Gene flow in the context of Allopatry

Evolutionary biology is a subfield of biology that analyzes the four mechanisms of evolution: natural selection, mutation, genetic drift, and gene flow. The purpose of evolutionary biology is to observe the diversity of life on Earth. The idea of natural selection was first researched by Charles Darwin as he studied bird beaks. The discipline of evolutionary biology emerged through what Julian Huxley called the modern synthesis of understanding, from previously unrelated fields of biological research, such as genetics and ecology, systematics, and paleontology. Huxley was able to take what Charles Darwin discovered and elaborate to build on his understandings.

The investigational range of current research has widened to encompass the genetic architecture of adaptation, molecular evolution, and the different forces that contribute to evolution, such as sexual selection, genetic drift, and biogeography. The newer field of evolutionary developmental biology ("evo-devo") investigates how embryogenesis is controlled, thus yielding a wider synthesis that integrates developmental biology with the fields of study covered by the earlier evolutionary synthesis.

Allopatric speciation (from Ancient Greek ἄλλος (állos) 'other' and πατρίς (patrís) 'fatherland') – also called 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.

Various geographic changes can arise such as the movement of continents, and the formation of mountains, islands, bodies of water, or glaciers. Human activity such as agriculture or developments can also change the distribution of species populations. These factors can substantially alter a region's geography, resulting in the separation of a species population into isolated subpopulations. The vicariant populations then undergo genetic changes as they become subjected to different selective pressures, experience genetic drift, and accumulate different mutations in the separated populations' gene pools. The barriers prevent the exchange of genetic information between the two populations leading to reproductive isolation. If the two populations come into contact they will be unable to reproduce—effectively speciating. Other isolating factors such as population dispersal leading to emigration can cause speciation (for instance, the dispersal and isolation of a species on an oceanic island) and is considered a special case of allopatric speciation called peripatric speciation.

In biology, a ring species is a connected series of neighbouring populations, each of which interbreeds with closely sited related populations, but for which there exist at least two end populations in the series which are too distantly related to interbreed, though there is a potential gene flow between linked neighbouring populations. Such non-breeding, though genetically connected, end populations may co-exist in the same region (sympatry) thus closing a ring. The German term Rassenkreis, meaning "circle of races", is also used.

Ring species represent speciation and have been cited as evidence of evolution. They illustrate what happens over time as populations genetically diverge, specifically because they represent, in living populations, what normally happens over time between long-deceased ancestor populations and living populations, in which the intermediates have become extinct. The evolutionary biologist Richard Dawkins remarks that ring species "are only showing us in the spatial dimension something that must always happen in the time dimension".

In biology, a mechanism is a system of causally interacting parts and processes that produce one or more effects. Phenomena can be explained by describing their mechanisms. For example, natural selection is a mechanism of evolution; other mechanisms of evolution include genetic drift, mutation, and gene flow. In ecology, mechanisms such as predation and host-parasite interactions produce change in ecological systems. In practice, no description of a mechanism is ever complete because not all details of the parts and processes of a mechanism are fully known. For example, natural selection is a mechanism of evolution that includes countless, inter-individual interactions with other individuals, components, and processes of the environment in which natural selection operates.

Genetic divergence is the process in which two or more populations of an ancestral species accumulate independent genetic changes (mutations) through time, often leading to reproductive isolation and continued mutation even after the populations have become reproductively isolated for some period of time, as there is not any genetic exchange anymore. In some cases, subpopulations cover living in ecologically distinct peripheral environments can exhibit genetic divergence from the remainder of a population, especially where the range of a population is very large (see parapatric speciation). The genetic differences among divergent populations can involve silent mutations (that have no effect on the phenotype) or give rise to significant morphological and/or physiological changes. Genetic divergence will always accompany reproductive isolation, either due to novel adaptations via selection and/or due to genetic drift, and is the principal mechanism underlying speciation.

On a molecular genetics level, genetic divergence is due to changes in a small number of genes in a species, resulting in speciation. However, researchers argue that it is unlikely that divergence is a result of a significant, single, dominant mutation in a genetic locus because if that were so, the individual with that mutation would have zero fitness. Consequently, they could not reproduce and pass the mutation on to further generations. Hence, it is more likely that divergence, and subsequently reproductive isolation, are the outcomes of multiple small mutations over evolutionary time accumulating in a population isolated from gene flow.

Genetic admixture occurs when previously isolated populations of organisms interbreed, resulting in a population with genetic ancestry from both sources. It can occur between species, such as with hybrids, or within species, such as when geographically distant individuals migrate to new regions. It results in a population with genetic backgrounds, or a gene pool, that is a mix of the source populations. Genetic admixture is recognized as an important contributor to rapid evolutionary responses, both at the level of gene flow between populations and between species.

Biological dispersal refers to both the movement of individuals (animals, plants, fungi, bacteria, etc.) from their birth site to their breeding site ('natal dispersal') and the movement from one breeding site to another ('breeding dispersal'). The term also encompasses the movement of propagules such as seeds and spores. Technically, dispersal is defined as any movement that has the potential to lead to gene flow. The act of dispersal involves three phases: departure, transfer, and settlement. Each phase is associated with distinct fitness costs and benefits. By simply moving from one habitat patch to another, an individual's dispersal can influence not only its own fitness but also broader processes such as population dynamics, population genetics, and species distribution. Understanding dispersal and its consequences, both for evolutionary strategies at a species level and for processes at an ecosystem level, requires understanding on the type of dispersal, the dispersal range of a given species, and the dispersal mechanisms involved. Biological dispersal can be correlated to population density. The range of variations of a species' location determines the expansion range.

Biological dispersal may be contrasted with geodispersal, which refers to the mixing of previously isolated populations (or entire biotas) following the erosion of geographic barriers to dispersal or gene flow.

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.

The mechanisms of reproductive isolation have been classified in a number of ways. Zoologist Ernst Mayr classified the mechanisms of reproductive isolation in two broad categories: pre-zygotic for those that act before fertilization (or before mating in the case of animals) and post-zygotic for those that act after it. The mechanisms are genetically controlled and can appear in species whose geographic distributions overlap (sympatric speciation) or are separate (allopatric 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 (other types of) 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 (e.g. sister species); their distribution being the result of peripatric speciation.

The concept of peripatric speciation was first outlined by the evolutionary biologist Ernst Mayr in 1954. Since then, other alternative models have been developed such as centrifugal speciation, that posits that a species' population experiences periods of geographic range expansion followed by shrinking periods, leaving behind small isolated populations on the periphery of the main population. Other models have involved the effects of sexual selection on limited population sizes. Other related models of peripherally isolated populations based on chromosomal rearrangements have been developed such as budding speciation and quantum speciation.

Microevolution is the change in allele frequencies that occurs over time within a population. This change is due to four different processes: mutation, selection (natural and artificial), gene flow and genetic drift. This change happens over a relatively short (in evolutionary terms) amount of time compared to the changes termed macroevolution.

Population genetics is the branch of biology that provides the mathematical structure for the study of the process of microevolution. Ecological genetics concerns itself with observing microevolution in the wild. Typically, observable instances of evolution are examples of microevolution; for example, bacterial strains that have antibiotic resistance.

Introgression, also known as introgressive hybridization, in genetics is the transfer of genetic material from one species into the gene pool of another by the repeated backcrossing of an interspecific hybrid with one of its parent species. Introgression is a long-term process, even when artificial; it may take many hybrid generations before significant backcrossing occurs. This process is distinct from most forms of gene flow in that it occurs between two populations of different species, rather than two populations of the same species.

Introgression also differs from simple hybridization. Simple hybridization results in a relatively even mixture; gene and allele frequencies in the first generation will be a uniform mix of two parental species, such as that observed in mules. Introgression, on the other hand, results in a complex, highly variable mixture of genes, and may only involve a minimal percentage of the donor genome.

The domestication of the dog was the process which led to the domestic dog. This included the dog's genetic divergence from the wolf, its domestication, and the emergence of the first dogs. Genetic studies suggest that all ancient and modern dogs share a common ancestry, descending from an ancient, now-extinct wolf population – or closely related wolf populations – which was distinct from the modern wolf lineage. The dog's similarity to the grey wolf is the result of substantial dog-into-wolf gene flow, with the modern grey wolf being the dog's nearest living relative. An extinct Late Pleistocene wolf may have been the ancestor of the dog.

The dog is a wolf-like canid. The genetic divergence between the dog's ancestor and modern wolves occurred between 20,000 and 40,000 years ago, just before or during the Last Glacial Maximum (20,000–27,000 years ago). This timespan represents the upper time-limit for the commencement of domestication because it is the time of divergence but not the time of domestication, which occurred later.

In biology, two closely related species or populations are considered sympatric when they exist in the same geographic area and thus frequently encounter each other. 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.

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.

In biogeography, the terms parapatric and parapatry are often used to describe the relationship between organisms whose ranges do not significantly overlap but are immediately adjacent to each other; they do not occur together except in a narrow contact zone. Parapatry is a geographical distribution opposed to sympatry (same area) and allopatry or peripatry (two similar cases of distinct areas).

Coalescent theory is a model of how alleles sampled from a population may have originated from a common ancestor. In the simplest case, coalescent theory assumes no recombination, no natural selection, and no gene flow or population structure, meaning that each variant is equally likely to have been passed from one generation to the next. The model looks backward in time, merging alleles into a single ancestral copy according to a random process in coalescence events. Under this model, the expected time between successive coalescence events increases almost exponentially back in time (with wide variance). Variance in the model comes from both the random passing of alleles from one generation to the next, and the random occurrence of mutations in these alleles.

The mathematical theory of the coalescent was developed independently by several groups in the early 1980s as a natural extension of classical population genetics theory and models, but can be primarily attributed to John Kingman. Advances in coalescent theory include recombination, selection, overlapping generations and virtually any arbitrarily complex evolutionary or demographic model in population genetic analysis.

During the Pleistocene, wolves were widely distributed across the Northern Hemisphere. Some Pleistocene wolves, such as Beringian wolves and those from Japan, exhibited large body size in comparison to modern gray wolf populations. Genetic analysis of the remains of Late Pleistocene wolves suggest that across their range populations of wolves maintained considerable gene flow between each other and thus there was limited genetic divergence between them. Modern wolves mostly draw their ancestry from some Siberian populations of Late Pleistocene gray wolves, which largely replaced other gray wolf populations after the Last Glacial Maximum.

It is widely agreed that the evolutionary lineage of the grey wolf can be traced back 2 million years to the Early Pleistocene species Canis etruscus, and its successor the Middle Pleistocene Canis mosbachensis. The grey wolf Canis lupus is a highly adaptable species that is able to exist in a range of environments and which possesses a wide distribution across the Holarctic. Studies of modern grey wolves have identified distinct sub-populations that live in close proximity to each other. This variation in sub-populations is closely linked to differences in habitat – precipitation, temperature, vegetation, and prey specialization – which affect cranio-dental plasticity.

The earliest specimens of the modern grey wolf date to around 400,000 years ago, or possibly earlier to 1 million years ago. Most modern wolves share most of their common ancestry within the last 25-23,000 years from earlier Siberian wolf populations. While some sources have suggested that this is the result of a population bottleneck, others suggest that this is a normal consequence of gene flow homogenising wolf genomes across their range.