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Introductory

SPECIES: Salix lanata ssp. richardsonii | Richardson Willow
ABBREVIATION : SALLANR SYNONYMS : NO-ENTRY SCS PLANT CODE : SALA4 COMMON NAMES : Richardson willow woolly willow TAXONOMY : The currently accepted scientific name for Richardson willow is Salix lanata L. ssp. richardsonii (Hook.) A. Skv. There are no varieties, forms, or natural hybrids; although hybridization and introgression with S. barclayi and with S. alaxensis var. alaxensis has been suggested [1]. S. lanata ssp. richardsonii is the western North American-eastern Asian race of S. lanata; other subspecies are S. lanata ssp. lanata L., the Eruasian race and S. lanata. ssp. calcicola (Fern and Weig.) Hult., the eastern Canadian Arctic race [1,5,14,15]. LIFE FORM : Shrub FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY COMPILED BY AND DATE : Lora L. Esser, September 1992 LAST REVISED BY AND DATE : NO-ENTRY AUTHORSHIP AND CITATION : Esser, Lora L. 1992. Salix lanata. In: Remainder of Citation

DISTRIBUTION AND OCCURRENCE

SPECIES: Salix lanata ssp. richardsonii | Richardson Willow
GENERAL DISTRIBUTION : Richardson willow occurs from the Arctic Coast southward through most of central and south-central Alaska; it does not occur in the western Alaskan Peninsula or Kenai Peninsula. It extends eastward across northern Canada to the Baffin Islands; southward to northwest Hudson Bay; and west to northern Britsh Columbia [1,5,19,35]. ECOSYSTEMS : FRES44 Alpine STATES : AK BC NT YT ADMINISTRATIVE UNITS : DENA GLBA BLM PHYSIOGRAPHIC REGIONS : NO-ENTRY KUCHLER PLANT ASSOCIATIONS : K052 Alpine meadows and barren SAF COVER TYPES : 201 White spruce 202 White spruce - paper birch 203 Balsam poplar 204 Black spruce 217 Aspen 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 : Richardson willow is a common, thicket-forming shrub of streambanks and moist slopes in the Arctic and above timberline where it is often associated with alders (Alnus spp.) and birch (Betula spp.); it is also found in open spruce (Picea spp.) stands and old burns at lower elevations [5,35]. Richardson willow can also be found in floodplain thickets on rivers and grows on newly exposed alluvial deposits that are periodically flooded [12,35]. Individual shrubs can be found on south-facing steppes, on pingos found in permafrost regions, and on dry, rock outcrops [13,39]. Published classifications describing Richardson willow as a codominant in community types are listed below: Arctic community types of Northwest Alaska [12].

VALUE AND USE

SPECIES: Salix lanata ssp. richardsonii | Richardson Willow
WOOD PRODUCTS VALUE : NO-ENTRY IMPORTANCE TO LIVESTOCK AND WILDLIFE : Richardson willow is an important food source for moose, caribou, mule deer, muskrat, and beaver [13,23,24,26,31]. During the winter in Alaska, moose feed primarily on shoots of current growth of willow (Salix spp.), quaking aspen (Populus tremuloides), paper birch (Betula papyrifera), and balsam poplar (Populus balsamifera) growing as shrubs or saplings in young, seral communities. Of the preceding species, willow is the most preferred by moose [23]. Of the more than 20 species of willow found in Denali National Park, Richardson willow was one of three species utilized the most [26]. In one study, willows comprised 94 percent of total biomass consumed by moose from January to April, with Richardson willow comprising 6.1 percent [31]. PALATABILITY : Richardson willow is considered moderately palatable. In one study, Richardson willow was consumed by moose to a greater extent when occurring in mixed stands with highly preferred species than when growing in pure stands [26]. Willow palatability increases as the season progresses [26]. NUTRITIONAL VALUE : Nutrient composition of Richardson willow consumed by moose in Denali National Park, Alaska, from January to April, 1984, was as follows [31]: gross energy: 5.08 kcal/g percent in vitro digestible organic matter: 39.4 percent of dry matter: 5.9 percent lignin: 19.4 percent ash: 2.8 percent ether extract: 8.3 The spring protein concentration of willow twigs is three times greater than that of willow bark. Calcium concentration is greater in bark than in twigs, and phosphorus concentration is greater in twigs than in bark [27]. COVER VALUE : Richardson willow characteristically produces dense thickets along streams and rivers, which provide thermal and hiding cover for mule deer. Branches are used by beaver in the construction of dams and lodges [24]. Richardson willow can stabilize streambanks when thickets are dense by moderately undercutting the bank, which provides hiding and resting cover for fish [24]. VALUE FOR REHABILITATION OF DISTURBED SITES : Richardson willow is useful in stabilizing streambanks and providing erosion control on severely disturbed sites [11]. Willow species are the most important colonizers of disturbed sites in the Alaskan taiga because of their ability to produce root and root crown shoots, which provide for quick recovery [13,37]. At an Alaskan arctic coastal plain site, Richardson willow colonized bare areas of tundra after removal of debris that had been there for 30 years. High percentage cover occurred on sites with favorable moisture and nutrient regimes [8]. Richardson willow was found to be a poor colonizer in areas where crude oil was spilled; plant recovery and establishment were extremely slow on these spills [16]. Willow planting, using stem cuttings, has been recognized as a valuable tool for restoring riparian habitat. Restoration of riparian habitat benefits a large number of wildlife species [24]. OTHER USES AND VALUES : Richardson willow is an important nectar producer for bees [29]. Tough, flexible shoots of Richardson willow can be woven into baskets and furniture [2]. Native Americans used the broth from boiled bark for sore throats and tuberculosis [2 ]. MANAGEMENT CONSIDERATIONS : Richardson willow is an important source of browse for moose in Alaska. If the management objective is to provide moose habitat and if environmental manipulation of species composition is possible, then only the growth of preferred species, such as Richardson willow, should be considered [26].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Salix lanata ssp. richardsonii | Richardson Willow
GENERAL BOTANICAL CHARACTERISTICS : Richardson willow is an erect, much-branched shrub usually forming dense clumps 3 to 6 feet (1-2 m) tall, sometimes to 15 feet (4.5 m) [5,35]. Young twigs are stout and densely hairy; older twigs are glabrous. The bark of Richardson willow is smooth [35]. Shrubs are composed of light wood that becomes brittle with age; a single trunk rarely survives 60 years [2]. In silty loam containing much organic matter, the roots of Richardson willow are numerous in the top 7.5 inches (19 cm) of soil, but become less abundant at 9 inches (23 cm). In frozen ground, roots of Richardson willow do not exceed 9 inches (23 cm), but as the ground thaws roots will grow up to 17 inches (43 cm) deep [12]. RAUNKIAER LIFE FORM : Phanerophyte REGENERATION PROCESSES : Sexual reproduction: Richardson willow is dioecious. The fruit of these plants is contained in a capsule that splits in half to release many seeds that are then dispersed by wind or water [11,33]. Optimum seed production occurs between 2 and 10 years [11]. Bees are the chief pollinating agents [11]. The seeds of Richardson willow are short-lived, germinating immediately on moist surfaces [21]. Seed germination occurs over a broad temperature range, 41 to 77 degrees Fahrenheit (5-25 degrees C). This appears to be a compensatory mechanism due to the short seed life [6]. Germination of Richardson willow seeds occurs best in moist, exposed mineral substrates that receive direct sunlight [11]. Vegetative reproduction: Richardson willow will sprout from the root crown or basal stem [11]. It will root readily from stem cuttings or from root and stem fragments buried in moist soil. Damaged and cut stems produce prolific sprouts from the stembase or root collar [11]. SITE CHARACTERISTICS : In Alaska and northern Canada, Richardson willow is found in wet areas such as heaths, riverbeds, and streams; it is also found in the open tundra, in pingos, and in mountains to at least 5,578 feet (1,700 m) [14,39]. In interior Alaska, Richardson willow occurs in glacial drift, outwash deposition areas, and on old river floodplains with considerable variation in habitat conditions [4]. Soils: Richardson willow grows best in moist, alluvial bottomlands but is also found in well-drained sandy or gravelly substrates. The general pH range for willows is 5.5 to 7.5 [11]. Growth of Richardson willow is severely reduced when water levels are maintained at or above the root crown for extended periods [11]. Plant associates: Richardson willow is commonly associated with the following species: quaking aspen, white spruce (Picea glauca), black spruce (P. mariana), Alaska paper birch (Betula resinifera), feltleaf willow (Salix alaxensis), diamondleaf willow (S. pulchra), netleaf willow (S. reticulata), American green alder (Alnus crispa), Sitka alder (A. fruticosa), bog birch (Betula glandulosa), lichens (Ericaceae spp.), huckleberry (Vaccinium spp.), bluejoint grass (Calamagrostis canadensis), bluegrass (Poa spp.), sedges (Carex spp.), and mosses (Polytrichum spp.) [12,27]. SUCCESSIONAL STATUS : Obligate Initial Community Species Richardson willow is an early successional species on moist sites and, once established, may persist in areas with frequent disturbances such as fires or flooding [7,8]. It also becomes important in the later stages of riparian succession. Successional studies have shown that once silt accumulates, Richardson willow will become established quickly [6]. Richardson willow was the first shrub to invade flood meadows, after grasses and horsetail, on sandy alluvium in the tundra [3]. Richardson willow has low shade tolerance and therefore loses dominance on sites that are heavily forested or succeeded by more shade-tolerant species [38]. SEASONAL DEVELOPMENT : Richardson willow flowers from May through July or August [6,20]. The fruit ripens soon after flowering, followed by seed dispersal in early to midsummer [11].

FIRE ECOLOGY

SPECIES: Salix lanata ssp. richardsonii | Richardson Willow
FIRE ECOLOGY OR ADAPTATIONS : Richardson willow sprouts rapidly from basal stems and roots after fire [28,32]. It produces minute, hairy seeds that are easily disseminated by wind, and which are important in reestablishment [21,32]. POSTFIRE REGENERATION STRATEGY : Ground residual colonizer (on-site, initial community) Secondary colonizer - off-site seed

FIRE EFFECTS

SPECIES: Salix lanata ssp. richardsonii | Richardson Willow
IMMEDIATE FIRE EFFECT ON PLANT : Richardson willow is a fire-tolerant species that sprouts readily from the root or root crown after being top-killed by fire [32]. If soil organic layers are completely removed by fire, then the roots of Richardson willow will not be able to sprout [11]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Richardson willow is an early successional species on burned sites because of its ability to sprout vigorously from the root crown or roots following fire [20,36]. Invasion by willows after fire depends on the time of year of the fire, weather, and the absence or presence of a mineral seedbed [36]. Richardson willow seeds need a nutrient-rich mineral seedbed to germinate. The chance of Richardson willow establishing after a fire lessens as available mineral soil seedbeds become occupied by faster growing herbaceous species and mosses [38]. Fire severity can affect willow postfire recovery. High-severity fires can damage the roots to the point of no recovery [20,38]. Following low-severity fires most willows will recover quickly because of their ability to send up new roots from the root crown [38]. Intense burning can completely kill willows [20]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : NO-ENTRY FIRE MANAGEMENT CONSIDERATIONS : Prescribed fire is widely used as a wildlife management tool to rejuvenate decadent willow stands and stimulate sprouting [11,25]. Early seral stage communities created by fire can increase the carrying capacity of winter range for moose in interior Alaska [40]. Recurring fires within some parts of the boreal forest have allowed aspen and willow to replace coniferous forests [32]. The tendency of willows to expand quickly following fires and other disturbances and to form dense thickets inhibits natural regeneration of conifers [11]. Prescribed burning can reduce initial competition from willow in areas to be planted with cultivated species [11].

REFERENCES

SPECIES: Salix lanata ssp. richardsonii | Richardson Willow
REFERENCES : 1. 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] 2. Arno, Stephen F.; Hammerly, Ramona P. 1977. Northwest trees. Seattle, WA: The Mountaineers. 222 p. [4208] 3. Bliss, L. C.; Cantlon, J. E. 1957. Succession on river alluvium in northern Alaska. American Midland Naturalist. 58(2): 452-469. [14931] 4. 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. [13877] 5. Brayshaw, T. Christopher. 1976. Catkin bearing plants of British Columbia. Occas. Pap. No. 18. Victoria, BC: The British Columbia Provincial Museum. 176 p. [6170] 6. Densmore, R. V.; Neiland, B. J.; Zasada, J. C.; Masters, M. A. 1987. Planting willow for moose habitat restoration on the North Slope of Alaska, U.S.A. Arctic and Alpine Research. 19(4): 537-543. [6080] 7. Dorn, Robert D. 1976. A synopsis of American Salix. Canadian Journal of Botany. 54: 2769-2789. [4457] 8. Ebersole, James J. 1987. Short-term vegetation recovery at an Alaskan arctic coastal plain site. Arctic and Alpine Research. 19(4): 442-450. [9476] 9. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905] 10. 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] 11. 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] 12. Hanson, Herbert C. 1953. Vegetation types in northwestern Alaska and comparisons with communities in other arctic regions. Ecology. 34(1): 111-140. [9781] 13. Henry, G. H. R.; Gunn, A. 1991. Recovery of tundra vegetation after overgrazing by caribou in arctic Canada. Arctic. 44(1): 38-42. [14747] 14. Hulten, Eric. 1968. Flora of Alaska and neighboring territories. Stanford, CA: Stanford University Press. 1008 p. [13403] 15. Kartesz, John T.; Kartesz, Rosemarie. 1980. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. Volume II: The biota of North America. Chapel Hill, NC: The University of North Carolina Press; in confederation with Anne H. Lindsey and C. Richie Bell, North Carolina Botanical Garden. 500 p. [6954] 16. Kershaw, G. Peter; Kershaw, Linda J. 1986. Ecological characteristics of 35-year-old crude-oil spills in tundra plant communities of the Mackenzie Mountains, N.W.T. Canadian Journal of Botany. 64: 2935-2947. [12972] 17. 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] 19. Little, Elbert L., Jr. 1979. Checklist of United States trees (native and naturalized). Agric. Handb. 541. Washington, DC: U.S. Department of Agriculture, Forest Service. 375 p. [2952] 20. Lotan, James E.; Alexander, Martin E.; Arno, Stephen F.; [and others]. 1981. Effects of fire on flora: A state-of-knowledge review. National fire effects workshop; 1978 April 10-14; Denver, CO. Gen. Tech. Rep. WO-16. Washington, DC: U.S. Department of Agriculture, Forest Service. 71 p. [1475] 21. 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] 22. 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] 23. Machida, Steven. 1979. Differential use of willow species by moose in Alaska. Fairbanks, AK: University of Alaska. 97 p. Thesis. [15098] 24. McCluskey, D. Cal; Brown, Jack; Bornholdt, Dave; [and others]. 1983. Willow planting for riparian habitat improvement. Tech. Note 363. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 21 p. [6408] 25. Meidinger, D.; Lewis, T.; Kowall, R. 1986. Biogeoclimatic zones and subzones of the northern portion of the Mackenzie Timber Supply Area, British Columbia. In: Northern Fire Ecology Project: Northern Mackenzie Timber Supply Area. Victoria, BC: Province of British Columbia, Ministry of Forests. 44 p. [9204] 26. Milke, Gary Clayton. 1969. Some moose-willow relationships in the interior of Alaska. College, AK: University of Alaska. 79 p. Thesis. [15801] 27. Miquelle, Dale G.; Van Ballenberghe, Victor. 1989. Impact of bark stripping by moose on aspen-spruce communities. Journal of Wildlife Management. 53(3): 577-586. [8911] 28. Parminter, John. 1984. Fire-ecological relationships for the biogeoclimatic zones of the northern portion of the Mackenzie Timber Supply Area: summary report. In: Northern Fire Ecology Project: Northern Mackenzie Timber Supply Area. Victoria, BC: Province of British Columbia, Ministry of Forests. 59 p. [9205] 29. Petersen, Stephen F. 1989. Beekeeping under northern lights. American Bee Journal. 129(1): 33-35. [12332] 30. Ferguson, Dennis E.; Boyd, Raymond J. 1988. Bracken fern inhibition of conifer regeneration in northern Idaho. Ogden, UT: U.S. Department of Agriculture, Forest Service, Intermountain Research Station. 11 p. [2834] 31. Risenhoover, Kenneth L. 1989. Composition and quality of moose winter diets in interior Alaska. Journal of Wildlife Management. 53(3): 568-577. [14930] 32. McCune, Bruce. 1982. Site, history and forest dynamics in the Bitterroot canyons, Montana. Madison, WI: University of Wisconsin. 166 p. Thesis. [7232] 33. Schopmeyer, C. S., tech. coord. 1974. Seeds of woody plants in the United States. Agriculture Handbook No. 450. Washington, DC: U.S. Department of Agriculture, Forest Service. 883 p. [2088] 34. 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] 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. 1973. Wildfire in the taiga of Alaska. Quaternary Research. 3: 465-495. [7247] 37. Viereck, Leslie A. 1975. Forest ecology of the Alaska taiga. In: Proceedings of the circumpolar conference on northern ecology; 1975 September 15-18; Ottawa, ON. Washington, DC: U.S. Department of Agriculture, Forest Service: 1-22. [7315] 38. 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] 39. Walker, Marilyn D.; Walker, Donald A.; Everett, Kaye R.; Short, Susan K. 1991. Steppe vegetation on south-facing slopes of pingos, central arctic coastal plain, Alaska, U.S.A. Arctic and Alpine Research. 23(2): 170-188. [14954] 40. Wolff, Jerry O. 1978. Burning and browsing effects on willow growth in interior Alaska. Journal of Wildlife Management. 42(1): 135-140. [3500]

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