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Peripatric speciation

Peripatric speciation is a mode of speciation in which a new species is formed from an isolated peripheral population. Since peripatric speciation resembles allopatric speciation, in that populations are isolated and prevented from exchanging genes, it can often be difficult to distinguish between them, and peripatric speciation may be considered one type or model of allopatric speciation. The primary distinguishing characteristic of peripatric speciation is that one of the populations is much smaller than the other, as opposed to allopatric speciation, in which similarly-sized populations become separated. The terms peripatric and peripatry are often used in biogeography, referring to organisms whose ranges are closely adjacent but do not overlap, being separated where these organisms do not occur—for example on an oceanic island compared to the mainland. Such organisms are usually closely related ; their distribution being the result of peripatric speciation.

History
Peripatric speciation was originally proposed by Ernst Mayr in 1954, It is related to the founder effect, where small living populations may undergo selection bottlenecks. The founder effect is based on models that suggest peripatric speciation can occur by the interaction of selection and genetic drift, Mayr first conceived of the idea by his observations of kingfisher populations in New Guinea and its surrounding islands. Around the same time, the botanist Verne Grant developed a model of quantum speciation very similar to Mayr's model in the context of plants. == Models ==
Models
Peripatric Peripatric speciation models are identical to models of vicariance (allopatric speciation). Peripatry can be distinguished from allopatric speciation by three key features: The peripatric model results in, what have been called, progenitor-derivative species pairs, whereby the derivative species (the peripherally isolated population)—geographically and genetically isolated from the progenitor species—diverges. A specific phylogenetic signature results from this mode of speciation: the geographically widespread progenitor species becomes paraphyletic (thereby becoming a paraspecies), with respect to the derivative species (the peripheral isolate). It is thought that the character traits of the peripherally isolated species become apomorphic, while the central population remains plesiomorphic. Modern cladistic methods have developed definitions that have incidentally removed derivative species by defining clades in a way that assumes that when a speciation event occurs, the original species no longer exists, while two new species arise; this is not the case in peripatric speciation. Loren H. Rieseberg and Luc Brouillet recognized the same dilemma in plant classification. Quantum and budding speciation The botanist Verne Grant proposed the term quantum speciation that combined the ideas of J. T. Gulick (his observation of the variation of species in semi-isolation), Sewall Wright (his models of genetic drift), Mayr (both his peripatric and genetic revolution models), and George Gaylord Simpson (his development of the idea of quantum evolution). Quantum speciation is a rapid process with large genotypic or phenotypic effects, whereby a new, cross-fertilizing plant species buds off from a larger population as a semi-isolated peripheral population. Evidence for the occurrence of this type of speciation has been found in several plant species pairs: Layia discoidea and L. glandulosa, Clarkia lingulata and C. biloba, and Stephanomeria malheurensis and S. exigua ssp. coronaria. The budding process, where a new species originates at the margins of an ancestral range, is thought to be common in plants Centrifugal speciation William Louis Brown, Jr. proposed an alternative model of peripatric speciation in 1957 called centrifugal speciation. This model contrasts with peripatric speciation by virtue of the origin of the genetic novelty that leads to reproductive isolation. A population of a species experiences periods of geographic range expansion followed by periods of contraction. During the contraction phase, fragments of the population become isolated as small refugial populations on the periphery of the central population. Because of the large size and potentially greater genetic variation within the central population, mutations arise more readily. These mutations are left in the isolated peripheral populations, promoting reproductive isolation. Consequently, Brown suggested that during another expansion phase, the central population would overwhelm the peripheral populations, hindering speciation. However, if the species finds a specialized ecological niche, the two may coexist. The phylogenetic signature of this model is that the central population becomes derived, while the peripheral isolates stay plesiomorphic Centrifugal speciation has been largely ignored in the scientific literature, often dominated by the traditional model of peripatric speciation. Centrifugal speciation may have taken place in tree kangaroos, South American frogs (Ceratophrys), shrews (Crocidura), and primates (Presbytis melalophos). or the Hawaiian cricket Laupala. However, while laboratory experiments are ongoing and yet to be completed in support of the model, there are field observations of shifts in the mating systems that undergo population bottlenecks which demonstrate that the dynamics of sexual selection is occurring in nature and therefore, it does represent a plausible process of peripatric speciation that takes place in nature. == Evidence ==
Evidence
Observational evidence and laboratory experiments support the occurrence of peripatric speciation. Islands and archipelagos are often the subject of speciation studies in that they represent isolated populations of organisms. Island species provide direct evidence of speciation occurring peripatrically in such that, "the presence of endemic species on oceanic islands whose closest relatives inhabit a nearby continent" must have originated by a colonization event. Peripatric speciation also occurs on continents, as isolation of small populations can occur through various geographic and dispersion events. Laboratory studies have been conducted where populations of Drosophila, for example, are separated from one another and evolve in reproductive isolation. Hawaiian archipelago Drosophila species on the Hawaiian archipelago have helped researchers understand speciation processes in great detail. It is well established that Drosophila has undergone an adaptive radiation into hundreds of endemic species on the Hawaiian island chain; originating from a single common ancestor (supported from molecular analysis). Studies consistently find that colonization of each island occurred from older to younger islands, and in Drosophila, speciating peripatrically at least fifty percent of the time. Other endemic species in Hawaii also provide evidence of peripatric speciation such as the endemic flightless crickets (Laupala). It has been estimated that, "17 species out of 36 well-studied cases of [Laupala] speciation were peripatric". Plant species in genera's such as Dubautia, Wilkesia, and Argyroxiphium have also radiated along the archipelago. Other animals besides insects show this same pattern such as the Hawaiian amber snail (Succinea caduca), and ʻElepaio flycatchers. Tetragnatha spiders have also speciated peripatrically on the Hawaiian islands, Numerous arthropods have been documented existing in patterns consistent with the geologic evolution of the island chain, in such that, phylogenetic reconstructions find younger species inhabiting the geologically younger islands and older species inhabiting the older islands (or in some cases, ancestors date back to when islands currently below sea level were exposed). Spiders such as those from the genus Orsonwelles exhibit patterns compatible with the old-to-young geology. Other endemic genera such as Argyrodes have been shown to have speciated along the island chain. Pagiopalus, Pedinopistha, and part of the family Thomisidae have adaptively radiated along the island chain, as well as the wolf spider family, Lycosidae. A host of other Hawaiian endemic arthropod species and genera have had their speciation and phylogeographical patterns studied: the Drosophila grimshawi species complex, damselflies (Megalagrion xanthomelas and Megalagrion pacificum), Doryonychus raptor, Littorophiloscia hawaiiensis, Anax strenuus, Nesogonia blackburni, Theridion grallator, Vanessa tameamea, Hyalopeplus pellucidus, Coleotichus blackburniae, Labula, Hawaiioscia, Banza (in the family Tettigoniidae), Caconemobius, Eupethicea, Ptycta, Megalagrion, Prognathogryllus, Nesosydne, Cephalops, Trupanea, and the tribe Platynini—all suggesting repeated radiations among the islands. Other islands Phylogenetic studies of a species of crab spider (Misumenops rapaensis) in the genus Thomisidae located on the Austral Islands have established the, "sequential colonization of [the] lineage down the Austral archipelago toward younger islands". M. rapaensis has been traditionally thought of as a single species; whereas this particular study found distinct genetic differences corresponding to the sequential age of the islands. The figwart plant species Scrophularia lowei is thought to have arisen through a peripatric speciation event, with the more widespread mainland species, Scrophularia arguta dispersing to the Macaronesian islands. Other members of the same genus have also arisen by single colonization events between the islands. Species patterns on continents The occurrence of peripatry on continents is more difficult to detect due to the possibility of vicariant explanations being equally likely. In addition, a great deal of research has been conducted on several species of land snails involving chirality that suggests peripatry (with some authors noting other possible interpretations). Red spruce (Picea rubens) has arisen from an isolated population of black spruce (Picea mariana). During the Pleistocene, a population of black spruce became geographically isolated, likely due to glaciation. The geographic range of the black spruce is much larger than the red spruce. The red spruce has significantly lower genetic diversity in both its DNA and its mitochondrial DNA than the black spruce. Furthermore, the genetic variation of the red spruce has no unique mitochondrial haplotypes, only subsets of those in the black spruce; suggesting that the red spruce speciated peripatrically from the black spruce population. It is thought that the entire genus Picea in North America has diversified by the process of peripatric speciation, as numerous pairs of closely related species in the genus have smaller southern population ranges; and those with overlapping ranges often exhibit weak reproductive isolation. This distribution and paleobiogeographic pattern correlates with other species expressing similar biographic range patterns Laboratory experiments Peripatric speciation has been researched in both laboratory studies and nature. Jerry Coyne and H. Allen Orr in Speciation suggest that most laboratory studies of allopatric speciation are also examples of peripatric speciation due to their small population sizes and the inevitable divergent selection that they undergo. The table is a non-exhaustive table of laboratory experiments focused explicitly on peripatric speciation. Most of the studies also conducted experiments on vicariant speciation as well. The "replicates" column signifies the number of lines used in the experiment—that is, how many independent populations were used (not the population size or the number of generations performed). == References ==
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