Evidence for speciation by reinforcement

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Reinforcement assists speciation by selecting against hybrids. Speciation by Reinforcement Schematic.svg
Reinforcement assists speciation by selecting against hybrids.

Reinforcement is a process within speciation where natural selection increases the reproductive isolation between two populations of species by reducing the production of hybrids. [1] [2] Evidence for speciation by reinforcement has been gathered since the 1990s, and along with data from comparative studies and laboratory experiments, has overcome many of the objections to the theory. [3] : 354  [4] [5] Differences in behavior or biology that inhibit formation of hybrid zygotes are termed prezygotic isolation. Reinforcement can be shown to be occurring (or to have occurred in the past) by measuring the strength of prezygotic isolation in a sympatric population in comparison to an allopatric population of the same species. [3] : 357  Comparative studies of this allow for determining large-scale patterns in nature across various taxa. [3] : 362  Mating patterns in hybrid zones can also be used to detect reinforcement. [6] Reproductive character displacement is seen as a result of reinforcement, [7] so many of the cases in nature express this pattern in sympatry. Reinforcement's prevalence is unknown, [4] but the patterns of reproductive character displacement are found across numerous taxa (vertebrates, invertebrates, plants, and fungi), and is considered to be a common occurrence in nature. [6] Studies of reinforcement in nature often prove difficult, as alternative explanations for the detected patterns can be asserted. [3] : 358  Nevertheless, empirical evidence exists for reinforcement occurring across various taxa [7] and its role in precipitating speciation is conclusive. [8]

Contents

Evidence from nature

Amphibians

The southern brown tree frog, Litoria ewingi Brown Tree Frog 2.jpg
The southern brown tree frog, Litoria ewingi

The two frog species Litoria ewingi and L. verreauxii live in southern Australia with their two ranges overlapping. The species have very similar calls in allopatry, but express clinal variation in sympatry, with notable distinctness in calls that generate female preference discrimination. [8] The zone of overlap sometimes forms hybrids and is thought to originate by secondary contact of once fully allopatric populations. [8]

Allopatric populations of Gastrophryne olivacea and G. carolinensis have recently come into secondary contact due to forest clearing. [9] The calls that the males make to attract females differ significantly in frequency and duration in the area where the two species overlap, despite them having similar calls where they do not. [3] : 359  Further, the hybrids that form in sympatry have calls that are intermediate between the two. [9] Similar patterns of reproductive character displacement involving acoustic displays have been found in Hyla cinerea and H. gratiosa , with greater female preference for conspecific males in areas of sympatry. [10]

Three species of true frogs ( Lithobates sphenocephalus , L. berlandieri , and L. blairi ) are temporally isolated in that their breeding seasons are spaced out in areas where they live in sympatry, but not where they live in allopatry. [11] Selection against interspecific mating due to low hybrid fitness and low hybrid fertility has reinforced the observed character displacement of breeding times. [11]

The rainforests of northeast Queensland, Australia were separated into north and south refugia by climate fluctuations of the Pliocene and Pleistocene. [12] About 6500 years ago, the rainforests reconnected, bringing the diverged, incipient populations of Litoria genimaculata into secondary contact. The species contact zones exhibit, "strong postzygotic selection against hybrids" and enhanced isolation from differences in mating call. [13]

An alternative to detecting reproductive character displacement in populations that overlap in sympatry is measuring rates of hybridization in contact zones. [9] The frog species Anaxyrus americanus and Anaxyrus woodhousii have shown a decrease in hybridization from 9% to 0% over approximately 30 years. [14] [9] A similar pattern was detected in the sympatric spadefoot toads Spea multiplicata and S. bombifrons have hybridized with decreasing frequency over a 27-year period (about 13 generations). [15]

Birds

The small ground finch, one of Darwin's finches of the Galapagos Geospiza fuliginosa.jpg
The small ground finch, one of Darwin's finches of the Galápagos

The Ficedula flycatchers exhibit a pattern that suggests premating isolation is being reinforced by sexual selection. [16] The pied flycatcher ( Ficedula hypoleuca ) has brown females, brown males, and black-and-white males. The related collard flycatcher ( Ficedula albicollis ) has brown females and only black-and-white males. The two species exist in separate populations that overlap in a zone of sympatry. [16] In the range of overlap, only brown males of F. hypoleuca exist and are thought to have evolved the brown plumage to prevent hybridization, [17] though there is also evidence indicating that such character displacement is explained by heterospecific competition for territory rather than reinforcement. [18] Mating choice tests of the species find that females of both species choose conspecific males in sympatry, but heterospecific males in allopatry [16] (see conspecific song preference). The patterns could suggest mimicry, driven by interspecific competition; [3] : 361  however, song divergence has been detected that shows a similar pattern to the mating preferences. [19]

Geospiza fuliginosa and G. difficilis males on the Galápagos Islands show a noted preference for conspecific females where they meet in sympatry, but not in allopatry. [20] Other birds such as the dark and light subspecies of the western grebe show enhanced prezygotic isolation. [21] It has been argued that reinforcement is extremely common in birds and has been documented in a wide range of bird species. [22]

Crustaceans

Reproductive character displacement in body size was detected in sympatric populations of Orconectes rusticus and O. sanbornii . [23]

Echinoderms

An example of gametic isolation involves the allopatric sea urchins ( Arbacia ) have minimal bindin differences (bindin is a protein involved in the process of sea urchin fertilization, used for species-specific recognition of the egg by the sperm) and have insufficient barriers to fertilization. [3] : 243  Comparison with the sympatric species Echinometra and Strongylocentrotus of the Indo-Pacific finds that they have significant differences in bindin proteins for fertilization and marked fertilization barriers. [24]

Echinometra viridis, one of several species in the genus that produce fertile hybrids Echinometra viridis (Reef Urchin).jpg
Echinometra viridis , one of several species in the genus that produce fertile hybrids

Laboratory matings of closely related sea urchin species Echinometra oblonga and E. sp. C (the species is unnamed, dubbed C) produce fertile and viable hybrids, but are unable to fertilize eggs of the parent species due to divergence of the alleles that code for bindin proteins: an example of post-zygotic isolation. [3] : 343–344  Populations in sympatry manifest this difference in bindin protein versus those in allopatry. [3] : 343–344  Selection actively acts against the formation of hybrids in both nature (as no documented cases of hybrids have been found) and in the laboratory. [25] Here, the evolution of female egg receptors is thought to pressure bindin evolution in a selective runaway process. [25] This example of reproductive character displacement is highly suggestive of being the result of—and has been cited as strong evidence for—reinforcement. [25] [3] : 343–344 

Fish

In British Columbia, benthic and limnetic morphs of Gasterosteus aculeatus exist together in sympatry in some lakes, while containing only one morph in other lakes. [26] Female benthic morphs in sympatric populations actively discriminate against limnetic males, resulting in low rates of crossing (some gene flow has occurred between the morphs) and low fitness hybrids. [3] : 360  Both selection against hybrids and reproductive character displacement in egg fertilization is observed in Etheostoma lepidum and E. spectabile . [27]

Fungi

A strong case of reinforcement occurring in fungi comes from studies of Neurospora . [28] In crosses between different species in the genera, sympatric pairs show low reproductive success, significantly lower than allopatric pairs. [28] This pattern is observed across small and large geographic scales, with distance correlating with reproductive success. [28] Further evidence of reinforcement in the species was the low fitness detected in the hybrids create from crosses, and that no hybrids have been found in nature, despite close proximity. [28]

Insects

Ethological isolation has been observed between some mosquito species in the Southeast Asian Aedes albopictus group, suggesting—from laboratory experiments of mating trials—that selection against hybrids is occurring, in the presence of reproductive character displacement. [29]

Female mate discrimination is increased with intermediate migration rates between allopatric populations of Timema cristinae (genus Timema ) compared to high rates of migration (where gene flow impedes selection) or low rates (where selection is not strong enough). [30] [31]

Distribution of the periodical cicadas in the U.S. The yellow area corresponds to the sympatric overlap of Magicicada neotredecim (blue) with Magicicada tredecim (red). Distribution magicicada usa fixed.png
Distribution of the periodical cicadas in the U.S. The yellow area corresponds to the sympatric overlap of Magicicada neotredecim (blue) with Magicicada tredecim (red).

Where the ranges of the cicada species Magicicada tredecim and M. neotredecim overlap (where they are sympatric), the pitch of M. neotredecim male calling songs is roughly 1.7 kHz compared to 1.1 kHz for those of M. tredecim, with corresponding female song pitch preference differences. [32] In allopatric M. neotredecim populations, the mating call pitch is 1.3–1.5 kHz. [32] The biogeography of the cicadas suggests that M. neotredecim originated after the retreat of the last glacial advance in North America. [32]

The song differences of Laupala crickets on the Hawaiian Islands appear to exhibit patterns consistent with character displacement in sympatric populations. [33] A similar pattern exists with Allonemobius fasciatus and A. socius , species of ground crickets in eastern North America. [34]

Males in sympatric populations of the damselflies Calopteryx maculata and C. aequabilis are able to discriminate between females of different species better than those in allopatric populations; with females of C. aequabilis in sympatric populations exhibiting lighter wing colors compared to allopatric females—an illustration of reproductive character displacement. [35] [36]

Fifteen species of sympatrically distributed Agrodiaetus butterflies with pronounced differences in wing color pattern likely arose as a result of speciation by reinforcement. [37] Phylogenetic patterns indicate the differences arose in allopatry and were reinforced when the distributions came into secondary contact. [38]

Drosophila

Drosophila fruit fly Drosophila melanogaster - side (aka).jpg
Drosophila fruit fly

Drosophila is one of the most studied species in speciation research. [39] Dobzhansky and Koller were the first to study isolation between Drosophila species. [3] : 358  Since then, other studies of natural populations such as the D. paulistorum races exhibiting stronger isolation in sympatry versus allopatry, [40] or the enhanced isolation found in sympatric populations of D. mojavensis and D. arizonae in southwest America. [41] Rare, sterile hybrids form between D. pseudoobscura and D. persimilis , with sympatric D. pseudoobscura females discriminating against D. persimilis males; more so than allopatric populations. [42] Other Drosophila research on reinforcement has been from laboratory experiments and is discussed below. On the east coast of Australia, D. serrata shares a zone of sympatric overlap with the closely related species D. birchii . [43] The species exhibits reproductive character displacement, with sexual selection operating on the hydrocarbons of the flies cuticle. [44] Reinforcement appears to be driving their speciation in nature, supported by simulated experimental laboratory populations. [45] [46]

Mammals

The deer mice Peromyscus leucopus and P. gossypinus exhibit reproductive character displacement in mating preferences, with heterospecific matings taking place between the species. [47]

Molluscs

Some gastropod shells, like those of Partula can coil in sinistral and dextral directions such as these, Neptunea angulata (left) and N. despecta (right) shells. Neptunea - links&rechts gewonden.jpg
Some gastropod shells, like those of Partula can coil in sinistral and dextral directions such as these, Neptunea angulata (left) and N. despecta (right) shells.

Partula suturalis is polymorphic for shell chirality in that it has two forms: sinistral (left-handed) and dextral (right-handed) shells, unlike other monomorphic species on the island of Mo'orea which have only one form (with the exception of P. otaheitana ). [48] This polymorphic trait has a direct effect on mate choice and mating behavior; as shown in laboratory mating tests that opposite-coil pairs mate much less often. [48] In areas where P. suturalis lives sympatrically with other sinistral and dextral Partula species, the opposite P. suturalis morph is typically present. [9] Butlin succinctly describes one example of this unique pattern:

P. suturalis is sympatric with the dextral P. aurantia and sinistral P. olympia, whose ranges abut but do not overlap; P. suturalis is sinestral in the range of P. aurantia and dextral in the range of P. olympia and does not normally hybridize with either species. However, where their ranges meet there is a sharp transition in the coil of P. suturalis and in this transition zone it hybridizes with both P. aurantia and P. olympia. [9]

The reversal in chirality to sinistrality must have evolved as an isolating mechanism, [49] with patterns of reproductive character displacement suggesting speciation by reinforcement. [48]

Satsuma largillierti lives on the western half of Okinawa Island while Satsuma eucosmia lives on the eastern half. Both populations overlap in sympatry along the middle of the island, where the penis length of the species differs significantly in sympatry (a case of reproductive character displacement [50] ), but not in allopatry. [51] A similar pattern in snails is found with Lymnaea peregra and L. ovata in the Swiss lake Seealpsee; with mating signal acting as the sympatrically displaced trait. [52]

The abalone genus Haliotis has 19 species that occur in sympatry and one that occurs in allopatry. Of the sympatric species, they all contain sperm lysin that drives gamete isolation, but the allopatric species does not. [53] [3] : 343  A similar pattern of sperm lysin differentiation is found in the mussel species Mytilus galloprovincialis and M. trossulus and has likely occurred within the last 200 years due to human-mediated distribution by ships. [3] : 343 

Plants

Plants are thought to provide suitable conditions for reinforcement to occur. [5] This is due to a number of factors such as the unpredictability of pollination, pollen vectors, hybridization, hybrid zones, among others. [5] The study of plants experiencing speciation by reinforcement has largely been overlooked by researchers; [3] : 364  however, there is evidence of its occurrence in them. [54]

In the Texas wildflower Phlox drummondii , cis-regulatory mutations of genes that code for anthocyanin pigmentation have caused genetic divergence of two populations. [55] Hybrids (between P. drummondii and P. cuspidata ) with maladaptive, intermediate characteristics are under-pollinated; increasing reproductive isolation through reinforcement. [55] The maintenance of the ancestral flower color in the allopatric population is favored weakly by selection, where the derived color in the sympatric population is being driven by strong selection. [56] Similarly, in P. pilosa and P. glaberrima , character displacement of petal color has been driven by selection, aided by pollen discrimination. [57] Displacement in flower size has also been observed in the nightshade species Solanum grayi and S. lymholtzianum in sympatry as well as S. rostratum and S. citrullifolium . [58]

Pinus muricata-1.jpg
Green southern population foliage of Pinus muricata
Pinus muricata foliage Mendocino.jpg
Blue northern population

The bishop pine is divided into two populations distinguished by monoterpene, stomata, and alloenzyme differences; flowering time; and needle color: blue foliage in the northern population and green foliage in the southern populations in California. [59] A small region exists where the species meet in a cline—sustained by selection due to a flowering time divergence, thought to represent reinforcement taking place. [9]

Similar patterns of both character displacement in sympatric populations of species have been documented in: [9] [3] : 361 

Comparative studies

Prezygotic isolation in allopatric (red) and sympatric (blue) species pairs of Drosophila. Gradients indicate the predictions of reinforcement for allopatric and sympatric populations. Enhanced pre-zygotic isolation in Drosophila (allopatric & sympatric plots-vert).png
Prezygotic isolation in allopatric (red) and sympatric (blue) species pairs of Drosophila . Gradients indicate the predictions of reinforcement for allopatric and sympatric populations.

Assortive mating is expected to increase among sympatric populations experiencing reinforcement. [8] This fact allows for the direct comparison of the strength of prezygotic isolation in sympatry and allopatry between different experiments and studies. [3] : 362  Jerry Coyne and H. Allen Orr surveyed 171 species pairs, collecting data on their geographic mode, genetic distance, and strength of both prezygotic and post-zygotic isolation; finding that prezygotic isolation was significantly stronger in sympatric pairs, correlating with the ages of the species. [3] : 362  Additionally, the strength of post-zygotic isolation was not different between sympatric and allopatric pairs. [8]

This finding lends support the predictions of speciation by reinforcement and correlates well with another later study by Daniel J. Howard. [3] : 363  In his study, 48 studies with observed reproductive character displacement (including plants, insects, crustaceans, molluscs, fish, amphibians, reptiles, birds, and mammals) were analyzed. [6] The cases met several criteria such as the trait in question serving as a reproductive barrier and if there existed clear patterns of sympatry versus allopatry. [6] Out of the 48 candidates, 69 percent (33 cases) found enhanced isolation in sympatry, suggesting that the pattern predicted by reinforcement is common in nature. [6] In addition to Howard's comparative study, he guarded against the potential for positive-result publication bias by surveying 37 studies of hybrid zones. A prediction of reinforcement is that assortive mating should be common in hybrid zones; a prediction that was confirmed in 19 of the 37 cases. [6]

A survey of the rates of speciation in fish and their associated hybrid zones found similar patterns in sympatry, supporting the occurrence of reinforcement. [68] One study in the plants Glycine and Silene ; however, did not find enhanced isolation. [69]

Laboratory experiments

Laboratory studies that explicitly test for reinforcement are limited. [3] : 357  In general, two types of experiments have been conducted: using artificial selection to mimic natural selection that eliminates the hybrids (often called "destroy-the-hybrids"), and using disruptive selection to select for a trait (regardless of its function in sexual reproduction). [3] : 355–357  Many experiments using the destroy-the-hybrids technique are generally cited as supportive of reinforcement; however, some researchers such as Coyne and Orr and William R. Rice and Ellen E. Hostert contend that they do not truly model reinforcement, as gene flow is completely restricted between two populations. [70] [3] : 356  The table below summarizes some of the laboratory experiments that are often cited as testing reinforcement in some form.

Table of laboratory studies of reinforcement [70] [3] : 354–357 
SpeciesExperimental designResultYear
D. paulistorum Destroyed hybridsPre-zygotic isolation1976 [71]
D.pseudoobscura &

D. persimilis

Destroyed hybridsPre-zygotic isolation; reproductive character displacement1950 [72]
D. melanogaster Destroyed hybridsPre-zygotic isolation; reproductive character displacement1974 [73]
D. melanogasterDestroyed hybridsPre-zygotic isolation; reproductive character displacement1956 [74]
D. melanogasterDestroyed hybridsNo pre-zygotic isolation detected1970 [75]
D. melanogasterDestroyed hybridsPre-zygotic isolation1953 [76]
D. melanogasterDestroyed hybridsPre-zygotic isolation1974 [77]
D. melanogasterAllopatric populations in secondary contactN/A1982 [78]
D. melanogasterN/A1991 [79]
D. melanogasterNo pre-zygotic isolation detected1966 [80] [81]
D. melanogasterAllowed gene flow between populationsNo pre-zygotic isolation detected1969 [82]
D. melanogasterN/ANo pre-zygotic isolation detected1984 [83]
D. melanogasterDestroyed some hybridsNo pre-zygotic isolation detected1983 [84]
D. melanogasterDisruptive selectionPre-zygotic isolation; assortive mating; all later replications of the experiment failed1962 [85]
D. melanogasterN/AN/A1997 [86]
D. melanogasterDestroyed hybridsPre-zygotic isolation1971 [87]
D. melanogasterDestroyed hybridsPre-zygotic isolation1973 [88]
D. melanogasterDestroyed hybridsPre-zygotic isolation1979 [89]
Zea mays Destroyed hybridsPre-zygotic isolation; reproductive character displacement1969 [90]

Related Research Articles

Speciation is the evolutionary process by which populations evolve to become distinct species. The biologist Orator F. Cook coined the term in 1906 for cladogenesis, the splitting of lineages, as opposed to anagenesis, phyletic evolution within lineages. Charles Darwin was the first to describe the role of natural selection in speciation in his 1859 book On the Origin of Species. He also identified sexual selection as a likely mechanism, but found it problematic.

Allopatric speciation – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from each other to an extent that prevents or interferes with gene flow.

<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">Haldane's rule</span> Observation in evolutionary biology

Haldane's rule is an observation about the early stage of speciation, formulated in 1922 by the British evolutionary biologist J. B. S. Haldane, that states that if — in a species hybrid — only one sex is inviable or sterile, that sex is more likely to be the heterogametic sex. The heterogametic sex is the one with two different sex chromosomes; in therian mammals, for example, this is the male.

<span class="mw-page-title-main">Peripatric speciation</span> Speciation in which a new species is formed from an isolated smaller peripheral population

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. Nevertheless, the primary characteristic of peripatric speciation proposes that one of the populations is much smaller than the other. 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.

<span class="mw-page-title-main">Sympatry</span> Biological concept

In biology, two related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter one another. An initially interbreeding population that splits into two or more distinct species sharing a common range exemplifies sympatric speciation. Such speciation may be a product of reproductive isolation – which prevents hybrid offspring from being viable or able to reproduce, thereby reducing gene flow – that results in genetic divergence. Sympatric speciation may, but need not, arise through secondary contact, which refers to speciation or divergence in allopatry followed by range expansions leading to an area of sympatry. Sympatric species or taxa in secondary contact may or may not interbreed.

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

In evolutionary biology, disruptive selection, also called diversifying selection, describes changes in population genetics in which extreme values for a trait are favored over intermediate values. In this case, the variance of the trait increases and the population is divided into two distinct groups. In this more individuals acquire peripheral character value at both ends of the distribution curve.

<span class="mw-page-title-main">Character displacement</span>

Character displacement is the phenomenon where differences among similar species whose distributions overlap geographically are accentuated in regions where the species co-occur, but are minimized or lost where the species' distributions do not overlap. This pattern results from evolutionary change driven by biological competition among species for a limited resource. The rationale for character displacement stems from the competitive exclusion principle, also called Gause's Law, which contends that to coexist in a stable environment two competing species must differ in their respective ecological niche; without differentiation, one species will eliminate or exclude the other through competition.

<span class="mw-page-title-main">Parapatric speciation</span> Speciation within a population where subpopulations are reproductively isolated

In parapatric speciation, two subpopulations of a species evolve reproductive isolation from one another while continuing to exchange genes. This mode of speciation has three distinguishing characteristics: 1) mating occurs non-randomly, 2) gene flow occurs unequally, and 3) populations exist in either continuous or discontinuous geographic ranges. This distribution pattern may be the result of unequal dispersal, incomplete geographical barriers, or divergent expressions of behavior, among other things. Parapatric speciation predicts that hybrid zones will often exist at the junction between the two populations.

The mechanisms of reproductive isolation are a collection of evolutionary mechanisms, behaviors and physiological processes critical for speciation. They prevent members of different species from producing offspring, or ensure that any offspring are sterile. These barriers maintain the integrity of a species by reducing gene flow between related species.

In biology, a cline is a measurable gradient in a single characteristic of a species across its geographical range. Clines usually have a genetic, or phenotypic character. They can show either smooth, continuous gradation in a character, or more abrupt changes in the trait from one geographic region to the next.

<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">Reinforcement (speciation)</span> Process of increasing reproductive isolation

Reinforcement is a process of speciation where natural selection increases the reproductive isolation between two populations of species. This occurs as a result of selection acting against the production of hybrid individuals of low fitness. The idea was originally developed by Alfred Russel Wallace and is sometimes referred to as the Wallace effect. The modern concept of reinforcement originates from Theodosius Dobzhansky. He envisioned a species separated allopatrically, where during secondary contact the two populations mate, producing hybrids with lower fitness. Natural selection results from the hybrid's inability to produce viable offspring; thus members of one species who do not mate with members of the other have greater reproductive success. This favors the evolution of greater prezygotic isolation. Reinforcement is one of the few cases in which selection can favor an increase in prezygotic isolation, influencing the process of speciation directly. This aspect has been particularly appealing among evolutionary biologists.

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

The scientific study of speciation — how species evolve to become new species — began around the time of Charles Darwin in the middle of the 19th century. Many naturalists at the time recognized the relationship between biogeography and the evolution of species. The 20th century saw the growth of the field of speciation, with major contributors such as Ernst Mayr researching and documenting species' geographic patterns and relationships. The field grew in prominence with the modern evolutionary synthesis in the early part of that century. Since then, research on speciation has expanded immensely.

<span class="mw-page-title-main">Laboratory experiments of speciation</span> Biological experiments

Laboratory experiments of speciation have been conducted for all four modes of speciation: allopatric, peripatric, parapatric, and sympatric; and various other processes involving speciation: hybridization, reinforcement, founder effects, among others. Most of the experiments have been done on flies, in particular Drosophila fruit flies. However, more recent studies have tested yeasts, fungi, and even viruses.

Maria R. Servedio is a Canadian-American professor at the University of North Carolina at Chapel Hill. Her research spans a wide range of topics in evolutionary biology from sexual selection to evolution of behavior. She largely approaches these topics using mathematical models. Her current research interests include speciation and reinforcement, mate choice, and learning with a particular focus on evolutionary mechanisms that promote premating (prezygotic) isolation. Through integrative approaches and collaborations, she uses mathematical models along with experimental, genetic, and comparative techniques to draw conclusions on how evolution occurs. She has published extensively on these topics and has more than 50 peer-reviewed articles. She served as Vice President in 2018 of the American Society of Naturalists, and has been elected to serve as President in 2023.

<i>Drosophila silvestris</i> Species of fly

Drosophila silvestris is a large species of fly in the family Drosophilidae that are primarily black with yellow spots. As a rare species of fruit fly endemic to Hawaii, the fly often experiences reproductive isolation. Despite barriers in nature, D. silvestris is able to breed with D. heteroneura to create hybrid flies in the laboratory.

<span class="mw-page-title-main">Etheostomatinae</span> Subfamily of fishes

Etheostomatinae is a species-rich subfamily of freshwater ray-finned fish, the members of which are commonly known as the darters. The subfamily is part of the family Percidae which also includes the perches, ruffes and pikeperches. The family is endemic to North America. It consists of three to five different genera and well over 200 species. A common name for these fish in southern Indiana is pollywog.

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.

References

  1. ↑ Hannes Schuler, Glen R. Hood, Scott P. Egan, and Jeffrey L. Feder (2016), Meyers, Robert A (ed.), "Modes and Mechanisms of Speciation", Reviews in Cell Biology and Molecular Medicine, 2 (3): 60–93, doi:10.1002/3527600906, ISBN   9783527600908 {{citation}}: CS1 maint: multiple names: authors list (link)
  2. ↑ Jeremy L. Marshall, Michael L. Arnold, and Daniel J. Howard (2002), "Reinforcement: the road not taken", Trends in Ecology & Evolution, 17 (12): 558–563, doi:10.1016/S0169-5347(02)02636-8 {{citation}}: CS1 maint: multiple names: authors list (link)
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