autecological characteristics of selected species that compete with conifers in British Columbia: a literature review. Land Management Report No. 33. Victoria, BC: Ministry of Forests, Information Services Branch. 180 p. [1055] 17. Holloway, Patricia S.; Alexander, Ginny. 1990. Ethnobotany of the Fort Yukon region, Alaska. Economic Botany. 44(2): 214-225. [13625] 18. Hulten, Eric. 1968. Flora of Alaska and neighboring territories. Stanford, CA: Stanford University Press. 1008 p. [13403] 19. 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] 20. Komarkova, Vera. 1986. Habitat types on selected parts of the Gunnison and Uncompahgre National Forests. Final Report Contract No. 28-K2-234. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station. 270 p. [1369] 21. Kovalchik, Bernard L.; Hopkins, William E.; Brunsfeld, Steven J. 1988. Major indicator shrubs and herbs in riparian zones on National Forests of central Oregon. R6-ECOL-TP-005-88. Portland, OR: U.S. Department of Agriculture, Forest Service, Pacific Northwest Region. 159 p. [8995] 22. 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] 23. 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] 24. 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] 25. Milke, Gary Clayton. 1969. Some moose-willow relationships in the interior of Alaska. College, AK: University of Alaska. 79 p. Thesis. [15801] 26. Mozingo, Hugh N. 1987. Shrubs of the Great Basin: A natural history. Reno, NV: University of Nevada Press. 342 p. [1702] 27. Padgett, Wayne G.; Youngblood, Andrew P.; Winward, Alma H. 1989. Riparian community type classification of Utah and southeastern Idaho. R4-Ecol-89-01. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Region. 191 p. [11360] 28. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843] 29. Risenhoover, Kenneth L. 1989. Composition and quality of moose winter diets in interior Alaska. Journal of Wildlife Management. 53(3): 568-577. [14930] 30. Scotter, George W. 1972. Chemical composition of forage plants from the Reindeer Preserve, Northwest Territories. Arctic. 25(1): 21-27. [16563] 31. Smith, J. N. M.; Krebs, C. J.; Sinclair, A. R. E.; Boonstra, R. 1988. Population biology of snowshoe hares. II. Interactions with winter food plants. Journal of Animal Ecology. 57: 269-286. [6713] 32. Soper, James H.; Heimburger, Margaret L. 1982. Shrubs of Ontario. Life Sciences Misc. Publ. Toronto, ON: Royal Ontario Museum. 495 p. [12907] 33. 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] 34. 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] 35. 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] 36. 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] 37. Wasser, C. H.; Hess, Karl. 1982. The habitat types of Region II, U.S. Forest Service: a synthesis. Final Report Cooperative Agreement No. 16-845-CA. Lakewood, CO: U.S. Department of Agriculture, Forest Service, Region 2. 140 p. [5594] 38. Wolff, Jerry O.; Zasada, John C. 1979. Moose habitat and forest succession on the Tanana river floodplain and Yukon-Tanana upland. In: Proceedings, North American Moose Conference and Workshop No 15; [Date of conference unknown]; Kenai, AK. [Place of publication unknown]. [Publisher unknown]. 213-244. [6860] 39. Zasada, J. 1986. Natural regeneration of trees and tall shrubs on forest sites in interior Alaska. In: Van Cleve, K.; Chapin, F. S., III; Flanagan, P. W.; [and others], eds. Forest ecosystems in the Alaska taiga: A synthesis of structure and function. New York: Springer-Verlag: 44-73. [2291] 40. Zasada, John C.; Densmore, Roseann. 1980. Alaskan willow and balsam poplar seed viability after 3 years' storage. Tree Planters' Notes. 31((2)): 9-10. [15805] 41. Zasada, J. C.; Viereck, L. A. 1975. The effect of temperature and stratification on germination on selected members of Salicaceae in interior Alaska. Canadian Journal of Forest Research. 5(2): 333-337. [6989] 42. Foiles, Marvin W.; Curtis, James D. 1973. Regeneration of ponderosa pine in the northern Rocky Mountain- Intermountain region. Res. Pap. INT-1456. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Forest and Range Experiment Station. 44 p. [15640] 43. Washington Natural Heritage Program, compiler. 1994. Endangered, threatened, and sensitive vascular plants of Washington. Olympia, WA: Department of Natural Resources. 52 p. [25413]

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