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Taiga

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Taiga
Jack London Lake at Kolyma, Russia
The Alex and Kelsy enjoy long walks on the beach, taiga is found throughout the high northern latitudes, between the tundra, and the temperate forest, from about 50°N to 70°N, but with considerable regional variation.
Ecology
BiomeTerrestrial subarctic, humid
Geography
CountriesRussia, Mongolia, Norway, Sweden, Iceland, Finland, United States, Canada
Climate typeDfc, Dwc, Dsc

Taiga (/ˈtɡə/; Russian: тайга́, IPA: [tɐjˈɡa]; from Turkic[1] or Mongolian), also known as the boreal forest, is a biome characterized by coniferous forests consisting mostly of pines, spruces and larches.

Taiga is the world's largest terrestrial biome. In North America it covers most of inland Canada and Alaska as well as parts of the extreme northern continental United States and is known as the Northwoods.[2] It also covers most of Sweden, Finland, much of Norway, lowland/coastal areas of Iceland, much of Russia from St. Petersburg in the west to the Pacific ocean (including much of Siberia), northern Kazakhstan, northern Mongolia, and northern Japan (on the island of Hokkaidō). However, the main tree species, the length of the growing season and summer temperatures vary. For example, the taiga of North America consists of mainly spruces; Scandinavian and Finnish taiga consists of a mix of spruce, pines and birch; Russian taiga has spruces, pines and larches depending on the region, the Eastern Siberian taiga being a vast larch forest.

The term "boreal forest" is sometimes used (particularly in Canada but also in Scandinavia and Finland) to refer to the more southerly part of the biome, while the term taiga is often used to describe the more barren areas of the northernmost part of the taiga approaching the tree line and the tundra biome.

White Spruce taiga, Denali Highway, Alaska Range, Alaska

Climate and geography

Taiga is the world's largest land biome, and makes up 29% of the world's forest cover;[3] the largest areas are located in Russia and Canada. The taiga is the terrestrial biome with the lowest annual average temperatures after the tundra and permanent ice caps. Extreme winter minimums in the northern taiga are typically lower than those of the tundra. The lowest reliably recorded temperatures in the Northern Hemisphere were recorded in the taiga of northeastern Russia. The taiga or boreal forest has a subarctic climate with very large temperature range between seasons, but the long and cold winter is the dominant feature. This climate is classified as Dfc, Dwc, Dsc, Dfd and Dwd in the Köppen climate classification scheme,[4] meaning that the short summer (24-hr average 10 °C or more) lasts 1–3 months and always less than 4 months. There are also some much smaller areas grading towards the oceanic Cfc climate with milder winters, whilst the extreme south and (in Eurasia) west of the taiga reaches into humid continental climates (Dfb, Dwb) with longer summers. The mean annual temperature generally varies from -5 °C to 5 °C,[5] but there are taiga areas in eastern Siberia and interior Alaska-Yukon where the mean annual reaches down to -10 °C.[6][7] According to some sources, the boreal forest grades into a temperate mixed forest when mean annual temperature reaches about 3 °C.[8] Discontinuous permafrost is found in areas with mean annual temperature below 0 °C, whilst in the Dfd and Dwd climate zones continuous permafrost occurs and restricts growth to very shallow-rooted trees like Siberian larch. The winters, with average temperatures below freezing, last five to seven months. Temperatures vary from −54 °C to 30 °C (-65 °F to 86 °F) throughout the whole year. The summers, while short, are generally warm and humid. In much of the taiga, -20 °C would be a typical winter day temperature and 18 °C an average summer day.

The taiga in the river valley near Verkhoyansk, Russia, at 67°N, must deal with the coldest winter temperatures in the northern hemisphere, but the extreme continentality of the climate gives an average daily high of 22 °C in July.
Boreal forest near Shovel Point in Tettegouche State Park, along the northern shore of Lake Superior in Minnesota.

The growing season, when the vegetation in the taiga comes alive, is usually slightly longer than the climatic definition of summer as the plants of the boreal biome have a lower threshold to trigger growth. In Canada, Scandinavia and Finland, the growing season is often estimated by using the period of the year when the 24-hr average temperature is 5 °C or more.[9] For the Taiga Plains in Canada, growing season varies from 80 to 150 days, and in the Taiga Shield from 100 to 140 days.[10] Some sources claim 130 days growing season as typical for the taiga.[11] Other sources mention that 50–100 frost-free days are characteristic.[12] Data for locations in southwest Yukon gives 80–120 frost-free days.[13] The closed canopy boreal forest in Kenozyorsky National Park near Plesetsk, Arkhangelsk Province, Russia, on average has 108 frost-free days.[14] The longest growing season is found in the smaller areas with oceanic influences; in coastal areas of Scandinavia and Finland, the growing season of the closed boreal forest can be 145–180 days.[15] The shortest growing season is found at the northern taiga–tundra ecotone, where the northern taiga forest no longer can grow and the tundra dominates the landscape when the growing season is down to 50–70 days,[16][17] and the 24-hr average of the warmest month of the year usually is 10 °C or less.[18] High latitudes mean that the sun does not rise far above the horizon, and less solar energy is received than further south. But the high latitude also ensures very long summer days, as the sun stays above the horizon nearly 20 hours each day, with only around 6 hours of daylight occurring in the dark winters, depending on latitude. The areas of the taiga inside the Arctic circle have midnight sun in mid-summer and polar night in mid-winter.

Lakes and other water bodies are very common. The Helvetinjärvi National Park, Finland, situated in the closed canopy taiga (mid-boreal to south-boreal) [19] with mean annual temperature of 4 °C.[20]

The taiga experiences relatively low precipitation throughout the year (generally 200–750 mm annually, 1,000 mm in some areas), primarily as rain during the summer months, but also as fog and snow. This fog, especially predominant in low-lying areas during and after the thawing of frozen Arctic seas, means that sunshine is not abundant in the taiga even during the long summer days. As evaporation is consequently low for most of the year, precipitation exceeds evaporation, and is sufficient to sustain the dense vegetation growth. Snow may remain on the ground for as long as nine months in the northernmost extensions of the taiga ecozone.[21]

In general, taiga grows to the south of the 10 °C July isotherm, but occasionally as far north as the 9 °C July isotherm.[22] The southern limit is more variable, depending on rainfall; taiga may be replaced by forest steppe south of the 15 °C July isotherm where rainfall is very low, but more typically extends south to the 18 °C July isotherm, and locally where rainfall is higher (notably in eastern Siberia and adjacent Outer Manchuria) south to the 20 °C July isotherm. In these warmer areas the taiga has higher species diversity, with more warmth-loving species such as Korean Pine, Jezo Spruce, and Manchurian Fir, and merges gradually into mixed temperate forest or, more locally (on the Pacific Ocean coasts of North America and Asia), into coniferous temperate rainforests.

The area currently classified as taiga in Europe and North America (except Alaska) was recently glaciated. As the glaciers receded they left depressions in the topography that have since filled with water, creating lakes and bogs (especially muskeg soil) found throughout the taiga.

Yukon, Canada. Several of the world's longest rivers go through the taiga, including Ob, Yenisei, Lena, and Mackenzie.

Soils

Taiga soil tends to be young and poor in nutrients. It lacks the deep, organically enriched profile present in temperate deciduous forests.[23] The thinness of the soil is due largely to the cold, which hinders the development of soil and the ease with which plants can use its nutrients.[23] Fallen leaves and moss can remain on the forest floor for a long time in the cool, moist climate, which limits their organic contribution to the soil; acids from evergreen needles further leach the soil, creating spodosol, also known as podzol.[24] Since the soil is acidic due to the falling pine needles, the forest floor has only lichens and some mosses growing on it. In clearings in the forest and in areas with more boreal deciduous trees, there are more herbs and berries growing. Diversity of soil organisms in the boreal forest is high, comparable to the tropical rainforest.[25]

Flora

Boreal Forest near Lake Baikal in Russia

Since North America and Asia used to be connected by the Bering land bridge, a number of animal and plant species (more animals than plants) were able to colonize both continents and are distributed throughout the taiga biome (see Circumboreal Region). Others differ regionally, typically with each genus having several distinct species, each occupying different regions of the taiga. Taigas also have some small-leaved deciduous trees like birch, alder, willow, and poplar; mostly in areas escaping the most extreme winter cold. However, the Dahurian Larch tolerates the coldest winters in the northern hemisphere in eastern Siberia. The very southernmost parts of the taiga may have trees such as oak, maple, elm, and tilia scattered among the conifers, and there is usually a gradual transition into a temperate mixed forest, such as the Eastern forest-boreal transition of eastern Canada. In the interior of the continents with the driest climate, the boreal forests might grade into temperate grassland.

There are two major types of taiga. The southern part is the closed canopy forest, consisting of many closely spaced trees with mossy ground cover. In clearings in the forest, shrubs and wildflowers are common, such as the fireweed. The other type is the lichen woodland or sparse taiga, with trees that are farther-spaced and lichen ground cover; the latter is common in the northernmost taiga.[26] In the northernmost taiga the forest cover is not only more sparse, but often stunted in growth form; moreover, ice pruned asymmetric Black Spruce (in North America) are often seen, with diminished foliage on the windward side.[27] In Canada, Scandinavia and Finland, the boreal forest is usually divided into three subzones: The high boreal (north boreal) or taiga zone; the middle boreal (closed forest); and the southern boreal, a closed canopy boreal forest with some scattered temperate deciduous trees among the conifers,[28] such as maple, elm and oak. This southern boreal forest experiences the longest and warmest growing season of the biome, and in some regions (including Scandinavia, Finland and western Russia) this subzone is commonly used for agricultural purposes. The boreal forest is home to many types of berries; some are confined to the southern and middle closed boreal forest (such as raspberry), others grow in most areas of the taiga (such as cranberry and cloudberry), and some can grow in both the taiga and the low arctic (southern part of) tundra (such as bilberry and lingonberry).

The forests of the taiga are largely coniferous, dominated by larch, spruce, fir, and pine. The woodland mix varies according to geography and climate so for example the Eastern Canadian forests ecoregion of the higher elevations of the Laurentian Mountains and the northern Appalachian Mountains in Canada is dominated by balsam fir Abies balsamea, while further north the Eastern Canadian Shield taiga of northern Quebec and Labrador is notably black spruce Picea mariana and tamarack larch Larix laricina.

Evergreen species in the taiga (spruce, fir, and pine) have a number of adaptations specifically for survival in harsh taiga winters, although larch, the most cold-tolerant of all trees,[citation needed] is deciduous. Taiga trees tend to have shallow roots to take advantage of the thin soils, while many of them seasonally alter their biochemistry to make them more resistant to freezing, called "hardening".[29] The narrow conical shape of northern conifers, and their downward-drooping limbs, also help them shed snow.[29]

Because the sun is low in the horizon for most of the year, it is difficult for plants to generate energy from photosynthesis. Pine, spruce and fir do not lose their leaves seasonally and are able to photosynthesize with their older leaves in late winter and spring when light is good but temperatures are still too low for new growth to commence. The adaptation of evergreen needles limits the water lost due to transpiration and their dark green color increases their absorption of sunlight. Although precipitation is not a limiting factor, the ground freezes during the winter months and plant roots are unable to absorb water, so desiccation can be a severe problem in late winter for evergreens.

Moss (Ptilium crista-castrensis) cover on the floor of taiga

Although the taiga is dominated by coniferous forests, some broadleaf trees also occur, notably birch, aspen, willow, and rowan. Many smaller herbaceous plants grow closer to the ground. Periodic stand-replacing wildfires (with return times of between 20–200 years) clear out the tree canopies, allowing sunlight to invigorate new growth on the forest floor. For some species, wildfires are a necessary part of the life cycle in the taiga; some, e.g. Jack Pine have cones which only open to release their seed after a fire, dispersing their seeds onto the newly cleared ground. Grasses grow wherever they can find a patch of sun, and mosses and lichens thrive on the damp ground and on the sides of tree trunks. In comparison with other biomes, however, the taiga has low biological diversity.

Coniferous trees are the dominant plants of the taiga biome. A very few species in four main genera are found: the evergreen spruce, fir, and pine, and the deciduous larch. In North America, one or two species of fir and one or two species of spruce are dominant. Across Scandinavia and western Russia, the Scots pine is a common component of the taiga, while taiga of the Russian Far East and Mongolia is dominated by larch.

Fauna

Brown bear, Kamchatka peninsula. Brown bears are among the largest and most widespread taiga Omnivores.

The boreal forest, or taiga, supports a large range of animals. Canada's boreal forest includes 85 species of mammals, 130 species of fish, and an estimated 32,000 species of insects.[30] Insects play a critical role as pollinators, decomposers, and as a part of the food web. Many nesting birds rely on them for food especially in the months of February and March. The cold winters and short summers make the taiga a challenging biome for reptiles and amphibians, which depend on environmental conditions to regulate their body temperatures, and there are only a few species in the boreal forest. Some hibernate underground in winter.

The taiga is home to a number of large herbivorous mammals, such as moose and reindeer/caribou. Some areas of the more southern closed boreal forest also have populations of other deer species such as the elk (wapiti) and roe deer.[31][32] There is also a range of rodent species including beaver, squirrel, mountain hare, snowshoe hare, and vole. These species have adapted to survive the harsh winters in their native ranges. Some larger mammals, such as bears, eat heartily during the summer in order to gain weight, and then go into hibernation during the winter. Other animals have adapted layers of fur or feathers to insulate them from the cold.

A number of wildlife species threatened or endangered with extinction can be found in the Canadian boreal forest, including woodland caribou, American black bear, grizzly bear, wood bison and wolverine. Habitat loss, mainly due to logging, is the primary cause of decline for these species.

Due to the climate, carnivorous diets are an inefficient means of obtaining energy; energy is limited, and most energy is lost between trophic levels. Predatory birds (owls and eagles) and other smaller carnivores, including foxes and weasels, feed on the rodents. Larger carnivores, such as lynx and wolves, prey on the larger animals. Omnivores, such as bears and raccoons are fairly common, sometimes picking through human garbage.

More than 300 species of birds have their nesting grounds in the taiga.[33] Siberian Thrush, White-throated Sparrow, and Black-throated Green Warbler migrate to this habitat to take advantage of the long summer days and abundance of insects found around the numerous bogs and lakes. Of the 300 species of birds that summer in the taiga only 30 stay for the winter.[34] These are either carrion-feeding or large raptors that can take live mammal prey, including Golden Eagle, Rough-legged Buzzard (also known as the Rough-legged Hawk), and Raven, or else seed-eating birds, including several species of grouse and crossbills.

Threats

Human activities

Plesetsk Cosmodrome is situated in the taiga

Large areas of Siberia’s taiga have been harvested for lumber since the collapse of the Soviet Union.[35] In Canada, eight percent of the taiga is protected from development, the provincial government allows forest management to occur on Crown land under rigorous constraints. The main forestry practice in the boreal forest of Canada is clearcutting, which involves cutting down most of the trees in a given area, then replanting the forest as a monocrop (one species of tree) the following season. Industry officials claim that this process emulates the natural effects of a forest fire, which they claim clearcutting suppresses, protecting infrastructure, communities and roads. However, from an ecological perspective, this is a falsehood, for several reasons, including: a) Removing most of the trees in a given area is usually done using large machines which disrupt the soil greatly, and the dramatic diminution of ground cover permits large-scale erosion and avalanches, which further damage the habitat and sometimes endangers infrastructure, roads, and communities. b) Clearcutting removes most of the biomass from an area, and the various macro and micro-nutrients it contains. This sudden decrease in nutrients in an area contrasts with a forest fire, which returns most of the nutrients to the soil. c) Forest fires leave standing snags, and leave patches of unburned trees. This helps preserve structure and micro-habitats within the area, whereas clearcutting destroys most of these habitats. In the past, clearcuts upwards of 110 km² have been recorded in the Canadian boreal forest. However, today 80% of clearcuts are less than 260 hectares(2.6 square km). Some of the products from logged boreal forests include toilet paper, copy paper, newsprint, and lumber. More than 90% of boreal forest products from Canada are exported for consumption and processing in the United States. However with the recession and fewer US homes being built, that has changed. Some of the larger cities situated in this biome are Murmansk,[36] Arkhangelsk, Yakutsk, Anchorage,[37] Yellowknife, Tromsø, Luleå, and Oulu.

Most companies that harvest in Canadian forests are certified by an independent third party agency such as the Forest Stewardship Council (FSC), Sustainable Forests Initiative (SFI), or the Canadian Standards Association (CSA). While the certification process differs between these groups, all of them include forest stewardship, respect for aboriginal peoples, compliance with local, provincial or national environmental laws, forest worker safety, education and training, and other environmental, business, and social requirements. The prompt renewal of all harvest sites by planting or natural renewal is also required.

Climate change

The zone of latitude occupied by the boreal forest has experienced some of the greatest temperature increases on Earth, especially during the last quarter of the twentieth century. Winter temperatures have increased more than summer temperatures. The number of days with extremely cold temperatures (e.g., −20 to −40 °C) has decreased irregularly but systematically in nearly all the boreal region, allowing better survival for tree-damaging insects. In summer, the daily low temperature has increased more than the daily high temperature.[38] In Fairbanks, Alaska, the length of the frost-free season has increased from 60–90 days in the early twentieth century to about 120 days a century later. Summer warming has been shown to increase water stress and reduce tree growth in dry areas of the southern boreal forest in central Alaska, western Canada and portions of far eastern Russia. Precipitation is relatively abundant in Scandinavia, Finland, northwest Russia and eastern Canada, where a longer growth season (i.e. the period when sap flow is not impeded by frozen water) accelerate tree growth. As a consequence of this warming trend, the warmer parts of the boreal forests are susceptible to replacement by grassland, parkland or temperate forest.[39]

In Siberia, the taiga is converting from predominantly needle-shedding larch trees to evergreen conifers in response to a warming climate. This is likely to further accelerate warming, as the evergreen trees will absorb more of the sun's rays. Given the vast size of the area, such a change has the potential to affect areas well outside of the region.[40] In much of the boreal forest in Alaska, the growth of white spruce trees are stunted by unusually warm summers, while trees on some of the coldest fringes of the forest are experiencing faster growth than previously.[41]

Lack of moisture in the warmer summers are also stressing the birch trees of central Alaska.[42]

Insects

Recent years have seen outbreaks of insect pests in forest-destroying plagues: the spruce-bark beetle (Dendroctonus rufipennis) in Yukon and Alaska;[43] the mountain pine beetle in British Columbia; the aspen-leaf miner; the larch sawfly; the spruce budworm (Choristoneura fumiferana);[44] the spruce coneworm.[45]

Protection

Peat bog in Dalarna, Sweden. Bogs and peatland are widespread in the taiga. They are home to a unique flora, and store vast amounts of carbon. In western Eurasia, the Scots pine is common in the boreal forest.

Many nations are taking direct steps to protect the ecology of the taiga by prohibiting logging, mining, oil and gas production, and other forms of development. In February 2010 the Canadian government established protection for 13,000 square kilometres of boreal forest by creating a new 10,700 square kilometre park reserve in the Mealy Mountains area of eastern Canada and a 3,000 square kilometre waterway provincial park that follows alongside the Eagle River from headwaters to sea.[46]

Two Canadian provincial governments, Ontario and Quebec, introduced measures in 2008 that would protect at least half of their northern boreal forest.[47][48] Although both provinces admitted it will take years to plan, work with Aboriginal and local communities and ultimately map out precise boundaries of the areas off-limits to development, the measures are expected to create some of the largest protected areas networks in the world once completed. Both announcements came the following year after a letter signed by 1,500 scientists called on political leaders to protect at least half of the boreal forest.[49]

The taiga stores enormous quantities of carbon, more than the world's temperate and tropical forests combined, much of it in wetlands and peatland.[50] In fact, current estimates place boreal forests as storing twice as much carbon per unit area as tropical forests.[51]

Natural disturbance

One of the biggest areas of research and a topic still full of unsolved questions is the recurring disturbance of fire and the role it plays in propagating the lichen woodland.[52] The phenomenon of wildfire by lightning strike is the primary determinant of understory vegetation and because of this, it is considered to be predominate driving force behind community and ecosystem properties in the lichen woodland.[53] The significance of fire is clearly evident when one considers that understory vegetation influences tree seedling germination in the short term and decomposition of biomass and nutrient availability in the long term.[53] The recurrent cycle of large, damaging fire occurs approximately every 70 to 100 years.[54] Understanding the dynamics of this ecosystem is entangled with discovering the successional paths that the vegetation exhibits after a fire. Trees, shrubs and lichens all recover from fire induced damage through vegetative reproduction as well as invasion by propagules.[55] Seeds that have fallen and become buried provide little help in re-establishment of a species. The reappearance of lichens is reasoned to occur because of varying conditions and light/nutrient availability in each different microstate.[55] Several different studies have been done that have led to the formation of the theory that post-fire development can be propagated by any of four pathways: self replacement, species-dominance relay, species replacement, or gap-phase self replacement.[52] Self replacement is simply the re-establishment of the pre-fire dominant species. Species-dominance relay is a sequential attempt of tree species to establish dominance in the canopy. Species replacement is when fires occur in sufficient frequency to interrupt species dominance relay. Gap-Phase Self-Replacement is the least common and so far has only been documented in Western Canada. It is a self replacement of the surviving species into the canopy gaps after a fire kills another species. The particular pathway taken after a fire disturbance depends on how the landscape is able to support trees as well as fire frequency.[56] Fire frequency has a large role in shaping the original inception of the lower forest line of the lichen woodland taiga.

It has been hypothesized by Serge Payette that the Spruce-Moss forest ecosystem was changed into the lichen woodland biome due to the initiation of two compounded strong disturbances: large fire and the appearance and attack of the spruce budworm.[57] The spruce budworm is a deadly insect to the spruce populations in the southern regions of the taiga. J.P. Jasinski confirmed this theory five years later stating “Their [lichen woodlands] persistence, along with their previous moss forest histories and current occurrence adjacent to closed moss forests, indicate that they are an alternative stable state to the spruce–moss forests”.[58]

Taiga ecoregions

East Siberian taiga Russia
Iceland boreal birch forests and alpine tundra Iceland
Kamchatka–Kurile meadows and sparse forests Russia
Kamchatka–Kurile taiga Russia
Northeast Siberian taiga Russia
Okhotsk–Manchurian taiga Russia
Sakhalin Island taiga Russia
Scandinavian and Russian taiga Finland, Norway, Russia, Sweden
Trans-Baikal conifer forests Mongolia, Russia
Urals montane tundra and taiga Russia
West Siberian taiga Russia
Alaska Peninsula montane taiga United States
Central Canadian Shield forests Canada
Cook Inlet taiga United States
Copper Plateau taiga United States
Eastern Canadian forests Canada
Eastern Canadian Shield taiga Canada
Interior Alaska–Yukon lowland taiga Canada, United States
Mid-Continental Canadian forests Canada
Midwestern Canadian Shield forests Canada
Muskwa–Slave Lake forests Canada
Newfoundland Highland forests Canada
Northern Canadian Shield taiga Canada
Northern Cordillera forests Canada
Northwest Territories taiga Canada
South Avalon–Burin oceanic barrens Canada, France (Saint Pierre and Miquelon)
Southern Appalachian spruce–fir forest United States
Southern Hudson Bay taiga Canada
Yukon Interior dry forests Canada

See also

References

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  53. ^ a b Nilsson, 421.
  54. ^ Johnson, 212.
  55. ^ a b Johnson, 200
  56. ^ Kurkowski, 1912.
  57. ^ Payette, 289.
  58. ^ Jasinski, 561.
General references
  • Arno, S. F. & Hammerly, R. P. 1984. Timberline. Mountain and Arctic Forest Frontiers. The Mountaineers, Seattle. ISBN 0-89886-085-7
  • Arno, S. F., Worral, J., & Carlson, C. E. (1995). Larix lyallii: Colonist of tree line and talus sites. Pp. 72–78 in Schmidt, W. C. & McDonald, K. J., eds., Ecology and Management of Larix Forests: A Look Ahead. USDA Forest Service General Technical Report GTR-INT-319.
  • Nilsson, M.C. "Understory vegetation as a forest ecosystem driver, evidence from the northern Swedish boreal forest." Frontiers in Ecology and the Environment. 3.8 (2005): 421-428.
  • Kurkowski, Thomas. "Relative Importance of Different Secondary Successional Pathways in an Alaskan Boreal Forest." Canadian Journal of Forest Research. 38. (2008): 1911-1923.
  • Payette, Serge. "Origin of the lichen woodland at its southern range limit in eastern Canada: the catastrophic impact of insect defoliators and fire on the spruce-moss forest." Canadian journal of forest research. 30.2 (2000): 288-305.
  • Johnson, E.A. "Vegetation Organization and Dynamics of Lichen Woodland Communities in the Northwest Territories." Ecology. 62.1 (1981): 200-215.
  • Jasinski, J.P. "The Creation of Alternative Stable States in Southern Boreal Forest: Quebec, Canada." Ecological Monographs. 75.4 (2005): 561-583.

Further reading