Ecotype

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Ecotypes are organisms which belong to the same species but possess different phenotypical features as a result of environmental factors such as elevation, climate and predation. Ecotypes can be seen in wide geographical distributions and may eventually lead to speciation.

Contents

Definition

In evolutionary ecology, an ecotype, [note 1] sometimes called ecospecies, describes a genetically distinct geographic variety, population, or race within a species, which is genotypically adapted to specific environmental conditions.

Typically, though ecotypes exhibit phenotypic differences (such as in morphology or physiology) stemming from environmental heterogeneity, they are capable of interbreeding with other geographically adjacent ecotypes without loss of fertility or vigor. [1] [2] [3] [4] [5]

Summary

An ecotype refers to organisms which belong to the same species but have different phenotypical characteristics as a result of their adaptations to different habitats. [6] Differences between these two groups is attributed to phenotypic plasticity and are too few for them to be termed as wholly different species. [7] Emergence of variants of the same species may occur in the same geographical region where different habitats provide distinct ecological niches for these organisms examples of these habitats include meadows, forests, swamps, and sand dunes. [8] Where similar ecological conditions occur in widely separated places, it is possible for a similar ecotype to occur in the separated locations. [9] [10] An ecotype is different from a subspecies, which may exist across a number of different habitats. [11] In animals, ecotypes owe their differing characteristics to the effects of a very local environment which has been hypothesized to lead to speciation through the emergence of reproductive barriers. [12] [13] [14] Therefore, ecotypes have no taxonomic rank. [15]

Terminology

Ecotypes are closely related to morphs or polymorphisms which is defined as the existence of distinct phenotypes among members of the same species. [16] Another term closely related is genetic polymorphism; and it is when species of the same population display variation in a specific DNA sequence, i.e. as a result of having more than one allele in a gene's locus. [17] . In order to be classified as such, morphs must occupy the same habitat at the same time and belong to a panmictic population (whose members can all potentially interbreed). [18] Polymorphism are maintained in populations of species by natural selection. [19] [20] In fact, Begon, Townsend, and Harper assert that

There is not always clear distinction between local ecotypes and genetic polymorphisms.

The notions "form" and "ecotype" may appear to correspond to a static phenomenon, however; this is not always the case. [21] Evolution occurs continuously both in time and space, so that ecotypes or forms may qualify as distinct species in a few generations. [22] Begon, Townsend, and Harper use an illuminating analogy on this:

... the origin of a species, whether allopatric or sympatric, is a process, not an event. For the formation of a new species, like the boiling of an egg, there is some freedom to argue about when it is completed.

Thus ecotypes and morphs can be thought of as precursory steps of potential speciation. [21]

Range and distribution

Panicum virgatum ecotypes and their distribution in North America Panicum virgatum ecotypes.png
Panicum virgatum ecotypes and their distribution in North America

Research indicates that sometimes ecotypes manifest when separated by great geographical distances as a result of genetic drift that may lead to significant genetic differences and hence variation. [23] Ecotypes may also emerge from local adaptation of species occupying small geographical scales (<1km), in such cases divergent selection due to selective pressure as a result of differences in microhabitats drive differentiation. [23] Hybridization among populations may increase population gene flow and reduce the effects of natural selection. [24] [25] Hybridization here is defined as when different but adjacent varieties of the same species (or generally of the same taxonomic rank) interbreed, which helps overcome local selection. [1] However other studies reveal that ecotypes may emerge even at very small scales (of the order of 10 m), within populations, and despite hybridization. [1] [26]

In ecotypes, it is common for continuous, gradual geographic variation to impose analogous phenotypic and genetic variation, a situation which leads to the emergence of clines. [1] A well-known example of a cline is the skin color gradation in indigenous human populations worldwide, which is related to latitude and amounts of sunlight. [27] [28] Ecotypes may display two or more distinct and discontinuous phenotypes even within the same population. [29] [30] Ecological systems may have a species abundance that can be either bimodal or multimodal. [31] Emergence of ecotypes may lead to speciation and can occur if conditions in a local environment change dramatically through space or time. [1]

Ecotype and speciation

Just as sunlight can appear as a dim crack in the sky before clouds part, the coarse boundaries of ecotypes may appear as a separation of principle component clusters before speciation.

— David B. Lowry, Ecotypes and the controversy over stages in the formation of new species, Biological Journal of the Linnean Society.

The birth of the term 'ecotype' originally came from early interest in understanding speciation. [21] Darwin argued that species evolved through natural selection from variations within population which he termed as 'varieties'. [32] Later on, through a series of experiments, Turresson studied the effect of the environment on heritable plant variation and came up with the term 'ecotype' to denote differences between groups occupying distinct habitats. [2] This, he argued, was a genotypical response of plants to habitat type and it denotes a first step toward isolating reproductive barriers that facilitate the emergence of 'species' via divergence and, ultimately, genetic isolation. [2] [33] [34] In his 1923 paper, Turesson states that variation among species in a population is not random, rather, it is driven by environmental selection pressure. [35] For example, the maturity of Trifolium subterraneum, a clover which was found to correlate to moisture condition; when sown in low rainfall areas of Adelaide after a few years the population would consist of genotypes that produced seeds early in the season (early genotype), however in higher rainfall areas the clover population would shift to mid-season genotypes, differences among population of Trifolium subterraneum is in response to the selective action of the habitat. [36] These adaptive differences were hereditary and would emerge in response to specific environmental conditions. [37] Heritable differences is a key feature in ecotypic variation. [38] Ecotypic variation is as a result of particular environmental trends. [36] Individuals, which are able to survive and reproduce successfully pass on their genes to the next generation and establish a population best adapted to the local environment. [39] Ecotypic variation is therefore described to have a genetic base, and are brought about by interactions between an individual's genes and the environment. [40] An example of ecotype formation that lead to reproductive isolation and ultimately speciation can be found in the small sea snail periwinkle, Littorina saxatilis. [41] It is distributes across different habitats such as lagoons, salt marshes and rocky shores the range of distribution is from Portugal to Novaya Zemlaya and Svalbard and from North Carolina to Greenland. [42] The polymorphic snail species have different heritable features such as size and shape depending on the habitat they occupy e.g. bare cliffs, boulders and barnacle belts. [42] Phenotypic evolution in these snails can be strongly attributed to different ecological factors present in their habitats. For example, in coastal regions of Sweden, Spain and UK, Littorina saxatilis posses different shell shape in response to predation by crabs or waves surges. [43] Predation by crabs, also called crab crushing, gives rise to snails with wary behavior having large and thick shells which can easily retract and avoid predation. Wave-surfs on the other hand, select for smaller sized snails with large apertures to increase grip and bold behavior. [43] All this provide the basis for the emergence of different snail ecotypes. Snail ecotypes on the basis of morphology and behavior pass these characteristic on to their offspring. [44]

Examples

Rangifer tarandus caribou, a member of the woodland ecotype Caribou.jpg
Rangifer tarandus caribou, a member of the woodland ecotype

See also

Explanatory notes

  1. ↑ Greek: οίκος = home and τύπος = type, coined by Göte Turesson in 1922
  2. ↑ Banfield, who worked with both the Canadian Wildlife Service and the National Museum of Canada, in his often-cited 1961 classification, identified five subspecies of Rangifer tarandus: 1) the largely migratory barren-ground caribou subspecies Rangifer tarandus groenlandicus, which are found mainly in the Canadian territories of Nunavut and the Northwest Territories, along with western Greenland; 2) the subspecies Rangifer tarandus caribou which is divided into ecotypes: boreal woodland caribou, (also known as forest-dwelling, woodland caribou (boreal), mountain woodland caribou and migratory woodland caribou) —the migratory George River Caribou Herd, for example in the Ungava region of Quebec; 3) Rangifer tarandus pearyi (Peary caribou), the smallest of the species, known as Tuktu in Inuktitut, found in the northern islands of Nunavut and the Northwest Territories; 4) Rangifer tarandus granti subspecies Grant's caribou, which are mainly migratory and live in Alaska and the northern Yukon and 5) the R. t. dawsoni subspecies; † Queen Charlotte Islands caribou from the Queen Charlotte Islands (extinct since 1910)

Related Research Articles

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<span class="mw-page-title-main">Sympatric speciation</span> Evolution of a new species from an ancestor in the same location

In evolutionary biology, sympatric speciation is the evolution of a new species from a surviving ancestral species while both continue to inhabit the same geographic region. In evolutionary biology and biogeography, sympatric and sympatry are terms referring to organisms whose ranges overlap so that they occur together at least in some places. If these organisms are closely related, such a distribution may be the result of sympatric speciation. Etymologically, sympatry is derived from Greek συν (sun-) 'together' and πατρίς (patrís) 'fatherland'. The term was coined by Edward Bagnall Poulton in 1904, who explains the derivation.

<span class="mw-page-title-main">Polymorphism (biology)</span> Occurrence of two or more clearly different morphs or forms in the population of a species

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<span class="mw-page-title-main">Disruptive selection</span> Natural selection for extreme trait values over intermediate ones

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<span class="mw-page-title-main">Migratory woodland caribou</span> Subspecies of deer

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<span class="mw-page-title-main">Barren-ground caribou</span> Subspecies of deer

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<span class="mw-page-title-main">Queen Charlotte Islands caribou</span> Extinct subspecies of deer

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<span class="mw-page-title-main">Finnish forest reindeer</span> Subspecies of deer

The Finnish forest reindeer(Rangifer tarandus fennicus, also known as Eurasian or European forest reindeer is a rare subspecies of the reindeer native to Finland and northwestern Russia. They are found primarily in Russian Karelia and the provinces of North Karelia, Savonia and Kainuu in Finland, though some range into central south Finland. They are distinct from the semi-domesticated mountain reindeer in their larger size, longer legs and preference for dense boreal forest habitat, where they are rarely seen by humans, over the open tundra. The Finnish herd migrates seasonally back and forth across the long Russo-Finnish border.

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<span class="mw-page-title-main">Ecological speciation</span>

Ecological speciation is a form of speciation arising from reproductive isolation that occurs due to an ecological factor that reduces or eliminates gene flow between two populations of a species. Ecological factors can include changes in the environmental conditions in which a species experiences, such as behavioral changes involving predation, predator avoidance, pollinator attraction, and foraging; as well as changes in mate choice due to sexual selection or communication systems. Ecologically-driven reproductive isolation under divergent natural selection leads to the formation of new species. This has been documented in many cases in nature and has been a major focus of research on speciation for the past few decades.

<span class="mw-page-title-main">Boreal woodland caribou</span> Subspecies of caribou or reindeer in North America

The boreal woodland caribou, also known as Eastern woodland caribou, boreal forest caribou and forest-dwelling caribou, is a North American subspecies of reindeer found primarily in Canada with small populations in the United States. Unlike the Porcupine caribou and barren-ground caribou, boreal woodland caribou are primarily sedentary.

Alexander William Francis Banfield, Frank Banfield, A. W. F, Banfield, was one of the small group of early Canadian mammalogists who worked with Canadian Wildlife Service (CWS) and the National Museum of Canada. His research and publications appeared repeatedly in publications on mammals in Canada. and in 1974 he published his book Mammals of Canada. His 1961 article "A Revision of the Reindeer and Caribou, Genus Rangifer" in the National Museum of Canada's Bulletin continues to be widely cited today in discussions on subspecies and ecotypes of caribou.

<span class="mw-page-title-main">Dolphin-Union caribou</span> Migratory population of caribou

Dolphin and Union Caribou, Dolphin and Union caribou herd, Dolphin-Union, locally known as Island Caribou, are a migratory population of barren-ground caribou, Rangifer tarandus groenlandicus, that occupy Victoria Island in the Canadian Arctic Archipelago and the nearby mainland. They are endemic to Canada. They migrate across the Dolphin and Union Strait from their summer grazing on Victoria Island to their winter grazing area on the Nunavut-Northwest Territories mainland in Canada. It is unusual for North American caribou to seasonally cross sea ice and the only other caribou to do so are the Peary caribou who are smaller in size and population. They were listed as Endangered by Committee on the Status of Endangered Wildlife in Canada (COSEWIC) since November 2017.

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<span class="mw-page-title-main">Reindeer distribution</span> Reindeer distribution around the world

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Allochronic speciation is a form of speciation arising from reproductive isolation that occurs due to a change in breeding time that reduces or eliminates gene flow between two populations of a species. The term allochrony is used to describe the general ecological phenomenon of the differences in phenology that arise between two or more species—speciation caused by allochrony is effectively allochronic speciation.

In biology, parallel speciation is a type of speciation where there is repeated evolution of reproductively isolating traits via the same mechanisms occurring between separate yet closely related species inhabiting different environments. This leads to a circumstance where independently evolved lineages have developed reproductive isolation from their ancestral lineage, but not from other independent lineages that inhabit similar environments. In order for parallel speciation to be confirmed, there is a set of three requirements that has been established that must be met: there must be phylogenetic independence between the separate populations inhabiting similar environments to ensure that the traits responsible for reproductive isolation evolved separately, there must be reproductive isolation not only between the ancestral population and the descendent population, but also between descendent populations that inhabit dissimilar environments, and descendent populations that inhabit similar environments must not be reproductively isolated from one another. To determine if natural selection specifically is the cause of parallel speciation, a fourth requirement has been established that includes identifying and testing an adaptive mechanism, which eliminates the possibility of a genetic factor such as polyploidy being the responsible agent.

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