Species distribution —or species dispersion — is the manner in which a biological
taxon
In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
is spatially arranged. The geographic limits of a particular taxon's distribution is its range, often represented as shaded areas on a map. Patterns of distribution change depending on the scale at which they are viewed, from the arrangement of individuals within a small family unit, to patterns within a population, or the distribution of the entire
species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
as a whole (range). Species distribution is not to be confused with
dispersal, which is the movement of individuals away from their
region of origin
In geography, regions, otherwise referred to as zones, lands or territories, are areas that are broadly divided by physical characteristics (physical geography), human impact characteristics (human geography), and the interaction of humanity and t ...
or from a population center of high
density
Density (volumetric mass density or specific mass) is the substance's mass per unit of volume. The symbol most often used for density is ''ρ'' (the lower case Greek letter rho), although the Latin letter ''D'' can also be used. Mathematical ...
.
Range
In
biology
Biology is the scientific study of life. It is a natural science with a broad scope but has several unifying themes that tie it together as a single, coherent field. For instance, all organisms are made up of cells that process hereditary i ...
, the range of a
species
In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
is the geographical area within which that species can be found. Within that range, distribution is the general structure of the species
population
Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction using a ...
, while dispersion is the variation in its
population density
Population density (in agriculture: standing stock or plant density) is a measurement of population per unit land area. It is mostly applied to humans, but sometimes to other living organisms too. It is a key geographical term.Matt RosenberPopul ...
.
Range is often described with the following qualities:
* Sometimes a distinction is made between a species' natural,
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 elsew ...
,
indigenous
Indigenous may refer to:
*Indigenous peoples
*Indigenous (ecology), presence in a region as the result of only natural processes, with no human intervention
*Indigenous (band), an American blues-rock band
*Indigenous (horse), a Hong Kong racehorse ...
, or native range, where it has historically originated and lived, and the range where a species has more recently established itself. Many terms are used to describe the new range, such as non-native, naturalized,
introduced, transplanted,
invasive, or colonized range.
''Introduced'' typically means that a species has been transported by humans (intentionally or accidentally) across a major geographical barrier.
* For species found in different regions at different times of year, especially
season
A season is a division of the year based on changes in weather, ecology, and the number of daylight hours in a given region. On Earth, seasons are the result of the axial parallelism of Earth's tilted orbit around the Sun. In temperate and pol ...
s, terms such as summer range and winter range are often employed.
* For species for which only part of their range is used for breeding activity, the terms breeding range and non-breeding range are used.
* For mobile animals, the term natural range is often used, as opposed to areas where it occurs as a
vagrant
Vagrancy is the condition of homelessness without regular employment or income. Vagrants (also known as bums, vagabonds, rogues, tramps or drifters) usually live in poverty and support themselves by begging, scavenging, petty theft, temporar ...
.
* Geographic or temporal qualifiers are often added, such as in ''British range'' or ''pre-1950 range''. The typical geographic ranges could be the latitudinal range and
elevation
The elevation of a geographic location is its height above or below a fixed reference point, most commonly a reference geoid, a mathematical model of the Earth's sea level as an equipotential gravitational surface (see Geodetic datum § Vert ...
al range.
Disjunct distribution
In biology, a taxon with a disjunct distribution is one that has two or more groups that are related but considerably separated from each other geographically. The causes are varied and might demonstrate either the expansion or contraction of a s ...
occurs when two or more areas of the range of a taxon are considerably separated from each other geographically.
Factors affecting species distribution
Distribution patterns may change by
season
A season is a division of the year based on changes in weather, ecology, and the number of daylight hours in a given region. On Earth, seasons are the result of the axial parallelism of Earth's tilted orbit around the Sun. In temperate and pol ...
, distribution by humans, in response to the availability of resources, and other
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 ...
and
biotic factors.
Abiotic
There are three main types of abiotic factors:
# climatic factors consist of sunlight, atmosphere, humidity, temperature, and salinity;
#
edaphic
Edaphology (from Greek , ''edaphos'', "ground",, ''-logia'') is concerned with the influence of soils on living beings, particularly plants.
It is one of two main divisions of soil science, the other being pedology. Edaphology includes the study ...
factors are abiotic factors regarding soil, such as the coarseness of soil, local geology,
soil pH
Soil pH is a measure of the acidity or basicity (alkalinity) of a soil. Soil pH is a key characteristic that can be used to make informative analysis both qualitative and quantitatively regarding soil characteristics. pH is defined as the neg ...
, and aeration; and
# social factors include land use and water availability.
An example of the effects of abiotic factors on species distribution can be seen in drier areas, where most individuals of a species will gather around water sources, forming a clumped distribution.
Researchers from the Arctic Ocean Diversity (ARCOD) project have documented rising numbers of warm-water crustaceans in the seas around Norway's Svalbard Islands. Arcod is part of the Census of Marine Life, a huge 10-year project involving researchers in more than 80 nations that aims to chart the diversity, distribution and abundance of life in the oceans. Marine Life has become largely affected by increasing
effects of global climate change
The effects of climate change impact the physical environment, ecosystems and human societies. The environmental effects of climate change are broad and far-reaching. They affect the water cycle, oceans, sea and land ice (glaciers), sea ...
. This study shows that as the ocean temperatures rise species are beginning to travel into the cold and harsh Arctic waters. Even the snow crab has extended its range 500 km north.
Biotic
Biotic factors such as predation, disease, and inter- and intra-specific competition for resources such as food, water, and mates can also affect how a species is distributed. For example, biotic factors in a quail’s environment would include their prey (insects and seeds), competition from other quail, and their predators, such as the coyote. An advantage of a herd, community, or other clumped distribution allows a population to detect predators earlier, at a greater distance, and potentially mount an effective defense. Due to limited resources, populations may be evenly distributed to minimize competition, as is found in forests, where competition for sunlight produces an even distribution of trees.
Humans
Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, culture, ...
are one of the largest distributors due to the current trends in
globalization
Globalization, or globalisation (Commonwealth English; see spelling differences), is the process of interaction and integration among people, companies, and governments worldwide. The term ''globalization'' first appeared in the early 20t ...
and the expanse of the
transportation
Transport (in British English), or transportation (in American English), is the intentional movement of humans, animals, and goods from one location to another. Modes of transport include air, land (rail and road), water, cable, pipeline, ...
industry. For example, large
tankers
Tanker may refer to:
Transportation
* Tanker, a tank crewman (US)
* Tanker (ship), a ship designed to carry bulk liquids
** Chemical tanker, a type of tanker designed to transport chemicals in bulk
** Oil tanker, also known as a petroleum tank ...
often fill their ballasts with water at one port and empty them in another, causing a wider distribution of aquatic species.
Patterns on large scales
On large scales, the pattern of distribution among individuals in a population is clumped.
Bird wildlife corridors
One common example of bird species' ranges are land mass areas bordering water bodies, such as oceans, rivers, or lakes; they are called a ''coastal strip''. A second example, some species of bird depend on water, usually a river, swamp, etc., or water related forest and live in a ''river corridor''. A separate example of a river corridor would be a river corridor that includes the entire drainage, having the edge of the range delimited by mountains, or higher elevations; the river itself would be a smaller percentage of this entire
wildlife corridor
A wildlife corridor, habitat corridor, or green corridor is an area of habitat connecting wildlife populations separated by human activities or structures (such as roads, development, or logging). This allows an exchange of individuals between ...
, but the corridor is created because of the river.
A further example of a bird wildlife corridor would be a mountain range corridor. In the U.S. of North America, the
Sierra Nevada
The Sierra Nevada () is a mountain range in the Western United States, between the Central Valley of California and the Great Basin. The vast majority of the range lies in the state of California, although the Carson Range spur lies primarily ...
range in the west, and the
Appalachian Mountains
The Appalachian Mountains, often called the Appalachians, (french: Appalaches), are a system of mountains in eastern to northeastern North America. The Appalachians first formed roughly 480 million years ago during the Ordovician Period. They ...
in the east are two examples of this habitat, used in summer, and winter, by separate species, for different reasons.
Bird species in these corridors are connected to a main range for the species (contiguous range) or are in an isolated geographic range and be a
disjunct range. Birds leaving the area, if they
migrate, would leave connected to the main range or have to fly over land not connected to the wildlife corridor; thus, they would be
passage migrant
Passage, The Passage or Le Passage may refer to:
Arts and entertainment Films
* ''Passage'' (2008 film), a documentary about Arctic explorers
* ''Passage'' (2009 film), a short movie about three sisters
* ''The Passage'' (1979 film), starring ...
s over land that they stop on for an intermittent, hit or miss, visit.
Patterns on small scales
On large scales, the pattern of distribution among individuals in a population is clumped. On small scales, the pattern may be clumped, regular, or random.
[
]
Clumped
Clumped distribution, also called aggregated distribution, clumped dispersion or patchiness, is the most common type of dispersion found in nature. In clumped distribution, the distance between neighboring individuals is minimized. This type of distribution is found in environments that are characterized by patchy resources. Animals need certain resources to survive, and when these resources become rare during certain parts of the year animals tend to “clump” together around these crucial resources. Individuals might be clustered together in an area due to social factors such as selfish herd
Herd behavior is the behavior of individuals in a group acting collectively without centralized direction. Herd behavior occurs in animals in herds, packs, bird flocks, fish schools and so on, as well as in humans. Voting, demonstrations, riot ...
s and family groups. Organisms that usually serve as prey form clumped distributions in areas where they can hide and detect predators easily.
Other causes of clumped distributions are the inability of offspring to independently move from their habitat. This is seen in juvenile animals that are immobile and strongly dependent upon parental care. For example, the bald eagle
The bald eagle (''Haliaeetus leucocephalus'') is a bird of prey found in North America. A sea eagle, it has two known subspecies and forms a species pair with the white-tailed eagle (''Haliaeetus albicilla''), which occupies the same niche as ...
's nest of eaglets exhibits a clumped species distribution because all the offspring are in a small subset of a survey area before they learn to fly. Clumped distribution can be beneficial to the individuals in that group. However, in some herbivore cases, such as cows and wildebeests, the vegetation around them can suffer, especially if animals target one plant in particular.
Clumped distribution in species acts as a mechanism against predation as well as an efficient mechanism to trap or corner prey. African wild dogs, ''Lycaon pictus
The African wild dog (''Lycaon pictus''), also called the painted dog or Cape hunting dog, is a wild canine which is a native species to sub-Saharan Africa. It is the largest wild canine in Africa, and the only extant member of the genus '' Lyc ...
'', use the technique of communal hunting to increase their success rate at catching prey. Studies have shown that larger packs of African wild dogs tend to have a greater number of successful kills. A prime example of clumped distribution due to patchy resources is the wildlife in Africa during the dry season; lions, hyenas, giraffes, elephants, gazelles, and many more animals are clumped by small water sources that are present in the severe dry season. It has also been observed that extinct and threatened species are more likely to be clumped in their distribution on a phylogeny. The reasoning behind this is that they share traits that increase vulnerability to extinction because related taxa are often located within the same broad geographical or habitat types where human-induced threats are concentrated. Using recently developed complete phylogenies for mammalian carnivores and primates it has been shown that the majority of instances threatened species are far from randomly distributed among taxa
In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
and phylogenetic
In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
clade
A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
s and display clumped distribution.
A contiguous distribution is one in which individuals are closer together than they would be if they were randomly or evenly distributed, i.e., it is clumped distribution with a single clump.
Regular or uniform
Less common than clumped distribution, uniform distribution, also known as even distribution, is evenly spaced. Uniform distributions are found in populations in which the distance between neighboring individuals is maximized. The need to maximize the space between individuals generally arises from competition for a resource such as moisture or nutrients, or as a result of direct social interactions between individuals within the population, such as territoriality. For example, penguins often exhibit uniform spacing by aggressively defending their territory among their neighbors. The burrows of great gerbil
The great gerbil (''Rhombomys opimus'') is a large rodent found throughout much of Central Asia.
Description
The largest of the gerbils, great gerbils have a head and body length of . Their skulls are distinctive by having two grooves in each i ...
s for example are also regularly distributed, which can be seen on satellite images. Plants also exhibit uniform distributions, like the creosote bushes in the southwestern region of the United States. ''Salvia leucophylla
''Salvia leucophylla'', the San Luis purple sage, purple sage, or gray sage, is an aromatic sage native to the southern coastal mountain ranges of the Californias.
Description
''S. leucophylla'' is an evergreen shrub that grows up to tall and ...
'' is a species in California that naturally grows in uniform spacing. This flower releases chemicals called terpene
Terpenes () are a class of natural products consisting of compounds with the formula (C5H8)n for n > 1. Comprising more than 30,000 compounds, these unsaturated hydrocarbons are produced predominantly by plants, particularly conifers. Terpenes ar ...
s which inhibit the growth of other plants around it and results in uniform distribution. This is an example of allelopathy
Allelopathy is a biological phenomenon by which an organism produces one or more biochemicals that influence the germination, growth, survival, and reproduction of other organisms. These biochemicals are known as allelochemicals and can have be ...
, which is the release of chemicals from plant parts by leaching, root exudation, volatilization, residue decomposition and other processes. Allelopathy can have beneficial, harmful, or neutral effects on surrounding organisms. Some allelochemicals even have selective effects on surrounding organisms; for example, the tree species ''Leucaena leucocephala
''Leucaena leucocephala'' is a small fast-growing Mimosoideae, mimosoid tree native to southern Mexico and northern Central America (Belize and Guatemala) and is now naturalized throughout the tropics including parts of Asia.
Common names inc ...
'' exudes a chemical that inhibits the growth of other plants but not those of its own species, and thus can affect the distribution of specific rival species. Allelopathy usually results in uniform distributions, and its potential to suppress weeds is being researched. Farming and agricultural practices often create uniform distribution in areas where it would not previously exist, for example, orange trees growing in rows on a plantation.
Random
Random distribution, also known as unpredictable spacing, is the least common form of distribution in nature and occurs when the members of a given species are found in environments in which the position of each individual is independent of the other individuals: they neither attract nor repel one another. Random distribution is rare in nature as biotic factors, such as the interactions with neighboring individuals, and abiotic factors, such as climate or soil conditions, generally cause organisms to be either clustered or spread. Random distribution usually occurs in habitats where environmental conditions and resources are consistent. This pattern of dispersion is characterized by the lack of any strong social interactions between species. For example; When dandelion
''Taraxacum'' () is a large genus of flowering plants in the family Asteraceae, which consists of species commonly known as dandelions. The scientific and hobby study of the genus is known as taraxacology. The genus is native to Eurasia and Nor ...
seeds
A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosperm pl ...
are dispersed by wind, random distribution will often occur as the seedlings land in random places determined by uncontrollable factors. Oyster larvae can also travel hundreds of kilometers powered by sea currents, which can result in their random distribution. Random distributions exhibit chance clumps (see Poisson clumping
Poisson may refer to:
People
*Siméon Denis Poisson, French mathematician
Places
* Poissons, a commune of Haute-Marne, France
* Poisson, Saône-et-Loire, a commune of Saône-et-Loire, France
Other uses
* Poisson (surname), a French surname
* Poi ...
).
Statistical determination of distribution patterns
There are various ways to determine the distribution pattern of species. The Clark–Evans nearest neighbor method can be used to determine if a distribution is clumped, uniform, or random.
To utilize the Clark–Evans nearest neighbor method, researchers examine a population of a single species. The distance of an individual to its nearest neighbor is recorded for each individual in the sample. For two individuals that are each other's nearest neighbor, the distance is recorded twice, once for each individual. To receive accurate results, it is suggested that the number of distance measurements is at least 50. The average distance between nearest neighbors is compared to the expected distance in the case of random distribution to give the ratio:
:
If this ratio ''R'' is equal to 1, then the population is randomly dispersed. If ''R'' is significantly greater than 1, the population is evenly dispersed. Lastly, if ''R'' is significantly less than 1, the population is clumped. Statistical tests (such as t-test, chi squared, etc.) can then be used to determine whether ''R'' is significantly different from 1.
The variance/mean ratio method focuses mainly on determining whether a species fits a randomly spaced distribution, but can also be used as evidence for either an even or clumped distribution. To utilize the Variance/Mean ratio method, data is collected from several random samples of a given population. In this analysis, it is imperative that data from at least 50 sample plots is considered. The number of individuals present in each sample is compared to the expected counts in the case of random distribution. The expected distribution can be found using Poisson distribution
In probability theory and statistics, the Poisson distribution is a discrete probability distribution that expresses the probability of a given number of events occurring in a fixed interval of time or space if these events occur with a known co ...
. If the variance/mean ratio is equal to 1, the population is found to be randomly distributed. If it is significantly greater than 1, the population is found to be clumped distribution. Finally, if the ratio is significantly less than 1, the population is found to be evenly distributed. Typical statistical tests used to find the significance of the variance/mean ratio include Student's t-test
A ''t''-test is any statistical hypothesis test in which the test statistic follows a Student's ''t''-distribution under the null hypothesis. It is most commonly applied when the test statistic would follow a normal distribution if the value of ...
and chi squared.
However, many researchers believe that species distribution models based on statistical analysis, without including ecological models and theories, are too incomplete for prediction. Instead of conclusions based on presence-absence data, probabilities that convey the likelihood a species will occupy a given area are more preferred because these models include an estimate of confidence in the likelihood of the species being present/absent. They are also more valuable than data collected based on simple presence or absence because models based on probability allow the formation of spatial maps that indicates how likely a species is to be found in a particular area. Similar areas can then be compared to see how likely it is that a species will occur there also; this leads to a relationship between habitat suitability and species occurrence.
Species distribution models
Species distribution can be predicted based on the pattern of biodiversity at spatial scales. A general hierarchical model can integrate disturbance, dispersal and population dynamics. Based on factors of dispersal, disturbance, resources limiting climate, and other species distribution, predictions of species distribution can create a bio-climate range, or bio-climate envelope. The envelope can range from a local to a global scale or from a density independence to dependence. The hierarchical model takes into consideration the requirements, impacts or resources as well as local extinctions in disturbance factors. Models can integrate the dispersal/migration model, the disturbance model, and abundance model. Species distribution models (SDMs) can be used to assess climate change impacts and conservation management issues. Species distribution models include: presence/absence models, the dispersal/migration models, disturbance models, and abundance models. A prevalent way of creating predicted distribution maps for different species is to reclassify a land cover layer depending on whether or not the species in question would be predicted to habit each cover type. This simple SDM is often modified through the use of range data or ancillary information, such as elevation or water distance.
Recent studies have indicated that the grid size used can have an effect on the output of these species distribution models. The standard 50x50 km grid size can select up to 2.89 times more area than when modeled with a 1x1 km grid for the same species. This has several effects on the species conservation planning under climate change predictions (global climate models, which are frequently used in the creation of species distribution models, usually consist of 50–100 km size grids) which could lead to over-prediction of future ranges in species distribution modeling. This can result in the misidentification of protected areas intended for a species future habitat.
Species Distribution Grids Project
The Species Distribution Grids Project is an effort led out of the University of Columbia to create maps and databases of the whereabouts of various animal species. This work is centered on preventing deforestation and prioritizing areas based on species richness. As of April 2009, data are available for global amphibian distributions, as well as birds and mammals in the Americas. The map galler
Gridded Species Distribution
contains sample maps for the Species Grids data set. These maps are not inclusive but rather contain a representative sample of the types of data available for download:
Image:North_America.jpg, Species richness map (amphibians)
Image:North_America_birds.jpg, Species richness map (birds)
Image:North_America_mammals.jpg, Species richness map (mammals)
See also
* Geographic range limit
The geographical limits to the distribution of a species are determined by biotic or abiotic factors. Core populations are those occurring within the centre of the range, and marginal populations (also called peripheral populations) are found at t ...
* Animal migration
Animal migration is the relatively long-distance movement of individual animals, usually on a seasonal basis. It is the most common form of migration in ecology. It is found in all major animal groups, including birds, mammals, fish, reptiles ...
* Biogeography
Biogeography is the study of the distribution of species and ecosystems in geographic space and through geological time. Organisms and biological communities often vary in a regular fashion along geographic gradients of latitude, elevation, ...
* Colonisation
Colonization, or colonisation, constitutes large-scale population movements wherein migrants maintain strong links with their, or their ancestors', former country – by such links, gain advantage over other inhabitants of the territory. When ...
* Cosmopolitan distribution
In biogeography, cosmopolitan distribution is the term for the range of a taxon that extends across all or most of the world in appropriate habitats. Such a taxon, usually a species, is said to exhibit cosmopolitanism or cosmopolitism. The ext ...
* Occupancy frequency distribution In macroecology and community ecology, an occupancy frequency distribution (OFD) is the distribution of the numbers of species occupying different numbers of areas. It was first reported in 1918 by the Danish botanist Christen C. Raunkiær in his st ...
Notes
External links
Livestock Grazing Distribution Patterns: Does Animal Age Matter?
{{Authority control
Animal migration
Biogeography
Ecology terminology
Population ecology
Population genetics