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Introductory

SPECIES: Vaccinium vitis-idaea | Mountain Cranberry
ABBREVIATION : VACVIT SYNONYMS : NO-ENTRY SCS PLANT CODE : VAVI VAVIM COMMON NAMES : mountain cranberry northern mountain cranberry lowbush cranberry TAXONOMY : The currently accepted scientific name of mountain cranberry is Vaccinium vitis-idaea Linnaeus (Ericaceae) [6,106,107]. Vaccinium vitis-idaea ssp. minus (Lodd.) Hulten is the only recognized subspecies occurring in North America [54]. In some areas, mountain cranberry hybridizes with dwarf bilberry (V. myrtillus) [1]. A naturally occurring hybrid (V. X intermedium Ruthe.) has been identified [87]. LIFE FORM : Shrub FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY COMPILED BY AND DATE : D. Tirmenstein, February, 1991. LAST REVISED BY AND DATE : NO-ENTRY AUTHORSHIP AND CITATION : Tirmenstein, D. 1991. Vaccinium vitis-idaea. In: Remainder of Citation

DISTRIBUTION AND OCCURRENCE

SPECIES: Vaccinium vitis-idaea | Mountain Cranberry
GENERAL DISTRIBUTION : Mountain cranberry is a circumpolar, circumboreal species that occurs throughout parts of North America, Eurasia, and Japan [101,106]. The New World subspecies (ssp. minus) extends from northwestern Greenland across the Canadian Arctic southward to New England [114]. It grows westward to the Great Lakes and British Columbia and reaches islands in the Bering Sea [42,114]. In North America, mountain cranberry is restricted to areas north of the glacial boundary [106]. The subspecies vitis-idaea occurs throughout northern Europe from Scandinavia to northern Italy and the Caucasus, across northern Siberia and Japan southward into northern China and Korea [42]. ECOSYSTEMS : FRES10 White - red - jack pine FRES11 Spruce - fir FRES19 Aspen - birch FRES23 Fir - spruce FRES26 Lodgepole pine FRES44 Alpine STATES : AK CT ME MA MN NH VT WI AB BC LB MB NB NF NT NS ON PE PQ SK YT ADMINISTRATIVE UNITS : ACAD DENA GLBA ISRO LACL WRST YUCH BLM PHYSIOGRAPHIC REGIONS : NO-ENTRY KUCHLER PLANT ASSOCIATIONS : K015 Western spruce - fir forest K093 Great Lakes spruce - fir forest K094 Conifer bog K095 Great Lakes pine forest K096 Northeastern spruce - fir forest K106 Northern hardwoods K107 Northern hardwoods - 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 17 Pin cherry 18 Paper birch 38 Tamarack 107 White spruce 201 White spruce 202 White spruce - paper birch 204 Black spruce 218 Lodgepole pine 251 White spruce - aspen 253 Black spruce - white spruce 254 Black spruce - paper birch SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Mountain cranberry grows as an understory dominant or codominant in a variety of forest communities including many dominated by jack pine (Pinus banksiana) and lodgepole pine (P. contorta). It also occurs as a dominant or indicator in dwarf shrub and shrub tundra communities. Common codominants include dwarf birch (Betula nana), alpine bearberry (Arctostaphylos alpina), Labrador tea (Ledum spp.), feathermoss (Pleuroiozium spp.), willow (Salix spp.), sedges (Carex spp.), lichen (Cladina spp.), and crowberry (Empetrum nigrum). Mountain cranberry is listed as a dominant or indicator in the following plant association, ecosystem association, habitat type, and community type classifications: Forest community types of west-central Alberta in relation to selected environmental factors [17] Field guide to forest ecosystems of west-cewntral Alberta [18] Vegetation types in northwestern Alaska and comparisons with communities in other arctic regions [39] Plant associates: Cloudberry (Rubus chamaemorus), Canada beadruby (Maianthemum canadense), prickly rose (Rosa acicularis), paper birch (B. papyrifera), sedge, mountain-laurel (Kalmia angustifolia), bearberry (Arctostaphylos uva-ursi), crowberry, twinflower (Linnaea borealis), willow, bog blueberry (Vaccinium ugliginosum), fireweed (Epilobium angustifolium), bluejoint reedgrass (Calamagrostis canadensis), bog Labrador tea, and feathermoss commonly occur with mountain cranberry in white and black spruce and jack pine communities [7,22,26,38,48,120]. Willows, bog Labrador tea, prickly rose, crowberry, bog blueberry, sedges, cottongrass (Eriophorum vaginatum), and cloudberry are common associates in treeless sphagnum bogs, cottongrass muskeg, and dwarf shrub marsh communities [84,111,114,120].

VALUE AND USE

SPECIES: Vaccinium vitis-idaea | Mountain Cranberry
WOOD PRODUCTS VALUE : NO-ENTRY IMPORTANCE TO LIVESTOCK AND WILDLIFE : Browse: Mountain cranberry browse is readily eaten by barren-ground caribou, black bear, moose, arctic hare, and snowshoe hare [38,42]. In parts of Alaska, it is an important if not key moose browse [3]. Utilization by moose is typically heaviest when available browse is limited and when light snow accumulations allow the animals to reach the plants easily [93]. On the Kenai Peninsula, it may comprise up to 25 percent of winter moose diets. Moose may dig through 20 inches (50 cm) of snow to feed on the foliage, but if winter snow depths are excessive, the animals rarely expend the energy necessary to reach the plants [77]. Generally, moose eat only trace amounts of mountain cranberry during the summer [10,77]. In some parts of Canada, mountain cranberry browse is a primary food of barren-ground caribou [73]. The evergreen leaves are an important item in the winter diet [38]. In the Mackenzie District of northwestern Saskatchewan, leaves of mountain cranberry and bog blueberry (V. uliginosum) accounted for 21.5 percent of the barren-ground caribou winter diet but only 3.8 percent of the summer diet [38]. However, in some areas, caribou continue to feed heavily on mountain cranberry browse throughout the summer [111]. In Newfoundland, snowshoe hares often consume large amounts of shoots during the winter [38]. Where snow depths prohibit winter use, hares may feed on leaves made available by melting snows. Seasonal percent composition of leaves of mountain cranberry and bog blueberry in the diet of snowshoe hares in Newfoundland was as follows [118]: winter April May summer fall 0.3 17.4 9.3 6.6 10.9 Mountain cranberry browse is of little value to domestic livestock but provides some winter browse for reindeer [23,73]. It is not eaten by domestic sheep if more preferred forage is available [88]. Fruit: Berries of mountain cranberry are an important food source for many species of birds and mammals. Many wildlife species feed on fruit left on the ground from the previous year [38,55]. Berries are an important spruce grouse food during spring, summer, and fall. Berries persisting from the previous year are eaten from late spring through early August. In interior Alaska, percent volume use of mountain cranberry by spruce grouse was 37.6 in July and August, 40.1 in September, and 17.3 in September [29]. In many areas, berries are an essential food source for birds migrating northward in the spring [38,55]. The common raven, ring-necked pheasant, rock ptarmigan, sea gulls, geese, grouse, partridges, and many species of songbirds, such as the scarlet tanager, eastern bluebird, and thrushes, readily consume mountain cranberry fruit [38,42,88]. Fruit of Vacciniums are readily eaten by the northern mockingbird, rufous-sided towhee, gray catbird, American robin, brown thrasher, ruffed grouse, spruce grouse, whimbrel, herring gull, and Canada goose [72,105,106]. The red-backed vole eats large quantities of mountain cranberry fruit during the fall. Berries are a primary winter food source as well; the rodents burrow under snow to reach the persistent fruit [117]. The red fox also consumes large amounts of fruit during late fall [38]. Mountain cranberry fruit is an important black bear food in many areas but is of particular importance in Alaska [40]. Berries remain on the plant over winter, and black bears begin feeding on berries during the early spring as soon as the snow has melted [38,40]. Fruit again assumes importance in black bear diets during the fall [40]. Many other mammals, including the polar bear, eastern chipmunk, and white-footed mouse, also feed on the fruit of mountain cranberry [38,55]. Fruits of many Vacciniums are readily eaten by species such as the red squirrel, gray fox, skunks, and chipmunks [72,106]. PALATABILITY : Mountain cranberry browse is at least seasonally palatable to many species of mammals including the barren-ground caribou, snowshoe and arctic hares, and moose. Berries are readily eaten by a variety of birds and mammals. Palatability of the fruit increases over winter [99]. NUTRITIONAL VALUE : Browse: Nutrient content of browse varies according to factors such as soils, phenological development, and proximity to smelters [42,45,95]. Calcium, manganese, aluminum, silver, lead, and boron tend to accumulate in plant tissue even at low soil levels [42]. Food value peaks in summer [38]. In winter, acid-detergent, fiber, and lignin content increase but levels of magnesium, zinc, manganese, calcium, potassium, sodium, copper, and iron decline. Protein content remains relatively constant throughout the year at 5 to 6 percent [77]. Energy content has been estimated at 509 kcal/100 g [73]. Nutritional value of browse from the Kenai Peninsula of Alaska was documented as follows [77]: August February Protein (%) 5.7 5.4 Ca (ppm) 4920.0 26.7 Mg (ppm) 1328.0 4.6 K (ppm) 438.3 29.8 Na (ppm) 55.0 22.8 Cu (ppm) 5.8 0.2 Fe (ppm) 51.3 3.2 Mg (ppm) 17.6 1.9 Zn (ppm) 8.3 0.3 Fruit: Berries are high in tannins and anthocyanins. The caloric content is moderate [38]. COVER VALUE : NO-ENTRY VALUE FOR REHABILITATION OF DISTURBED SITES : Potential rehabilitation value of mountain cranberry has not been well documented. Plants are able to survive on extremely harsh sites, and some rehabilitation potential is possible. On the Arctic Coastal Plain, sprouts have been observed on and under debris left from oil exploration activities [28]. Mountain cranberry can be readily propagated from seed and stem or rhizome cuttings [32,42]. Meristem propagation techniques have also been described [101]. Stem cuttings root easily if planted in the spring or early fall but exhibit slow rhizome development and poor subsequent vegetative spread [32]. Clumps of wild mountain cranberry can be divided and transplanted onto disturbed sites [42]. Survival of these transplants is variable, ranging from 30 to 90 percent [32]. Propagation techniques have been examined in detail [25,32,42,61,63]. OTHER USES AND VALUES : Mountain cranberry fruit can be eaten raw or cooked to make a tart sauce [6,99]. Berries are used to make preserves, jam, jelly, candy, syrup, pickles, juice beverages, and wine [42,47]. Fruit can be added to rose hips to make a tasty jelly [38], or added to various ice cream products [42]. In some areas, berry-picking is an important recreational activity [59]. Fruit is widely processed and marketed in Japan and Europe [42] and is harvested commercially in parts of Alaska, Scandinavia, the U.S.S.R., and Canada [42,43]. Considerable amounts of fruit are imported into the United States annually [11]. Much of this imported fruit is consumed by peoples of Scandinavian descent who use the so-called "Swedish lingenberry" in traditional dishes [6]. Mountain cranberry has the potential for more extensive commercial development [15,37,74]. Some native stands could be managed with a minimum of cultivation, as are those of low sweet blueberry [see VACANG] [74]. The feasibility of expanded commercial operations is currently being tested in parts of North America [42]. Many Native Americans and indigenous peoples of Eurasia used the leaves and fruit of mountain cranberry as food or medicine [57,106]. Preparations made from the leaves were used to treat bladder problems, gout, and rheumatism [90]. Medicinal fruit jellies were used to treat sore throats and colds [106]. The Slave, Athabaska, Cree, and Inuit people ate the fruit fresh and preserved them for winter use [38,106]. Berries were often boiled and mixed with oil to facilitate storage for long periods [106]. Arbutin, which is obtained from the leaves and stems, is used by the pharmaceutical industry in preparations used to treat intestinal disorders. Mountain cranberry forms a dense, attractive mat and has been planted as an ornamental ground cover [24]. It was first cultivated in 1789 [42]. Mountain cranberry has shown promise for use in developing hardy fruit-producing cultivars [64]. MANAGEMENT CONSIDERATIONS : Fruit production: Fruit production in mountain cranberry varies widely according to geographic location, site factors such as shade and soil, annual weather conditions, and the genetic make-up of the individual clone [62,63,71,78,81]. Poor fruit production may be due to a lack of pollinators, cold damp weather during flowering, late spring frosts, or hail [42,43,57]. Plants growing in the shade rarely produce fruit or flowers, but plants growing in full sun commonly bear an abundance of fruit [62]. Some geographic variation in this pattern has been noted. On dry sunny sites in Alberta, flower bud production may be greatest in partial shade of aspen (Populus tremuloides) [38]. In the cool, rainy climates of the Maritime Provinces, flower bud production is typically best on exposed sites [38]. Kuchko [57] reported poor yields beneath forest canopy, although yields were often good in adjacent gaps created by timber harvest. Fruit yields are generally greater on peat than on mineral soil [63]. Under experimental conditions, plants produced 82 kg/100 m sq on peat but produced only 14 kg/100 m sq on mineral soil [63]. Temperatures of 30 degrees F (-1.5 degrees C) can kill 50 percent of all flowers, and exposure to 26 degrees F (-3.5 degrees C) can destroy 50 percent of the buds and unripe fruit [62]. In harsh arctic environments, only plants in protected areas, such as on south-facing rock crevices, flower [42]. Maximum yields in cultivated stands may reach 9,140 pounds per acre (8,150 kg/ha) [38]. Elsewhere, yields may range from 19.5 pounds per acre (17.4 kg/ha) [51] in Swedish peatlands to 560 pounds per acre (500 kg/ha) in some Finnish forests [38]. Yields are generally highest where mountain cranberry cover is greatest and competitors are few [71]. Details on fruit yields are available [32,57,81]. Cultivation: Mountain cranberry generally responds more favorably to fertilizer and irrigation than do other members of the genus [56]. However, the application of fertilizer does not always increase fruit yields. Comparatively little fertilizer is required for good growth and development [42]; if too much is added, vegetative growth may be promoted at the expense of fruit production [101]. Where weeds are a problem, fertilizer may increase competitors at the expense of mountain cranberry [62]. Mulches such as milled peat can increase fruit production in some instances [42]. The effects of mulch, fertilizers, and irrigation have been examined in detail [32,42,46,53,62,63]. Fruit yields may be increased by various means. Herbicides have been used to reduce weeds in commercially managed fields of mountain cranberry [37,63]. Honeybees can be used to supplement native bee populations when pollinator availability is low [74]. Fruit is generally harvested by hand [42]. Small comb-sieves or rakes are commonly used [38,101]. Chemical response: Mountain cranberry is susceptible to herbicides such as 2,4-D and 2,4,5-T [38]. These herbicides cause browning of stems and leaves and at high concentrations can kill the plants [38]. The effect of herbicides has been documented [8,38]. Damage/disease: Plants can be killed by exposure to cold temperatures in the absence of a protective snow cover [83]. Unacclimated plants can be killed by exposure to temperatures of 28 degrees F (-2.5 degrees C) or below;, acclimated plants can survive exposure to temperatures as low as 8 degrees F (-22 degrees C) [42]. Mountain cranberry is susceptible to several diseases and insect infestations [38,42]. Environmental considerations: Mountain cranberry growing near smelters can accumulate high concentrations of heavy metals [95]. Plants growing near a zinc smelter in Poland exhibited reduced leaf size and other types of damage [20]. Mountain cranberry can also accumulate a wide range of radionuclides such as radium-226, lead-210, and uranium [97]. Tests indicate that summer oil spills are more damaging to mountain cranberry than those that occur in February [38]. Predisturbance cover of 48 percent was reduced to 0 by a summer crude oil spill. A low-intensity winter spill reduced cover to 12 percent while a high-intensity oil winter spill reduced cover to 6 percent [38]. Recovery of mountain cranberry can occur 10 to 15 years after an oil spill [38]. Timber harvest: After some types of logging treatments in a mature white spruce (Picea glauca) forest in Alaska, cover and frequency of mountain cranberry increased fairly rapidly [26]. Biomass: Mountain cranberry biomass is strongly correlated with canopy cover [44,77]. Maximum dry matter accumulation occurs in full sunlight [43]. Holloway [42] observed that 80 percent of the total biomass of mature plants is underground. Biomass has been examined in detail [38,42,94].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Vaccinium vitis-idaea | Mountain Cranberry
GENERAL BOTANICAL CHARACTERISTICS : Mountain cranberry is a low, creeping, evergreen subshrub that commonly reaches 2 to 6 inches (5-15 cm) in height [4,90,114]. It typically grows in dense rhizomatous colonies and frequently forms mats [106]. Stems are slender and trailing [38,114]; stem morhpology has been examined in detail [76]. The root system is variable [6]. Plants have a network of fine, shallow, fibrous roots, and may possess a taproot [32,38]. The dichotomously branched rhizomes possess numerous hairlike roots [98]. Maximum rooting depths of 2 to 11 inches (5-28 cm) have been reported [42,100]. The thick, simple, leaves are obovate, oblong, or elliptic [38,90]. The green leaves turn purplish in fall [38]. Flowers occur on terminal racemes singly or in groups of up to 15 [90]. Floral morphology has been examined in detail [79]. Fruit is a bright to dark red, globular berry approximately 0.2 to 0.4 inch (6-10 mm) in diameter [4,55,106,114]. The four-celled berries are acidic to sour or bitter [6,90,106]. Yellow, short-beaked seeds average 0.04 inch (1 mm) in length [42,106]. RAUNKIAER LIFE FORM : Chamaephyte Geophyte REGENERATION PROCESSES : Mountain cranberry reproduces through seed and by vegetative means [38]. Seed: In many areas, seedlings first bear fruit at 3 or 4 years of age [32,63]. However, British studies suggest that few flowers are produced until plants reach 5 to 10 years of age [88]. Flowers are pollinated by bumblebees and bee flies (syrphid flies) [38,42]. Plants may be self- or cross-pollinated, but fruit set is much greater after cross-pollination [6]. Berries are often produced in abundance. In parts of North America, berries average 3 to 15 seeds per berry [43]. Seeds are dispersed by birds and mammals [38]. Germination: Seed can germinate on bare ground, but only if conditions are favorable [38]. Fresh seed generally exhibits best germination [37,38]. Germination declined from an average of 76.5 percent for seed extracted from fresh fruit and then planted immediately to less than 10 percent for seed stored 12 to 16 months before planting [38]. In laboratory tests, good germination was reported after stratification at 32 to 41 degrees F (0-5 degrees C) for up to 5 months [42,61]. Seeds typically germinate within 3 weeks after exposure to temperatures of -7 to -4 degrees F (20-25 degrees C) in light or dark [32,42]. Germination characteristics of mountain cranberry have been examined in detail [19,62,63]. Seedling establishment: Seedlings are rarely observed in the field [42,62,75,102]. In Estonia, seedlings are generally observed only in protected areas such as near tree stumps, fallen logs, or stones [71]. Some seedlings do develop in favorable years in parts of Nova Scotia and Newfoundland [38]. Seedbanking: Seed of mountain cranberry has been detected in soil samples [75,106], but seed banking potentials for this species are unknown. In black spruce (Picea mariana)-jack pine forests of the Northwest Territories, seeds of mountain cranberry and bog blueberry were found in 71 percent of the soil samples tested [50]. Studies near the Great Slave Lake revealed mountain cranberry and bog blueberry seed in 65 percent of the cores sampled. Seed densities averaged 3.3 per 1,000 cc, but only 21 percent were viable. A second study revealed 4.8 seeds per 1,000 cc, but none of the seeds were viable [38]. Vegetative regeneration: Vegetative regeneration is of primary importance in the mountain cranberry [88]. Plants commonly expand through horizontal rhizomes [88]. Rhizomes may sprout singly or in groups of 1 or 2 per square meter [42]. Large, older clones may be separated into numerous daughter clones by disturbances such as frost, fire, or burrowing mammals [38,88]. Rhizome length, depth, and the location of shoots on the rhizomes are greatly influenced by soil and other site characteristics [98]. Rhizome depth is inversely related to the thickness of soil organic layers [98]. Rhizomes grow well in peat but can also penetrate to mineral soil. In Britain, rhizomes are generally confined to the humus layer [38] and are estimated to average 4 to 8 inches (10-20 cm) deep [88]. Smith [98] reported that rhizome depth in Alberta varied from 8 to 11 inches (19-28 cm). He found that 22.1 percent of the shoots were located terminally and 77.9 percent arose at midrhizome locations. Rhizome characteristics as related to various site characteristics have been examined in detail. The trailing or creeping stems of mountain cranberry also root at the nodes [38,114]. This mode of regeneration may be important on some harsh, subarctic sites [38]. SITE CHARACTERISTICS : Mountain cranberry is widely distributed in northern temperate forests and in many arctic and alpine communities [38,60,114]. It commonly grows on exposed sites, such as windswept crags, bare headlands, rocky ledges, scree, sea cliffs, hilly rocky barrens, and mountain summits [21,38,81,88,91]. At the southern edge of its range, mountain cranberry occurs primarily in bogs, but in the north it grows on both wet and dry sites [38]. Mountain cranberry occurs on high moors, heath barrens, sand dunes, and in peatlands, forest swamps, and bogs [38,41,114]. In mature forests, plants often grow on top of decaying tree stumps [42]. Climate: Mountain cranberry grows under a variety of climatic regimes. In much of Canada, it occurs in areas characterized by short cool summers and long cold winters [38]. In black spruce-white spruce-jack pine forests of northern Canada, its distribution may be correlated with arctic air masses. However, in harsh rockfield and tussock communities of the far North, it may be related to the influence of moist Pacific air masses [38,60]. In taiga communities of Alaska, winters are long and cold, but summers are short and hot [110]. Mean annual precipitation is 8 inches (21 cm), and average annual temperature is 20 degrees F (-6.7 degrees C) [84]. In parts of the Northwest Territories, annual precipitation averages 12 inches (30.4 cm) [68]. Soils: Mountain cranberry grows on shallow, poorly developed mineral soil as well as on drained peat [51,88]. Soils are often of low fertility and have little calcium but may be high in decaying organics [42,101]. Mountain cranberry commonly grows on acidic sandy loams or loamy clays [42,57]. Holloway and others [45] reported poorest vegetative growth on sandy soils. Soil pH ranges from 2.7 to 8.2, but best growth has been reported at 4.0 to 4.9 [38,42,49]. Soils are often characterized by low base saturation and low lime content [45]. Soils may be derived from a variety of parent materials, including sandstone, gneiss, granite, and glacial outwash sands and gravel [38]. Elevation: In New England, mountain cranberry is generally restricted to higher mountains [55]. In the Northwest Territories, plants often occur at lower elevations (to 4,950 feet [1,500 m]) [38]. Generalized elevational ranges by geographic location are as follows: Location Elevation Authority Adirondacks up to 5,300 ft (1,615 m) Keeler 1969 e Canada sea level to 4,250 ft (0-1,290 m) Hall and Shay 1981 AB to 7,400 ft (2,250 m) Hall and Shay 1981 Yukon 6,900 to 7,900 ft (2,100-2,400 m) Hall and Shay 1981 SUCCESSIONAL STATUS : Mountain cranberry is noted for its wide ecological amplitude [69]. It is not generally considered a pioneer species but does occur in early seral stages in some communities [38,65]. Mountain cranberry persists indefinitely, unless shaded out by conifers, and assumes a climax role in various rockfield communities of the far North [38]. Mountain cranberry commonly invades tundra bog communities dominated by species such as alpine sweetgrass (Hierochloe alpina), lichens (Alectoria ochroleuca, A. nitidula), and woodrush (Luzula confusa) from adjacent summit rockfields. Mountain cranberry also invades senescent cottongrass tussock communities and areas of frost activity after the establishment of initial pioneers. However, on some sites, seral mat communities made up of mountain cranberry, crowberry, and lichens eventually give rise to white spruce stands. In barrens of Newfoundland, mountain cranberry grows as a seral species which is displaced by black spruce and balsam fir [38]. It also occurs in some early seral communities dominated by paper birch [65]. Black spruce: Mountain cranberry is important in stable climax black spruce communities but also dominates many seral stages [34,65,80]. Stands are initially colonized by bryophytes and herbaceous species such as fireweed and willow [7,27]. Mountain cranberry generally reaches stable levels within 25 years after fire or other disturbances [38]. However, maximum cover and frequency were attained at 144 years in certain black spruce/mountain cranberry communities [38]. In black spruce stands in interior Alaska, mountain cranberry is present within 5 to 30 years after disturbance and persists for many years. It is common in stands 200 years old or older and represents the most abundant low shrub in tree-dominated stages [34]. In black spruce stands of the Northwest Territories, mountain cranberry remains abundant in 200- to 300-year-old stands despite the decline of most vascular plants [7]. Chapin and others [14] reported that mountain cranberry becomes more prominent as succession progresses from immature black spruce to muskeg. White spruce: Mountain cranberry occurs in many climax white spruce forests on uplands of interior Alaska [65]. It is present during the moss-herb stage which occurs 1 to 5 years after fire [27,34]. Mountain cranberry peaks and declines after the dense tree stage, which occurs from 15 to 40-46 years or longer after fire, but remains present in later stages [34]. Dyrness and others [27] reported that in interior Alaska, mountain cranberry was common in 150-year-old white spruce stands. Jack pine: In jack pine-lichen woodlands of the northern Canada, mountain cranberry is an early colonizer on recently burned sites [13]. It persists after "the cessation of major successional changes" at 25 to 45 years [13] and remains common in stands up to 280 years of age [38]. Tundra communities: In sedge-tussock tundra and shrub tundra communities of Alaska's Seward Peninsula, bryophytes initially reestablished burned sites. Bryophytes often reach maximum cover within 2 to 4 years after fire, but the recovery of shrubs such as mountain cranberry is often much slower. In shrub-tundra communities, mountain cranberry may not recover to preburn levels even by 5 to 6 years after fire or other disturbance [84,85]. SEASONAL DEVELOPMENT : In Alaska, vegetative buds began growth during the first week of June and underwent rapid elongation throughout June. The growth rate of terminal vegetative buds decreased by July 1. Leaf expansion began during the last week of May and the first week of June; all leaves had expanded within 1 month [42]. Karlsson [52] observed that old leaves became photosynthetically active approximately 2 weeks after bud break. Near Mt. Washington, New Hampshire, and in parts of Nova Scotia, vegetative growth began in late June [38,42]. In Britain and perhaps elsewhere, leaf expansion can begin as early as March, although it usually occurs from mid-May to mid-June. Shoot growth generally ends in mid-July. Leaves may persist for up to 3 years. However, some old leaves may be shed by August of the second year [38]. Plants become dormant by fall [42]. Flowers develop from buds initiated the previous year [101]. In interior Alaska, reproductive bud growth begins in mid-May [42]. In parts of Britain, two periods of flowering (spring and summer) have been observed at certain low-elevation sites [88]. Flowering may last 9 to 18 days [57] or as long as 19 to 27 days. Fruit ripens approximately 78 to 84 days after full bloom [42]. Phenological development may be related to the timing of snowmelt [38]. In interior Alaska, plants were in maximum full bloom approximately 6 weeks after snowmelt and exhibited first visible signs of growth 2 weeks after snowmelt. Unusually cool temperatures can delay phenological development. Roots and rhizomes undergo two periods of active growth annually in early spring and fall [42].

FIRE ECOLOGY

SPECIES: Vaccinium vitis-idaea | Mountain Cranberry
FIRE ECOLOGY OR ADAPTATIONS : Mountain cranberry occurs in a variety of communities across a wide climatic range. It persists under a regime of relatively frequent fires but also grows in areas that rarely burn. Black spruce communities are dependent on frequent fires, and most associated species, including mountain cranberry, are well adapted to fire [110]. Fires in black spruce communities of Alaska and northern Canada are commonly lightning caused and tend to be large [68,110]. Fire frequencies average 80 to 200 years [96,110]. In moister black spruce/mountain cranberry communities in eastern Canada, fires may occur at 500-year intervals [35]. Mountain cranberry remains important in jack pine stands that burn at 20- to 40-year intervals and in Swedish pine forests that burn every 40 years [13,30]. Fire may be an important factor in treeline communities of the North. In Siberia, past extensive fires may have destroyed forest communities. Trees may have been unable to reestablish on these harsh sites under the current climatic regimes. Low-shrub-dominated tundra communities composed of species such as mountain cranberry may have eventually replaced these forest stands [108]. Fire intervals in shrub subzones of forest-tundra communities have been estimated at 1,460 years [96]. Mountain cranberry continues to be abundant on these infrequently burned sites. In many forest communities, mountain cranberry requires fire for its maintenance [30]. Increases in cover and vigor after fire are commonly observed [9]. Mountain cranberry generally reestablishes a site through sprouting from rhizomes and aerial stems. Very limited reestablishment may occur on exceptional sites in good years by seed transported from off-site. POSTFIRE REGENERATION STRATEGY : Small shrub, adventitious-bud root crown Rhizomatous shrub, rhizome in soil Initial-offsite colonizer (off-site, initial community)

FIRE EFFECTS

SPECIES: Vaccinium vitis-idaea | Mountain Cranberry
IMMEDIATE FIRE EFFECT ON PLANT : Underground regenerative structures of mountain cranberry generally survive light fires [102,115]. Plants often survive even when aerial portions are consumed by fire [92]. However, plants may be killed by moderate to heavy, duff-consuming fires [115]. Survival is related to many factors including soil moisture levels, season of burn, fire severity and intensity, and rhizome depth [38]. Rhizomes can sometimes survive soil surface temperatures of 820 degrees F (438 degrees C) [102]. In arctic tussock communities, plants often survive severe fires which remove all aboveground material [116]. The heat-sensitive seeds of mountain cranberry are usually destroyed by fire [115]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Mountain cranberry commonly sprouts from rhizomes or buds located on surviving portions of aerial stems after fire damages or consumes aboveground material [115,116]. Sprouting from stumps, or "rootstocks" has also been reported [13,65]. Reestablishment through seed is extremely rare [102]. Surviving portions of the aerial stems sprout within a short time, but rhizome sprouting may be delayed until the following year [115]. The speed of reestablishment varies according to the season of burn, site characteristics, and fire intensity and severity. Reestablishment is generally rapid after light fires [27]; plants are often common on lightly burned sites [102]. Regeneration may be slow after hot fires that damage or destroy underground regenerative structures [27,102,111]. In northern spruce communities, intense, stand-destroying, late summer fires which consume the organic layer [110] can be particularly damaging to mountain cranberry [65]. On some sites, plants may sprout within months after a light burn and regain preburn cover within a few years [102]. Mountain cranberry generally appears within the first 6 years after fire in black spruce-lichen, jack pine-lichen, and white spruce-birch communities [38]. Residual survivors were observed at the end of the fifth growing season in black spruce communities of southeastern Manitoba [16]. In a severely burned black spruce community of interior Alaska, it became abundant within 5 years after fire but set little fruit [117]. Viereck [109] observed slow recovery after fire in a black spruce/feathermoss- lichen community of interior Alaska. Recovery was as follows: percent cover (1971 fire) unburned 1972 1975 1980 18 1 1 7 Mountain cranberry was present within 1 to 5 years after fire in white spruce communities of Alaska [34]. Mountain cranberry is a common early colonizer in jack pine communities, although reestablishment generally takes at least several years [13]. In North America, postfire recovery may be more rapid in moister, eastern boreal forests [68]. Reestablishment of mountain cranberry is often slow in tundra communities [85]. Plants attained preburn coverage by the end of two full growing seasons in arctic tussock communities [116]. In northwestern Alaska, production was still significantly lower on sites burned 13 years earlier than on unburned sites [33]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : Fire severity: Recovery is typically much more rapid after light fires. Postfire recovery in Alaska has been documented as follows [111]: percent cover 1971 1972 1973 1974 control heavy burn .05 .50 .30 .90 ---- light burn 3.45 3.55 1.65 6.10 20.35 heavy burn 0 .15 .20 .35 6.90 In sedge-shrub tundra on the Seward Peninsula of Alaska, little or no sprouting was observed within 2 years on severely burned sites where mountain cranberry was a preburn dominant. After light to moderate severity burns in sedge-tussock communities, mountain cranberry resprouted and recovered relatively quickly. Shoot densities increased significantly on two of the four sites, from 43 shoots per meter square to 126 per meter square and from 25 per meter square to 43 per meter square. However, sprouts were generally located on the surface or sides of tussocks, suggesting that they escaped burning within the tussock mass [84]. Recovery of mountain cranberry may be relatively slow in many types of tundra shrub communities. Recovery of mountain cranberry by tundra community was as follows [84]: sedge tussock-shrub tundra (burned 1977) Frequency (no. of plots)* Cover (%) 1973 1978 1979 1973 1978 1979 site 2 10 10 10 6.9 0.8 1.3 site 3 -- 10 10 --- 0.5 0.5 site 4 -- 10 10 --- 0.5 0.5 site 5 -- 9 9 --- 0.5 0.5 *Number of 1-m2 plots in which species occurs (ten plots sampled) birch and ericaceous shrub tundra (burned 1977) 1973 1978 1979 Cover Cover Cover _ _ _ Freq.* (x %) Freq.* (x %) Freq.* (x %) nonfrost boils 10 7 0 0 0 0 frost boils 10 4 0 0 0 0 *Number of 1-m2 quadrats in which species occurs/no. of quadrats on that site X 10 sedge-shrub tundra (burned 1977) Prefire (1973) 1 yr after (1978) 2 yrs after (1979) Freq.* Cover** Freq.* Cover** Dens.*** Freq.* Cover** Dens.*** site 8 10 2.8 3 0.2 2 4 0.2 3 site 9 10 15.5 6 0.3 2 7 0.3 5 * Number of 1-m2 plots in which the species occurs (ten plots sampled) ** Mean percent cover averaged over 10 plots *** shoots/m2 Posfire frequencies of mountain cranberry 1 year after a summer fire in sedge tussock-shrub communities of the Seward Peninsula of Alaska were greatly reduced [119]: sampling date late May 1978 mid-June 1978 freq. % freq. % burned 0.23 0.05 unburned 1.00 1.00 FIRE MANAGEMENT CONSIDERATIONS : Postfire biomass: Postfire reduction in mountain cranberry production was as follows after a fire in an arctic tussock community [116]: mean annual production (g/m sq) burned unburned site 1 0.1 5.8 site 2 0.6 7.5 site 3 0.6 1.8 site 4 4.0 9.5 Biomass: Biomass following a late June wildfire in interior Alaska was measured at 0.04 grams per square meter during postfire year 1, 0.08 grams per square meter during postfire year 2, and 1.4 grams per square meter during postfire year, compared to a control measurement of 5.1 grams per square meter [108]. Fuels and flammability: Engelmark [30] reported that Vacciniums are highly flammable due to specific chemical properties. In northern Sweden, species such as mountain cranberry can serve as ignition points and as a continuous fuel mat for surface fires. In many black spruce stands of Alaska and northern Canada, an open, highly flammable, ericaceous shrub layer can carry a fire [110]. However, Quintilio and others [82] observed that an extensive mat of mountain cranberry and alpine bearberry served as an effective fire barrier in a jack pine stand near Darwin Lake, Alberta. Fire seldom penetrated more than a few centimeters into the vegetative mat. The extensive ground mat noticeably reduced the fire spread rate and coverage [82].

FIRE CASE STUDIES

SPECIES: Vaccinium vitis-idaea | Mountain Cranberry
CASE NAME : Black spruce, Fairbanks, Alaska REFERENCE : Viereck, L. A.; Foote, J.; Dyrness, C. T.; [and others]. 1979 [113] SEASON/SEVERITY CLASSIFICATION : summer/moderately severe Unit 2 - burned July 22, 1976 Units 1L, 3, and 4L - burned August 26, 1976 STUDY LOCATION : The study site is within the Washington Creek Fire Study and Training Area located approximately 22 miles (40 km) north of Fairbanks, Alaska. PREFIRE VEGETATIVE COMMUNITY : The preburn community was an unevenly spaced stand of black spruce (Picea mariana) approximately 70 years old. The forested area was interspersed with shrub-dominated openings. The forest understory was made up of various shrubs, herbs, mosses, and lichens. Green alder (Alnus viridis ssp. crispa), paper birch (Betula papyrifera), and willows (Salix spp.) were scattered throughout the area. Mountain cranberry (Vaccinium vitis-idaea), bog blueberry (V. uglinosum), and bog Labrador tea (Ledum groenlandicum) were common in the small clearings. Lichen mats carpeted the open areas. Moss was abundant throughout the community; cover ranged from 72 to 85 percent. TARGET SPECIES PHENOLOGICAL STATE : Not reported. SITE DESCRIPTION : Elevation: 1,700 feet (520 m) Aspect: slopes to the southeast Soil: shallow Fairplay silt loam with shattered bedrock and stones at 8 to 20 inches (20-50 cm) in depth Climate: average monthly precipitation- April - 0.3 inch (6.4 mm) May - 0.7 inch (18.6 mm) June - 1.3 inches (35.3 mm) July - 1.8 inches (46.7 mm) August - 2.2 inches (55.9 mm) No permafrost was encountered. FIRE DESCRIPTION : Unit 2: fuel stick moisture - 6.6 percent relative humidity - 32 percent wind speed/direction - 0-5 miles/hour (0-2.2 m/s) from SSE temperature - 76 degrees F (24 degrees C) Unit 2 was ignited at the southeastern corner. The fire soon began to crown and moved throughout the unit within 7 minutes. The fire was particularly intense at the center. The fire was hot at the 0 to 9 foot (0-3 m) level, but did not burn hot within the forest floor. Temperatures reached approximately 249 degrees F (121 degrees C) at 2 inches (5 cm) above the ground. Units 1L, 3, and 4L: fuel stick moisture - 6.6 percent relative humidity - 40 percent wind speed/direction - 6 to 8 miles/hour (2.7-3.6 m/s) from the SSE temperature - 66 degrees F (19 degrees C) Unit 1L was ignited at the southeastern corner. The fire began to crown within 1 minute, but the burning of crowns was spotty. The unit was covered by the fire within 9 minutes. The fire burned into the organic layer for several hours, and in spots, smoldered and flamed for several days. Fuels were added to the unit prior to the burn. Loaded fuel was estimated at 29,720 pounds per acres (26,500 kg/ha). Unit 3 was ignited at the eastern corner. Spotty torching occurred but the fire was mainly a slow-moving ground fire. The fire burned across the unit in 29 minutes. The ground fire was not very intense, but did continue for several days. Approximately 40 to 50 percent of the ground vegetation was moderately to heavily burned. Although much of the area was lightly burned, an estimated 40 percent of the unit burned to mineral soil. Unit 4L was ignited at the middle of the southeastern boundary. Fire completed its run within 6 minutes. Several parts of the unit experienced a very hot crown fire. The fire burned into the organic layer in some areas, and soil temperatures reached 163 degrees F (73 degrees C). Approximately 70 percent of the ground vegetation was heavily or moderately burned. Supplemental fuels were added to the unit prior to the fire. Loaded fuel was estimated at 15,600 kg/ha. FIRE EFFECTS ON TARGET SPECIES : Aboveground vegetation, including mountain cranberry, was totally destroyed in units 2, 3, and 4L. Some unburned portions of aboveground vegetation remained on unit 1L. An inventory conducted in September, revealed no regrowth on plots burned in August (Units 1L, 3, and 4). Sprouts of bluejoint reedgrass (Calamagrostis canadensis) and Scouler willow (Salix scouleriana) were observed on unit 2 which had been burned in July. No mountain cranberry was reported. FIRE MANAGEMENT IMPLICATIONS : Summer fires can temporarily remove ericaceous shrubs such as mountain cranberry. At least several years may be required before preburn cover is reached.

REFERENCES

SPECIES: Vaccinium vitis-idaea | Mountain Cranberry
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