ecology Ecology () is the study of the relationships between living organisms, including humans, and their physical environment. Ecology considers organisms at the individual, population, community, ecosystem, and biosphere level. Ecology overl ...

, a disturbance is a temporary change in environmental conditions that causes a pronounced change in an

ecosystem An ecosystem (or ecological system) consists of all the organisms and the physical environment with which they interact. These biotic and abiotic components are linked together through nutrient cycles and energy flows. Energy enters the syste ...

. Disturbances often act quickly and with great effect, to alter the physical structure or arrangement of biotic and

abiotic In biology and ecology, abiotic components or abiotic factors are non-living chemical and physical parts of the environment that affect living organisms and the functioning of ecosystems. Abiotic factors and the phenomena associated with them under ...

elements. A disturbance can also occur over a long period of time and can impact the

biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') l ...

within an ecosystem. Major ecological disturbances may include

fire Fire is the rapid oxidation of a material (the fuel) in the exothermic chemical process of combustion, releasing heat, light, and various reaction products. At a certain point in the combustion reaction, called the ignition point, flames a ...

flood A flood is an overflow of water ( or rarely other fluids) that submerges land that is usually dry. In the sense of "flowing water", the word may also be applied to the inflow of the tide. Floods are an area of study of the discipline hydrol ...

ing,

storm A storm is any disturbed state of the natural environment or the atmosphere of an astronomical body. It may be marked by significant disruptions to normal conditions such as strong wind, tornadoes, hail, thunder and lightning (a thunderstorm), ...

insect Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body ( head, thorax and abdomen), three ...

outbreak In epidemiology, an outbreak is a sudden increase in occurrences of a disease when cases are in excess of normal expectancy for the location or season. It may affect a small and localized group or impact upon thousands of people across an entire ...

s and trampling.

Earthquake An earthquake (also known as a quake, tremor or temblor) is the shaking of the surface of the Earth resulting from a sudden release of energy in the Earth's lithosphere that creates seismic waves. Earthquakes can range in intensity, fr ...

s, various

types of volcanic eruptions Several types of volcanic eruptions—during which lava, tephra (ash, lapilli, volcanic bombs and volcanic blocks), and assorted gases are expelled from a volcanic vent or fissure—have been distinguished by volcanologists. These are oft ...

tsunami A tsunami ( ; from ja, 津波, lit=harbour wave, ) is a series of waves in a water body caused by the displacement of a large volume of water, generally in an ocean or a large lake. Earthquakes, volcanic eruptions and other underwater exp ...

firestorm A firestorm is a conflagration which attains such intensity that it creates and sustains its own wind system. It is most commonly a natural phenomenon, created during some of the largest bushfires and wildfires. Although the term has been used ...

s, impact events,

climate change In common usage, climate change describes global warming—the ongoing increase in global average temperature—and its effects on Earth's climate system. Climate change in a broader sense also includes previous long-term changes to ...

, and the devastating effects of

human impact on the environment Human impact on the environment (or anthropogenic impact) refers to changes to biophysical environments and to ecosystems, biodiversity, and natural resources caused directly or indirectly by humans. Modifying the environment to fit the need ...

( anthropogenic disturbances) such as clearcutting, forest clearing and the introduction of invasive species can be considered major disturbances. Not only invasive species can have a profound effect on an ecosystem, but also naturally occurring species can cause disturbance by their behavior. Disturbance forces can have profound immediate effects on ecosystems and can, accordingly, greatly alter the

natural community In ecology, a community is a group or association of populations of two or more different species occupying the same geographical area at the same time, also known as a biocoenosis, biotic community, biological community, ecological communit ...

. Because of these and the impacts on populations, disturbance determines the future shifts in dominance, various species successively becoming dominant as their life history characteristics, and associated life-forms, are exhibited over time.

Criteria

Conditions under which natural disturbances occur are influenced mainly by

climate Climate is the long-term weather pattern in an area, typically averaged over 30 years. More rigorously, it is the mean and variability of meteorological variables over a time spanning from months to millions of years. Some of the meteorologi ...

, weather, and location. Natural fire disturbances for example occur more often in areas with a higher incidence of lightning and flammable biomass, such as

longleaf pine ecosystem The longleaf pine ecosystem is a climax savanna habitat found within the Southern United States; it includes many rare plant and animal species, and is one of the most biodiverse in North America. Once the largest ecosystem in North America, it n ...

s in the southeastern United States. Conditions often occur as part of a cycle and disturbances may be periodic. Other disturbances, such as those caused by humans, invasive species or impact events, can occur anywhere and are not necessarily cyclic. Extinction vortices may result in multiple disturbances or a greater frequency of a single disturbance. Immediately after a disturbance there is a pulse of recruitment or regrowth under conditions of little

competition Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, indiv ...

for space or other resources. After the initial pulse, recruitment slows since once an individual plant is established it is very difficult to displace. Due to the varying forms of disturbance this directly impacts the organisms which will exploit the disturbance and create diversity within an ecosystem. Because scale-dependent relationships are ubiquitous in ecology, the spatial scale modulates the effect of disturbance on natural communities. For example,

seed dispersal In Spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vectors ...

and

herbivory A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpart ...

may decrease with distance from the edge of a burn. Consequently, plant communities in the interior areas of large fires respond differently than those in smaller fires. Although disturbance types have varied on ecosystems, spatial scale likely influences ecological interactions and community recovery from all cases because organisms differ in dispersal and movement capabilities.

Cyclic disturbance

Often, when disturbances occur naturally, they provide conditions that favor the success of different species over pre-disturbance organisms. This can be attributed to physical changes in the biotic and abiotic conditions of an ecosystem. Because of this, a disturbance force can change an ecosystem for significantly longer than the period over which the immediate effects persist. With the passage of time following a disturbance, shifts in dominance may occur with ephemeral herbaceous life-forms progressively becoming over topped by taller perennials herbs, shrubs and trees. However, in the absence of further disturbance forces, many ecosystems trend back toward pre-disturbance conditions. Long lived species and those that can regenerate in the presence of their own adults finally become dominant. Such alteration, accompanied by changes in the abundance of different species over time, is called ecological succession. Succession often leads to conditions that will once again predispose an ecosystem to disturbance. Pine forests in western North America provide a good example of such a cycle involving insect outbreaks. The

mountain pine beetle The mountain pine beetle (''Dendroctonus ponderosae'') is a species of bark beetle native to the forests of western North America from Mexico to central British Columbia. It has a hard black exoskeleton, and measures approximately , about the siz ...

(''Dendroctonus ponderosae'') plays an important role in limiting pine trees like

lodgepole pine ''Pinus contorta'', with the common names lodgepole pine and shore pine, and also known as twisted pine, and contorta pine, is a common tree in western North America. It is common near the ocean shore and in dry montane forests to the subalpin ...

in forests of western North America. In 2004 the beetles affected more than 90,000 square kilometres. The beetles exist in

endemic Endemism is the state of a species being found in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also found else ...

and

epidemic An epidemic (from Greek ἐπί ''epi'' "upon or above" and δῆμος ''demos'' "people") is the rapid spread of disease to a large number of patients among a given population within an area in a short period of time. Epidemics of infectious ...

phases. During epidemic phases swarms of beetles kill large numbers of old pines. This mortality creates openings in the forest for new vegetation. Spruce, fir, and younger pines, which are unaffected by the beetles, thrive in canopy openings. Eventually pines grow into the canopy and replace those lost. Younger pines are often able to ward off beetle attacks but, as they grow older, pines become less vigorous and more susceptible to infestation. This cycle of death and re-growth creates a temporal mosaic of pines in the forest. Similar cycles occur in association with other disturbances such as fire and windstorms. When multiple disturbance events affect the same location in quick succession, this often results in a "compound disturbance," an event which, due to the combination of forces, creates a new situation which is more than the sum of its parts. For example, windstorms followed by fire can create fire temperatures and durations that are not expected in even severe wildfires, and may have surprising effects on post-fire succession. Environmental stresses can be described as pressure on the environment, with compounding variables such as extreme temperature or precipitation changes—which all play a role in the diversity and succession of an ecosystem. With environmental moderation, diversity increases because of the intermediate-disturbance effect, decreases because of the competitive-exclusion effect, increases because of the prevention of competitive exclusion by moderate predation, and decreases because of the local extinction of prey by severe predation. A reduction in recruitment density reduces the importance of competition for a given level of environmental stress.

Species adapted to disturbance (eurytopy)

A disturbance may change a forest significantly. Afterwards, the forest floor is often littered with dead material. This decaying matter and abundant sunlight promote an abundance of new growth. In the case of forest fires a portion of the nutrients previously held in plant biomass is returned quickly to the soil as biomass burns. Many plants and animals benefit from disturbance conditions.Pringle, L. 1979. Natural Fire: Its Ecology in Forests. William Morrow and Company, New York. 27-29. Some species are particularly suited for exploiting recently disturbed sites. Vegetation with the potential for rapid growth can quickly take advantage of the lack of competition. In the northeastern United States, shade-intolerant trees (trees stenotopic to shade) like pin cherry and

aspen Aspen is a common name for certain tree species; some, but not all, are classified by botanists in the section ''Populus'', of the '' Populus'' genus. Species These species are called aspens: *'' Populus adenopoda'' – Chinese aspen (Chin ...

quickly fill in forest gaps created by fire or windstorm (or human disturbance). Silver maple and eastern sycamore are similarly well adapted to floodplains. They are highly tolerant of standing water and will frequently dominate floodplains where other species are periodically wiped out. When a tree is blown over, gaps typically are filled with small herbaceous

seedling A seedling is a young sporophyte developing out of a plant embryo from a seed. Seedling development starts with germination of the seed. A typical young seedling consists of three main parts: the radicle (embryonic root), the hypocotyl (embryo ...

s but, this is not always the case; shoots from the fallen tree can develop and take over the gap. The sprouting ability can have major impacts on the plant population, plant populations that typically would have exploited the tree fall gap get over run and can not compete against the shoots of the fallen tree. Species adaptation to disturbances is species specific but how each organism adapts affects all the species around them. Another species well adapted to a particular disturbance is the

Jack pine Jack pine (''Pinus banksiana'') is an eastern North American pine. Its native range in Canada is east of the Rocky Mountains from the Mackenzie River in the Northwest Territories to Cape Breton Island in Nova Scotia, and the north-central and ...

in boreal forests exposed to crown fires. They, as well as some other pine species, have specialized

serotinous Serotiny in botany simply means 'following' or 'later'. In the case of serotinous flowers, it means flowers which grow following the growth of leaves, or even more simply, flowering later in the season than is customary with allied species. Havi ...

cones that only open and disperse seeds with sufficient heat generated by fire. As a result, this species often dominates in areas where competition has been reduced by fire. Species that are well adapted for exploiting disturbance sites are referred to as pioneers or early successional species. These shade-intolerant species are able to photosynthesize at high rates and as a result grow quickly. Their fast growth is usually balanced by short life spans. Furthermore, although these species often dominate immediately following a disturbance, they are unable to compete with shade-tolerant species later on and replaced by these species through succession. However these shifts may not reflect the progressive entry to the community of the taller long-lived forms, but instead, the gradual emergence and dominance of species that may have been present, but inconspicuous directly after the disturbance. Disturbances have also been shown to be important facilitators of non-native plant invasions. While plants must deal directly with disturbances, many animals are not as immediately affected by them. Most can successfully evade fires, and many thrive afterwards on abundant new growth on the forest floor. New conditions support a wider variety of plants, often rich in nutrients compared to pre-disturbance vegetation. The plants in turn support a variety of wildlife, temporarily increasing biological diversity in the forest.

Importance

Biological diversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic ('' genetic variability''), species ('' species diversity''), and ecosystem ('' ecosystem diversity'') ...

is dependent on natural disturbance. The success of a wide range of species from all taxonomic groups is closely tied to natural disturbance events such as fire, flooding, and windstorm. As an example, many shade-intolerant plant species rely on disturbances for successful establishment and to limit competition. Without this perpetual thinning, diversity of forest flora can decline, affecting animals dependent on those plants as well. A good example of this role of disturbance is in

ponderosa pine ''Pinus ponderosa'', commonly known as the ponderosa pine, bull pine, blackjack pine, western yellow-pine, or filipinus pine is a very large pine tree species of variable habitat native to mountainous regions of western North America. It is the ...

(''Pinus ponderosa'') forests in the western United States, where surface fires frequently thin existing vegetation allowing for new growth. If fire is suppressed, douglas fir (''Pesudotsuga menziesii''), a shade tolerant species, eventually replaces the pines. Douglas firs, having dense crowns, severely limit the amount of sunlight reaching the forest floor. Without sufficient light new growth is severely limited. As the diversity of surface plants decreases, animal species that rely on them diminish as well. Fire, in this case, is important not only to the species directly affected but also to many other organisms whose survival depends on those key plants. Diversity is low in harsh environments because of the intolerance of all but opportunistic and highly resistant species to such conditions. The interplay between disturbance and these biological processes seems to account for a major portion of the organization and spatial patterning of natural communities. Disturbance variability and species diversity are heavily linked, and as a result require adaptations that help increase plant fitness necessary for survival.

References

External links