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The geographical limits to the
distribution Distribution may refer to: Mathematics *Distribution (mathematics), generalized functions used to formulate solutions of partial differential equations * Probability distribution, the probability of a particular value or value range of a vari ...
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 ...
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 the boundary of the range. The inability of a species to expand its range beyond a certain geographic area is because of some
limiting factor A limiting factor is a variable of a system that causes a noticeable change in output or another measure of a type of system. The limiting factor is in a pyramid shape of organisms going up from the producers to consumers and so on. A factor not l ...
or factors to which the species cannot successfully adapt. In some cases, geographical range limits are entirely predictable, such as the physical barrier of an ocean for a terrestrial species. In other cases the specific reasons why species do not pass these boundaries are unknown, however,
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 overlaps wi ...
is the main determinant of the distribution of a species. The fitness of a species falls at the edges of its distributional range, with population growth and fitness falling to zero beyond where a species can survive. For many species of invertebrate animals, the exact geographic range limits have never been precisely ascertained, because not enough scientific field work has been carried in many parts of the world to map distribution more precisely, therefore finding a range extension for species, especially marine species, is not an uncommon occurrence. Marginal distributions can have conservation implications.


Terminology

The science of understanding the distributions of organisms is known as
chorology Chorology (from Greek , ''khōros'', "place, space"; and , ''-logia'') can mean * the study of the causal relations between geographical phenomena occurring within a particular region * the study of the spatial distribution of organisms ( biogeogra ...
a branch of
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, ...
. The core population of a species are those individuals occurring within the centre of the range. Although one cannot ever truly know the ideal
niche Niche may refer to: Science *Developmental niche, a concept for understanding the cultural context of child development *Ecological niche, a term describing the relational position of an organism's species *Niche differentiation, in ecology, the ...
of a particular species, it can be approximated from the core of the distribution, this is known as the "realized ecological niche". Marginal or peripheral populations are those found at the boundary of the range. When the distribution of a species is changing, the leading edge populations are at the expanding geographic edge of the distribution range whilst rear edge populations are undergoing retreat. The central‐marginal hypothesis, also sometimes called the "central-peripheral population hypothesis", posits that there is less genetic diversity and greater inter‐population genetic differentiation at the range margins, as compared to the range cores. This is based on the assumption that the
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
is most ideal at the centre of a distribution and ecological conditions decline towards the margin. Because the
population size In population genetics and population ecology, population size (usually denoted ''N'') is the number of individual organisms in a population. Population size is directly associated with amount of genetic drift, and is the underlying cause of effect ...
at the margin is likely to be smaller,
genetic drift Genetic drift, also known as allelic drift or the Wright effect, is the change in the frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene variants to disappear completely and there ...
can have a larger effect and reduce the
genetic variation Genetic variation is the difference in DNA among individuals or the differences between populations. The multiple sources of genetic variation include mutation and genetic recombination. Mutations are the ultimate sources of genetic variation, ...
of marginal populations. Reduced
gene flow In population genetics, gene flow (also known as gene migration or geneflow and allele flow) is the transfer of genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent a ...
between central and peripheral populations also limits the genetic diversity at the margins. High
selection pressure Any cause that reduces or increases reproductive success in a portion of a population potentially exerts evolutionary pressure, selective pressure or selection pressure, driving natural selection. It is a quantitative description of the amount of ...
, due to a less than ideal habitat at the margin, furthermore reduces genetic diversity. Although exceptions to this hypothesis are common, in general this rule appears to hold empirically true. The
spatial distribution A spatial distribution in statistics is the arrangement of a phenomenon across the Earth's surface and a graphical display of such an arrangement is an important tool in geographical and environmental statistics. A graphical display of a spatial d ...
often differs, with the population being more dense in the centre as opposed to the margins, this can often have a simple
probability distribution In probability theory and statistics, a probability distribution is the mathematical function that gives the probabilities of occurrence of different possible outcomes for an experiment. It is a mathematical description of a random phenomenon i ...
pattern. The gene flow between central and peripheral populations may prevent range expansion when it does not allow the
gene pool The gene pool is the set of all genes, or genetic information, in any population, usually of a particular species. Description A large gene pool indicates extensive genetic diversity, which is associated with robust populations that can surv ...
at margin to differentiate. Conditions at the centre of the range differ from those at the periphery, therefore adapted alleles at the centre may not benefit marginal populations experiencing different conditions. The asymmetrical gene flow hypothesis posits that there is more
gene flow In population genetics, gene flow (also known as gene migration or geneflow and allele flow) is the transfer of genetic material from one population to another. If the rate of gene flow is high enough, then two populations will have equivalent a ...
from central to peripheral populations. Empirical data supporting this theory is less robust. When circumstances, usually climatic, restrict the distribution to a small area, this is known as a refugium. In Europe, for example, the geographical spokes sticking out of the continent in the south - the
Iberian Peninsula The Iberian Peninsula (), ** * Aragonese and Occitan: ''Peninsula Iberica'' ** ** * french: Péninsule Ibérique * mwl, Península Eibérica * eu, Iberiar penintsula also known as Iberia, is a peninsula in southwestern Europe, defi ...
,
Italy Italy ( it, Italia ), officially the Italian Republic, ) or the Republic of Italy, is a country in Southern Europe. It is located in the middle of the Mediterranean Sea, and its territory largely coincides with the homonymous geographical re ...
and the
Balkans The Balkans ( ), also known as the Balkan Peninsula, is a geographical area in southeastern Europe with various geographical and historical definitions. The region takes its name from the Balkan Mountains that stretch throughout the who ...
served as refugia for warmth-adapted species during the
Ice Ages An ice age is a long period of reduction in the temperature of Earth's surface and atmosphere, resulting in the presence or expansion of continental and polar ice sheets and alpine glaciers. Earth's climate alternates between ice ages and gree ...
.


Abiotic factors

Gradients in any
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 ...
factor, such as climate, create physiological barriers to dispersal. All species have limits of tolerance to abiotic factors. Too much or too little of anything can lower their survival and
reproductive success Reproductive success is an individual's production of offspring per breeding event or lifetime. This is not limited by the number of offspring produced by one individual, but also the reproductive success of these offspring themselves. Reproduct ...
and cause reduced fitness. Changes in temperature resulting from
global warming 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 E ...
, for example, may cause a species to change its geographical distribution northward.
Precipitation In meteorology, precipitation is any product of the condensation of atmospheric water vapor that falls under gravitational pull from clouds. The main forms of precipitation include drizzle, rain, sleet, snow, ice pellets, graupel and hail. ...
can also be a key determinant in limiting the geographic range edges of species. This is often seen in organisms with high water demands, whose survival and reproduction is limited by dry conditions. Moisture in the soil can also put limits on the distribution of an organism. There are many other abiotic factors that can determine a species range, including
dissolved oxygen Oxygen saturation (symbol SO2) is a relative measure of the concentration of oxygen that is dissolved or carried in a given medium as a proportion of the maximal concentration that can be dissolved in that medium at the given temperature. It ca ...
,
conductivity Conductivity may refer to: *Electrical conductivity, a measure of a material's ability to conduct an electric current **Conductivity (electrolytic), the electrical conductivity of an electrolyte in solution **Ionic conductivity (solid state), elec ...
,
alkalinity Alkalinity (from ar, القلوي, al-qaly, lit=ashes of the saltwort) is the capacity of water to resist acidification. It should not be confused with basicity, which is an absolute measurement on the pH scale. Alkalinity is the strength of ...
and pH.


Biotic factors

Interactions between organisms can cause limitations to the distribution of a species. One interaction that may limit a distribution is
predation Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill the ...
, where prey species are limited from a particular area by very efficient predators, or where these predators may permit certain prey to have larger ranges.
Interspecific competition Interspecific competition, in ecology, is a form of competition in which individuals of ''different'' species compete for the same resources in an ecosystem (e.g. food or living space). This can be contrasted with mutualism, a type of symbiosis. ...
is another common determinant of the distribution of individual species. Where two similar species share an overlapping range, competition often causes the distributions to shift to exclude one of the two. The geographic range of one species may be linked to another, where the range of one species cannot extend independent of the other. This is seen in
parasitism Parasitism is a Symbiosis, close relationship between species, where one organism, the parasite, lives on or inside another organism, the Host (biology), host, causing it some harm, and is Adaptation, adapted structurally to this way of lif ...
or mutualism, where survival is not possible without the
hosts A host is a person responsible for guests at an event or for providing hospitality during it. Host may also refer to: Places *Host, Pennsylvania, a village in Berks County People *Jim Host (born 1937), American businessman *Michel Host ( ...
. Parasitism can also play another role in determining the distribution of a species: marginal populations with suboptimal habitats can carry a higher
parasite load Parasite load is a measure of the number and virulence of the parasites that a host organism harbours. Quantitative parasitology deals with measures to quantify parasite loads in samples of hosts and to make statistical comparisons of parasitism ac ...
. This may be because less favourable conditions at the margins of a distribution lead to lower resistance to infection.


Anthropogenic factors

Humans can cause changes to the environment and alter distributions.
Deforestation Deforestation or forest clearance is the removal of a forest or stand of trees from land that is then converted to non-forest use. Deforestation can involve conversion of forest land to farms, ranches, or urban use. The most concentrated d ...
can increase the habitat of certain species and allow them to expand their distribution, or change distributions in response to a decrease in habitat. Recent changes in average temperatures, which may be caused by humans, are causing changes in the distribution of some species, such as northward expansion. Humans have also initiated many range expansions by introducing species to new locations both intentionally and accidentally. These species may survive and reproduce in these new locations and thus expand their distribution. These species may also cause changes in the distributions of
native species In biogeography, a native species is indigenous to a given region or ecosystem if its presence in that region is the result of only local natural evolution (though often popularised as "with no human intervention") during history. The term is equ ...
that cannot tolerate the novel competition.


Combined influences

In most cases combinations of factors are responsible for limiting the geographic range edge of species. Abiotic and biotic factors may work together in determining the range of a species. An example might be some obligate seeder plants where the distribution is limited by the presence of wildfires, which are needed to allow their
seed bank A seed bank (also seed banks or seeds bank) stores seeds to preserve genetic diversity; hence it is a type of gene bank. There are many reasons to store seeds. One is to preserve the genes that plant breeders need to increase yield, disease res ...
to germinate, and also use dispersal of their seeds mediated by ants.


See also

*
Edge effects In ecology, edge effects are changes in population or community structures that occur at the boundary of two or more habitats. Areas with small habitat fragments exhibit especially pronounced edge effects that may extend throughout the range. As ...
*
Species distribution Species distribution —or species dispersion — is the manner in which a biological taxon 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 ...
*
Niche construction Niche construction is the process by which an organism alters its own (or another species') local environment. These alterations can be a physical change to the organism’s environment or encompass when an organism actively moves from one habita ...
*
Mutualism Parasitism Continuum The hypothesis or paradigm of Mutualism Parasitism Continuum postulates that compatible host-symbiont associations can occupy a broad continuum of interactions with different fitness outcomes for each member. At one end of the continuum lies obliga ...


References

{{reflist, 32em Population ecology