| 词条 | Picea glauca | |||||||||
| 释义 |
| name = White spruce | image = Picea glauca Fairbanks.jpg | image_caption = Mature white spruce in Fairbanks, Alaska | status = LC | status_system = IUCN3.1 | status_ref = [1] | genus = Picea | species = glauca | authority = (Moench) Voss | range_map = Picea glauca levila.PNG | range_map_caption = Natural range | synonyms ={{hidden begin|toggle=left|title=List}}
| synonyms_ref = [1] }}Picea glauca, the white spruce,[2] is a species of spruce native to the northern temperate and boreal forests in North America. Picea glauca was originally native from central Alaska all through the east, across southern/central Canada to the Avalon Peninsula in Newfoundland. It now has become naturalized southward into the far northern United States border states like Montana, Minnesota, Wisconsin, Michigan, Vermont, New Hampshire and Maine; there is also an isolated population in the Black Hills of South Dakota and Wyoming.[3][4][5][6] It is also known as Canadian spruce, skunk spruce, cat spruce, Black Hills spruce, western white spruce, Alberta white spruce, and Porsild spruce.[7] DescriptionThe white spruce is a large coniferous evergreen tree which grows normally to {{Convert|15|to|30|m|ft|round=5|abbr=on}} tall, but can grow up to {{Convert|40|m|ft|round=5|abbr=on}} tall with a trunk diameter of up to {{Convert|1|m|ft|abbr=on}}. The bark is thin and scaly, flaking off in small circular plates {{convert|5|to|10|cm|in|0|abbr=on}} across. The crown is narrow – conic in young trees, becoming cylindric in older trees. The shoots are pale buff-brown, glabrous (hairless) in the east of the range, but often pubescent in the west, and with prominent pulvini. The leaves are needle-like, {{Convert|12|to|20|mm|in|frac=16|abbr=on}} long, rhombic (diamond-shaped) in cross-section, glaucous blue-green above (whence glauca) with several thin lines of stomata, and blue-white below with two broad bands of stomata.[3] The cones are pendulous, slender, cylindrical, {{Convert|3|to|7|cm|in|frac=4|abbr=on}} long and {{convert|1.5|cm|in|frac=8|abbr=on}} wide when closed, opening to {{Convert|2.5|cm|in|frac=8|abbr=on}} broad. They have thin, flexible scales {{Convert|15|mm|in|abbr=on}} long, with a smoothly rounded margin. They are green or reddish, maturing to pale brown 4 to 8 months after pollination. The seeds are black, {{Convert|2|to|3|mm|in|frac=32|abbr=on}} long, with a slender, {{Convert|5|to|8|mm|in|frac=32|abbr=on}} long pale brown wing.[3] SeedsSeeds are small ({{convert|2.5|to|5.0|mm|in|disp=or|frac=32|abbr=in}} long), oblong, and acute at the base. Determinations of the average number of sound seeds per white spruce cone have ranged from 32 to 130 (Waldron 1965, Zasada and Viereck 1970).[8][9]Common causes of empty seed are lack of pollination, abortion of the ovule, and insect damage. The average weight per individual seed varies from 1.1 mg to 3.2 mg (Hellum 1976, Zasada et al. 1978).[10] Each seed is clasped by a thin wing 2 to 4 times as long as the seed. Seed and wing are appressed to the cone scale. Embryo and megagametophyte are soft and translucent at first; later the endosperm becomes firm and milky white, while the embryo becomes cream-colored or light yellow. At maturity, the testa darkens rapidly from light brown to dark brown or black (Crossley 1953).[11] Mature seed “snaps in two” when cut by a sharp knife on a firm surface (Crossley 1953).[11] White spruce cones reach their maximum size after 800 growing degree days (GDD). Cone moisture content decreases gradually after about 1000 GDD (Cram and Worden 1957)[12] Cone colour also can be used to help determine the degree of maturation, but cones may be red, pink or green (Teich 1970).[13] Collection and storage dates and conditions influence germination requirements and early seedling growth (Zasada 1973, Edwards 1977, Winston and Haddon 1981).[14][15][16] A bushel (35 L) of cones, which may contain 6500 to 8000 cones, yields {{convert|6|to|20|oz|g}} of clean seed (USDA Forest Service 1948).[17] Seed dispersal begins after cone scales reflex with cone maturation in the late summer or early fall of the year of formation. Cones open at moisture contents of 45% to 70% and specific gravities of 0.6 to 0.8 (Cram and Worden 1957, Zasada 1973, Winston and Haddon 1981).[12][14][16] Weather affects both the initiation and pattern of seed dispersal (Nienstaedt and Zasada 1990),[7] but cone opening and the pattern of seed dispersal can vary among trees in the same stand (Zasada 1986). Even after dispersal has begun, cold, damp weather will cause cone scales to close; they will reopen during dry weather. Most seed falls early rather than late, but dispersal may continue through fall and winter (Zasada 1986),[18] even into the next growing season (Rowe 1953).[19] Seed dispersal occurs mainly in late summer or early fall (Waldron 1965).[8] White spruce seed is initially dispersed through the air by wind. Both the initiation and pattern of seed dispersal depend on the weather (Nienstaedt and Zasada 1990),[7] but these can vary among trees in the same stand (Zasada 1986).[18] Small amounts of white spruce seed are normally dispersed beyond 100 m from the seed source, but exceptionally seeds have been found more than 300–400 m from the nearest seed source (Zasada 1986).[18] Root systemThe root system of white spruce is highly variable and adaptable (Wagg 1964, 1967),[20][21] responding to a variety of edaphic factors, especially soil moisture, soil fertility, and mechanical impedance. On soils that limit rooting depth, the root system is plate-like, but it is a common misconception to assume that white spruce is genetically constrained to develop plate-like root systems irrespective of soil conditions (Sutton 1969).[22] In the nursery, or naturally in the forest, white spruce usually develops several long “running” roots just below the ground surface (Mullin 1957).[23] The structure of the tracheids in the long lateral roots of white spruce varies with soil nitrogen availability (Krasowski and Owens 1999).[24] StemWhite spruce can live for several hundred years. Ages of 200 to 300 years are commonly attained throughout much of the range, and Dallimore and Jackson (1961)[25] estimated the normal lifespan of white spruce at 250 to 300 years. Slow-growing trees in rigorous climates are also capable of great longevity. White spruce {{convert|6|to|10|m|ft|abbr=on}} high on the shore of Urquhart Lake, Northwest Territories, were found to be more than 300 years old (Hare and Ritchie 1972),[26] BarkThe bark of mature white spruce is scaly or flaky, grey-brown (Brayshaw 1960)[27] or ash-brown (Harlow and Harrar 1950),[28] but silvery when freshly exposed. Resin blisters are normally lacking, but the Porsild spruce Picea glauca var. porsildii Raup has been credited with having smooth resin-blistered bark (Hosie 1969).[29] White spruce bark is mostly less than {{convert|8|mm|in|disp=or|frac=16|abbr=on}} (Hale 1955)[30] and not more than {{convert|9.5|mm|in|disp=or|frac=64|abbr=on}} thick (Chang 1954).[31] DistributionWhite spruce has a transcontinental range in North America. In Canada, its contiguous distribution encompasses virtually the whole of the Boreal, Subalpine, Montane, Columbia, Great Lakes – St. Lawrence, and Acadian Forest Regions (Rowe 1972),[32] extending into every province and territory (Forestry Branch 1961).[33] On the west coast of Hudson Bay, it extends to Seal River, about 59°N, "from which the northward limit runs apparently almost directly north-west to near the mouth of the Mackenzie River, or about latitude 68°" (Bell 1881).[34] Collins and Sumner (1953)[35] reported finding white spruce within 13 km of the Arctic coast in the Firth Valley, Yukon, at about 69°30′ N, 139°30′ W. It reaches within 100 km of the Pacific Ocean in the Skeena Valley, overlapping with the range of Sitka spruce (Picea sitchensis [Bong.] Carr.), and almost reaching the Arctic Ocean at latitude 69° N in the District of Mackenzie, with white spruce up to 15 m high occurring on some of the islands in the Delta near Inuvik.[36] The wide variety of ecological conditions in which 4 Quebec conifers, including white spruce, are able to establish themselves, was noted by Lafond (1966),[37] but white spruce was more exacting than black spruce. In the United States, the range of white spruce extends into Maine, Vermont, New Hampshire, New York, Michigan, Wisconsin, Minnesota, and Alaska (Sargent 1922, Harlow and Harrar 1950),[28][38] where it reaches the Bering Strait in 66°44′ N" at Norton Bay and the Gulf of Alaska at Cook Inlet (Nienstaedt and Zasada 1990).[7] Southern outliers have been reported in southern Saskatchewan and the Cypress Hills of southwestern Saskatchewan (Scoggan 1957, Nienstaedt and Zasada 1990)[7][39] and southeastern Alberta,[40] northwestern Montana (Munns 1938, Harlow and Harrar 1950),[28][41] south-central Montana (Munns 1938),[41] in the Black Hills on the Wyoming–South Dakota boundary (Munns 1938, Fernald 1950),[41][42] on the Manitoba–North Dakota boundary (Munns 1938),[41] and at Shushan, New York (Cook and Smith 1959).[43] White spruce is the northernmost tree species in North America, reaching just north of 69°N latitude in the Mackenzie River delta.[44] It grows between sea level and an elevation of {{Convert|1520|m|ft|abbr=on}}. Its northern distribution roughly correlates to the location of the tree line, which includes an isothermic value of {{Convert|10|C|F}} for mean temperature in July, as well as the position of the Arctic front; cumulative summer degree days, mean net radiation, and the amount of light intensities also figure. White spruce is generally found in regions where the growing season exceeds 60 days annually.[7] The southern edge of the zone in which white spruce forms 60% or more of the total stand corresponds more or less to the July isotherm of {{Convert|18|C|F}} around the Great Lakes; in the Prairie Provinces its limit is north of this isotherm. During the summer solstice, day length values range from 17 hours at its southern limits to 24 hours above the Arctic Circle.[7] One of the hardiest conifers, white spruce in parts of its range withstands mean daily January temperature of {{convert|-6.7|C|F}} and extreme minimum temperatures as low as {{convert|-56.5|C|F}}; minimum temperatures of {{convert|-50|C|F}} are general throughout much of the range except the southernmost and southeasternmost parts (Fowells 1965).[45] By itself, or with black spruce and tamarack (Larix laricina), white spruce forms the northern boundary of tree-form growth (Sutton 1969).[46] White spruce up to 15 m in height occur at 69°N on islands in the Mackenzie Delta near Inuvik in the Northwest Territories. Hustich (1966)[47] depicted Picea spp. as forming the northernmost limit of tree growth in North America. The arctic or northern timberline in North America forms a broad transition zone from Labrador to northern Alaska. In Labrador, white spruce is not abundant and constitutes less than 5% of the forest, with a range that coincides very closely with that of black spruce but extending slightly further north (Wilton 1964).[48] The range of white spruce extends westwards from Newfoundland and Labrador, and along the northern limit of trees to Hudson Bay, Northwest Territories, Yukon, and into northwestern Alaska (Fowells 1965).[45] Although Bell (1881)[34] was emphatic that it “nowhere reaches the Atlantic coast [from which it recedes] further and further in going north”. Across western Canada and Alaska, white spruce occurs further north than black spruce, and, while poplar (Populus), willow, and birch may occur along streams well into the tundra beyond the limits of spruce, the hardwoods are usually no more than scrub (Hustich 1953).[49] Spruce characteristically occurs in fingers of tree-form forest, extending far down the northern rivers and as scattered clumps of dwarfed “bush” spruce on intervening lands (Munns 1938, Halliday and Brown 1943).[41][50] In Manitoba, Scoggan (1957, citing a 1951 but otherwise unreferenced report by Baldwin)[39] noted that the northernmost collection of white spruce was at latitude 59°48’N, but Bryson et al. (1965)[51] found white spruce in the northern edge of continuous forest in central Canada at Ennadai Lake, about 60°45′ N, 101°’W, just north of the northwest corner of Manitoba. Bryson et al. (1965)[51] noted that the forest retained “the same general characteristics as when it was first described [by Tyrrell (1897)[52]] in 1896”. Collins and Sumner (1953)[35] reported finding white spruce within 13 km of the Arctic coast in the Firth valley, Yukon, at about 69°30′ N, 139°30′ W, and Sargent (1922)[38] noted that white spruce in Alaska “reached Behring Strait in 66°44′ N”. Climate, especially temperature, is obviously a factor in determining distributions of northern flora. Halliday and Brown (1943)[50] suggested that white spruce’s northern limit corresponds “very closely” with the July mean monthly isotherm of 10 °C in Ungava, but that the northern limit west of Hudson Bay was south of that isotherm. Other climatic factors that have been suggested as affecting the northern limit of white spruce include: cumulative summer degree days, position of the Arctic front in July, mean net radiation especially during the growing season, and low light intensities (Nienstaedt and Zasada 1990).[7] Topography, soil conditions, and glaciation may also be important in controlling northern limits of spruce (Drew and Shanks 1965).[53]The southern limit of distribution of white spruce is more complex. From east of the main range of coastal mountains in British Columbia, the southern continuous limit of white spruce is the forest/prairie interface through Alberta, Saskatchewan, Manitoba, the northern parts of Minnesota and Wisconsin, central Michigan, northeastern New York, and Maine (Fowells 1965).[45] Sargent (1922)[38] and Harlow and Harrar (1950)[28] also included Vermont and New Hampshire; and, while Dame and Brooks (1901)[54] excluded New York and states further west, they included Massachusetts as far south as Amherst and Northampton, “probably the southern limit of the species” in that area. Nisbet (1905)[55] gave the range of white spruce as extending to “Carolina”, but he did not recognize red spruce as a species and presumably included it with white spruce. Towards the southern parts of its range, white spruce encounters increasingly effective ecological competition from hardwoods, some of which may reinforce their growth-rate or sprouting competitiveness with allelopathic depredation of coniferous regeneration (Tubbs 1976).[56] Further southward extension of the distribution is inhibited by white spruce’s cold requirement. White spruce as an exotic speciesAs an exotic, white spruce is widespread but uncommon. It was introduced into England (Hereman 1868)[57] and parts of continental Europe (Nisbet 1905, Jackson 1948)[55][58] in or soon after the year 1700, into Denmark about 1790 (Sabroe 1954),[59] and into Tasmania and Ceylon shortly before 1932 (Troup 1932).[60] Nisbet (1905)[55] noted that firmly-rooted white spruce served very well to stabilize windswept edges of woods in Germany. In a narrow belt of mixed Norway and white spruces over an extremely exposed hilltop crest at high elevation in northern England, the Norway spruce were “completely dwarfed” whereas the white spruce had reached heights of between 3 and 4.3 m (Guillebaud et al. 1920).[61] The age of the belt was not recorded, but adjoining 66-year-old stands may have been of the same vintage. White spruce has also been used as a minor plantation species in England (Selby 1842, Anon. 1879)[62][63] and Scotland (United Kingdom Forestry Commission 1920).[64] In Scotland, at Corrour, Inverness-shire, Sir John Stirling Maxwell in 1907 began using white spruce in his pioneering plantations at high elevations on deep peat. However, plantations in Britain have generally been unsatisfactory (Edlin 1962),[65] mainly because of damage by spring frosts after mild weather had induced flushing earlier in the season. However, the species is held in high regard in the Belgian peat region, where it grows better than do the other spruces (Fraser 1933).[66] EcologyWhite spruce is a climax canopy tree in the boreal forests of Canada and Alaska. It generally occurs on well-drained soils in alluvial and riparian zones, although it also occurs in soils of glacial and lacustrine origin.[7] The understory is dominated by feather mosses (Hylocomium splendens and Pleurozium schreberi, Ptilium crista-castrensis, and Dicranum spp.), and occasionally peat moss.[67] In the far north, the total depth of the moss and underlying humus is normally between {{Convert|25|to|46|cm|in|abbr=on}}, although it tends to be shallower when hardwoods are present in the stand.[7] White spruce grows in soils with pH values of 4.7–7.0, although they have been found in soils as acidic as 4.0 in subalpine fir forests in the Northwest Territories. A presence of calcium in the soil is common to white spruce found in northern New York. White spruce most commonly grows in the soil orders of Alfisols and Inceptisols. Soil properties such as fertility, temperature, and structural stability are partial determinants of the ability of white spruce to grow in the extreme northern latitudes.[7] In the northern limits of its range, white spruce is the climax species along with black spruce; birch and aspen are the early succession species.[67] Wildfires typically occur every 60 to 200 years, although they have been known to occur as infrequently as every 300 years.[67]White spruce will grow in USDA Growing Zones 3-7, but is not adapted to heat and humidity and will perform poorly in a hot climate.{{citation needed|date=February 2014}} The tree attains its greatest longevity and growth potential in Zones 3-4. SoilsWhite spruce occurs on a wide variety of soils, including soils of glacial, lacustrine, marine, and alluvial origins; overlying basic dolomites, limestones and acidic Precambrian and Devonian granites and gneisses; and Silurian sedimentary schists, shales, slates, and conglomerates (Halliday 1937).[68] The wide range of textures accommodated includes clays (Wilde et al. 1949, 1954; Nienstaedt 1957, Rowe 1972),[32][69][70][71] even those that are massive when wet and columnar when dry (Jameson 1963),[72] and sand flats, and coarse soils (Forest Section L 4d, Rowe 1972).[32] Its occurrence on some organic soils is not characteristic, except perhaps on shallow mesic organic soils in Saskatchewan and in association with black spruce on organic soils in central Yukon (Nienstaedt and Zasada 1990).[7] Podzolized, brunisolic, luvisolic, gleysolic, and regosolic (immature) soils are typical of those supporting white spruce throughout the range of the species (Nienstaedt 1957).[71] Soils supporting white spruce are most commonly Alfisols or Inceptisols (Nienstaedt and Zasada 1990).[7] In the podzol region of Wisconsin, white spruce occurs on loam podzols, podzolized gley loams, strongly podzolized clays, gley-podzol clays, stream-bottom soils, and wood peat (Wilde et al. 1949).[69] Moist sandy loams also support good growth (Harlow and Harrar 1950).[28] On sandy podzols (Wilde et al. 1949),[69] it is usually a minor species (Nienstaedt and Zasada 1990).[7] Good development occurs on moist alluvium (Seeley, cited by Nienstaedt 1957; Jeffrey 1961, 1964; Lacate et al. 1965; Viereck 1973)[71][73][74][75][76] on the banks of streams and borders of swamps (Sargent 1898, Kenety 1917, Rowe 1972).[32][77][78] White spruce makes good growth on well-drained lacustrine soils in Alberta Mixedwoods (Heger 1971),[79] on moderately-well-drained clay loams in Saskatchewan (Kabzems 1971),[80] and on melanized loams and clays (with sparse litter and a dark-coloured organically-enriched mineral horizon) in the Algoma district of Ontario (Wilde et al. 1954).[70] White spruce becomes less accommodating of soil with increasing severity of climate. The distribution of white spruce in Labrador seems to depend almost entirely on the character of the soil (Sargent 1898),[77] and between the southwestern shores of Hudson Bay and the northeastern regions of Saskatchewan, white spruce is confined to very local physiographic features, characterized by well-drained or fertile soils (Ritchie 1956).[81] On dry, deep, outwash deposits in northern Ontario, both white spruce and aspen grow slowly (MacLean 1960).[82] But, broadly, white spruce is able to tolerate considerable droughtiness of sites that are fertile, and no fertile site is too moist unless soil moisture is stagnant (Sutton 1968).[83] Soil fertility holds the key not just to white spruce growth but to the distribution of the species. At least moderate fertility is needed for good growth, but white spruce occurs on many sites where nutrient deficiencies depress its growth more than that of black spruce, red spruce, Norway spruce, and the pines generally (Heiberg and White 1951, Lafond 1954, McLeod 1956, MacArthur 1957, Paine 1960, Swan 1960).[84] Minimum soil-fertility standards recommended for white spruce sufficient to produce 126 to 157 m3/ha of wood at 40 years are much higher than for pine species commonly planted in the Lake States (Wilde 1966):[85] 3.5% organic matter, 12.0 meq/100 g exchange capacity, 0.12% total N, 44.8 kg/ha available P, 145.7 kg/ha available K, 3.00 meq/100 g exchangeable Ca, and 0.70 meq/100 g exchangeable Mg. Forest floors under stands dominated by white spruce respond in ways that vary with site conditions, including the disturbance history of the site (Nienstaedt and Zasada 1990).[7] Composition, biomass, and mineral soil physical and chemical properties are affected. In Alaska, the accumulation of organic layers (to greater thicknesses in mature stands of spruce than those in hardwood stands on similar sites) leads to decreased soil temperatures, in some cases leading to the development of permafrost (Viereck 1970a, b, Viereck et al. 1983).[86][87][88] Acidity of the mineral soil sampled at an average depth of 17 cm in 13 white spruce stands on abandoned farmland in Ontario increased by 1.2 pH units over a period of 46 years (Brand et al. 1986).[89] A considerable range of soil pH is tolerated by white spruce (Nienstaedt 1957).[71] Thrifty stands of white spruce in Manitoba have developed on soils of pH 7.6 at only 10 cm below the surface, and pH 8.4 at 43 cm below the surface (Stoeckeler 1938, USDA Forest Service 1938);[90][91] rooting depth in those soils was at least 81 cm. An abundant calcium supply is common to most white spruce locations in New York state (Nienstaedt and Zasada 1990).[7] Chlorosis was observed in young white spruce in heavily limed nursery soils at about pH 8.3 (Stone, cited by Nienstaedt 1957).[71] Wilde (1966)[85] gave 4.7 to 6.5 as the approximate optimum range of pH for white spruce in Wisconsin, but optimum growth seems possible at pH levels up to 7.0 and perhaps higher (Sutton 1968).[83] Alluvium on the floodplains of northern rivers shows pH levels from 5.0 to 8.2 (Zasada et al. 1977).[92] High-lime ecotypes may exist (Pelletier 1966),[93] and in Canada Forest Section B8 the presence of balsam poplar and white spruce on some of the moulded moraines and clays seems to be correlated with the considerable lime content of these materials (Rowe 1972, Stiell 1976),[32][94] while calcareous soils are favourable sites for northern outliers of white spruce (Hustich 1953).[49] Mature stands of white spruce in boreal regions often have well-developed moss layers dominated by feather mosses, e.g., Hylocomium splendens (Hedw.) B.S.G., Pleurozium schreberi (Brid.) Mitt., Ptlium crista-castrensis (Hedw.) De Not., and Dicranum Hedw. spp. rather than Sphagnum Dill. spp. (La Roi and Stringer 1976, Viereck 1987).[95][96] The thickness of the moss–organic layer commonly exceeds 25 cm in the far north and may approach twice that figure. The mosses compete for nutrients and have a major influence on soil temperatures in the rooting zone. Permafrost development in parts of Alaska, Yukon, and the Northwest Territories is facilitated by the insulative organic layer (Viereck 1970a, b, Gill 1975, Van Cleve and Yarie 1986).[86][87][97][98] The role of windthrow in maintaining diversification of the bryophyte flora in boreal spruce forests has been described by Jonsson et al. (1990)[99] and Jonsson and Dynesius (1993).[100] Cold hardinessWhite spruce is extremely hardy to low temperatures, provided the plant is in a state of winter dormancy. Throughout the greater part of its range, white spruce routinely survives and is undamaged by winter temperatures of {{convert|−50|C|F}}, and even lower temperatures occur in parts of the range (Fowells 1965, Nienstaedt and Zasada 1990).[7][45] Boreal Picea are among the few extremely hardy conifers in which the bud primordia are able to survive temperatures down to {{convert|-70|C|F|disp=comma}} (Sakai and Larcher 1987).[101] Especially important in determining the response of white spruce to low temperatures is the physiological state of the various tissues, notably the degree of "hardening" or dormancy. A natural progression of hardening and dehardening occurs in concert with the seasons (Glerum 1985).[102] While different tissues vary in ability to tolerate exposure to stressful temperatures, white spruce, as with woody plants in general, has necessarily developed sufficient winter hardiness in its various tissues to enable them to survive the minimum temperatures experienced in the distribution range. White spruce is subject to severe damage from spring frosts. Newly flushed shoots of white spruce are very sensitive to spring frost (Smith 1949, Rowe 1955, McLeod 1964).[103][104][105] This sensitivity is a major constraint affecting young trees planted without overstorey nurses in boreal climates (Sutton 1992).[106] Forest successionForest succession in its traditional sense implies two important features that resist direct examination (Solomon et al. 1981).[107] First, classical definitions generally connote directional changes in species composition and community structure through time, yet the time frame needed for documentation of change far exceeds an average lifespan (Solomon et al. 1981).[107] The second feature that defies quantitative description is the end point or climax. Floodplain deposits in the Northwest Territory, Canada, are important in relation to the development of productive forest types with a component of white spruce (Jeffrey 1964).[74] The most recently exposed surfaces are occupied by sandbar vegetation or riparian shrub willows and alder (Alnus incana); with increasing elevation, the shrubs give way successively to balsam poplar and white spruce forest. In contrast, older floodplains, with predominantly brown wooded soils, typically carry white spruce–trembling aspen mixedwood forest. Interrelationships among nutrient cycling, regeneration, and subsequent forest development on floodplains in interior Alaska were addressed by Van Cleve et al. (1980),[108] who pointed out that the various stages in primary succession reflect physical, chemical, and biological controls of ecosystem structure and function. Thus, each successional stage has a species combination in harmony with site quality. Short-circuiting succession by planting a late successional species such as white spruce on an early successional surface may result in markedly reduced growth rates because of nitrogen insufficiency. Without application of substantial amounts of fertilizer, use would have to be made of early successional alder and its site-ameliorating additions of nitrogen. Neiland and Viereck noted that “the slow establishment and growth of spruce under birch stands [in Alaska] may be partially due to effects of shading and general competition for water and nutrients, but may also be more directly related to the birch itself. Heikinheimo (1915 cited by Lutz 1956),[109][110] found that birch ash inhibited white spruce seedlings, and Gregory (1966)[111] found that birch litter has a smothering effect on spruce seedlings” (Neiland and Viereck 1977).[112] On dry upland sites, especially south-facing slopes, the mature vegetation is white spruce, white birch, trembling aspen, or a combination of these species. Succession follows in one of two general patterns. In most cases, aspen and birch develop as a successional stage after fire before reaching the spruce stage. But, occasionally, with optimal site conditions and a source of seed, white spruce will invade with the hardwoods or within a few years thereafter, thereby producing even-aged white spruce stands without an intervening hardwood stage. Associated forest coverThe White Spruce Cover Type may include other species in small numbers. In Alaska, associates include paper birch, trembling aspen, balsam poplar, and black spruce; in western Canada, additional associates are subalpine fir, balsam fir, Douglas-fir, jack pine, and lodgepole pine (Dyrness 1980).[113] Seral species giving way to white spruce include paper birch, aspen, balsam poplar, jack pine, and lodgepole pine. On certain river bottom sites, however, black spruce may replace white spruce (Viereck 1970a).[113] Earlier successional stages leading to the white spruce climax are the white spruce–paper birch, white spruce–aspen, balsam poplar, jack pine, and lodgepole pine types. The type shows little variation. The forest is generally closed and the trees well formed, other than those close to the timberline. Lesser vegetation in mature stands is dominated by mosses. Vascular plants are typically few, but shrubs and herbs that occur “with a degree of regularity” include: alder, willows, mountain cranberry, red-fruit bearberry, black crowberry, prickly rose, currant, buffaloberry, blueberry species, bunchberry, twinflower, tall lungwort, northern comandra, horsetail, bluejoint grass, sedge species, as well as ground-dwelling mosses and lichens. Several white spruce communities have been identified in interior Alaska: white spruce/feathermoss; white spruce/dwarf birch/feathermoss; white spruce/dwarf birch/sphagnum; white spruce/avens/moss; and white spruce/alder/bluejoint (Viereck 1975, Dyrness 1980).[114][113] Of the Eastern Forest Cover Types recognized by the Society of American Foresters (Eyre 1980),[184] only one, White Spruce, names that species in its title. The eastern White Spruce Cover Type, as defined, encompasses white spruce both in pure stands, and in mixed stands “in which white spruce is the major [undefined] component” (Payette 1980).[115] In most of its range, white spruce occurs more typically in association with trees of other species than in pure stands. White spruce is an associated species in the following Eastern Forest cover types, by the Society of American Foresters: In the Boreal Forest Region: (1) jack pine, (5) balsam fir, (12) black spruce, (16) aspen, (18) paper birch, and (38) tamarack. In the Northern Forest Region: (15) red pine, (21) eastern white pine, (24) hemlock-yellow birch, (25) sugar maple-beech-yellow birch, (27) sugar maple, (30) red spruce-yellow birch, (32) red spruce, (33) red spruce-balsam fir, (37) northern white-cedar, and (39) black ash-American elm-red maple (Nienstaedt and Zasada 1990, Eyre 1980).[7][116] PredatorsOutbreaks of spruce beetles have destroyed over {{convert|2300000|acre|km2}} of forests in Alaska.[117] Although sometimes described, e.g., by Switzer (1960),[118] as relatively resistant to attack by insects and disease, white spruce is far from immune to depredation. Important insect pests of white spruce include the spruce budworm (Choristoneura fumiferana [Clemens]), the yellow-headed spruce sawfly (Pikonema alaskensis Rohwer), the European spruce sawfly (Gilpinia hercyniae [Hartig]), the spruce bud moth (Zeiraphera canadensis)[190], and spruce beetle (Dendroctonus rufipennis [Kirby]) (Fowells 1965, Rose and Lindquist 1985, Ives and Wong 1988).[119][120][121] As well, other budworms, sawflies, and bark beetles, gall formers, bud midges, leaf miners, aphids, leaf eaters, leaf rollers, loopers, mites, scales, weevils, borers, pitch moths, and spittlebugs cause varying degrees of damage to white spruce (Ives and Wong 1988).[121] A number of sawflies feed on spruce trees. Among them European spruce sawfly, yellow-headed spruce sawfly, green-headed spruce sawfly and the spruce webspinning sawfly (Rose and Lindquist 1985).[120] More than a dozen kinds of looper feed on the spruces, fir, and hemlock in eastern Canada. The full-grown larvae of the larvae vary in length from 15 mm to 35 mm. Some feed briefly in the fall and complete their feeding in the spring; others feed mainly in the summer; still others feed mainly in the late summer and fall. The fall and spring feeding group includes the dash-lined looper (Protoboarmia porcelaria indicataria Walker), the diamond-backed looper (Hypagyrtis piniata Packard), the fringed looper (Campaea perlata Guenée), and the false loopers (Syngrapha species). The summer feeding group includes the false hemlock looper (Nepytia canosaria Walker), occasionally occurring in large numbers and usually in conjunction with the hemlock looper (Lambdina fiscellaria fiscellaria Guenée), the small spruce loopers Eupithecia species, the yellowlined conifer looper (Cladara limitaria Walker), and the saddleback looper (Ectropis crepuscularia Denis and Schiffermüller). The late summer and fall group includes the common spruce-fir looper (Semiothisa signaria dispuncta Walker) and the similar Semiothisa fissinotata Walker on hemlock, the small spruce loopers Eupithecia species, the spruce looper Caripeta divisata Walker, occasionally abundant, the transversebanded looper (Hydriomena divisaria Walker), and the whitelined looper (Eufidonia notataria Walker). UsesWhite spruce is of major economic importance in Canada for its wood, harvested for paper-making and construction. It is also used to a small extent as a Christmas tree. The wood is also exported to Japan where, known as "shin-kaya", it is used to make go boards as a substitute for the rare kaya wood. Black Hills Spruce (Picea glauca var. densata) is used for bonsai (see image). White spruce is the provincial tree of Manitoba and the state tree of South Dakota. VarietiesSeveral geographical varieties have been described, but are not accepted as distinct by all authors. These comprise, from east to west:[3]
The two western varieties are distinguished by pubescent (downy) shoots, and may be related to extensive hybridisation and/or intergradation with the closely related Engelmann spruce found further south in the Rocky Mountains. White spruce also hybridises readily with the closely related Sitka spruce where they meet in southern Alaska; this hybrid is known as Picea × lutzii.[3] CultivarsNumerous cultivars of various sizes, colours and shapes have been selected for use in parks and gardens. The following have gained the Royal Horticultural Society's Award of Garden Merit.[122]
'Conica' is a dwarf conifer with very slender leaves, like those normally found only on one-year-old seedlings, and very slow growth, typically only {{Convert|2|-|10|cm|in|abbr=on}} per year. Older specimens commonly 'revert', developing normal adult foliage and starting to grow much faster; this 'reverted' growth must be pruned if the plant is to be kept dwarf. ChemistryIsorhapontin can be found in spruce species such as the white spruce.[126]References1. ^{{ThePlantList |id=kew-2563473 |taxon=Picea glauca}} 2. ^{{PLANTS|id=PIGL|taxon=Picea glauca|accessdate=2 October 2015}} 3. ^1 2 3 4 {{cite book |last=Farjon |first=A. |date=1990 |title=Pinaceae. Drawings and Descriptions of the Genera |publisher=Koeltz Scientific Books |isbn=3-87429-298-3}} 4. ^{{cite book |last=Rushforth |first=K. |date=1987 |title=Conifers |publisher=Helm |isbn=0-7470-2801-X}} 5. ^1 {{IUCN2006|assessor=Conifer Specialist Group|year=1998|id=42323|title=Picea glauca|downloaded=12 May 2006}} 6. ^{{Gymnosperm Database |family=Pinaceae |genus=Picea |species=glauca}} 7. ^1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 {{Silvics |first1=Hans |last1=Nienstaedt |first2=John C. |last2=Zasada |volume=1 |genus=Picea |species=glauca | accessdate = 14 November 2010}} 8. ^1 {{cite journal |last=Waldron |first=R.M. |date=1965 |title=Cone production and seedfall in a mature white spruce stand |journal=For. Chron. |volume=41 |issue=3 |pages=314–329}} 9. ^Zasada, J.C.; Viereck, L.A. 1970. White spruce cone and seed production in interior Alaska, 1957–68. USDA, For. Serv., Pacific NW For. Range Exp. Sta., Portland OR, Res. Note PNW-129. 11 p. [Coates et al. 1994] 10. ^Hellum, A.K. 1976. Grading seed by weight in white spruce. USDA, For. Serv., Tree Plant. Notes 27(1):16–17, 23–24. (Cited in Coates et al. 1994). - Zasada, J.C.; Foote, M.J.; Deneke, F.J.; Parkerson, R.H. 1978. Case history of an excellent white spruce cone and seed crop in interior Alaska: cone and seed production, germination and seedling survival. USDA, For. Serv., Pacific NW For. Range Exp. Sta., Portland OR, Gen. Tech. Rep. PNW-65. 53 p. 11. ^1 Crossley, D.I. 1953. Seed maturity in white spruce. Canada Dep. Resour. and Devel., For. Branch, For. Res. Div., Ottawa ON, Silv. Res. Note 104. 16 p. 12. ^1 Cram, W.H.; Worden, H.A. 1957. Maturity of white spruce cones and seed. For. Sci. 3:263–269. 13. ^Teich, A.H. 1970. Genetic control of female flower color and random mating in white spruce. Can. Dep. Fish. For., Can. For. Serv., Ottawa ON, Bi-mo. Res. Notes 26:2. 14. ^1 Zasada, J.C. 1973. 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Forest Cover Types of the United States and Canada. Soc. Amer. Foresters, Washington DC. 148 p. 117. ^{{cite web|url=http://www.na.fs.fed.us/spfo/pubs/fidls/sprucebeetle/sprucebeetle.htm |title=The Spruce Beetle – FIDL |publisher=Na.fs.fed.us |accessdate=2013-07-13}} 118. ^Switzer, A.L.K. 1960. Spruce management for the future. For. Chron. 36(2):163–165. 119. ^Fowells, H.A. 1965. Picea (spruces). p. 287–327 in Silvics of Forest Trees of the United States. H.A. Fowells (Compiler), USDA, Forest Service, Washington DC, Agric. Handbook No. 271. 120. ^1 2 Rose, A.H.; Lindquist, O.H. 1985. Insects of eastern spruces, fir and, hemlock, revised edition. Gov’t Can., Can. For. Serv., Ottawa, For. Tech. Rep. 23. 159 p. (cited in Coates et al. 1994, cited orig ed 1977) 121. ^1 Ives, W.G.H.; Wong, H.R. 1988. Tree and shrub insects of the prairie provinces. Gov’t Can., Can. For. Serv., Edmonton AB, Inf. Rep. NOR-X-292. 327 p.[Coates et al. 1994] 122. ^{{cite web | url = https://www.rhs.org.uk/plants/pdfs/agm-lists/agm-ornamentals.pdf | title = AGM Plants - Ornamental | date = July 2017 | page = 78 | publisher = Royal Horticultural Society | accessdate = 25 April 2018}} 123. ^{{cite web | url = https://www.rhs.org.uk/Plants/90592/i-Picea-glauca-i-Echiniformis/Details| title = RHS Plantfinder - Picea glauca ‘Echiniformis’ | accessdate = 25 April 2018}} 124. ^{{cite web | url = https://www.rhs.org.uk/Plants/59793/i-Picea-glauca-i-var-i-albertiana-i-Alberta-Globe/Details | title = RHS Plantfinder - Picea glauca var. albertiana ‘Alberta Globe’ | accessdate = 25 April 2018}} 125. ^{{cite web | url = https://www.rhs.org.uk/Plants/83270/i-Picea-glauca-i-var-i-albertiana-i-Conica/Details | title = RHS Plantfinder - Picea glauca var. albertiana ‘Conica’ | accessdate = 25 April 2018}} 126. ^{{Cite journal | last1 = Hammerbacher | first1 = A. | last2 = Ralph | first2 = S. G. | last3 = Bohlmann | first3 = J. | last4 = Fenning | first4 = T. M. | last5 = Gershenzon | first5 = J. | last6 = Schmidt | first6 = A. | doi = 10.1104/pp.111.181420 | title = Biosynthesis of the Major Tetrahydroxystilbenes in Spruce, Astringin and Isorhapontin, Proceeds via Resveratrol and is Enhanced by Fungal Infection | journal = Plant Physiology | volume = 157 | issue = 2 | pages = 876–890 | year = 2011 | pmid = 21865488| pmc =3192583 }} External links
23 : Least concern plants|Ornamental trees|Picea|Provincial symbols of Manitoba|Taiga and boreal forests|Trees of humid continental climate|Trees of continental subarctic climate|Trees of Alberta|Trees of British Columbia|Trees of Canada|Trees of Manitoba|Trees of Ontario|Trees of Saskatchewan|Trees of Western Canada|Trees of Subarctic America|Trees of Eastern Canada|Trees of Alaska|Trees of the Northeastern United States|Trees of the Northwestern United States|Trees of the North-Central United States|Trees of the United States|Trees of the Great Lakes region (North America)|Symbols of South Dakota |
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