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''Populus tremuloides'' is a
deciduous In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...
tree native to cooler areas of
North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...
, one of several species referred to by the common name
aspen Aspen is a common name for certain tree species in the Populus sect. Populus, of the ''Populus'' (poplar) genus. Species These species are called aspens: * ''Populus adenopoda'' – Chinese aspen (China, south of ''P. tremula'') * ''Populus da ...
. It is commonly called quaking aspen,Quaking Aspen
by the Bryce Canyon
National Park Service The National Park Service (NPS) is an List of federal agencies in the United States, agency of the Federal government of the United States, United States federal government, within the US Department of the Interior. The service manages all List ...
trembling aspen, American aspen, mountain or golden aspen, trembling poplar, white poplar, and popple, as well as others. The trees have tall trunks, up to tall, with smooth pale bark, scarred with black. The glossy green leaves, dull beneath, become golden to yellow, rarely red, in autumn. The species often propagates through its roots to form large clonal groves originating from a shared root system. These roots are not
rhizome In botany and dendrology, a rhizome ( ) is a modified subterranean plant stem that sends out roots and Shoot (botany), shoots from its Node (botany), nodes. Rhizomes are also called creeping rootstalks or just rootstalks. Rhizomes develop from ...
s, as new growth develops from adventitious buds on the parent root system (the ortet). ''Populus tremuloides'' is the most widely distributed tree in North America, being found from Canada to central Mexico. It is the defining species of the
aspen parkland Aspen parkland refers to a very large area of ecotone, transitional biome between prairie and boreal forest in two sections, namely the Peace River Country of northwestern Alberta crossing the border into British Columbia, and a much larger area ...
biome A biome () is a distinct geographical region with specific climate, vegetation, and animal life. It consists of a biological community that has formed in response to its physical environment and regional climate. In 1935, Tansley added the ...
in the
Prairie Provinces The Canadian Prairies (usually referred to as simply the Prairies in Canada) is a region in Western Canada. It includes the Canadian portion of the Great Plains and the Prairie provinces, namely Alberta, Saskatchewan, and Manitoba. These provin ...
of Canada and extreme northwest
Minnesota Minnesota ( ) is a U.S. state, state in the Upper Midwestern region of the United States. It is bordered by the Canadian provinces of Manitoba and Ontario to the north and east and by the U.S. states of Wisconsin to the east, Iowa to the so ...
.


Description

The quaking aspen is a tall, fast-growing tree, usually at maturity, with a trunk in diameter; records are in height and in diameter. The
bark Bark may refer to: Common meanings * Bark (botany), an outer layer of a woody plant such as a tree or stick * Bark (sound), a vocalization of some animals (which is commonly the dog) Arts and entertainment * ''Bark'' (Jefferson Airplane album), ...
is relatively smooth, whitish (light green when young), and is marked by thick black horizontal scars and prominent black knots. Parallel vertical scars are tell-tale signs of elk, which strip off aspen bark with their front teeth. The
leaves A leaf (: leaves) is a principal appendage of the stem of a vascular plant, usually borne laterally above ground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, stem, ...
on mature trees are nearly round, in diameter with small rounded teeth, and a long flattened petiole. The leaves are green above and gray below. Young trees and root sprouts have much larger ( long), nearly triangular leaves. (Some species of ''Populus'' have petioles flattened partially along their length, while the aspens and some other poplars have them flattened from side to side along the entire length of the petiole.) Aspens are
dioecious Dioecy ( ; ; adj. dioecious, ) is a characteristic of certain species that have distinct unisexual individuals, each producing either male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproduction is ...
, with separate male and female clones. The flowers are catkins long, produced in early spring before the leaves. The fruit is a pendulous string of capsules, each capsule containing about ten minute seeds embedded in cottony fluff, which aids wind dispersal of the seeds when they are mature in early summer. Trees as young as 2–3 years old may begin seed production, but significant output starts at 10 years of age. Best seed production is obtained between the ages of 50 and 70 years. Quaking aspen grows more slowly in the dry conditions of western North America than it does in the more humid east and also lives longer—ages of 80–100 years are typical, with some individuals living 200 years; the root system can live much longer. In the east, stands decay faster, sometimes in 60 years or less depending on the region.


Name

The quaking or trembling of the leaves that is referred to in the common names is due to the flexible flattened petioles. The specific epithet, ''tremuloides'', evokes this trembling behavior and can be literally translated as "like ''(Populus) tremula''", the European trembling aspen.


Distribution

Quaking aspen occurs across
Canada Canada is a country in North America. Its Provinces and territories of Canada, ten provinces and three territories extend from the Atlantic Ocean to the Pacific Ocean and northward into the Arctic Ocean, making it the world's List of coun ...
in all provinces and territories, with the possible exception of regions of
Nunavut Nunavut is the largest and northernmost Provinces and territories of Canada#Territories, territory of Canada. It was separated officially from the Northwest Territories on April 1, 1999, via the ''Nunavut Act'' and the Nunavut Land Claims Agr ...
north of the James Bay islands. In the
United States The United States of America (USA), also known as the United States (U.S.) or America, is a country primarily located in North America. It is a federal republic of 50 U.S. state, states and a federal capital district, Washington, D.C. The 48 ...
, it can be found as far north as the northern foothills of the
Brooks Range The Brooks Range (Gwich’in language, Gwich'in: ''Gwazhał'') is a mountain range in far northern North America stretching some from west to east across northern Alaska into Canada's Yukon Territory. Reaching a peak elevation of on Mount Isto, ...
in
Alaska Alaska ( ) is a non-contiguous U.S. state on the northwest extremity of North America. Part of the Western United States region, it is one of the two non-contiguous U.S. states, alongside Hawaii. Alaska is also considered to be the north ...
, where road margins and gravel pads provide islands of well-drained habitat in a region where soils are often waterlogged due to underlying
permafrost Permafrost () is soil or underwater sediment which continuously remains below for two years or more; the oldest permafrost has been continuously frozen for around 700,000 years. Whilst the shallowest permafrost has a vertical extent of below ...
. It occurs at low elevations as far south as northern
Nebraska Nebraska ( ) is a landlocked U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders South Dakota to the north; Iowa to the east and Missouri to the southeast, both across the Missouri River; Ka ...
and central
Indiana Indiana ( ) is a U.S. state, state in the Midwestern United States, Midwestern region of the United States. It borders Lake Michigan to the northwest, Michigan to the north and northeast, Ohio to the east, the Ohio River and Kentucky to the s ...
. In the Western United States, this tree rarely survives at elevations lower than due to hot summers experienced below that elevation, and is generally found at . It grows at high elevations as far south as
Guanajuato Guanajuato, officially the Free and Sovereign State of Guanajuato, is one of the 32 states that make up the Political divisions of Mexico, Federal Entities of Mexico. It is divided into Municipalities of Guanajuato, 46 municipalities and its cap ...
, Mexico. It grows in isolated areas in northeastern Mexico as well as
Baja California Baja California, officially the Free and Sovereign State of Baja California, is a state in Mexico. It is the northwesternmost of the 32 federal entities of Mexico. Before becoming a state in 1952, the area was known as the North Territory of B ...
,
Jalisco Jalisco, officially the Free and Sovereign State of Jalisco, is one of the 31 states which, along with Mexico City, comprise the 32 Political divisions of Mexico, Federal Entities of Mexico. It is located in western Mexico and is bordered by s ...
, the
State of Mexico The State of Mexico, officially just Mexico, is one of the 32 federal entities of the United Mexican States. Colloquially known as Edomex (from , the abbreviation of , and ), to distinguish it from the name of the whole country, it is the mo ...
,
Michoacán Michoacán, formally Michoacán de Ocampo, officially the Free and Sovereign State of Michoacán de Ocampo, is one of the 31 states which, together with Mexico City, compose the Political divisions of Mexico, Federal Entities of Mexico. The stat ...
,
Sinaloa Sinaloa (), officially the (), is one of the 31 states which, along with Mexico City, compose the Federal Entities of Mexico. It is divided into 18 municipalities, and its capital city is Culiacán Rosales. It is located in northwest Mexic ...
,
Sonora Sonora (), officially Estado Libre y Soberano de Sonora (), is one of the 31 states which, along with Mexico City, comprise the Administrative divisions of Mexico, Federal Entities of Mexico. The state is divided into Municipalities of Sonora, 72 ...
, and
Veracruz Veracruz, formally Veracruz de Ignacio de la Llave, officially the Free and Sovereign State of Veracruz de Ignacio de la Llave, is one of the 31 states which, along with Mexico City, comprise the 32 Political divisions of Mexico, Federal Entit ...
. Quaking aspen grows in a wide variety of climatic conditions. January and July average temperatures range from and in the Alaska Interior to and in
Fort Wayne, Indiana Fort Wayne is a city in Allen County, Indiana, United States, and its county seat. Located in northeastern Indiana, the city is west of the Ohio border and south of the Michigan border. The city's population was 263,886 at the 2020 census ...
. Average annual precipitation ranges from in Gander, Newfoundland and Labrador to as little as in the Alaska Interior. The southern limit of the species' range roughly follows the mean July isotherm. In the sagebrush steppe, aspens occur with chokecherry, serviceberry, and hawthorn, forming a habitable haven for animal life. Shrub-like dwarf clones exist in marginal environments too cold and dry to be hospitable to full-size trees, for example at the species' upper elevation limits in the White Mountains.


Ecology

Quaking aspen propagates itself primarily through root sprouts, and extensive
clonal colonies A clonal colony or genet is a group of genetically identical individuals, such as plants, fungi, or bacteria, that have grown in a given location, all originating vegetatively, not sexually, from a single ancestor. In plants, an individual in ...
are common. Each colony is its own clone, and all trees in the clone have identical characteristics and share a single root structure. A clone may turn color earlier or later in the fall than its neighboring aspen clones. Fall colors are usually bright tones of yellow; in some areas, red blushes may be occasionally seen. As all trees in a given clonal colony are considered part of the same organism, one clonal colony, named Pando, is considered the heaviest and oldest living organism on the planet. Pando spans across 43 hectares and weighs six million kilograms. Most scientists agree that the Pando seed set down sometime between 8,000 and 12,000 years ago when climate currents in the region shifted at the end of the last ice age. Aspens do produce seeds, but seldom grow from them. Pollination is inhibited by the fact that aspens are either male or female, and large stands are usually all clones of the same sex. Even if pollinated, the small seeds (three million per pound) are only viable a short time as they lack a stored food source or a protective coating.Ewing, Susan. The Great Alaska Nature Factbook. Portland: Alaska Northwest Books, 1996. The buds and bark supply food for
snowshoe hare The snowshoe hare (''Lepus americanus''), also called the varying hare or snowshoe rabbit, is a species of hare found in North America. It has the name "snowshoe" because of the large size of its hind feet. The animal's feet prevent it from sink ...
s,
moose The moose (: 'moose'; used in North America) or elk (: 'elk' or 'elks'; used in Eurasia) (''Alces alces'') is the world's tallest, largest and heaviest extant species of deer and the only species in the genus ''Alces''. It is also the tal ...
, black bears,
cottontail rabbit Cottontail rabbits are in the ''Sylvilagus'' genus, which is in the family Leporidae. They are found in the Americas. Most ''Sylvilagus'' species have stub tails with white undersides that show when they retreat, giving them their characterist ...
s,
porcupine Porcupines are large rodents with coats of sharp Spine (zoology), spines, or quills, that protect them against predation. The term covers two Family (biology), families of animals: the Old World porcupines of the family Hystricidae, and the New ...
s, deer,
grouse Grouse are a group of birds from the order (biology), order Galliformes, in the family (biology), family Phasianidae. Grouse are presently assigned to the Tribe (biology), tribe Tetraonini (formerly the subfamily Tetraoninae and the family Tetr ...
, and
mountain beaver The mountain beaver (''Aplodontia rufa'')Other names include boomer, mountain boomer, ground bear, giant mole, gehalis, sewellel, suwellel, showhurll, showtl, and showte, as well as a number of other Native American terms. "Mountain beaver" is a ...
s. The shoots are eaten by
sheep Sheep (: sheep) or domestic sheep (''Ovis aries'') are a domesticated, ruminant mammal typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus '' Ovis'', in everyday usage it almost always refers to d ...
,
goat The goat or domestic goat (''Capra hircus'') is a species of Caprinae, goat-antelope that is mostly kept as livestock. It was domesticated from the wild goat (''C. aegagrus'') of Southwest Asia and Eastern Europe. The goat is a member of the ...
s, and
cattle Cattle (''Bos taurus'') are large, domesticated, bovid ungulates widely kept as livestock. They are prominent modern members of the subfamily Bovinae and the most widespread species of the genus '' Bos''. Mature female cattle are calle ...
. Sheep and goats also browse the foliage, as do game animals including elk. Grouse and quail especially eat the buds in winter. Mammals such as
beaver Beavers (genus ''Castor'') are large, semiaquatic rodents of the Northern Hemisphere. There are two existing species: the North American beaver (''Castor canadensis'') and the Eurasian beaver (''C. fiber''). Beavers are the second-large ...
s and rabbits eat the bark, foliage, and buds. Beavers also store aspen logs for winter food. Other animals nest in aspen groves. The leaves of the quaking aspen and other species in the genus ''Populus'' serve as food for
caterpillar Caterpillars ( ) are the larval stage of members of the order Lepidoptera (the insect order comprising butterflies and moths). As with most common names, the application of the word is arbitrary, since the larvae of sawflies (suborder ...
s of various moths and butterflies. Quaking aspen trees also serve as hosts to certain damaging insects such as the large aspen tortrix.


Dieback

Increased mortality in trembling aspen stands have been reported since the early 1990s across North America. As this accelerated in 2004, a debate over causes began. This increased dieback has been linked to multiple stressors, such as defoliation by the forest tent caterpillar (''Malacosoma disstria''), wood-boring beetles such as the poplar borer (''Saperda calcarata'') and the bronze poplar borer (''Agrilus liragus''), and fungal disturbances such as those by the Cytospora canker (''Valsa sordida''). Many areas of the Western US have experienced increased diebacks which are often attributed to ungulate grazing and
wildfire suppression Wildfire suppression is a range of firefighting tactics used to suppress wildfires. Firefighting efforts depend on many factors such as the available fuel, the local atmospheric conditions, the features of the terrain, and the size of the wildfir ...
. At high elevations where grasses can be rare,
ungulate Ungulates ( ) are members of the diverse clade Euungulata ("true ungulates"), which primarily consists of large mammals with Hoof, hooves. Once part of the clade "Ungulata" along with the clade Paenungulata, "Ungulata" has since been determined ...
s can browse young aspen sprouts and prevent those young trees from reaching maturity. As a result, some aspen groves close to cattle or other grazing animals, such as deer or elk, have very few young trees and can be invaded by
conifers Conifers () are a group of cone-bearing seed plants, a subset of gymnosperms. Scientifically, they make up the division Pinophyta (), also known as Coniferophyta () or Coniferae. The division contains a single extant class, Pinopsida. All e ...
, which are not typically browsed. Another possible deterrent to aspen regeneration is widespread wildfire suppression. Aspens are vigorous
resprouter Resprouters are plant species that are Fire adaptations, adapted to survive fire by the activation of dormant vegetative buds to produce regrowth. Plants may resprout from a bud bank that can be located in different places, including in the trun ...
s and even though the above-ground portion of the organism may die in a wild-fire, the roots, which are often protected from lethal temperatures during a fire, will sprout new trees soon after a fire. Disturbances such as fires seem to be a necessary ecological event in order for aspens to compete with conifers, which tend to replace aspens over long, disturbance-free intervals. The current dieback in the American West may have roots in the strict fire suppression policy in the United States. On the other hand, the widespread decimation of conifer forests by the mountain pine beetle may provide increased opportunities for aspen groves to proliferate under the right conditions. Increased mortality has also been linked in turn to climate change. Thaw-freeze events and light snowfall in late winter as a result of increased temperatures has led to increased dieback in Southern and Western Canada. Furthermore, climate records show that historically, most periods of aspen decline have been paired with periods of severe drought, which has worsened in recent years due to a changing climate. Many stands of aspen that have been affected by climate change in recent years have poor regeneration potential, leading to concerns of widespread loss of aspen cover in the future. Because of vegetative regeneration by aspen, where an entire group of trees are essentially clones, there is a concern that something that hits one will eventually kill all of the trees, presuming they share the same vulnerability. A conference was held in
Utah Utah is a landlocked state in the Mountain states, Mountain West subregion of the Western United States. It is one of the Four Corners states, sharing a border with Arizona, Colorado, and New Mexico. It also borders Wyoming to the northea ...
in September 2006 to share notes and consider investigative methodology.


Uses

Aspen bark contains a substance that was extracted by indigenous North Americans and European settlers of the western U.S. as a
quinine Quinine is a medication used to treat malaria and babesiosis. This includes the treatment of malaria due to ''Plasmodium falciparum'' that is resistant to chloroquine when artesunate is not available. While sometimes used for nocturnal leg ...
substitute. Like other poplars, aspens make poor fuel wood, as they dry slowly, rot quickly, and do not give off much heat. Yet they are still widely used in campgrounds because they are cheap and plentiful and not widely used in building lumber. Pioneers in the North American west used them to create log cabins and dugouts, though they were not the preferred species. Aspen wood is used for pulp products (its main application in Canada) such as books, newsprint, and fine printing paper. It is especially good for panel products such as
oriented strand board Oriented strand board (OSB) is a type of engineered wood, formed by adding adhesives and then compressing layers of wood strands (flakes) in specific orientations. It was invented by Armin Elmendorf in California in 1963. OSB may have a rough and ...
and waferboard. It is light in weight and is used for furniture, boxes and crates, core stock in plywood, and wall panels.


Culture and Cultivation

The quaking aspen is the state tree of Utah. The
cultivar A cultivar is a kind of Horticulture, cultivated plant that people have selected for desired phenotypic trait, traits and which retains those traits when Plant propagation, propagated. Methods used to propagate cultivars include division, root a ...
'Prairie Gold' was introduced to the plant trade by the Nebraska Statewide Arboretum and a local nursery with a claim that it is better adapted the warmer conditions and lower humidity of lower elevations. It is rated as hardy only to USDA zone 4. Typically, quaking aspens in cultivation are hardy in USDA zones 1 to 6. They require good drainage and moist or occasionally wet soils. With good drainage they adapt to clay, silt, sand, or shallowly rocky soil conditions.


See also

* Pando, an exceptionally large clonal colony of ''P. tremuloides'' determined to be a single male tree * Largest organisms


References


External links


US Forest Service Fire Effects Information System: ''Populus tremuloides''
* ttp://www.plantmaps.com/nrm/populus-tremuloides-quaking-aspen-native-range-map.php Interactive Distribution Map for ''Populus tremuloides'' *Farrar, John Laird. ''Trees In Canada''. Fitzhenry and Whiteside, 1995 *Hickman, James C., ed.
The Jepson Manual: Higher Plants of California
', . University of California Press, 1993. * {{Authority control tremuloides Trees of Northern America Taxa named by André Michaux Plants used in traditional Native American medicine Flora of Western Canada Flora of Eastern Canada Flora of the Northwestern United States Flora of the North-Central United States Flora of the Northeastern United States Flora of the Southwestern United States Flora of the South-Central United States Flora of Northeastern Mexico Flora of Baja California Flora of Delaware Flora of Jalisco Flora of Maryland Flora of the State of Mexico Flora of Michoacán Flora of North Carolina Flora of Sinaloa Flora of Sonora Flora of Veracruz Flora of Virginia Flora of Alaska Flora of the Northwest Territories Flora of Nunavut Flora of Yukon