Picea glauca

From Wikipedia, the free encyclopedia
Jump to: navigation, search
Picea glauca
Picea glauca Fairbanks.jpg
Mature white spruce in Alaska
Scientific classification
Kingdom: Plantae
Division: Pinophyta
Class: Pinopsida
Order: Pinales
Family: Pinaceae
Genus: Picea
Species: P. glauca
Binomial name
Picea glauca
(Moench) Voss
Picea glauca levila.PNG
Synonyms[1]

Picea glauca (white spruce) is a species of spruce native to the northern temperate and boreal forests in North America, from central Alaska to as far east as the Avalon Peninsula in Newfoundland, and south to northern Montana, Minnesota, Wisconsin, Michigan, northwestern Pennsylvania, upstate New York, Vermont, New Hampshire, and Maine; there is also an isolated population in the Black Hills of South Dakota and Wyoming.[2][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]

Description[edit]

Foliage and cones
Seeds

The white spruce is a large coniferous evergreen tree which grows normally to 15 to 30 metres (49 to 98 ft) tall, but can grow up to 40 metres (130 ft) tall with a trunk diameter of up to 1 metre (3.3 ft). The bark is thin and scaly, flaking off in small circular plates 5 to 10 centimetres (2.0 to 3.9 in) 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, 12 to 20 millimetres (0.47 to 0.79 in) long, rhombic in cross-section, glaucous blue-green above with several thin lines of stomata, and blue-white below with two broad bands of stomata.[2][3]

The cones are pendulous, slender, cylindrical, 3 to 7 centimetres (1.2 to 2.8 in) long and 1.5 centimetres (0.59 in) wide when closed, opening to 2.5 centimetres (0.98 in) broad. They have thin, flexible scales 15 millimetres (0.59 in) 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, 2 to 3 millimetres (0.079 to 0.118 in) long, with a slender, 5 to 8 millimetres (0.20 to 0.31 in) long pale brown wing.[2][3]

Root system[edit]

The root system of white spruce is highly variable and adaptable (Wagg 1964, 1967),[8][9] 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).[10] In the nursery, or naturally in the forest, white spruce usually develops several long “running” roots just below the ground surface (Mullin 1957).[11]

The structure of the tracheids in the long lateral roots of white spruce varies with soil nitrogen availability (Krasowski and Owens 1999).[12]

Seeds[edit]

Seeds are small (2.5 mm to 5.0 mm 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).[13][14]

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).[15][16]

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).[17] Mature seed “snaps in two” when cut by a sharp knife on a firm surface (Crossley 1953).[17]

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)[18]

Cone colour also can be used to help determine the degree of maturation, but cones may be red, pink or green (Teich 1970).[19] Collection and storage dates and conditions influence germination requirements and early seedling growth (Zasada 1973, Edwards 1977, Winston and Haddon 1981).[20][21][22]

A bushel (35 L) of cones, which may contain 6500 to 8000 cones, yields 6 to 20 ounces (170g to 567 g) of clean seed (USDA Forest Service 1948).[23]

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).[18][20][22] Weather affects both the initiation and pattern of seed dispersal (Nienstaedt and Zasada 1990),[24] 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),[25] even into the next growing season (Rowe 1953).[26] Seed dispersal occurs mainly in late summer or early fall (Waldron 1965).[13]

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),[24] but these can vary among trees in the same stand (Zasada 1986).[25] 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).[25]

Stem[edit]

White 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)[27] 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 6 m to 10 m high on the shore of Urquhart Lake, Northwest Territories, were found to be more than 300 years old (Hare and Ritchie 1972),[28]

Bark[edit]

The bark of mature white spruce is scaly or flaky, gray-brown (Brayshaw 1960)[29] or ash-brown (Harlow and Harrar 1950),[30] 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).[31]

White spruce bark is mostly less than 8 mm (Hale 1955)[32] and not more than 9.5 mm thick (Chang 1954).[33]

Distribution[edit]

White 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),[34] extending into every province and territory (Forestry Branch 1961).[35] 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).[36] Collins and Sumner (1953)[37] 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 (Northwest Territories webpage). The wide variety of ecological conditions in which 4 Quebec conifers, including white spruce, are able to establish themselves, was noted by Lafond (1966),[38] 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),[39][30] 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).[40]

Southern outliers have been reported in southern Saskatchewan and the Cypress Hills of southwestern Saskatchewan (Scoggan 1957, Nienstaedt and Zasada 1990)[41][40] and southeastern Alberta (Cypress Hills Alberta Web site), northwestern Montana (Munns 1938, Harlow and Harrar 1950),[42][30] south-central Montana (Munns 1938),[42] in the Black Hills on the Wyoming–South Dakota boundary (Munns 1938, Fernald 1950),[42][43] on the Manitoba–North Dakota boundary (Munns 1938),[42] and at Shushan, New York (Cook and Smith 1959).[44]

White spruce taiga along the Denali Highway in the Alaska Range

White spruce is the northernmost tree species in North America, reaching just north of 69°N latitude in the Mackenzie River delta.[45] It grows between sea level and an elevation of 1,520 metres (4,990 ft). Its northern distribution roughly correlates to the location of the tree line, which includes an isothermic value of 10 °C (50 °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 generally is found in regions where the growing season exceeds 60 days annually.[7]

The southern distribution corresponds to the July isotherm of 18 °C (64 °F) around the Great Lakes; in the Prairie Provinces its limit is north of this isotherm. During the summer solstice, photoperiod 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 -6.7°C and extreme minimum temperatures as low as -56.5°C; minimum temperatures of -50°C are general throughout much of the range except the southernmost and southeasternmost parts (Fowells 1965).[46] By itself, or with black spruce and tamarack (Larix laricina (Du Roi) K. Koch), white spruce forms the northern boundary of tree-form growth (Sutton 1969).[47] 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)[48] 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).[49]

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).[46] Although Bell (1881)[36] 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).[50] 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).[42][51] In Manitoba, Scoggan (1957, citing a 1951 but otherwise unreferenced report by Baldwin)[41] noted that the northernmost collection of white spruce was at latitude 59°48’N, but Bryson et al. (1965)[52] 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)[52] noted that the forest retained “the same general characteristics as when it was first described [by Tyrrell (1897)[53]] in 1896”. Collins and Sumner (1953)[37] 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)[39] 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)[51] 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).[40] Topography, soil conditions, and glaciation may also be important in controlling northern limits of spruce (Drew and Shanks 1965).[54]

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).[46] Sargent (1922)[39] and Harlow and Harrar (1950)[30] also included Vermont and New Hampshire; and, while Dame and Brooks (1901)[55] 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)[56] 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/sprouting competitiveness with allelopathic depredation of coniferous regeneration (Tubbs 1976).[57] Further southward extension of the distribution is inhibited by white spruce’s cold requirement.

White spruce as an exotic species[edit]

As an exotic, white spruce is widespread but uncommon. It was introduced into England (Hereman 1868)[58] and parts of continental Europe (Nisbet 1905, Jackson 1948)[56][59] in or soon after the year 1700, into Denmark about 1790 (Sabroe 1954),[60] and into Tasmania and Ceylon shortly before 1932 (Troup 1932).[61]

Nisbet (1905)[56] 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).[62] 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)[63][64] and Scotland (United Kingdom Forestry Commission 1920).[65] 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),[66] 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).[67]

Ecology[edit]

White 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.[68] In the far north, the total depth of the moss and underlying humus is normally between 25 to 46 centimetres (9.8 to 18.1 in), 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.[68] Wildfires typically occur every 60 to 200 years, although they have been known to occur as infrequently as every 300 years.[68]

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] The tree attains its greatest longevity and growth potential in Zones 3-4.

Soils[edit]

White 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).[69] The wide range of textures accommodated includes clays (Wilde et al. 1949, 1954; Nienstaedt 1957, Rowe 1972),[70][71][72][34] even those that are massive when wet and columnar when dry (Jameson 1963),[73] and sand flats, and coarse soils (Forest Section L 4d, Rowe 1972).[34] 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).[40]

Podzolized, brunisolic, luvisolic, gleysolic, and regosolic (immature) soils are typical of those supporting white spruce throughout the range of the species (Nienstaedt 1957).[72] Soils supporting white spruce are most commonly Alfisols or Inceptisols (Nienstaedt and Zasada 1990).[40] 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).[70] Moist sandy loams also support good growth (Harlow and Harrar 1950).[30] On sandy podzols (Wilde et al. 1949),[70] it is usually a minor species (Nienstaedt and Zasada 1990).[40] Good development occurs on moist alluvium (Seeley, cited by Nienstaedt 1957; Jeffrey 1961, 1964; Lacate et al. 1965; Viereck 1973)[72][74][75][76][77] on the banks of streams and borders of swamps (Sargent 1898, Kenety 1917, Rowe 1972).[78][79][34] White spruce makes good growth on well-drained lacustrine soils in Alberta Mixedwoods (Heger 1971),[80] on moderately-well-drained clay loams in Saskatchewan (Kabzems 1971),[81] 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).[71]

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),[78] 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).[82]

On dry, deep, outwash deposits in northern Ontario, both white spruce and aspen grow slowly (MacLean 1960).[83] 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).[84] 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).[85][86][87][88][89][90] 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):[91] 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).[40] 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).[92][93][94] 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).[95]

A considerable range of soil pH is tolerated by white spruce (Nienstaedt 1957).[72] 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);[96][23] 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).[40] Chlorosis was observed in young white spruce in heavily-limed nursery soils at about pH 8.3 (Stone, cited by Nienstaedt 1957).[72] Wilde (1966)[91] 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).[84] Alluvium on the floodplains of northern rivers shows pH levels from 5.0 to 8.2 (Zasada et al. 1977).[97] High-lime ecotypes may exist (Pelletier 1966),[98] 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),[34][99] while calcareous soils are favourable sites for northern outliers of white spruce (Hustich 1953).[50]

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).[100][101] 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).[92][93][102][103] The role of windthrow in maintaining diversification of the bryophyte flora in boreal spruce forests has been described by Jonsson et al. (1990)[104] and Jonsson and Dynesius (1993).[105]

Cold hardiness[edit]

White 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 -50°C, and even lower temperatures occur in parts of the range (Fowells 1965, Nienstaedt and Zasada 1990).[106][107] Boreal Picea are among the few extremely hardy conifers in which the bud primordia are able to survive temperatures down to -70°C (Sakai and Larcher 1987).[108]

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).[109] 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).[110][34][87] This sensitivity is a major constraint affecting young trees planted without overstorey nurses in boreal climates (Sutton 1992).[111]

Forest succession[edit]

Forest succession in its traditional sense implies two important features that resist direct examination (Solomon et al. 1981).[112] 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).[112] 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).[113] 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),[114] 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),[115][116] found that birch ash inhibited white spruce seedlings, and Gregory (1966)[117] found that birch litter has a smothering effect on spruce seedlings” (Neiland and Viereck 1977).[118]

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 cover[edit]

The 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).[119] 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).[120] 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).[77][119]

Of the Eastern Forest Cover Types recognized by the Society of American Foresters (Eyre 1980),[121] 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).[122]

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).[123][121]

Varieties[edit]

A dwarf Alberta white spruce in Los Angeles County Arboretum and Botanic Garden

Several geographical varieties have been described, but are not accepted as distinct by all authors. These comprise, from east to west:[2][3]

  • Picea glauca var. glauca (Typical or Eastern white spruce). From Newfoundland west to eastern Alberta, on lowland plains.
  • Picea glauca var. densata (Black Hills white spruce). The Black Hills in South Dakota.
  • Picea glauca var. albertiana (Alberta white spruce). The Rocky Mountains in Alberta, British Columbia and northwest Montana.
  • Picea glauca var. porsildii (Alaska white spruce). Alaska and Yukon.

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.[2][3]

Predators[edit]

Outbreaks of spruce beetles have destroyed over 2,300,000 acres (9,300 km2) of forests in Alaska.[124]

Although sometimes described, e.g., by Switzer (1960),[125] 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]), and spruce beetle (Dendroctonus rufipennis [Kirby]) (Fowells 1965, Rose and Lindquist 1985, Ives and Wong 1988).[126][127][128] 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).[128]

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).[127]

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).

Uses[edit]

A dwarf Alberta spruce, with reversion in one branch
Black Hills spruce grown as bonsai

White 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.

White spruce is the provincial tree of Manitoba and the state tree of South Dakota.

Cultivars[edit]

Numerous cultivars have been selected for use in parks and gardens. The cultivar P. glauca 'Echiniformis' has gained the Royal Horticultural Society's Award of Garden Merit.[129]

A dwarf cultivar, P. glauca var. albertiana 'Conica', is a popular garden plant. It has very slender leaves, like those normally found only on one-year-old seedlings, and very slow growth, typically only 2–10 centimetres (0.79–3.94 in) 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.

Chemistry[edit]

Isorhapontin can be found in spruce species such as the white spruce.[130]

See also[edit]

References[edit]

  1. ^ "The Plant List: A Working List of All Plant Species". 
  2. ^ a b c d e Farjon, A. (1990). Pinaceae. Drawings and Descriptions of the Genera. Koeltz Scientific Books ISBN 3-87429-298-3.
  3. ^ a b c d e Rushforth, K. (1987). Conifers. Helm ISBN 0-7470-2801-X.
  4. ^ Conifer Specialist Group (1998). Picea glauca. 2006. IUCN Red List of Threatened Species. IUCN 2006. www.iucnredlist.org. Retrieved on 12 May 2006.
  5. ^ Gymnosperm Database: Picea glauca
  6. ^ Flora of North America: Picea glauca
  7. ^ a b c d e f Nienstaedt, Hans; Hans Nienstaedt; John C. Zasada (1990). "Picea glauca (Moench) Voss". Silvics of North America, Volume 1:Conifers. United States Forest Service. Retrieved 2010-11-14. 
  8. ^ Wagg, J.W.G. 1964. White spruce regeneration on the Peace and Slave River lowlands. Can. Dep. For., For. Res. Branch, Ottawa ON, Publ. 1069. 35 p.
  9. ^ Wagg, J.W.B. 1967. Origin and development of white spruce root-forms. Can. Dep. For. Rural Devel., For. Branch, Ottawa ON, Publ. 1192. 45 p.
  10. ^ Sutton, R.F. 1969. Form and development of conifer root systems. Commonw. For. Bureau, Oxford, U.K., Tech. Communication No. 7. 131 p.
  11. ^ Mullin, R.E. 1957. Experiments with root and top pruning of white spruce nursery stock. Ont. Dep. Lands For., Res. Div., Toronto ON, Res. Rep. 36. 31 p.
  12. ^ Krasowski, M.J.; Owens, J.N. 1999. Tracheids in white spruce seedling’s long lateral roots in response to nitrogen availability. Plant and Soil 217(1/2):215–228.
  13. ^ a b Waldron, R.M. 1965. Cone production and seedfall in a mature white spruce stand. For. Chron. 41(3):314–329.
  14. ^ 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]
  15. ^ 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).
  16. ^ 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.
  17. ^ a b 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.
  18. ^ a b Cram, W.H.; Worden, H.A. 1957. Maturity of white spruce cones and seed. For. Sci. 3:263–269.
  19. ^ 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.
  20. ^ a b Zasada, J.C. 1973. Effect of cone storage method and collection date on Alaskan white spruce (Picea glauca) seed quality. p. 1–10 (paper 19) in Proc. Seed Problems. IUFRO Symp. Seed Processing, Bergen, Norway. Working Party S2.01, Royal Coll. For., Bergen, Norway, Vol. 1. [Coates et al. 1994]
  21. ^ Edwards, I.K. 1977. Fertility of transplant fields at the Prince Albert Forest Nursery. Can. Dep. Fish. Environ., Can. For. Serv., Northern For. Res. Centre, Edmonton AB, Inf. Rep. NOR-X-189. 21 p.
  22. ^ a b Winston, D.A.; Haddon, B.D. 1981. Effects of early cone collection and artificial ripening on white spruce and red pine germination. Can. J. For. Res. 11:817–826.
  23. ^ a b USDA Forest Service. 1948. Woody-plant Seed Manual. USDA, For. Serv., Washington DC, Misc. Publ. 654. 416 p.
  24. ^ a b Nienstaedt, H.; Zasada, J.C. 1990. Picea glauca (Moench) Voss. p. 204–226 in Burns, R.M. and Honkala, B.H. (Tech. Coord.), Silvics of North America, Vol. 1, Conifers. USDA, For. Serv., Washington DC, Agric. Handbook 654.
  25. ^ a b c Zasada, J. 1986. Natural regeneration of trees and tall shrubs on forest sites in interior Alaska. p. 44–73 in Van Cleve, K.; Chapin, F.S.; Flanagan, P.W.; Viereck, L.A.; Dyrness, C.T. (Eds.). Forest Ecosystems in the Alaskan Taiga: a Synthesis of Structure and Function. Springer-Verlag New York NY.
  26. ^ Rowe, J.S. 1953. Viable seed on white spruce trees in midsummer. Can. Dep. Northern Affairs and National Resources, For. Branch, For. Res. Div., Ottawa ON, Silv. Leafl. 99. 2 p.
  27. ^ Dallimore, W.; Jackson, A.B. 1961. A Handbook of Coniferae including Ginkgoaceae, 3rd (1948) ed. reprinted with corrections. Arnold, London, U.K. 686 p.
  28. ^ Hare, F.K.; Ritchie, J. 1972. The boreal bioclimates. Geogr. Rev. 62:333–365.
  29. ^ Brayshaw, T.C. 1960. Key to the native trees of Canada. Canada Dep. For., Bull. 125. 43 p.
  30. ^ a b c d e Harlow, W.M.; Harrar, E.S. 1950. Textbook of Dendrology, 3rd ed. McGraw-Hill, New York NY. 555 p.
  31. ^ Hosie, R.C. 1969. Native Trees of Canada, 7th ed. Can. Dep. Fish. For., Can. For. Serv., Ottawa ON. 380 p.
  32. ^ Hale, J.D. 1955. Thickness and density of bark. Pulp and Paper Mag. Canada, Dec.:3–7.
  33. ^ Chang, Y.P. 1954. Bark structure of North American conifers. USDA, For. Serv., Tech. Bull. 1095. 86 p.
  34. ^ a b c d e f Rowe, J.S 1972. Forest regions of Canada. Can. Dep. Environ., Can. For. Serv., Ottawa ON, Publ. 1300. 172 p.
  35. ^ Forestry Branch. 1961. Native Trees of Canada, 6th ed. Canada Dep. Northern Affairs and National Resour., For. Branch, Ottawa ON, Bull. 61. 291 p.
  36. ^ a b Bell, R. 1881. The northern limits of the principal forest trees of Canada east of the Rocky Mountains. p.38c–56c in Geological and Natural History Survey of Canada, Ottawa ON, Report 1879/1880.
  37. ^ a b Collins, G.L.; Sumner, L. 1953. Northeast Arctic: the last great wilderness. Sierra Club Bull. 38:13–26.
  38. ^ Lafond, A. 1966. Notes sur l’écologie de quatre conifères du Québec: Picea mariana, P. glauca, Abies balsamea, et Pinus banksiana. Naturaliste Canadien, Québec 93:823–842.
  39. ^ a b c Sargent, C.S. 1922. Manual of the Trees of North America, 2nd corrected ed. Houghton and Mifflin, Boston, 510 p., reprinted 1961 in 2 volumes, Dover Publications, New York NY, Vol. 1. 433 p. [E3999 bib gives 910 p.]
  40. ^ a b c d e f g h Nienstaedt, H.; Zasada, J.C. 1990. Picea glauca (Moench) Voss. p. 204–226 in Burns, R.M. and Honkala, B.H. (Tech. Coord.), Silvics of North America, Vol. 1, Conifers. USDA, For. Serv., Washington DC, Agric. Handbook 654.
  41. ^ a b Scoggan, H.J. 1957. Flora of Manitoba. Can. Dep. Northern Affairs and National Resources, Nat. Museum Can., Ottawa ON, Bull. 140. 619 p.
  42. ^ a b c d e Munns, E.N. 1938. The distribution of important forest trees of the United States. USDA, For. Serv., Washington DC, Misc. Publ. 287. 176 p.
  43. ^ Fernald, M.L. 1950. Gray’s Manual of Botany, 8th ed. Amer. Book, New York NY. 1632 p.
  44. ^ Cook, D.B.; Smith, R.H. 1959. A white spruce outlier at Shushan, New York. Ecology 40:333–337.
  45. ^ Arno, S. F. & Hammerly, R. P. (1984). Timberline. Mountain and Arctic Forest Frontiers. The Mountaineers, Seattle. ISBN 0-89886-085-7.
  46. ^ a b c 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.
  47. ^ Sutton, R.F. 1969. Silvics of white spruce (Picea glauca [Moench] Voss). Can. Dep. Fish. For., For. Branch, Ottawa ON, Publ. 1250. 57 p. (Cited in Coates et al. 1994).
  48. ^ Hustich, I. 1966. On the forest–tundra and the northern tree-lines. Annales Univ. Turku A.II, Vol. 36:7–47.
  49. ^ Wilton, R.F. 1964. The forests of Labrador. Can. Dep. For., For. Res. Branch, Ottawa ON, Publ. 1066. 72 p.
  50. ^ a b Hustich, I. 1953. The boreal limits of conifers. Arctic 6:149–162.
  51. ^ a b Halliday, W.E.D.; Brown, A.W.A. 1943. The distribution of some important forest trees in Canada. Ecology 24:353–373.
  52. ^ a b Bryson, R.A.; Irving, W.H.; Larson, J.A. 1965. Radiocarbon and soil evidence of former forest in the southern Canadian tundra. Science 147(3653):46–48.
  53. ^ Tyrell, J.B. 1897. Geol. Surv. Can., Ottawa ON, Ann. Rep. 1896, Vol. 9. (Cited by Bryson et al. 1965, orig. not seen)
  54. ^ Drew, J.V.; Shanks, R.E. 1965. Landscape relationships of soils and vegetation in the forest–tundra ecotone, Upper Firth River Valley, Alaska–Canada. Ecol. Monogr. 35:285–306.
  55. ^ Dame, L.L.; Brooks, H. 1901. Handbook of the Trees of New England. Ginn, Boston MA. 196 p.
  56. ^ a b c Nisbet, J. 1905. The Forester. Blackwood and Sons, Edinburgh and London, U.K., Vol. 1. 506 p.
  57. ^ Tubbs, C.H. 1976. Effect of sugar maple root exudate on seedlings of northern conifer species. USDA, For. Serv., Res. Note NC-213. 2 p.
  58. ^ Hereman, S. 1868. Paxton’s Botanical Dictionary (Revised and corrected), Bradbury, Evans, London, U.K. 623 p.
  59. ^ Jackson, A.B. 1948. The Identification of Conifers. Arnold, London, U.K. 152 p.
  60. ^ Sabroe, A.S. 1954. Forestry in Denmark, 3rd ed. Danish Heath Soc., Copenhagen. 118 p.
  61. ^ Troup, H.S. 1932. Exotic Forest Trees in the British Empire. Clarendon Press, Oxford, U.K. 268 p.
  62. ^ Guillebaud, W.H.; Steven, H.M.; Marsden, R.E. 1920. Rate of growth of conifers in the British Isles. Forestry Commission, HMSO, London, U.K., Bull. 3. 84 p.
  63. ^ Selby, P.J. 1842. A history of British forest-trees. Van Voorst, London. 540 p.
  64. ^ Anon. (C.P.J.) 1879. Fir. pp. 222–225 in vol IX. Encyclopedia Britannica, 9th ed.
  65. ^ United Kingdom Forestry Commission. 1920. Beaufort estate. p. 57–62 in Programme, British Empire For. Conf., London, U.K.
  66. ^ Edlin, H.L. 1962. A modern sylva or a discourse of forest trees. 3. The spruces. Quart J. For. 56:292–300.
  67. ^ Fraser, G.K. 1933. Studies of certain Scottish Moorlands in relation to tree growth. For. Commission, HMSO, London, U.K. 112 p.
  68. ^ a b c Trainor, Sarah (2010-11-02). "Meeting Alaska’s Fire Science and Climate Information Needs for Forest Managers" (PDF). Forest Wisdom (Santa Fe, NM: Forestry Guild) (16): 4–5. Retrieved 2010-11-11. 
  69. ^ Halliday, W.E.D. 1937. A forest classification for Canada. Can. Dep. Mines and Resources, Dominion For. Serv., Ottawa ON, Bull. 89. 50 p.
  70. ^ a b c Wilde, S.A.; Wilson, F.G.; White, D.P. 1949. Soils of Wisconsin in relation to silviculture. Wisconsin Conserv. Dep.,Madison WI, Publ. 525–49. 171 p.
  71. ^ a b Wilde, S.A.; Voigt, G.K.; Pierce, R.S. 1954. The relationship of soils and forest growth in the Algoma district of Ontario, Canada. J. Soil Sci. 5:22–38.
  72. ^ a b c d e Nienstaedt, H. 1957. Silvical characteristics of white spruce (Picea glauca). USDA, For. Serv., Lake States For. Exp. Sta., St. Paul MN, Pap. 55. 24 p.
  73. ^ Jameson, J.S. 1963. Comparison of tree growth on two sites in the Riding Mountain Forest Experimental Area. Can. Dep. For., For. Res. Branch, Ottawa ON, Publ. 1019. 36 p.
  74. ^ Jeffrey, W.W. 1961. Origin and structure of some white spruce stands on the lower Peace River. Can. Dep. For., For. Res. Branch, Ottawa ON, Tech. Note 103. 20 p.
  75. ^ Jeffrey, W.W. 1964. Forest types along lower Liard River, Northwest Territories. Can. Dep. For., For. Res. Branch, Ottawa ON, Publ. 1035. 103 p.
  76. ^ Lacate, D.S.; Horton, K.W.; Blyth, A.W. 1965. Forest conditions on the Lower Peace River. Can. Dep. For., For. Res. Branch, Ottawa ON, Publ. 1094. 53 p.
  77. ^ a b Viereck, L.A. 1973. Wildfire in the taiga of Alaska. Quaternary Res. 3:465–495.
  78. ^ a b Sargent, C.S. 1898. The Silva of North America. A description of the trees which grow naturally in North America exclusive of Mexico. Vol. XII. Coniferae. Houghton Mifflin, Riverside Press, Cambridge, Boston MA. 144 p.
  79. ^ Kenety, W.H. 1917. Preliminary study of white spruce in Minnesota. Univ. Minnesota, Cloquet Exp. Sta. MN, Bull. 168. 30 p.
  80. ^ Heger, L. 1971. Site-index/soil relationships for white spruce in Alberta mixedwoods. Can. Dep. Environ., Can. For. Serv., For. Manage. Instit., Ottawa ON, Inf. Rep. FMR-X-32. 15 p.
  81. ^ Kabzems, A 1971. Growth and yield of well stocked white spruce in the mixedwood section , Saskatchewan. Saskatchewan Dep. Nat. Resour., For. Branch, Prince Albert SK, Tech. Bull. 5. 75 p. (Cited in Coates et al. 1994).
  82. ^ Ritchie, J.C. 1956. The vegetation of northern Manitoba. I. Studies in the southern spruce forest zone. Can. J. Bot. 34(4):523–561.
  83. ^ MacLean, D.W. 1960. Some aspects of the aspen–birch–spruce–fir type in Ontario. Can. Dep. Northern Affairs National Resources, For. Branch, For. Res. Div., Ottawa ON, Tech. Note 94. 24 p.
  84. ^ a b Sutton, R.F. 1968. Ecology of young white spruce (Picea glauca [Moench] Voss). Ph.D. thesis, Cornell Univ., Ithaca NY, Univ. Microfilms, Ann Arbor, Michigan MI, 68–11645. 500 p.
  85. ^ Heiberg, S.O.; White, D.P. 1951. Potassium deficiency of reforested pine and spruce stands in northern New York. Soil Sci. Soc. Amer. Proc. 15:369–376.
  86. ^ Lafond, A. 1954. Les déficiences en potassium et magnésium des plantations de Pinus strobus, Pinus resinosa et Picea glauca de la province de Québec. Assoc. Ing. For. Prov. Québec, Texte des Conf. 34 Assemb. Ann.:65–82.
  87. ^ a b McLeod, J.W. 1956. Plantations of the Acadia Forest Experiment Station. Can. Dep. Northern Affairs National Resour., For. Branch, For. Res. Div., Ottawa ON, Tech. Note 31. 25 p.
  88. ^ MacArthur, J.D. 1957. The effects of manure on a white and Norway spruce plantation at Grand’Mère, Quebec. Can. Dep. Northern Affairs National Resour., For. Branch, For. Res. Div., Ottawa ON, Tech. Note 64. 15 p.
  89. ^ Paine, L.A. 1960. Studies in forest pathology. XXII. Nutrient deficiencies and climatic factors causing low volume production and active deterioration in white spruce. Can. Dep. Agric., For. Biol. Div., Ottawa ON, Publ. 1067. 29 p.
  90. ^ Swan, H.S.D. 1960. The mineral nutrition of Canadian pulpwood species. 1. The influence of nitrogen, phosphorus, potassium and magnesium deficiencies on the growth and development of white spruce, black spruce, jack pine and western hemlock seedlings grown in a controlled environment. Pulp Paper Res. Instit. Can., Montreal QC, Woodlands Res. Index No. 116, Tech. Rep. 168. 66 p.
  91. ^ a b Wilde, S.A. 1966. Soil standards for planting Wisconsin conifers. J. For. 64(6):389–391.
  92. ^ a b Viereck, L.A. 1970a. Forest succession and soil development adjacent to the Chena River in interior Alaska. Arctic Alp. Res. 2(1):1–26. [wS. BA51:76183]
  93. ^ a b Viereck, L.A. 1970b. Soil temperatures in river bottom stands in interior Alaska. p. 223–233 in Proc. Ecology of the Subarctic Regions, July–Aug. 1966, Helsinki, Finland, UNESCO. [Nienstaedt and Zasada 1990]
  94. ^ Viereck, L.A.; Dyrness, C.T.; Van Cleve, K.; Foote, M.J. 1983. Vegetation, soils, and forest productivity in selected forest types in interior Alaska. Can. J. For. Res. 13(5):703–720.
  95. ^ Brand, D.G.; Kehoe, P.; Connors, M. 1986. Coniferous afforestation leads to soil acidification in central Ontario. Can. J. For. Res. 16(6):1389–1391.
  96. ^ Stoeckeler, J.H. 1938. Soil adaptability of white spruce. J. For. 36:1145–1147.
  97. ^ Zasada, J.C.; Van Cleve, K.; Werner, R.A.; McQueen, J.A.; Nyland, E. 1977. Forest biology and management in high-latitude North American forests. p. 137–195 in Proc. North American Forest Lands at Latitudes North of 60 degrees. Sympos., Univ. Alaska, Fairbanks AK, Sept. 19–22, 1977.
  98. ^ Pelletier, J.R. 1966. Tree breeding in Canada. Commonw. For. Rev. 45(1):9–10.
  99. ^ Stiell, W.M. 1976. White spruce: artificial regeneration in Canada. Dep. Environ., Can. For. Serv., Ottawa ON, Inf. Rep. FMR-X-85. 275 p.
  100. ^ La Roi, G.H.; Stringer, M.H. 1976. Ecological studies in the boreal spruce–fir forests of the North American taiga. II. Analysis of the bryophyte flora. Can. J. Bot. 54:619–643. [Nienstaedt and Zasada 1990]
  101. ^ Viereck, E.G. 1987. Alaska’s wilderness medicines – healthful plants of the North. Alaska Publishing, Edmonds, Washington WA. 107 p. [Nienstaedt and Zasada 1990]
  102. ^ Gill, D. 1975. Influence of white spruce trees on permafrost-table microtopography, Mackenzie River Delta. Can. J. Earth Sci. 12(2):263–272.
  103. ^ Van Cleve, K.; Yarie, J. 1986. Interaction of temperature, moisture, and soil chemistry in controlling nutrient cycling and ecosystem development in the taiga of Alaska. p. 160–189 in Van Cleve, K.; Chapin, F.S.; Flanagan, P.W.; Viereck, L.A.; Dyrness, C.T. (Eds.). 1986. Forest Ecosystems in the Alaskan Taiga. Springer-Verlag, New York NY.
  104. ^ Jonsson, B.G.; Esseen, P.A.; Jonsson, B. 1990. Treefall disturbance maintains high bryophyte diversity in a boreal spruce forest. J. Ecology 78(4):924–936.
  105. ^ Jonsson, B.G.; Dynesius, M. 1993. Uprooting in boreal spruce forests: long-term variation in disturbance rate. Can. J. For. Res. 23(11):2383–2388.
  106. ^ 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.
  107. ^ Nienstaedt, H.; Zasada, J.C. 1990. Picea glauca (Moench) Voss. p. 204–226 in Burns, R.M. and Honkala, B.H. (Tech. Coord.), Silvics of North America, Vol. 1, Conifers. USDA, For. Serv., Washington DC, Agric. Handbook 654.
  108. ^ Sakai, A.; Larcher, W. (Eds.) 1987. Frost Survival of Plants. Springer-Verlag, New York NY. 321 p.
  109. ^ Glerum, C. 1985. Frost hardiness of coniferous seedlings: principles and applications. p. 107–123 in Duryea, M.L. (Ed.). Proceedings: Evaluating seedling quality: principles, procedures, and predictive abilities of major tests. Workshop, October 1984, Oregon State Univ., For. Res. Lab., Corvallis OR.
  110. ^ Smith, B.J. 1949. Silvicultural work at the Sault Ste. Marie Division [of Abitibi Power and Paper Co. Ltd.]. Can. Pulp Paper Assoc., Woodlands Section, Woodlands Section Index No. 1050 (F-2). 4 p.
  111. ^ Sutton, R.F. 1992. White spruce (Picea glauca [Moench] Voss): stagnating boreal old-field plantations unresponsive to fertilization and weed control. For. Chron. 68:249–258.
  112. ^ a b Solomon, A.M.; West, D.C., and Solomon, J.A. 1981. Simulating the role of climate change and species imiigration in forest succession. p. 154–178 in West, D.C.; Shugart, H.H.; Botkin, D.B. (Eds.). Forest Succession: Concepts and Application. Springer-Verlag, New York NY.
  113. ^ Jeffrey, W.W. 1964. Forest types along lower Liard River, Northwest Territories. Can. Dep. For., For. Res. Branch, Ottawa ON, Publ. 1035. 103 p.
  114. ^ Van Cleve, K.; Dyrness, R.; Viereck, L. 1980. Nutrient cycling in interior Alaska flood plains. p. 11–18 in Murray, M.; Van Veldhuizen, R.M. (Eds.). Proc. Workshop, Fairbanks, Alaska, Nov. 1979. USDA, For. Serv., Pacific Northwest For. Exp. Sta., Portland OR, Gen. Tech. Rep., PNW-107. 52 p.
  115. ^ Heikinheimo, O. 1915. Der einfluss der brandwirtschaft auf die Wälder Finnlands. Kaskiviljelyksen Vaikutus Suomen Metsin. Acta Forest. Fenn. 4:1–264, 1–149, 1–59 [German summary p 1–59]
  116. ^ Lutz, H.J. 1956. Ecological effects of forest fires in the interior of Alaska. USDA, For. Serv., Washington DC, Tech. Bull. 1133. 121 p.
  117. ^ Gregory, R.A. 1966. The effect of leaf litter upon establishment of white spruce beneath paper birch. For. Chron. 42:251–255.
  118. ^ Neiland, B.J.; Viereck, L.A. 1977. Forest types and ecosystems. p. 109–136 in North American Forest Lands at Latitudes North of 60 Degress, Proc. sympos., Univ. Alaska, Fairbanks AK, Sept. 1977.
  119. ^ a b Dyrness, C.T. 1980. Western forest cover types, Northern Interior (Boreal): White spruce. p.81; White spruce–Aspen. p.82; Black spruce–White spruce. p.84; and Black spruce–Paper birch. p.85. in Eyre, F.H. (Ed.). Forest Cover Types of the United States and Canada. Soc. Amer. Foresters, Washington DC.
  120. ^ Dyrness, C.T. 1980. Western forest cover types, Northern Interior (Boreal): White spruce. p.81; White spruce–Aspen. p.82; Black spruce–White spruce. p.84; and Black spruce–Paper birch. p.85. in Eyre, F.H. (Ed.). Forest Cover Types of the United States and Canada. Soc. Amer. Foresters, Washington DC.
  121. ^ a b Eyre, F.H. (Ed.) 1980. Forest Cover Types of the United States and Canada. Soc. Amer. Foresters, Washington DC. 148 p.
  122. ^ Payette, S. 1980. Eastern forest cover types, Boreal Forest Region: white spruce. p.15 in Eyre, F.H. (Ed.). Forest Cover Types of the United States and Canada. Soc. Amer. Foresters, Washington DC. 148 p.
  123. ^ Nienstaedt, H.; Zasada, J.C. 1990. Picea glauca (Moench) Voss. p. 204–226 in Burns, R.M. and Honkala, B.H. (Tech. Coord.), Silvics of North America, Vol. 1, Conifers. USDA, For. Serv., Washington DC, Agric. Handbook 654.
  124. ^ "The Spruce Beetle - FIDL". Na.fs.fed.us. Retrieved 2013-07-13. 
  125. ^ Switzer, A.L.K. 1960. Spruce management for the future. For. Chron. 36(2):163–165.
  126. ^ 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.
  127. ^ a b 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)
  128. ^ a b 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]
  129. ^ "RHS Plant Selector - Picea glauca 'Echiniformis'". Retrieved 30 June 2013. 
  130. ^ Hammerbacher, A.; Ralph, S. G.; Bohlmann, J.; Fenning, T. M.; Gershenzon, J.; Schmidt, A. (2011). "Biosynthesis of the Major Tetrahydroxystilbenes in Spruce, Astringin and Isorhapontin, Proceeds via Resveratrol and is Enhanced by Fungal Infection". Plant Physiology 157 (2): 876–890. doi:10.1104/pp.111.181420. PMC 3192583. PMID 21865488.  edit