“boreal forest”的意思、由来-百科全书

boreal forest

northern forest

Introduction

also called taiga

vegetation composed primarily of cone-bearing, needle-leaved, or scale-leaved evergreen trees, found in regions that have long winters and moderate to high annual precipitation.

The boreal (meaning northern) forest region occupies about 17 percent of the Earth's land surface area in a circumpolar belt of the far Northern Hemisphere. Northward beyond this limit, the boreal forest merges into the circumpolar tundra. The boreal forest is characterized predominantly by a limited number of conifer species—i.e., pine (Pinus), spruce (Picea), larch (Larix), fir (Abies)—and to a lesser degree by some deciduous genera such as birch (Betula) and poplar (Populus). These trees reach the highest latitudes of any trees on the Earth. Boreal plants and animals are adapted to short growing seasons of long days that vary from cool to warm. Winters are long and very cold, the days are short, and a persistent snowpack is the norm. The boreal forests of North America and Eurasia display a number of similarities, even sharing some plant and animal species. The northern forests of Russia, especially Siberia, are referred to as taiga, meaning “little sticks,” a term now widely accepted as an alternative to boreal forest.

Origin

During the final period of maximum cold temperatures (23,000 to 16,500 years ago), in the latter part of the Pleistocene (Pleistocene Epoch) Ice Age (which ended 10,000 years ago), species that now constitute the boreal forest were displaced as far south as 30° N latitude by the continental glaciers of Europe, Asia, and North America and by the hyperarid and extremely cold environments of unglaciated Asia and North America. As the glaciers began to retreat gradually about 18,000 years ago, species of the boreal forest began to move northward in Europe and North America. In eastern and central North America the northward movement of the forest was relatively steady and gradual. An exception to this progression occurred about 9,000 years ago in western Canada when white spruce spread rapidly northward across 2,000 kilometres (1,240 miles) of newly deglaciated land in only 1,000 years. This rapid migration resulted from seed dispersal facilitated by strong northward winds caused by clockwise atmospheric circulation around the remnant ice cap of northern Quebec and the western part of Hudson Bay.

Because so much of the Earth's water was bound up in ice at this time, sea levels were lower than they are today, and this allowed migrations of various terrestrial species to occur. Many areas that are now islands were then connected to the nearby mainland—e.g., the British Isles were linked to Europe. As the climate warmed during the last stages of the glacial period, but before the sea level rose to its current position, some plants and animals of the mainland European boreal forest migrated to Britain. This biota exists today as part of the boreal forest in the highlands of Scotland. The areas of lowland central Alaska, the central Yukon Territory (Yukon), and far eastern Russia, which had climates too arid to permit the formation of ice sheets, were connected by the Bering Land Bridge, across which many species migrated. As a result, today across Alaska a gradient in plant characteristics can be observed, ranging from typical North American forms in the east to those with Eurasian characteristics in the west.

Distribution

The boreal forests of North America and Eurasia (Figure 1-->

) are broad belts of vegetation that span their respective continents from Atlantic to Pacific coasts. In North America the boreal forest occupies much of Canada and Alaska. Although related transition forest types are present in the northern tier of the lower 48 United States, true boreal forest stops just north of the southern Canadian border. The vast taiga of Asia extends across Russia and southward into northeastern China and Mongolia. In Europe most of Finland, Sweden, and Norway are covered with boreal forest. A small, isolated area of boreal forest in the Scottish Highlands lacks some continental species but does contain the most widespread conifer of the Eurasian boreal forest, Scotch pine (Pinus sylvestris).

The position of the boreal forest zone generally is controlled by the degree of warmth experienced during the growing season, the temperature of the soil, and the extreme minimum winter temperature. The boreal forest belt consists of three roughly parallel zones: closed canopy forest, lichen woodland or sparse taiga, and forest-tundra. The closed canopy forest is the southernmost portion of the taiga. It contains the greatest richness of species, the warmest soils, the highest productivity, and the longest growing season within the boreal zone. North of the closed canopy forest is the lichen woodland—a smaller parallel zone of sparse forest or woodland in which tree crowns do not form a closed canopy. Lichen mats and tundralike vegetation make up a significant portion of the ground cover. To the north of the lichen woodland lies forest-tundra, which occurs along the northern edge of tree growth (tree line). Patches of trees consisting of only a few species dot restricted portions of the landscape, forming a complex mosaic with tundra. Many trees in the forest-tundra zone have never been known to produce viable seeds or have done so only sporadically. These trees were established during warmer climatic episodes from a few hundred to a few thousand years ago and have persisted since, usually by vegetative (asexual) reproduction. Forest fires in this zone remove trees, and because of the lack of reproduction, only unburned patches of trees remain.

The closed forest, or southern taiga zone, on both continents is not distributed along a strictly east-west axis. At the western margin of Europe the warming influence of the Gulf Stream allows the closed canopy forest to grow at its northernmost location, generally between about 60° and 70° N. In western North America the Kuroshio and North Pacific currents likewise warm the climate and cause the northward deflection of the forest into Alaska and the Yukon Territory in Canada. On the eastern margin of the continents the boreal forest is deflected southward to between about 50° and 60° N by the cold polar air masses that flow south along these coasts. This is the southernmost limit of the boreal forest, to the south of which, in humid eastern North America and Europe, lies a northern deciduous broad-leaved transition forest. In this forest small stands of boreal conifers are distributed on cooler or less productive sites such as peaty wetlands. In the arid centre of both continents the closed canopy boreal forest is bordered to the south by a forest parkland of trees and grassland.

The central portions of Eurasia and North America are regions of flat or gently rolling topography. There, the northern and southern boundaries of the boreal forest are broad and gradual; they have fluctuated by as much as 200 kilometres (125 miles) during the past few thousand years. A well-defined but complex boundary is formed between taiga and alpine tundra on the mountains of the Pacific edge in western North America and the far eastern region of Russia. Generally the boreal forest does not come into contact with the humid temperate or subpolar rainforest of coastal Alaska and British Columbia because of high mountain barriers, but some low-elevation regions have a transition zone often characterized by trees that are a hybrid of Sitka spruce (Picea sitchensis) and white spruce (P. glauca). In Norway and Scotland a variant form of the boreal forest occupies extremely humid environments.

Practically all the large river systems of the taiga of Siberia, including the Ob, Yenisey, and Lena rivers, are northward-flowing. The Ob in western Siberia forms a great lowland basin with a considerable percentage of the land surface covered with poorly drained peaty wetlands. In such situations within the boreal zone a closed canopy forest is generally absent.

Environmental conditions

climate

Coldness is the dominant climatic factor in boreal forest regions, although a surprising diversity of climates exists. Several factors—namely, the solar elevation angle, day length, and snow cover—conspire to produce this cold climate. In the boreal region the Sun is never directly overhead (90°) as it can be in the tropics. The maximum solar angle decreases with increasing latitude. At latitude 50° N in the southern boreal region the maximum solar angle is 63.5° and at the Arctic Circle it is only 47°. As a result solar energy is less intense in the boreal regions because it is spread out over a greater area of the Earth's surface than it is in equatorial regions. Day length also affects temperature. Long winter nights at high latitudes allow radiation emitted by the surface of the Earth to escape into the atmosphere, especially in continental interiors where cloud cover is less abundant than it is near the coast. Snow cover, too, affects the climate, because it reflects incoming solar radiation and amplifies cooling. During winter, a snowpack persists for at least five months in the southern portion of boreal forest regions and for seven or eight months in the northern reaches. The boreal forest actually mitigates this cooling because it roughens and darkens what would otherwise be a smooth, snow-covered, energy-reflecting surface for much of the year. It has been estimated that the Earth would be significantly colder without the boreal forest.

The northern limit of the North American boreal forest coincides with the mean position of the Arctic front—the boundary between Arctic and mid-continental air masses—in the summer; its southern limit coincides with the mean frontal position in the winter. Mean annual temperatures in the boreal forest range from a few degrees Celsius above freezing to −10° C (14° F) or more. Areas with a mean annual temperature below freezing are susceptible to the formation of permafrost soils (frozen ground; see below Soils (boreal forest)).

The mean temperature of January, the coldest month, is generally less than −10 °C (14° F) across the boreal region. The boreal forest includes areas that experience some of the lowest temperatures on the Earth, excluding Antarctica. At the height of winter an intensely cold pocket of air develops over inland areas of far eastern Siberia; mean temperatures of −50° C (−58° F) have been recorded in this region. As this Siberian cold air flows over the unfrozen northern Pacific Ocean a great temperature contrast develops that results in strong, westward-moving storm systems. The movement, position, and strength of these storms control much of the weather in the Northern Hemisphere.

Boreal forests do not grow on areas surrounding the Bering Strait. A rigorous cold climate with a very short snow-free season precludes the growth of trees on the Russian side of the Bering Strait in the Chukotka region of the Russian Far East. On the North American side in western Alaska summers are too cool for trees to grow because of cold air masses moving off the Bering Sea.

The growing season in the boreal forest is generally cool; the mean temperature of the warmest month, July, is generally between 15° and 20° C (59° and 68° F). Summer daytime high temperatures are typically cool to warm—20° to 25° C (68° to 77° F)—for much of the growing season in the boreal forest. Cool summer temperatures can actually produce higher photosynthetic efficiency in plants than can warmer conditions. Species adapted to cold respire less (use up less of their food stores) when photosynthesizing at cool temperatures in intense summer light than they do at higher temperatures, allowing a greater net gain in biomass (dry mass of organic matter).

Areas of the boreal forest located in the centre of continents generally receive 30 to 50 centimetres (12 to 20 inches) of annual precipitation. Precipitation totals are relatively modest in these locations because they are a significant distance from unfrozen oceans that supply moisture. Some boreal forests are semiarid and may even include grasslands interspersed with the forest. These forests are found in regions of the Yukon Territory and Alaska that occur on the leeward side of mountains which are sheltered from moisture-bearing winds, as well as in some portions of the interior of far eastern Russia. Annual precipitation in low elevations of these regions is 30 centimetres or less. The highest annual precipitation total in the boreal forest, which can exceed 100 centimetres, is in eastern North America and northern Europe. During ancient eras of colder climate these regions also received relatively abundant precipitation, which resulted in the buildup of glacial ice sheets. Today these once heavily glaciated regions support extensive lakes, streams, and wetlands.

Extended periods of clear, dry weather in the boreal region are caused by persistent strong polar high pressure systems. If strong high pressure persists during the long days near the summer solstice, temperatures can warm to 30° C (86° F) or higher. Intense heating at the ground surface often produces convective storms with lightning but little rain, causing forest fires.

Soils (soil)

Boreal forest conifer litter is highly acidic. Soils of the more humid and southern boreal forest are highly leached spodosols (Spodosol), which are characterized by the leaching of iron, aluminum, and organic matter from the chemically and biologically distinct surface layer—horizon A—to the next layer—horizon B. Much of the soil of central and eastern Canada—granitic Canadian Shield—has been repeatedly scraped clean by glacial advances. Thus, productive forests often are restricted to portions of the landscape where soil material has been deposited by glaciers. Peaty wetlands occur where surface drainage is impeded by permafrost, youthful glacial topography, or aggraded rivers; their soils are characteristically organic soils, or histosols (Histosol). Soils in much of boreal western North America and Asia are inceptisols (Inceptisol), which have little horizon development. Very thin surface salt deposits are found in the most arid portions of the boreal forest.

Cold soils are characteristic of the boreal forest region, which overlaps the zone of permafrost. Permafrost is soil or earth material that remains below 0° C (32° F) for at least two years. The surface, or active, layer of permafrost thaws in the warm season and freezes in the winter, but the soil below the active layer remains continuously frozen. Because the plant rooting zone is restricted to the active layer, nutrient supply is limited and secure anchoring for roots is lacking. Some trees and other plants of the taiga (especially black spruce 【Picea mariana】 and tamarack 【Larix laricina】 in North America and larches in Siberia) can grow on permafrost if the active layer is sufficiently deep, but several species are eliminated from permafrost.

The boreal forest itself is an important contributing factor to the development of permafrost. The latter stages of forest growth—characterized by development of an intact forest canopy, growth of an insulating moss cover in summer, and accumulation of forest litter—may cool the soil to such an extent that permafrost develops. Warming of the soil is promoted by forest fires, which remove the canopy, moss, and forest litter layers. In the absence of an intact canopy, a deeper and more effective insulating layer of snow accumulates in the winter. The presence of dark ash following a fire increases solar energy absorption on the site for several years.

The boreal forest of Europe generally lacks permafrost, but east of the Ural Mountains and from central Canada northward permafrost is common. In southern and central boreal forests, permafrost occurs sporadically and occupies only a small percentage of the landscape that experiences the coldest temperatures. The northern portion of closed canopy forest and the lichen woodland zone are in a region of discontinuous permafrost, where permafrost is found on north-facing slopes and in cold air drainage basins but is absent from south-facing slopes and newly deposited alluvial sites. Most of the forest-tundra is within the continuous permafrost zone.

Forest productivity in the middle and northern taiga zones is directly related to soil temperature. Warmer soils decompose organic matter more quickly, releasing nutrients for new plant growth and creating a more productive site. Productive forest types occupy warmer, south-facing slopes and river terraces, and less productive dwarf or sparse forest occupies the north-facing and basin permafrost sites.

Floodplains (floodplain) throughout the boreal forest regions are free of permafrost, high in soil fertility, and repeatedly disturbed in ways that renew the early, rapid growth stages of forest succession. Floodplains are a mosaic of productive shrubland and forest that serve as a major habitat for moose (Alces alces), which influence ecosystem structure and function.

South-central Alaska and the adjacent Yukon Territory and British Columbia support the most extensive ice sheets and glaciers (glacier) in the world outside the polar desert regions of Antarctica and Greenland. Glacial meltwater is a large part of the flow of larger rivers such as the Yukon and Tanana in Alaska and the Yukon Territory. Glacial meltwater (outwash) carries a heavy load of suspended sediment that deposits in riverbeds and causes frequent channel shifts. Glacial river floodplains are extensive, very dynamic, and constantly renewed with fertile soil material. In the ancient past exposed deposits of glacial silt were picked up by strong winds and deposited on surrounding hillsides. Fertile soils, known as loess, resulted, on which highly productive upland forests are found today. Because the beds of glacially fed rivers are rising, the landscape through which they flow is partially drowned from the impeded drainage, often preventing forest growth and favouring the development of marshes and mires.

The biota and its adaptations

Nearly all major taxonomic groups have fewer species of animals and plants in the boreal forest than they have in other terrestrial ecosystems at lower latitudes. This accords with the species diversity gradient that is observed from lower to higher latitudes, with numbers of species decreasing in a poleward direction.

Trees (tree)

Scotch pine is the most widely distributed pine species in the world, growing from northern Scotland to the Russian Pacific shore. The relatively humid and productive taiga of northern Europe and south-central Siberia is dominated by this species. Forest management has greatly favoured this species in Scandinavia and Finland. It is a thick-barked species and easily survives light ground fires, often reaching ages of 350 to 400 years, with some individuals being older than 700 years. European aspen and Siberian spruce are essentially transcontinental in distribution as well.

The species composition of Eurasian taiga is different east of central Siberia from that which prevails westward into Europe. Distinctive European species include Norway spruce (Picea abies), a large dominant species of the productive humid boreal forest, and Sukaczev larch (Larix sukaczewii), an early successional species (one of the first species to colonize an area after a disturbance) of European Russia. Gray (Betula populifolia) and white birch (B. pendula) occur across northern Europe and well into central Siberia. The birches often form dense stands of light or white barked trees that are considered a characteristic feature of the boreal forest. Siberian larch (Larix sibirica) and Siberian fir (Abies sibirica) are restricted to north-central Asia. Species restricted to northeastern Asia include chosenia (Chosenia arbutifolia), an early successional broad-leaved tree of floodplains; Siberian stone pine (Pinus sibirica), a short shrub or tree; and Asian spruce (Picea obovata).

All North American tree species are distributed across the continent except jack pine (Pinus banksiana), lodgepole pine (Pinus contorta), and balsam fir (Abies balsamea). Jack pine is a relatively small, short-lived, early successional tree occurring in the eastern and central parts of boreal forests east of the Rocky Mountains. Lodgepole pine is a longer-lived, early successional species growing in western Canada and along the Rocky Mountain axis from the central Yukon Territory southward to well south of the boreal forest limit. Balsam fir is a shade-tolerant, late successional, but relatively short-lived tree that occurs only in the eastern and central boreal forest.

Major taiga tree species are well-adapted to extreme winter cold. The northernmost trees in North America are white spruce that grow along the Mackenzie River delta in Canada, near the shore of the Arctic Ocean. The northernmost trees in the world are Gmelin larch (Larix gmelinii) found at latitude 72°40′ N on the Taymyr Peninsula in the central Arctic region of Russia.

A representative profile of the vegetation is shown in Figure 2-->

Other plants (plant)

A distinctive feature of the flora of boreal forests is the abundance and diversity of mosses (moss); about one-third of the ground cover under boreal forest is dominated by moss. Much of the ground cover in older conifer stands is moss, which grows on rocks, on tree trunks, and in the pits formed by upturned trees. Extensive peaty wetlands in the boreal region are often thick accumulations of dead sphagnum and other mosses, sedges, and other plants; a living moss layer continually grows at the surface.

Lichens (lichen) (a symbiotic association of a fungus and algae) constitute a significant part of the ground cover in the lichen woodland or sparse taiga. Lichens are also generally well distributed on tree trunks and especially in the canopy of older conifers throughout the boreal forest. Because lichens and mosses are dispersed by airborne spores that can travel long distances, many species of both groups are found across the entire circumpolar boreal forest.

Many vascular plants are also widespread across the circumpolar north. Some forest understory species dominate their habitats; they include twinflower (Linnaea borealis), lingonberry (Vaccinium vitis-idaea), baneberry (Actaea rubra), and Swedish and Canadian dwarf cornel (Cornus suecica and C. canadensis). Several boreal forest plants are adapted to rapid colonization and growth in recently burned areas, such as fireweed (Epilobium angustifolium). The extensive peatlands of the boreal north support a typical flora that usually includes species such as Labrador tea (Ledum palustre), cloudberry (Rubus chamaemorus), cotton grass (Eriophorum species), and crowberry (Empetrum nigrum or E. hermaphroditum). In northern Europe crowberry also grows as shrub mats under Scotch pine forests or woodlands. Crowberry has been shown to produce secondary chemical compounds that inhibit or kill Scotch pine seedlings. Periodic light ground fires reduce the abundance and vigour of crowberry and allow tree regeneration.

Specialized orchids (orchid) in the forest understory include calypso (Calypso bulbosa), coral root (Corallorrhiza trifida), and lady's slipper (Cypripedum species). The roots of these plants form particular associations with fungi (mycorrhizae). Willow shrubs (Salix species) are one of the first plants to emerge following disturbances on floodplains and occasionally on uplands as well. Important grasses across the boreal region include species of bromegrass (Bromus species), bluegrass (Poa species), reed bent grass (Calamagrostis species), and vanilla grass (Hierochloe odorata). Many freshwater aquatic plants such as sedges (Carex species) and pondweeds (Potamogeton species) are distributed widely across the boreal zone of both continents because migratory waterfowl and shorebirds are effective in dispersing their seeds. Several species of ferns are common to the boreal forests of the two continents, especially in regions of higher precipitation.

Mammals (mammal)

Because a winter snowpack is a dependable feature of the taiga, several mammals display obvious adaptations to it. The snowshoe, or varying, hare (Lepus americanus), for example, undergoes an annual change in colour of its pelage, or fur, from brownish or grayish in the summer to pure white in the winter, providing effective camouflage. Its feet are large in proportion to its body size, a snowshoelike adaptation for weight distribution that allows the hare to travel over the surface of snow rather than sink down into it. The lynx (Lynx canadensis) is the principal predator of the snowshoe hare (Figure 2-->

from the population ecology article ). It, too, has large feet, with fur between the toes, enabling the lynx to remain on the snow's surface. Most animals of the boreal forest are well adapted to the cold and survive it easily if they have enough food to maintain an energy balance through the winter.

moose are the largest browsing animals in the boreal forest. In the summer they eat willow and broad-leaved trees and also wade in lakes and ponds to consume aquatic plants. Throughout the winter moose eat large quantities of woody twigs and buds. Moose depend on high-quality feeding areas in the shrub zone along river floodplains and on the early successional growth of woody plants in burned or cutover forest. Intensive browsing by moose can alter the composition of the forest in its early stages of growth, often increasing the dominance of conifers, which they do not consume in as great amounts as they do broad-leaved trees. Harvesting a moose for winter food is an important and even critical element of survival for humans living in isolated rural areas of the taiga.

Moose populations are controlled by various means. Wolves (wolf) (Canis lupus) prey on moose across most of the boreal forest, and some scientists and game managers believe that once moose numbers are depressed wolf predation can keep moose populations low. As a result, wolf trapping or shooting programs are carried out as a game management measure to increase prey numbers. The natural regulation of moose populations by wolf predation and the presence of wolves themselves is valued as well. As a result programs to control wolf populations are often the subject of intense debate. Other factors control moose numbers, such as the restriction of access to plants during years of deep snow and lack of early successional woody plant growth caused by forest maturation. Where the boreal forest is extensively cut for forest products, moose numbers have increased greatly, often to levels that are considered undesirable for forest regeneration. Subsistence and sport hunting of moose are widely used tools of moose population management.

Another large-hoofed browsing mammal that is present seasonally in the boreal forest is the reindeer (Rangifer tarandus (reindeer)) in Eurasia and the closely related caribou in North America. A large portion of the reindeer population is semidomesticated and herded (migration) by nomadic peoples such as the Lapps of Scandinavia and several native peoples in northern Russia. Caribou migrate the greatest distances of any large land mammal in North America. They often move in vast herds of 500,000 animals or more, seldom stopping or pausing because they must constantly forage in these environments of generally low productivity. During the early winter, reindeer and caribou migrate south from their summer ranges in the tundra to the forest-tundra or lichen woodland, where they graze primarily on lichens. Later in winter caribou typically move to open forests and sedge-rich lake margins or to snow-free wind-swept mountains. In April and May caribou form long columns and migrate back north to the tundra.

Several mammals of the boreal region are valued for their furs (fur), and trapping and trade in furs has been an important part of the culture, economy, and history of the region as long as humans have lived there. Important furbearing species include lynx and marten (Martes americana) and in wetland habitats beaver (Castor canadensis), mink (Mustela vison; see photograph-->

), and muskrat (Ondatra zibethica).

In the North American boreal forest the northern flying squirrel (Glaucomys sabrinus) is adapted to consume fungi, especially underground fruiting bodies (sporocarps) of fungi that form mutually beneficial relationships (mutualism) with trees by colonizing their roots. The flying squirrel's consumption and dispersal of these underground fungi provide a significant benefit to the forest as a whole. (For further information on mutualism, see community ecology: Interspecific interactions and the organization of communities: Mutualism (community ecology).)

Birds (bird)

The boreal forest is the migratory destination of large numbers of birds for the summer breeding season. These include several passerine songbirds typical of shrub and forest habitats, such as thrushes, flycatchers, and warblers. Many of these species consume insects in the canopy of the boreal forest and other habitats. Predators of these birds occur in the forest as well, such as the sharp-shinned hawk (Accipiter striatus) and the northern goshawk (A. gentilis). Populations of several boreal forest-breeding migratory thrushes, flycatchers, and warblers may be declining because of the loss of their wintering habitats in the tropical forests of the world and the changes to or loss of forest habitats in the temperate zones along their migratory routes.

Birds of the boreal forest fill a variety of niches. Some are seed consumers or dispersers, others are insect consumers. They carry out other specialized roles as well. For example, the yellow-bellied sapsucker (Sphyrapicus varius) drills evenly spaced rows of small holes in the bark of trees and then visits these “wells” to obtain sap and the insects it attracts. Various other birds, mammals, and insects benefit from the sap wells, too.

Woodpeckers (woodpecker) excavate tree cavities, which subsequently are used by many species of birds and mammals. Woodpeckers are specialized predators of wood- and bark-inhabiting insects; they are thought to be important in the control of the spruce beetle (Dendroctonus rufipennis) population. In searching for insects, woodpeckers chisel (see photograph-->

) or strip the bark off dead or dying trees, promoting more rapid decay and the release of nutrients from dead trees. As large old trees have become rarer through forest cutting, some year-round resident woodpeckers such as the northern three-toed woodpecker (Picoides tridactylus) and the great spotted woodpecker (Dendrocopos major) have lost their habitats and declined in numbers.

Few bird species remain in boreal forests through the long cold winters because of limited opportunities for food, although some undertake only a short migration south. Resident bird species include the common raven (Corvus corax) and the boreal and black-capped chickadees of North America and the Siberian tit (Parus species).

The extensive areas of lakes, ponds, and wetlands (Figure 2-->

from the article boundary ecosystem )—especially in the glaciated part of the boreal forest—provide a large habitat for waterfowl and shorebirds, although the birds primarily occur in low densities across the landscape. North American shorebirds that breed in forested peatlands include common snipe (Gallinago gallinago) and yellowlegs (Tringa species). Commonly encountered waterfowl are northern pintail (Anas acuta), scaup (Aythya species), and scoters (Melanitta species).

Insects (insect)

The boreal forest is the home of relatively few species of insects, but extensive and usually uniform areas of habitat periodically support high populations of species that do live there. The boreal forest lacks the elaborate complexes of invertebrate predators and parasites that serve as stabilizers of the insect populations in warmer regions. As a result, boreal insect populations occasionally increase rapidly and cause outbreaks. Some outbreaks can injure or kill trees across widespread areas of the boreal forest. Once an outbreak reaches a certain size it can become self-sustaining, much like a forest fire; the effects of the spruce budworm and spruce beetle in North America are well-documented examples. Outbreaks can be triggered by unusual weather or physical injuries that stress trees and make them vulnerable to the insects; they can end for a variety of reasons, including production of defensive chemicals by the host plants or depletion of susceptible host plants.

Perhaps the insects most noticeable to humans in the boreal forest are mosquitoes (mosquito), which belong to several species. Mosquitoes feed on and are fed upon by many of the birds of the boreal forest. Wetland areas of the boreal region, such as sites having poor drainage because of permafrost, provide extensive mosquito breeding sites. Where well-oxygenated, flowing water is found, biting flies are abundant. Almost all food webs that support fish in boreal forest streams are dependent on insects.

Conifers serve as hosts for a variety of wood-boring beetles, spruce beetles, bark beetles, and ips beetles (Ips species). These insects aid in wood decomposition and nutrient release. Some beetles have outer shells with specialized indentations specifically matched to the shape and size of the spores of wood-decomposing fungi (fungus). Fungal spores become securely lodged in these cuplike structures. As the beetles burrow into wood they inoculate it with fungi.

A variety of lepidopterans (moths and butterflies) are adapted to feeding on the leaves of boreal trees. These include defoliators and leaf rollers.

Soil organisms (soil organism)

The species richness and total biomass of soil organisms are significantly lower in the boreal forest than they are at lower latitudes. Dominant soil organisms are protozoans, nematodes, rotifers, and tardigrades. These organisms live primarily in soil water film and soil pore water. The soil fauna of the boreal forest region is distinctive because it generally lacks large invertebrates such as millipedes, isopods (springtails), and earthworms, especially in the middle and northern taiga. Larger soil invertebrate animals perform the function of biting off (shredding) pieces of leaf litter in forest soils and passing them through their guts. As a result of this activity thick layer of several years' accumulation of only partially decomposed plant material is characteristic of boreal forest soils.

Fungi are the dominant organisms in the task of decomposition of boreal forest litter, but flushes of bacterial growth occur in response to triggering factors. The soil animals generally do not attack the forest litter directly but instead exert their influence by grazing on the fungi and bacteria. The rate of decomposition in boreal forest soils does not keep pace with the rate of production, causing the progressive accumulation of organic matter. At middle depths of the forest floor small invertebrates, especially dipteran larvae, partially consume or skeletonize leaf litter before emerging as adults.

community structure

Natural disturbances

The boreal forest is well adapted to development following natural disturbances, which include fire, floods, snow breakage, and insect outbreaks. Characteristic of the boreal forest is the general lack of late successional species that develop under an intact forest canopy. (For further information on succession see community ecology: Patterns of community structure: Ecological succession (community ecology).)

fire is the primary agent responsible for natural disturbances in the boreal forest. It can result from natural causes, such as lightning, or it can be set by humans. Large-scale insect outbreaks can weaken or kill trees over vast areas, thus creating an environment less resistant to fire. In the period between 1981 and 1989 an estimated 3 million hectares (7.4 million acres) burned annually in the former Soviet Union, almost all of which occurred within the taiga region of Russia. The so-called Black Dragon Fire of 1987 in China and Russia may have been the largest single fire in the world in the past several hundred years. During the 20th century, about 1 million hectares of taiga in Canada have burned annually; a great majority of the burning occurred in the less accessible boreal forest of the northern and western parts of the country. In Alaska in years that have prolonged hot and dry periods of summer weather, millions of hectares burn, primarily in a few very large fires. Intervals of about 200 years occur between fires in the uplands of northwestern Canada and in the interior of Alaska. In much of the central and western boreal forest of North America replacement of vegetation on upland sites, presumably by fire, appears to be necessary for forest regeneration. Floodplain islands usually do not burn and contain white spruce trees as old as 400 years. In the northern boreal forest of Europe a pattern of periodic light ground fires in Scotch pine forests was typical before the era of fire control. The thick bark of these mature trees allowed them to survive these fires. In much of the boreal forest only wildland fires that threaten high-value resources are actively suppressed. Complete fire suppression would cause soil temperature to decline gradually, promoting permafrost development that would cause a significant decrease in site productivity.

Jack pine and lodgepole pine have cones that remain closed on the tree (serotinous), and black spruce has semiserotinous cones; these cones do not open to release their seeds until a wax layer is melted by the heat of fire. White spruce seedlings require the bare mineral soil produced by burning of thick organic layers of the forest floor for proper establishment; they may time their periodic production of seed to dry periods when fire is more likely.

Effects of human use and management of the boreal forest

Different degrees of forest (forestry) development have had various effects on biodiversity around the circumpolar boreal forest zone.

A highly developed forest industry based on intensive forest utilization is maintained in boreal Scandinavian (Scandinavia) countries and Finland. About 95 percent of the productive forest types of Finland and the Scandinavian countries have been harvested at least once. Finland is located almost entirely within the boreal region and is one of the most forested nations in the world. About 6.5 percent of Finnish land, which includes large areas of marginal forest, woodland, and tundra, is protected from human modification. Only about 3 percent of Swedish forests are protected, most of which is concentrated in marginal forests of mountainous regions. Between 10 and 15 percent of species in Swedish forests are threatened.

The Canadian boreal forest represents nearly 7.5 percent of the Earth's forested area. Much of the harvesting of Canadian forest has been carried out in primary (previously unlogged) forest. Nearly all the mature first-growth timber, especially of southern, central, and eastern boreal Canada, is anticipated to be removed by the late 1990s. Considerable effort has been devoted to forest regeneration and tending of new stands, although a certain amount of land does not meet reforestation goals.

In Alaska the amount of land with at least 10 percent forest cover in the boreal region is estimated at about 46 million hectares, or 12 percent of the state, only 5.5 million hectares of which is considered productive timberland. Of all areas in the world, Alaska probably has the largest percentage of its surface area, about 40 percent, devoted to strict protection of natural habitats and species. Local-scale logging traditionally has been carried out for much of the 20th century. Plans to accelerate logging are being considered.

The taiga of Siberia covers 680 million hectares and represents nearly 19 percent of the world's forested area and possibly 25 percent of the world's forest volume. About 400,000 hectares of the Russian (Russia) taiga are logged annually, and nearly an equal area is burned, with perhaps half of the burned area resulting from destructive fires of human origin. Social and economic problems in the early postcommunist era have slowed the amount of logging by one-third to one-half; however, several large-scale joint ventures between Russian organizations and foreign partners to harvest the forests are under way. The fate of the Siberian boreal forest has become a matter of international concern.

Large areas, perhaps exceeding 2 million hectares, of the Russian taiga near Norilsk and the Kola Peninsula have been destroyed by air pollution. Many oil pipelines (pipeline) are leaking in Siberia, and repairs and maintenance are minimal. In July through September 1994 more than 150,000 metric tons of crude oil were spilled in the Kolva, Usa, and Pechora river basins of the republic of Komi in Russia. Spring meltwater could carry spilled oil in this region into the northward-flowing rivers that empty into the Arctic Ocean.

Biological productivity

Primary productivity (the rate at which photosynthesis occurs) of boreal forest ecosystems often is limited by cold soil temperatures (see above Environmental conditions: Soils (boreal forest)). Net annual primary production (the total amount of productivity less that used by photosynthetic organisms in cellular respiration) in boreal forest types varies greatly, from slightly more than 2 metric tons per hectare near the polar tree limit to about 10 metric tons per hectare along its southern margin. Boreal forests are estimated to contain about 18 percent of the Earth's total biomass (the dry weight of organic matter). The boreal forest or taiga of Siberia alone represents 57 percent of the Earth's coniferous wood volume. Ecosystems and soils of the boreal region store a significant amount of the Earth's carbon in the form of dead but undecomposed or partially decomposed organic matter. Global warming or land use changes could enhance decomposition, leading to the release of increased amounts of stored carbon into the atmosphere in the form of the greenhouse (greenhouse effect) gas carbon dioxide. (For further discussion of biological productivity see biosphere: The organism and the environment: Resources of the biosphere: The flow of energy (biosphere).)

Glenn Patrick Juday

Additional Reading

Herman H. Shugart, Rik Leemans, and Gordon B. Bonan (eds.), A Systems Analysis of the Global Boreal Forest (1992), covers ecosystem processes, forest patterns in space and time, and computer models, including chapters on the Eurasian boreal forest, tree and shrub reproduction, and fire in the boreal forest. The Plant Cover of Sweden (1965), is the most complete reference on the original condition of the boreal forest of Sweden, focusing on the landscape pattern of native vegetation and on plant indicators of various forest regions and ecosystem types. Deborah L. Elliott-Fisk, “The Boreal Forest,” in Michael G. Barbour and William Dwight Billings (eds.), North American Terrestrial Vegetation (1988), pp. 33–62, is a basic reference on the forest types and communities found across the North American boreal region and includes the historical development of the forest and the ecological characteristics of individual tree species. K. Van Cleve et al. (eds.), Forest Ecosystems in the Alaskan Taiga: A Synthesis of Structure and Function (1986), looks at recent investigations of one of the best-studied parts of the boreal forest, describing both upland and floodplain ecosystems; an update in a special issue of Canadian Journal of Forest Research, vol. 23, no. 5 (May 1993), contains the results of a multidisciplinary research project on ecological succession of a productive river floodplain in central Alaska, including a detailed discussion on soil. J.S. Rowe, Forest Regions of Canada (1972), well illustrated, is the best reference on the overall distribution and classification of boreal forest types in Canada. James A. Larsen, The Boreal Ecosystem (1980), focuses on properties of the middle boreal forest of central Canada. Edward A. Johnson, Fire and Vegetation Dynamics: Studies From the North American Boreal Forest (1992), is a well-illustrated treatment of fire weather, fire behaviour, and fire effects in the North American boreal forest. Lennart Hansson (ed.), The Ecological Principles of Nature Conservation: Applications in Temperate and Boreal Environments (1992), also well illustrated, discusses the effects of forest management on biodiversity, focusing particularly on the boreal forest of northern Europe.Glenn Patrick Juday

词条	boreal forest
释义	boreal forest northern forest Introduction also called taiga vegetation composed primarily of cone-bearing, needle-leaved, or scale-leaved evergreen trees, found in regions that have long winters and moderate to high annual precipitation. The boreal (meaning northern) forest region occupies about 17 percent of the Earth's land surface area in a circumpolar belt of the far Northern Hemisphere. Northward beyond this limit, the boreal forest merges into the circumpolar tundra. The boreal forest is characterized predominantly by a limited number of conifer species—i.e., pine (Pinus), spruce (Picea), larch (Larix), fir (Abies)—and to a lesser degree by some deciduous genera such as birch (Betula) and poplar (Populus). These trees reach the highest latitudes of any trees on the Earth. Boreal plants and animals are adapted to short growing seasons of long days that vary from cool to warm. Winters are long and very cold, the days are short, and a persistent snowpack is the norm. The boreal forests of North America and Eurasia display a number of similarities, even sharing some plant and animal species. The northern forests of Russia, especially Siberia, are referred to as taiga, meaning “little sticks,” a term now widely accepted as an alternative to boreal forest. Origin During the final period of maximum cold temperatures (23,000 to 16,500 years ago), in the latter part of the Pleistocene (Pleistocene Epoch) Ice Age (which ended 10,000 years ago), species that now constitute the boreal forest were displaced as far south as 30° N latitude by the continental glaciers of Europe, Asia, and North America and by the hyperarid and extremely cold environments of unglaciated Asia and North America. As the glaciers began to retreat gradually about 18,000 years ago, species of the boreal forest began to move northward in Europe and North America. In eastern and central North America the northward movement of the forest was relatively steady and gradual. An exception to this progression occurred about 9,000 years ago in western Canada when white spruce spread rapidly northward across 2,000 kilometres (1,240 miles) of newly deglaciated land in only 1,000 years. This rapid migration resulted from seed dispersal facilitated by strong northward winds caused by clockwise atmospheric circulation around the remnant ice cap of northern Quebec and the western part of Hudson Bay. Because so much of the Earth's water was bound up in ice at this time, sea levels were lower than they are today, and this allowed migrations of various terrestrial species to occur. Many areas that are now islands were then connected to the nearby mainland—e.g., the British Isles were linked to Europe. As the climate warmed during the last stages of the glacial period, but before the sea level rose to its current position, some plants and animals of the mainland European boreal forest migrated to Britain. This biota exists today as part of the boreal forest in the highlands of Scotland. The areas of lowland central Alaska, the central Yukon Territory (Yukon), and far eastern Russia, which had climates too arid to permit the formation of ice sheets, were connected by the Bering Land Bridge, across which many species migrated. As a result, today across Alaska a gradient in plant characteristics can be observed, ranging from typical North American forms in the east to those with Eurasian characteristics in the west. Distribution The boreal forests of North America and Eurasia (Figure 1-->) are broad belts of vegetation that span their respective continents from Atlantic to Pacific coasts. In North America the boreal forest occupies much of Canada and Alaska. Although related transition forest types are present in the northern tier of the lower 48 United States, true boreal forest stops just north of the southern Canadian border. The vast taiga of Asia extends across Russia and southward into northeastern China and Mongolia. In Europe most of Finland, Sweden, and Norway are covered with boreal forest. A small, isolated area of boreal forest in the Scottish Highlands lacks some continental species but does contain the most widespread conifer of the Eurasian boreal forest, Scotch pine (Pinus sylvestris). The position of the boreal forest zone generally is controlled by the degree of warmth experienced during the growing season, the temperature of the soil, and the extreme minimum winter temperature. The boreal forest belt consists of three roughly parallel zones: closed canopy forest, lichen woodland or sparse taiga, and forest-tundra. The closed canopy forest is the southernmost portion of the taiga. It contains the greatest richness of species, the warmest soils, the highest productivity, and the longest growing season within the boreal zone. North of the closed canopy forest is the lichen woodland—a smaller parallel zone of sparse forest or woodland in which tree crowns do not form a closed canopy. Lichen mats and tundralike vegetation make up a significant portion of the ground cover. To the north of the lichen woodland lies forest-tundra, which occurs along the northern edge of tree growth (tree line). Patches of trees consisting of only a few species dot restricted portions of the landscape, forming a complex mosaic with tundra. Many trees in the forest-tundra zone have never been known to produce viable seeds or have done so only sporadically. These trees were established during warmer climatic episodes from a few hundred to a few thousand years ago and have persisted since, usually by vegetative (asexual) reproduction. Forest fires in this zone remove trees, and because of the lack of reproduction, only unburned patches of trees remain. The closed forest, or southern taiga zone, on both continents is not distributed along a strictly east-west axis. At the western margin of Europe the warming influence of the Gulf Stream allows the closed canopy forest to grow at its northernmost location, generally between about 60° and 70° N. In western North America the Kuroshio and North Pacific currents likewise warm the climate and cause the northward deflection of the forest into Alaska and the Yukon Territory in Canada. On the eastern margin of the continents the boreal forest is deflected southward to between about 50° and 60° N by the cold polar air masses that flow south along these coasts. This is the southernmost limit of the boreal forest, to the south of which, in humid eastern North America and Europe, lies a northern deciduous broad-leaved transition forest. In this forest small stands of boreal conifers are distributed on cooler or less productive sites such as peaty wetlands. In the arid centre of both continents the closed canopy boreal forest is bordered to the south by a forest parkland of trees and grassland. The central portions of Eurasia and North America are regions of flat or gently rolling topography. There, the northern and southern boundaries of the boreal forest are broad and gradual; they have fluctuated by as much as 200 kilometres (125 miles) during the past few thousand years. A well-defined but complex boundary is formed between taiga and alpine tundra on the mountains of the Pacific edge in western North America and the far eastern region of Russia. Generally the boreal forest does not come into contact with the humid temperate or subpolar rainforest of coastal Alaska and British Columbia because of high mountain barriers, but some low-elevation regions have a transition zone often characterized by trees that are a hybrid of Sitka spruce (Picea sitchensis) and white spruce (P. glauca). In Norway and Scotland a variant form of the boreal forest occupies extremely humid environments. Practically all the large river systems of the taiga of Siberia, including the Ob, Yenisey, and Lena rivers, are northward-flowing. The Ob in western Siberia forms a great lowland basin with a considerable percentage of the land surface covered with poorly drained peaty wetlands. In such situations within the boreal zone a closed canopy forest is generally absent. Environmental conditions climate Coldness is the dominant climatic factor in boreal forest regions, although a surprising diversity of climates exists. Several factors—namely, the solar elevation angle, day length, and snow cover—conspire to produce this cold climate. In the boreal region the Sun is never directly overhead (90°) as it can be in the tropics. The maximum solar angle decreases with increasing latitude. At latitude 50° N in the southern boreal region the maximum solar angle is 63.5° and at the Arctic Circle it is only 47°. As a result solar energy is less intense in the boreal regions because it is spread out over a greater area of the Earth's surface than it is in equatorial regions. Day length also affects temperature. Long winter nights at high latitudes allow radiation emitted by the surface of the Earth to escape into the atmosphere, especially in continental interiors where cloud cover is less abundant than it is near the coast. Snow cover, too, affects the climate, because it reflects incoming solar radiation and amplifies cooling. During winter, a snowpack persists for at least five months in the southern portion of boreal forest regions and for seven or eight months in the northern reaches. The boreal forest actually mitigates this cooling because it roughens and darkens what would otherwise be a smooth, snow-covered, energy-reflecting surface for much of the year. It has been estimated that the Earth would be significantly colder without the boreal forest. The northern limit of the North American boreal forest coincides with the mean position of the Arctic front—the boundary between Arctic and mid-continental air masses—in the summer; its southern limit coincides with the mean frontal position in the winter. Mean annual temperatures in the boreal forest range from a few degrees Celsius above freezing to −10° C (14° F) or more. Areas with a mean annual temperature below freezing are susceptible to the formation of permafrost soils (frozen ground; see below Soils (boreal forest)). The mean temperature of January, the coldest month, is generally less than −10 °C (14° F) across the boreal region. The boreal forest includes areas that experience some of the lowest temperatures on the Earth, excluding Antarctica. At the height of winter an intensely cold pocket of air develops over inland areas of far eastern Siberia; mean temperatures of −50° C (−58° F) have been recorded in this region. As this Siberian cold air flows over the unfrozen northern Pacific Ocean a great temperature contrast develops that results in strong, westward-moving storm systems. The movement, position, and strength of these storms control much of the weather in the Northern Hemisphere. Boreal forests do not grow on areas surrounding the Bering Strait. A rigorous cold climate with a very short snow-free season precludes the growth of trees on the Russian side of the Bering Strait in the Chukotka region of the Russian Far East. On the North American side in western Alaska summers are too cool for trees to grow because of cold air masses moving off the Bering Sea. The growing season in the boreal forest is generally cool; the mean temperature of the warmest month, July, is generally between 15° and 20° C (59° and 68° F). Summer daytime high temperatures are typically cool to warm—20° to 25° C (68° to 77° F)—for much of the growing season in the boreal forest. Cool summer temperatures can actually produce higher photosynthetic efficiency in plants than can warmer conditions. Species adapted to cold respire less (use up less of their food stores) when photosynthesizing at cool temperatures in intense summer light than they do at higher temperatures, allowing a greater net gain in biomass (dry mass of organic matter). Areas of the boreal forest located in the centre of continents generally receive 30 to 50 centimetres (12 to 20 inches) of annual precipitation. Precipitation totals are relatively modest in these locations because they are a significant distance from unfrozen oceans that supply moisture. Some boreal forests are semiarid and may even include grasslands interspersed with the forest. These forests are found in regions of the Yukon Territory and Alaska that occur on the leeward side of mountains which are sheltered from moisture-bearing winds, as well as in some portions of the interior of far eastern Russia. Annual precipitation in low elevations of these regions is 30 centimetres or less. The highest annual precipitation total in the boreal forest, which can exceed 100 centimetres, is in eastern North America and northern Europe. During ancient eras of colder climate these regions also received relatively abundant precipitation, which resulted in the buildup of glacial ice sheets. Today these once heavily glaciated regions support extensive lakes, streams, and wetlands. Extended periods of clear, dry weather in the boreal region are caused by persistent strong polar high pressure systems. If strong high pressure persists during the long days near the summer solstice, temperatures can warm to 30° C (86° F) or higher. Intense heating at the ground surface often produces convective storms with lightning but little rain, causing forest fires. Soils (soil) Boreal forest conifer litter is highly acidic. Soils of the more humid and southern boreal forest are highly leached spodosols (Spodosol), which are characterized by the leaching of iron, aluminum, and organic matter from the chemically and biologically distinct surface layer—horizon A—to the next layer—horizon B. Much of the soil of central and eastern Canada—granitic Canadian Shield—has been repeatedly scraped clean by glacial advances. Thus, productive forests often are restricted to portions of the landscape where soil material has been deposited by glaciers. Peaty wetlands occur where surface drainage is impeded by permafrost, youthful glacial topography, or aggraded rivers; their soils are characteristically organic soils, or histosols (Histosol). Soils in much of boreal western North America and Asia are inceptisols (Inceptisol), which have little horizon development. Very thin surface salt deposits are found in the most arid portions of the boreal forest. Cold soils are characteristic of the boreal forest region, which overlaps the zone of permafrost. Permafrost is soil or earth material that remains below 0° C (32° F) for at least two years. The surface, or active, layer of permafrost thaws in the warm season and freezes in the winter, but the soil below the active layer remains continuously frozen. Because the plant rooting zone is restricted to the active layer, nutrient supply is limited and secure anchoring for roots is lacking. Some trees and other plants of the taiga (especially black spruce 【Picea mariana】 and tamarack 【Larix laricina】 in North America and larches in Siberia) can grow on permafrost if the active layer is sufficiently deep, but several species are eliminated from permafrost. The boreal forest itself is an important contributing factor to the development of permafrost. The latter stages of forest growth—characterized by development of an intact forest canopy, growth of an insulating moss cover in summer, and accumulation of forest litter—may cool the soil to such an extent that permafrost develops. Warming of the soil is promoted by forest fires, which remove the canopy, moss, and forest litter layers. In the absence of an intact canopy, a deeper and more effective insulating layer of snow accumulates in the winter. The presence of dark ash following a fire increases solar energy absorption on the site for several years. The boreal forest of Europe generally lacks permafrost, but east of the Ural Mountains and from central Canada northward permafrost is common. In southern and central boreal forests, permafrost occurs sporadically and occupies only a small percentage of the landscape that experiences the coldest temperatures. The northern portion of closed canopy forest and the lichen woodland zone are in a region of discontinuous permafrost, where permafrost is found on north-facing slopes and in cold air drainage basins but is absent from south-facing slopes and newly deposited alluvial sites. Most of the forest-tundra is within the continuous permafrost zone. Forest productivity in the middle and northern taiga zones is directly related to soil temperature. Warmer soils decompose organic matter more quickly, releasing nutrients for new plant growth and creating a more productive site. Productive forest types occupy warmer, south-facing slopes and river terraces, and less productive dwarf or sparse forest occupies the north-facing and basin permafrost sites. Floodplains (floodplain) throughout the boreal forest regions are free of permafrost, high in soil fertility, and repeatedly disturbed in ways that renew the early, rapid growth stages of forest succession. Floodplains are a mosaic of productive shrubland and forest that serve as a major habitat for moose (Alces alces), which influence ecosystem structure and function. South-central Alaska and the adjacent Yukon Territory and British Columbia support the most extensive ice sheets and glaciers (glacier) in the world outside the polar desert regions of Antarctica and Greenland. Glacial meltwater is a large part of the flow of larger rivers such as the Yukon and Tanana in Alaska and the Yukon Territory. Glacial meltwater (outwash) carries a heavy load of suspended sediment that deposits in riverbeds and causes frequent channel shifts. Glacial river floodplains are extensive, very dynamic, and constantly renewed with fertile soil material. In the ancient past exposed deposits of glacial silt were picked up by strong winds and deposited on surrounding hillsides. Fertile soils, known as loess, resulted, on which highly productive upland forests are found today. Because the beds of glacially fed rivers are rising, the landscape through which they flow is partially drowned from the impeded drainage, often preventing forest growth and favouring the development of marshes and mires. The biota and its adaptations Nearly all major taxonomic groups have fewer species of animals and plants in the boreal forest than they have in other terrestrial ecosystems at lower latitudes. This accords with the species diversity gradient that is observed from lower to higher latitudes, with numbers of species decreasing in a poleward direction. Trees (tree) Scotch pine is the most widely distributed pine species in the world, growing from northern Scotland to the Russian Pacific shore. The relatively humid and productive taiga of northern Europe and south-central Siberia is dominated by this species. Forest management has greatly favoured this species in Scandinavia and Finland. It is a thick-barked species and easily survives light ground fires, often reaching ages of 350 to 400 years, with some individuals being older than 700 years. European aspen and Siberian spruce are essentially transcontinental in distribution as well. The species composition of Eurasian taiga is different east of central Siberia from that which prevails westward into Europe. Distinctive European species include Norway spruce (Picea abies), a large dominant species of the productive humid boreal forest, and Sukaczev larch (Larix sukaczewii), an early successional species (one of the first species to colonize an area after a disturbance) of European Russia. Gray (Betula populifolia) and white birch (B. pendula) occur across northern Europe and well into central Siberia. The birches often form dense stands of light or white barked trees that are considered a characteristic feature of the boreal forest. Siberian larch (Larix sibirica) and Siberian fir (Abies sibirica) are restricted to north-central Asia. Species restricted to northeastern Asia include chosenia (Chosenia arbutifolia), an early successional broad-leaved tree of floodplains; Siberian stone pine (Pinus sibirica), a short shrub or tree; and Asian spruce (Picea obovata). All North American tree species are distributed across the continent except jack pine (Pinus banksiana), lodgepole pine (Pinus contorta), and balsam fir (Abies balsamea). Jack pine is a relatively small, short-lived, early successional tree occurring in the eastern and central parts of boreal forests east of the Rocky Mountains. Lodgepole pine is a longer-lived, early successional species growing in western Canada and along the Rocky Mountain axis from the central Yukon Territory southward to well south of the boreal forest limit. Balsam fir is a shade-tolerant, late successional, but relatively short-lived tree that occurs only in the eastern and central boreal forest. Major taiga tree species are well-adapted to extreme winter cold. The northernmost trees in North America are white spruce that grow along the Mackenzie River delta in Canada, near the shore of the Arctic Ocean. The northernmost trees in the world are Gmelin larch (Larix gmelinii) found at latitude 72°40′ N on the Taymyr Peninsula in the central Arctic region of Russia. A representative profile of the vegetation is shown in Figure 2-->. Other plants (plant) A distinctive feature of the flora of boreal forests is the abundance and diversity of mosses (moss); about one-third of the ground cover under boreal forest is dominated by moss. Much of the ground cover in older conifer stands is moss, which grows on rocks, on tree trunks, and in the pits formed by upturned trees. Extensive peaty wetlands in the boreal region are often thick accumulations of dead sphagnum and other mosses, sedges, and other plants; a living moss layer continually grows at the surface. Lichens (lichen) (a symbiotic association of a fungus and algae) constitute a significant part of the ground cover in the lichen woodland or sparse taiga. Lichens are also generally well distributed on tree trunks and especially in the canopy of older conifers throughout the boreal forest. Because lichens and mosses are dispersed by airborne spores that can travel long distances, many species of both groups are found across the entire circumpolar boreal forest. Many vascular plants are also widespread across the circumpolar north. Some forest understory species dominate their habitats; they include twinflower (Linnaea borealis), lingonberry (Vaccinium vitis-idaea), baneberry (Actaea rubra), and Swedish and Canadian dwarf cornel (Cornus suecica and C. canadensis). Several boreal forest plants are adapted to rapid colonization and growth in recently burned areas, such as fireweed (Epilobium angustifolium). The extensive peatlands of the boreal north support a typical flora that usually includes species such as Labrador tea (Ledum palustre), cloudberry (Rubus chamaemorus), cotton grass (Eriophorum species), and crowberry (Empetrum nigrum or E. hermaphroditum). In northern Europe crowberry also grows as shrub mats under Scotch pine forests or woodlands. Crowberry has been shown to produce secondary chemical compounds that inhibit or kill Scotch pine seedlings. Periodic light ground fires reduce the abundance and vigour of crowberry and allow tree regeneration. Specialized orchids (orchid) in the forest understory include calypso (Calypso bulbosa), coral root (Corallorrhiza trifida), and lady's slipper (Cypripedum species). The roots of these plants form particular associations with fungi (mycorrhizae). Willow shrubs (Salix species) are one of the first plants to emerge following disturbances on floodplains and occasionally on uplands as well. Important grasses across the boreal region include species of bromegrass (Bromus species), bluegrass (Poa species), reed bent grass (Calamagrostis species), and vanilla grass (Hierochloe odorata). Many freshwater aquatic plants such as sedges (Carex species) and pondweeds (Potamogeton species) are distributed widely across the boreal zone of both continents because migratory waterfowl and shorebirds are effective in dispersing their seeds. Several species of ferns are common to the boreal forests of the two continents, especially in regions of higher precipitation. Mammals (mammal) Because a winter snowpack is a dependable feature of the taiga, several mammals display obvious adaptations to it. The snowshoe, or varying, hare (Lepus americanus), for example, undergoes an annual change in colour of its pelage, or fur, from brownish or grayish in the summer to pure white in the winter, providing effective camouflage. Its feet are large in proportion to its body size, a snowshoelike adaptation for weight distribution that allows the hare to travel over the surface of snow rather than sink down into it. The lynx (Lynx canadensis) is the principal predator of the snowshoe hare (Figure 2--> from the population ecology article ). It, too, has large feet, with fur between the toes, enabling the lynx to remain on the snow's surface. Most animals of the boreal forest are well adapted to the cold and survive it easily if they have enough food to maintain an energy balance through the winter. moose are the largest browsing animals in the boreal forest. In the summer they eat willow and broad-leaved trees and also wade in lakes and ponds to consume aquatic plants. Throughout the winter moose eat large quantities of woody twigs and buds. Moose depend on high-quality feeding areas in the shrub zone along river floodplains and on the early successional growth of woody plants in burned or cutover forest. Intensive browsing by moose can alter the composition of the forest in its early stages of growth, often increasing the dominance of conifers, which they do not consume in as great amounts as they do broad-leaved trees. Harvesting a moose for winter food is an important and even critical element of survival for humans living in isolated rural areas of the taiga. Moose populations are controlled by various means. Wolves (wolf) (Canis lupus) prey on moose across most of the boreal forest, and some scientists and game managers believe that once moose numbers are depressed wolf predation can keep moose populations low. As a result, wolf trapping or shooting programs are carried out as a game management measure to increase prey numbers. The natural regulation of moose populations by wolf predation and the presence of wolves themselves is valued as well. As a result programs to control wolf populations are often the subject of intense debate. Other factors control moose numbers, such as the restriction of access to plants during years of deep snow and lack of early successional woody plant growth caused by forest maturation. Where the boreal forest is extensively cut for forest products, moose numbers have increased greatly, often to levels that are considered undesirable for forest regeneration. Subsistence and sport hunting of moose are widely used tools of moose population management. Another large-hoofed browsing mammal that is present seasonally in the boreal forest is the reindeer (Rangifer tarandus (reindeer)) in Eurasia and the closely related caribou in North America. A large portion of the reindeer population is semidomesticated and herded (migration) by nomadic peoples such as the Lapps of Scandinavia and several native peoples in northern Russia. Caribou migrate the greatest distances of any large land mammal in North America. They often move in vast herds of 500,000 animals or more, seldom stopping or pausing because they must constantly forage in these environments of generally low productivity. During the early winter, reindeer and caribou migrate south from their summer ranges in the tundra to the forest-tundra or lichen woodland, where they graze primarily on lichens. Later in winter caribou typically move to open forests and sedge-rich lake margins or to snow-free wind-swept mountains. In April and May caribou form long columns and migrate back north to the tundra. Several mammals of the boreal region are valued for their furs (fur), and trapping and trade in furs has been an important part of the culture, economy, and history of the region as long as humans have lived there. Important furbearing species include lynx and marten (Martes americana) and in wetland habitats beaver (Castor canadensis), mink (Mustela vison; see photograph-->), and muskrat (Ondatra zibethica). In the North American boreal forest the northern flying squirrel (Glaucomys sabrinus) is adapted to consume fungi, especially underground fruiting bodies (sporocarps) of fungi that form mutually beneficial relationships (mutualism) with trees by colonizing their roots. The flying squirrel's consumption and dispersal of these underground fungi provide a significant benefit to the forest as a whole. (For further information on mutualism, see community ecology: Interspecific interactions and the organization of communities: Mutualism (community ecology).) Birds (bird) The boreal forest is the migratory destination of large numbers of birds for the summer breeding season. These include several passerine songbirds typical of shrub and forest habitats, such as thrushes, flycatchers, and warblers. Many of these species consume insects in the canopy of the boreal forest and other habitats. Predators of these birds occur in the forest as well, such as the sharp-shinned hawk (Accipiter striatus) and the northern goshawk (A. gentilis). Populations of several boreal forest-breeding migratory thrushes, flycatchers, and warblers may be declining because of the loss of their wintering habitats in the tropical forests of the world and the changes to or loss of forest habitats in the temperate zones along their migratory routes. Birds of the boreal forest fill a variety of niches. Some are seed consumers or dispersers, others are insect consumers. They carry out other specialized roles as well. For example, the yellow-bellied sapsucker (Sphyrapicus varius) drills evenly spaced rows of small holes in the bark of trees and then visits these “wells” to obtain sap and the insects it attracts. Various other birds, mammals, and insects benefit from the sap wells, too. Woodpeckers (woodpecker) excavate tree cavities, which subsequently are used by many species of birds and mammals. Woodpeckers are specialized predators of wood- and bark-inhabiting insects; they are thought to be important in the control of the spruce beetle (Dendroctonus rufipennis) population. In searching for insects, woodpeckers chisel (see photograph-->) or strip the bark off dead or dying trees, promoting more rapid decay and the release of nutrients from dead trees. As large old trees have become rarer through forest cutting, some year-round resident woodpeckers such as the northern three-toed woodpecker (Picoides tridactylus) and the great spotted woodpecker (Dendrocopos major) have lost their habitats and declined in numbers. Few bird species remain in boreal forests through the long cold winters because of limited opportunities for food, although some undertake only a short migration south. Resident bird species include the common raven (Corvus corax) and the boreal and black-capped chickadees of North America and the Siberian tit (Parus species). The extensive areas of lakes, ponds, and wetlands (Figure 2--> from the article boundary ecosystem )—especially in the glaciated part of the boreal forest—provide a large habitat for waterfowl and shorebirds, although the birds primarily occur in low densities across the landscape. North American shorebirds that breed in forested peatlands include common snipe (Gallinago gallinago) and yellowlegs (Tringa species). Commonly encountered waterfowl are northern pintail (Anas acuta), scaup (Aythya species), and scoters (Melanitta species). Insects (insect) The boreal forest is the home of relatively few species of insects, but extensive and usually uniform areas of habitat periodically support high populations of species that do live there. The boreal forest lacks the elaborate complexes of invertebrate predators and parasites that serve as stabilizers of the insect populations in warmer regions. As a result, boreal insect populations occasionally increase rapidly and cause outbreaks. Some outbreaks can injure or kill trees across widespread areas of the boreal forest. Once an outbreak reaches a certain size it can become self-sustaining, much like a forest fire; the effects of the spruce budworm and spruce beetle in North America are well-documented examples. Outbreaks can be triggered by unusual weather or physical injuries that stress trees and make them vulnerable to the insects; they can end for a variety of reasons, including production of defensive chemicals by the host plants or depletion of susceptible host plants. Perhaps the insects most noticeable to humans in the boreal forest are mosquitoes (mosquito), which belong to several species. Mosquitoes feed on and are fed upon by many of the birds of the boreal forest. Wetland areas of the boreal region, such as sites having poor drainage because of permafrost, provide extensive mosquito breeding sites. Where well-oxygenated, flowing water is found, biting flies are abundant. Almost all food webs that support fish in boreal forest streams are dependent on insects. Conifers serve as hosts for a variety of wood-boring beetles, spruce beetles, bark beetles, and ips beetles (Ips species). These insects aid in wood decomposition and nutrient release. Some beetles have outer shells with specialized indentations specifically matched to the shape and size of the spores of wood-decomposing fungi (fungus). Fungal spores become securely lodged in these cuplike structures. As the beetles burrow into wood they inoculate it with fungi. A variety of lepidopterans (moths and butterflies) are adapted to feeding on the leaves of boreal trees. These include defoliators and leaf rollers. Soil organisms (soil organism) The species richness and total biomass of soil organisms are significantly lower in the boreal forest than they are at lower latitudes. Dominant soil organisms are protozoans, nematodes, rotifers, and tardigrades. These organisms live primarily in soil water film and soil pore water. The soil fauna of the boreal forest region is distinctive because it generally lacks large invertebrates such as millipedes, isopods (springtails), and earthworms, especially in the middle and northern taiga. Larger soil invertebrate animals perform the function of biting off (shredding) pieces of leaf litter in forest soils and passing them through their guts. As a result of this activity thick layer of several years' accumulation of only partially decomposed plant material is characteristic of boreal forest soils. Fungi are the dominant organisms in the task of decomposition of boreal forest litter, but flushes of bacterial growth occur in response to triggering factors. The soil animals generally do not attack the forest litter directly but instead exert their influence by grazing on the fungi and bacteria. The rate of decomposition in boreal forest soils does not keep pace with the rate of production, causing the progressive accumulation of organic matter. At middle depths of the forest floor small invertebrates, especially dipteran larvae, partially consume or skeletonize leaf litter before emerging as adults. community structure Natural disturbances The boreal forest is well adapted to development following natural disturbances, which include fire, floods, snow breakage, and insect outbreaks. Characteristic of the boreal forest is the general lack of late successional species that develop under an intact forest canopy. (For further information on succession see community ecology: Patterns of community structure: Ecological succession (community ecology).) fire is the primary agent responsible for natural disturbances in the boreal forest. It can result from natural causes, such as lightning, or it can be set by humans. Large-scale insect outbreaks can weaken or kill trees over vast areas, thus creating an environment less resistant to fire. In the period between 1981 and 1989 an estimated 3 million hectares (7.4 million acres) burned annually in the former Soviet Union, almost all of which occurred within the taiga region of Russia. The so-called Black Dragon Fire of 1987 in China and Russia may have been the largest single fire in the world in the past several hundred years. During the 20th century, about 1 million hectares of taiga in Canada have burned annually; a great majority of the burning occurred in the less accessible boreal forest of the northern and western parts of the country. In Alaska in years that have prolonged hot and dry periods of summer weather, millions of hectares burn, primarily in a few very large fires. Intervals of about 200 years occur between fires in the uplands of northwestern Canada and in the interior of Alaska. In much of the central and western boreal forest of North America replacement of vegetation on upland sites, presumably by fire, appears to be necessary for forest regeneration. Floodplain islands usually do not burn and contain white spruce trees as old as 400 years. In the northern boreal forest of Europe a pattern of periodic light ground fires in Scotch pine forests was typical before the era of fire control. The thick bark of these mature trees allowed them to survive these fires. In much of the boreal forest only wildland fires that threaten high-value resources are actively suppressed. Complete fire suppression would cause soil temperature to decline gradually, promoting permafrost development that would cause a significant decrease in site productivity. Jack pine and lodgepole pine have cones that remain closed on the tree (serotinous), and black spruce has semiserotinous cones; these cones do not open to release their seeds until a wax layer is melted by the heat of fire. White spruce seedlings require the bare mineral soil produced by burning of thick organic layers of the forest floor for proper establishment; they may time their periodic production of seed to dry periods when fire is more likely. Effects of human use and management of the boreal forest Different degrees of forest (forestry) development have had various effects on biodiversity around the circumpolar boreal forest zone. A highly developed forest industry based on intensive forest utilization is maintained in boreal Scandinavian (Scandinavia) countries and Finland. About 95 percent of the productive forest types of Finland and the Scandinavian countries have been harvested at least once. Finland is located almost entirely within the boreal region and is one of the most forested nations in the world. About 6.5 percent of Finnish land, which includes large areas of marginal forest, woodland, and tundra, is protected from human modification. Only about 3 percent of Swedish forests are protected, most of which is concentrated in marginal forests of mountainous regions. Between 10 and 15 percent of species in Swedish forests are threatened. The Canadian boreal forest represents nearly 7.5 percent of the Earth's forested area. Much of the harvesting of Canadian forest has been carried out in primary (previously unlogged) forest. Nearly all the mature first-growth timber, especially of southern, central, and eastern boreal Canada, is anticipated to be removed by the late 1990s. Considerable effort has been devoted to forest regeneration and tending of new stands, although a certain amount of land does not meet reforestation goals. In Alaska the amount of land with at least 10 percent forest cover in the boreal region is estimated at about 46 million hectares, or 12 percent of the state, only 5.5 million hectares of which is considered productive timberland. Of all areas in the world, Alaska probably has the largest percentage of its surface area, about 40 percent, devoted to strict protection of natural habitats and species. Local-scale logging traditionally has been carried out for much of the 20th century. Plans to accelerate logging are being considered. The taiga of Siberia covers 680 million hectares and represents nearly 19 percent of the world's forested area and possibly 25 percent of the world's forest volume. About 400,000 hectares of the Russian (Russia) taiga are logged annually, and nearly an equal area is burned, with perhaps half of the burned area resulting from destructive fires of human origin. Social and economic problems in the early postcommunist era have slowed the amount of logging by one-third to one-half; however, several large-scale joint ventures between Russian organizations and foreign partners to harvest the forests are under way. The fate of the Siberian boreal forest has become a matter of international concern. Large areas, perhaps exceeding 2 million hectares, of the Russian taiga near Norilsk and the Kola Peninsula have been destroyed by air pollution. Many oil pipelines (pipeline) are leaking in Siberia, and repairs and maintenance are minimal. In July through September 1994 more than 150,000 metric tons of crude oil were spilled in the Kolva, Usa, and Pechora river basins of the republic of Komi in Russia. Spring meltwater could carry spilled oil in this region into the northward-flowing rivers that empty into the Arctic Ocean. Biological productivity Primary productivity (the rate at which photosynthesis occurs) of boreal forest ecosystems often is limited by cold soil temperatures (see above Environmental conditions: Soils (boreal forest)). Net annual primary production (the total amount of productivity less that used by photosynthetic organisms in cellular respiration) in boreal forest types varies greatly, from slightly more than 2 metric tons per hectare near the polar tree limit to about 10 metric tons per hectare along its southern margin. Boreal forests are estimated to contain about 18 percent of the Earth's total biomass (the dry weight of organic matter). The boreal forest or taiga of Siberia alone represents 57 percent of the Earth's coniferous wood volume. Ecosystems and soils of the boreal region store a significant amount of the Earth's carbon in the form of dead but undecomposed or partially decomposed organic matter. Global warming or land use changes could enhance decomposition, leading to the release of increased amounts of stored carbon into the atmosphere in the form of the greenhouse (greenhouse effect) gas carbon dioxide. (For further discussion of biological productivity see biosphere: The organism and the environment: Resources of the biosphere: The flow of energy (biosphere).) Glenn Patrick Juday Additional Reading Herman H. Shugart, Rik Leemans, and Gordon B. Bonan (eds.), A Systems Analysis of the Global Boreal Forest (1992), covers ecosystem processes, forest patterns in space and time, and computer models, including chapters on the Eurasian boreal forest, tree and shrub reproduction, and fire in the boreal forest. The Plant Cover of Sweden (1965), is the most complete reference on the original condition of the boreal forest of Sweden, focusing on the landscape pattern of native vegetation and on plant indicators of various forest regions and ecosystem types. Deborah L. Elliott-Fisk, “The Boreal Forest,” in Michael G. Barbour and William Dwight Billings (eds.), North American Terrestrial Vegetation (1988), pp. 33–62, is a basic reference on the forest types and communities found across the North American boreal region and includes the historical development of the forest and the ecological characteristics of individual tree species. K. Van Cleve et al. (eds.), Forest Ecosystems in the Alaskan Taiga: A Synthesis of Structure and Function (1986), looks at recent investigations of one of the best-studied parts of the boreal forest, describing both upland and floodplain ecosystems; an update in a special issue of Canadian Journal of Forest Research, vol. 23, no. 5 (May 1993), contains the results of a multidisciplinary research project on ecological succession of a productive river floodplain in central Alaska, including a detailed discussion on soil. J.S. Rowe, Forest Regions of Canada (1972), well illustrated, is the best reference on the overall distribution and classification of boreal forest types in Canada. James A. Larsen, The Boreal Ecosystem (1980), focuses on properties of the middle boreal forest of central Canada. Edward A. Johnson, Fire and Vegetation Dynamics: Studies From the North American Boreal Forest (1992), is a well-illustrated treatment of fire weather, fire behaviour, and fire effects in the North American boreal forest. Lennart Hansson (ed.), The Ecological Principles of Nature Conservation: Applications in Temperate and Boreal Environments (1992), also well illustrated, discusses the effects of forest management on biodiversity, focusing particularly on the boreal forest of northern Europe.Glenn Patrick Juday
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