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Speciation is the
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
ary process by which populations evolve to become distinct
species In biology, a species is the basic unit of Taxonomy (biology), 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 biologist Orator F. Cook coined the term in 1906 for
cladogenesis Cladogenesis is an evolutionary splitting of a parent species into two distinct species, forming a clade. This event usually occurs when a few organisms end up in new, often distant areas or when environmental changes cause several extinctions, ...
, the splitting of lineages, as opposed to
anagenesis Anagenesis is the gradual evolution of a species that continues to exist as an interbreeding population. This contrasts with cladogenesis, which occurs when there is branching or splitting, leading to two or more lineages and resulting in separate ...
, phyletic evolution within lineages.
Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended fr ...
was the first to describe the role of
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Charl ...
in speciation in his 1859 book ''
On the Origin of Species ''On the Origin of Species'' (or, more completely, ''On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life''),The book's full original title was ''On the Origin of Species by Me ...
''. He also identified
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (int ...
as a likely mechanism, but found it problematic. There are four geographic modes of speciation in nature, based on the extent to which speciating
population Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Governments typically quantify the size of the resident population within their jurisdiction using a ...
s are isolated from one another:
allopatric 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 ...
, peripatric,
parapatric 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 ...
, and
sympatric 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 s ...
. Speciation may also be induced artificially, through
animal husbandry Animal husbandry is the branch of agriculture concerned with animals that are raised for meat, fibre, milk, or other products. It includes day-to-day care, selective breeding, and the raising of livestock. Husbandry has a long history, startin ...
, agriculture, or laboratory experiments. Whether
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 ...
is a minor or major contributor to speciation is the subject of much ongoing discussion. Rapid sympatric speciation can take place through
polyploid Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
y, such as by doubling of chromosome number; the result is progeny which are immediately
reproductively isolated 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 ...
from the parent population. New species can also be created through hybridization, followed by reproductive isolation, if the hybrid is favoured by natural selection.


Historical background

In addressing the origin of species, there are two key issues: # the evolutionary mechanisms of speciation # how the separateness and individuality of species is maintained Since Charles Darwin's time, efforts to understand the nature of species have primarily focused on the first aspect, and it is now widely agreed that the critical factor behind the origin of new species is reproductive isolation.


Darwin's dilemma: why species exist

In ''
On the Origin of Species ''On the Origin of Species'' (or, more completely, ''On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life''),The book's full original title was ''On the Origin of Species by Me ...
'' (1859), Darwin interpreted biological evolution in terms of natural selection, but was perplexed by the clustering of organisms into species. Chapter 6 of Darwin's book is entitled "Difficulties of the Theory". In discussing these "difficulties" he noted This dilemma can be described as the absence or rarity of transitional varieties in habitat space. Another dilemma, related to the first one, is the absence or rarity of transitional varieties in time. Darwin pointed out that by the theory of natural selection "innumerable transitional forms must have existed", and wondered "why do we not find them embedded in countless numbers in the crust of the earth". That clearly defined species actually do exist in nature in both space and time implies that some fundamental feature of natural selection operates to generate and maintain species.


Effect of sexual reproduction on species formation

It has been argued that the resolution of Darwin's first dilemma lies in the fact that
out-crossing Out-crossing or out-breeding is the technique of crossing between different breeds. This is the practice of introducing distantly related genetic material into a breeding line, thereby increasing genetic diversity. Outcrossing can be a useful ...
sexual reproduction Sexual reproduction is a type of reproduction that involves a complex life cycle in which a gamete ( haploid reproductive cells, such as a sperm or egg cell) with a single set of chromosomes combines with another gamete to produce a zygote tha ...
has an intrinsic cost of rarity. The cost of rarity arises as follows. If, on a resource gradient, a large number of separate species evolve, each exquisitely adapted to a very narrow band on that gradient, each species will, of necessity, consist of very few members. Finding a mate under these circumstances may present difficulties when many of the individuals in the neighborhood belong to other species. Under these circumstances, if any species' population size happens, by chance, to increase (at the expense of one or other of its neighboring species, if the environment is saturated), this will immediately make it easier for its members to find sexual partners. The members of the neighboring species, whose population sizes have decreased, experience greater difficulty in finding mates, and therefore form pairs less frequently than the larger species. This has a snowball effect, with large species growing at the expense of the smaller, rarer species, eventually driving them to
extinction Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
. Eventually, only a few species remain, each distinctly different from the other. The cost of rarity not only involves the costs of failure to find a mate, but also indirect costs such as the cost of communication in seeking out a partner at low population densities. Rarity brings with it other costs. Rare and unusual features are very seldom advantageous. In most instances, they indicate a ( non-silent)
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, mitos ...
, which is almost certain to be deleterious. It therefore behooves sexual creatures to avoid mates sporting rare or unusual features (
koinophilia Koinophilia is an evolutionary hypothesis proposing that during sexual selection, animals preferentially seek mates with a minimum of unusual or mutant features, including functionality, appearance and behavior. Koinophilia intends to explain th ...
). Sexual populations therefore rapidly shed rare or peripheral phenotypic features, thus canalizing the entire external appearance, as illustrated in the accompanying illustration of the
African pygmy kingfisher The African pygmy kingfisher (''Ispidina picta'') is a small insectivorous kingfisher found in the Afrotropics, mostly in woodland habitats. Taxonomy The African pygmy kingfisher was described by the French polymath Georges-Louis Leclerc, Comte ...
, ''Ispidina picta''. This uniformity of all the adult members of a sexual species has stimulated the proliferation of
field guide A field guide is a book designed to help the reader identify wildlife (flora or fauna) or other objects of natural occurrence (e.g. rocks and minerals). It is generally designed to be brought into the " field" or local area where such objects e ...
s on birds, mammals, reptiles, insects, and many other
taxa In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...
, in which a species can be described with a single illustration (or two, in the case of
sexual dimorphism Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most anim ...
). Once a population has become as homogeneous in appearance as is typical of most species (and is illustrated in the photograph of the African pygmy kingfisher), its members will avoid mating with members of other populations that look different from themselves. Thus, the avoidance of mates displaying rare and unusual phenotypic features inevitably leads to reproductive isolation, one of the hallmarks of speciation. In the contrasting case of organisms that reproduce asexually, there is no cost of rarity; consequently, there are only benefits to fine-scale adaptation. Thus, asexual organisms very frequently show the continuous variation in form (often in many different directions) that Darwin expected evolution to produce, making their classification into "species" (more correctly, morphospecies) very difficult.


Modes

All forms of natural speciation have taken place over the course of
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
; however, debate persists as to the relative importance of each mechanism in driving
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') le ...
. One example of natural speciation is the diversity of the
three-spined stickleback The three-spined stickleback (''Gasterosteus aculeatus'') is a fish native to most inland and coastal waters north of 30°N. It has long been a subject of scientific study for many reasons. It shows great morphological variation throughout its ra ...
, a marine fish that, after the last glacial period, has undergone speciation into new
freshwater Fresh water or freshwater is any naturally occurring liquid or frozen water containing low concentrations of dissolved salts and other total dissolved solids. Although the term specifically excludes seawater and brackish water, it does inclu ...
colonies in isolated lakes and streams. Over an estimated 10,000 generations, the sticklebacks show structural differences that are greater than those seen between different
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomencl ...
of fish including variations in fins, changes in the number or size of their bony plates, variable jaw structure, and color differences.


Allopatric

During allopatric (from the ancient Greek ''allos'', "other" + ''patrā'', "fatherland") speciation, a population splits into two geographically isolated populations (for example, by
habitat fragmentation Habitat fragmentation describes the emergence of discontinuities (fragmentation) in an organism's preferred environment (habitat), causing population fragmentation and ecosystem decay. Causes of habitat fragmentation include geological process ...
due to geographical change such as
mountain formation Mountain formation refers to the geological processes that underlie the formation of mountains. These processes are associated with large-scale movements of the Earth's crust (tectonic plates). Folding, faulting, volcanic activity, igneous int ...
). The isolated populations then undergo genotypic or
phenotypic In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology or physical form and structure, its developmental processes, its biochemical and physiological prope ...
divergence as: (a) they become subjected to dissimilar selective pressures; (b) they independently undergo genetic drift; (c) different
mutation In biology, a mutation is an alteration in the nucleic acid sequence of the genome of an organism, virus, or extrachromosomal DNA. Viral genomes contain either DNA or RNA. Mutations result from errors during DNA or viral replication, mitos ...
s arise in the two populations. When the populations come back into contact, they have evolved such that they are reproductively isolated and are no longer capable of exchanging
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
s.
Island genetics Small populations can behave differently from larger populations. They are often the result of population bottlenecks from larger populations, leading to loss of heterozygosity and reduced genetic diversity and loss or fixation of alleles and s ...
is the term associated with the tendency of small, isolated genetic pools to produce unusual traits. Examples include
insular dwarfism Insular dwarfism, a form of phyletic dwarfism, is the process and condition of large animals evolving or having a reduced body size when their population's range is limited to a small environment, primarily islands. This natural process is disti ...
and the radical changes among certain famous island chains, for example on Komodo. The
Galápagos Islands The Galápagos Islands ( Spanish: , , ) are an archipelago of volcanic islands. They are distributed on each side of the equator in the Pacific Ocean, surrounding the centre of the Western Hemisphere, and are part of the Republic of Ecuador ...
are particularly famous for their influence on Charles Darwin. During his five weeks there he heard that
Galápagos tortoise The Galápagos tortoise or Galápagos giant tortoise (''Chelonoidis niger'') is a species of very large tortoise in the genus '' Chelonoidis'' (which also contains three smaller species from mainland South America). It comprises 15 subspecies (1 ...
s could be identified by island, and noticed that
finch The true finches are small to medium-sized passerine birds in the family Fringillidae. Finches have stout conical bills adapted for eating seeds and nuts and often have colourful plumage. They occupy a great range of habitats where they are usu ...
es differed from one island to another, but it was only nine months later that he reflected that such facts could show that species were changeable. When he returned to England, his speculation on evolution deepened after experts informed him that these were separate species, not just varieties, and famously that other differing Galápagos birds were all species of finches. Though the finches were less important for Darwin, more recent research has shown the birds now known as
Darwin's finches Darwin's finches (also known as the Galápagos finches) are a group of about 18 species of passerine birds. They are well known for their remarkable diversity in beak form and function. They are often classified as the subfamily Geospizinae or t ...
to be a classic case of adaptive evolutionary radiation.


Peripatric

In peripatric speciation, a subform of allopatric speciation, new species are formed in isolated, smaller peripheral populations that are prevented from exchanging genes with the main population. It is related to the concept of a
founder effect In population genetics, the founder effect is the loss of genetic variation that occurs when a new population is established by a very small number of individuals from a larger population. It was first fully outlined by Ernst Mayr in 1942, using ...
, since small populations often undergo
bottleneck Bottleneck literally refers to the narrowed portion (neck) of a bottle near its opening, which limit the rate of outflow, and may describe any object of a similar shape. The literal neck of a bottle was originally used to play what is now known as ...
s. Genetic drift is often proposed to play a significant role in peripatric speciation. Case studies include Mayr's investigation of bird fauna; the Australian bird '' Petroica multicolor''; and reproductive isolation in populations of ''
Drosophila ''Drosophila'' () is a genus of flies, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or (less frequently) pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species ...
'' subject to population bottlenecking.


Parapatric

In parapatric speciation, there is only partial separation of the zones of two diverging populations afforded by geography; individuals of each species may come in contact or cross habitats from time to time, but reduced fitness of the
heterozygote Zygosity (the noun, zygote, is from the Greek "yoked," from "yoke") () is the degree to which both copies of a chromosome or gene have the same genetic sequence. In other words, it is the degree of similarity of the alleles in an organism. Mo ...
leads to selection for behaviours or mechanisms that prevent their
interbreeding In biology, a hybrid is the offspring resulting from combining the qualities of two organisms of different breeds, varieties, species or genera through sexual reproduction. Hybrids are not always intermediates between their parents (such as in ...
. Parapatric speciation is modelled on continuous variation within a "single", connected habitat acting as a source of natural selection rather than the effects of isolation of habitats produced in peripatric and allopatric speciation. Parapatric speciation may be associated with differential landscape-dependent
selection Selection may refer to: Science * Selection (biology), also called natural selection, selection in evolution ** Sex selection, in genetics ** Mate selection, in mating ** Sexual selection in humans, in human sexuality ** Human mating strategi ...
. Even if there is a
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 two populations, strong differential selection may impede assimilation and different species may eventually develop. Habitat differences may be more important in the development of reproductive isolation than the isolation time. Caucasian rock lizards ''
Darevskia ''Darevskia'' is a genus of wall lizards of the family Lacertidae living in the Caucasus, Iran and Turkey in forest and grassy habitats with numerous rock outcrops. Among rock lizards, 7 parthenogenetic species are known. Description These ar ...
rudis'', ''D. valentini'' and ''D. portschinskii'' all
hybrid Hybrid may refer to: Science * Hybrid (biology), an offspring resulting from cross-breeding ** Hybrid grape, grape varieties produced by cross-breeding two ''Vitis'' species ** Hybridity, the property of a hybrid plant which is a union of two dif ...
ize with each other in their
hybrid zone A hybrid zone exists where the ranges of two interbreeding species or diverged intraspecific lineages meet and cross-fertilize. Hybrid zones can form ''in situ'' due to the evolution of a new lineage but generally they result from secondary contact ...
; however, hybridization is stronger between ''D. portschinskii'' and ''D. rudis'', which separated earlier but live in similar habitats than between ''D. valentini'' and two other species, which separated later but live in climatically different habitats. Ecologists refer to parapatric and peripatric speciation in terms of
ecological niche In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for ...
s. A niche must be available in order for a new species to be successful.
Ring species 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 relate ...
such as ''
Larus ''Larus'' is a large genus of gulls with worldwide distribution (by far the greatest species diversity is in the Northern Hemisphere). Many of its species are abundant and well-known birds in their ranges. Until about 2005–2007, most gulls ...
'' gulls have been claimed to illustrate speciation in progress, though the situation may be more complex. The grass ''
Anthoxanthum odoratum ''Anthoxanthum odoratum'', known as sweet vernal grass, is a short-lived perennial grass that is native to acidic grassland in Eurasia and northern Africa. It is grown as a lawn grass and a house plant, due to its sweet scent, and can also be f ...
'' may be starting parapatric speciation in areas of mine contamination.


Sympatric

Sympatric speciation is the formation of two or more descendant species from a single ancestral species all occupying the same geographic location. Often-cited examples of sympatric speciation are found in insects that become dependent on different
host 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 ...
plants in the same area. The best known example of sympatric speciation is that of the
cichlid Cichlids are fish from the family Cichlidae in the order Cichliformes. Cichlids were traditionally classed in a suborder, the Labroidei, along with the wrasses ( Labridae), in the order Perciformes, but molecular studies have contradicted this ...
s of
East Africa East Africa, Eastern Africa, or East of Africa, is the eastern subregion of the African continent. In the United Nations Statistics Division scheme of geographic regions, 10-11-(16*) territories make up Eastern Africa: Due to the historical ...
inhabiting the
Rift Valley lakes The Rift Valley lakes are a series of lakes in the East African Rift valley that runs through eastern Africa from Ethiopia in the north to Malawi in the south, and includes the African Great Lakes in the south. These include some of the world' ...
, particularly
Lake Victoria Lake Victoria is one of the African Great Lakes. With a surface area of approximately , Lake Victoria is Africa's largest lake by area, the world's largest tropical lake, and the world's second-largest fresh water lake by surface area after L ...
,
Lake Malawi Lake Malawi, also known as Lake Nyasa in Tanzania and Lago Niassa in Mozambique, is an African Great Lake and the southernmost lake in the East African Rift system, located between Malawi, Mozambique and Tanzania. It is the fifth largest fre ...
and
Lake Tanganyika Lake Tanganyika () is an African Great Lake. It is the second-oldest freshwater lake in the world, the second-largest by volume, and the second-deepest, in all cases after Lake Baikal in Siberia. It is the world's longest freshwater lake. T ...
. There are over 800 described species, and according to estimates, there could be well over 1,600 species in the region. Their evolution is cited as an example of both
natural Nature, in the broadest sense, is the physics, physical world or universe. "Nature" can refer to the phenomenon, phenomena of the physical world, and also to life in general. The study of nature is a large, if not the only, part of science. ...
and
sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (int ...
. A 2008 study suggests that sympatric speciation has occurred in Tennessee cave salamanders. Sympatric speciation driven by ecological factors may also account for the extraordinary diversity of crustaceans living in the depths of Siberia's
Lake Baikal Lake Baikal (, russian: Oзеро Байкал, Ozero Baykal ); mn, Байгал нуур, Baigal nuur) is a rift lake in Russia. It is situated in southern Siberia, between the federal subjects of Irkutsk Oblast to the northwest and the Rep ...
. Budding speciation has been proposed as a particular form of sympatric speciation, whereby small groups of individuals become progressively more isolated from the ancestral stock by breeding preferentially with one another. This type of speciation would be driven by the conjunction of various advantages of inbreeding such as the expression of advantageous recessive phenotypes, reducing the recombination load, and reducing the cost of sex. The hawthorn fly (''
Rhagoletis pomonella The apple maggot (''Rhagoletis pomonella''), also known as the railroad worm (but distinct from the '' Phrixothrix'' beetle larva, also called railroad worm), is a species of fruit fly, and a pest of several types of fruits, especially apples. ...
''), also known as the apple maggot fly, appears to be undergoing sympatric speciation. Different populations of hawthorn fly feed on different fruits. A distinct population emerged in North America in the 19th century some time after apples, a non-native species, were introduced. This apple-feeding population normally feeds only on apples and not on the historically preferred fruit of hawthorns. The current hawthorn feeding population does not normally feed on apples. Some evidence, such as that six out of thirteen
allozyme Alloenzymes (or also called allozymes) are variant forms of an enzyme which differ structurally but not functionally from other allozymes coded for by different alleles at the same locus. These are opposed to isozymes, which are enzymes that perf ...
loci are different, that hawthorn flies mature later in the season and take longer to mature than apple flies; and that there is little evidence of interbreeding (researchers have documented a 4–6% hybridization rate) suggests that sympatric speciation is occurring.


Methods of selection


Reinforcement

Reinforcement, also called the Wallace effect, is the process by which natural selection increases reproductive isolation. Adapted from ''Evolution'' (2004), 3rd edition (Malden, MA:
Blackwell Publishing Wiley-Blackwell is an international scientific, technical, medical, and scholarly publishing business of John Wiley & Sons. It was formed by the merger of John Wiley & Sons Global Scientific, Technical, and Medical business with Blackwell Publis ...
), .
It may occur after two populations of the same species are separated and then come back into contact. If their reproductive isolation was complete, then they will have already developed into two separate incompatible species. If their reproductive isolation is incomplete, then further mating between the populations will produce hybrids, which may or may not be fertile. If the hybrids are infertile, or fertile but less fit than their ancestors, then there will be further reproductive isolation and speciation has essentially occurred, as in
horse The horse (''Equus ferus caballus'') is a Domestication, domesticated, odd-toed ungulate, one-toed, ungulate, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two Extant taxon, extant subspecies of wild horse, ''Equus fer ...
s and
donkey The domestic donkey is a hoofed mammal in the family Equidae, the same family as the horse. It derives from the African wild ass, ''Equus africanus'', and may be classified either as a subspecies thereof, ''Equus africanus asinus'', or as a ...
s. One reasoning behind this is that if the parents of the hybrid offspring each have naturally selected traits for their own certain environments, the hybrid offspring will bear traits from both, therefore would not fit either ecological niche as well as either parent (ecological speciation). The low fitness of the hybrids would cause selection to favor
assortative mating Assortative mating (also referred to as positive assortative mating or homogamy) is a mating pattern and a form of sexual selection in which individuals with similar phenotypes or genotypes mate with one another more frequently than would be exp ...
, which would control hybridization. This is sometimes called the Wallace effect after the evolutionary biologist
Alfred Russel Wallace Alfred Russel Wallace (8 January 1823 – 7 November 1913) was a British naturalist, explorer, geographer, anthropologist, biologist and illustrator. He is best known for independently conceiving the theory of evolution through natural sele ...
who suggested in the late 19th century that it might be an important factor in speciation. Conversely, if the hybrid offspring are more fit than their ancestors, then the populations will merge back into the same species within the area they are in contact. Another important theoretical mechanism is the arise of intrinsic genetic incompatibilities, addressed in the Bateson-Dobzhansky-Muller model. Genes from allopatric populations will have different evolutionary backgrounds and are never tested together until hybridization at secondary contact, when negative epistatic interactions will be exposed. In other words, new alleles will emerge in a population and only pass through selection if they work well together with other genes in the same population, but it may not be compatible with genes in an allopatric population, be those other newly derived alleles or retained ancestral alleles. This is only revealed through new hybridization. Such incompatibilities cause lower fitness in hybrids regardless of the ecological environment, and are thus intrinsic, although they can originate from the adaptation to different environments. The accumulation of such incompatibilities increases faster and faster with time, creating a "snowball" effect. There is a large amount of evidence supporting this theory, primarily from laboratory populations such as ''Drosophila'' and ''Mus'', and some genes involved in incompatibilities have been identified. Reinforcement favoring reproductive isolation is required for both parapatric and sympatric speciation. Without reinforcement, the geographic area of contact between different forms of the same species, called their "hybrid zone", will not develop into a boundary between the different species. Hybrid zones are regions where diverged populations meet and interbreed. Hybrid offspring are common in these regions, which are usually created by diverged species coming into
secondary contact Secondary contact is the process in which two allopatricaly distributed populations of a species are geographically reunited. This contact allows for the potential for the exchange of genes, dependent on how reproductively isolated the two popul ...
. Without reinforcement, the two species would have uncontrollable
inbreeding Inbreeding is the production of offspring from the mating or breeding of individuals or organisms that are closely related genetically. By analogy, the term is used in human reproduction, but more commonly refers to the genetic disorders and ...
. Reinforcement may be induced in artificial selection experiments as described below.


Ecological

Ecological selection is "the interaction of individuals with their environment during resource acquisition". Natural selection is inherently involved in the process of speciation, whereby, "under ecological speciation, populations in different environments, or populations exploiting different resources, experience contrasting natural selection pressures on the traits that directly or indirectly bring about the evolution of reproductive isolation". Evidence for the role ecology plays in the process of speciation exists. Studies of stickleback populations support ecologically-linked speciation arising as a by-product, alongside numerous studies of parallel speciation, where isolation evolves between independent populations of species adapting to contrasting environments than between independent populations adapting to similar environments. Ecological speciation occurs with much of the evidence, "...accumulated from top-down studies of adaptation and reproductive isolation".


Sexual selection

Sexual selection Sexual selection is a mode of natural selection in which members of one biological sex choose mates of the other sex to mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex (int ...
can drive speciation in a clade, independently of
natural selection Natural selection is the differential survival and reproduction of individuals due to differences in phenotype. It is a key mechanism of evolution, the change in the heritable traits characteristic of a population over generations. Charl ...
. However the term "speciation", in this context, tends to be used in two different, but not mutually exclusive senses. The first and most commonly used sense refers to the "birth" of new species. That is, the splitting of an existing species into two separate species, or the budding off of a new species from a parent species, both driven by a biological "fashion fad" (a preference for a feature, or features, in one or both sexes, that do not necessarily have any adaptive qualities). In the second sense, "speciation" refers to the wide-spread tendency of sexual creatures to be grouped into clearly defined species, rather than forming a continuum of
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology or physical form and structure, its developmental processes, its biochemical and physiological prope ...
s both in time and space – which would be the more obvious or logical consequence of natural selection. This was indeed recognized by Darwin as problematic, and included in his ''
On the Origin of Species ''On the Origin of Species'' (or, more completely, ''On the Origin of Species by Means of Natural Selection, or the Preservation of Favoured Races in the Struggle for Life''),The book's full original title was ''On the Origin of Species by Me ...
'' (1859), under the heading "Difficulties with the Theory". There are several suggestions as to how
mate choice Mate choice is one of the primary mechanisms under which evolution can occur. It is characterized by a "selective response by animals to particular stimuli" which can be observed as behavior.Bateson, Paul Patrick Gordon. "Mate Choice." Mate Choic ...
might play a significant role in resolving Darwin's dilemma. — Commentary by Mazhuvancherry K. Unnikrishnan and H. S. Akhila If speciation takes place in the absence of natural selection, it might be referred to as nonecological speciation.


Artificial speciation

New species have been created by
animal husbandry Animal husbandry is the branch of agriculture concerned with animals that are raised for meat, fibre, milk, or other products. It includes day-to-day care, selective breeding, and the raising of livestock. Husbandry has a long history, startin ...
, but the dates and methods of the initiation of such species are not clear. Often, the domestic counterpart can still interbreed and produce fertile offspring with its wild ancestor. This is the case with domestic
cattle Cattle (''Bos taurus'') are large, domesticated, cloven-hooved, herbivores. They are a prominent modern member of the subfamily Bovinae and the most widespread species of the genus ''Bos''. Adult females are referred to as cows and adult ma ...
, which can be considered the same species as several varieties of wild ox,
gaur The gaur (''Bos gaurus''; ), also known as the Indian bison, is a bovine native to South Asia and Southeast Asia, and has been listed as Vulnerable on the IUCN Red List since 1986. The global population was estimated at a maximum of 21,000 m ...
, and
yak The domestic yak (''Bos grunniens''), also known as the Tartary ox, grunting ox or hairy cattle, is a species of long-haired domesticated cattle found throughout the Himalayan region of the Indian subcontinent, the Tibetan Plateau, Kachin Sta ...
; and with domestic
sheep Sheep or domestic sheep (''Ovis aries'') are domesticated, ruminant mammals typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus ''Ovis'', in everyday usage it almost always refers to domesticated sh ...
that can interbreed with the
mouflon The mouflon (''Ovis gmelini'') is a wild sheep native to Cyprus, the Caspian region from eastern Turkey, Armenia, Azerbaijan, and Iran. It is thought to be the ancestor of all modern domestic sheep breeds. Taxonomy ''Ovis gmelini'' was the sc ...
. The best-documented creations of new species in the laboratory were performed in the late 1980s. William R. Rice and George W. Salt bred ''
Drosophila melanogaster ''Drosophila melanogaster'' is a species of fly (the taxonomic order Diptera) in the family Drosophilidae. The species is often referred to as the fruit fly or lesser fruit fly, or less commonly the "vinegar fly" or "pomace fly". Starting with Ch ...
''
fruit flies Fruit fly may refer to: Organisms * Drosophilidae, a family of small flies, including: ** ''Drosophila'', the genus of small fruit flies and vinegar flies ** ''Drosophila melanogaster'' or common fruit fly ** '' Drosophila suzukii'' or Asian fruit ...
using a maze with three different choices of habitat such as light/dark and wet/dry. Each generation was placed into the maze, and the groups of flies that came out of two of the eight exits were set apart to breed with each other in their respective groups. After thirty-five generations, the two groups and their offspring were isolated reproductively because of their strong habitat preferences: they mated only within the areas they preferred, and so did not mate with flies that preferred the other areas. The history of such attempts is described by Rice and Elen E. Hostert (1993). Diane Dodd used a laboratory experiment to show how reproductive isolation can develop in ''
Drosophila pseudoobscura ''Drosophila'' () is a genus of flies, belonging to the family Drosophilidae, whose members are often called "small fruit flies" or (less frequently) pomace flies, vinegar flies, or wine flies, a reference to the characteristic of many species ...
'' fruit flies after several generations by placing them in different media, starch- and maltose-based media. Dodd's experiment has been replicated many times, including with other kinds of fruit flies and foods. Such rapid evolution of reproductive isolation may sometimes be a relic of infection by ''
Wolbachia ''Wolbachia'' is a genus of intracellular bacteria that infects mainly arthropod species, including a high proportion of insects, and also some nematodes. It is one of the most common parasitic microbes, and is possibly the most common reprodu ...
'' bacteria. An alternative explanation is that these observations are consistent with sexually-reproducing animals being inherently reluctant to mate with individuals whose appearance or behavior is different from the norm. The risk that such deviations are due to heritable maladaptations is high. Thus, if an animal, unable to predict natural selection's future direction, is conditioned to produce the fittest offspring possible, it will avoid mates with unusual habits or features. Sexual creatures then inevitably group themselves into reproductively isolated species.


Genetics

Few speciation genes have been found. They usually involve the reinforcement process of late stages of speciation. In 2008, a speciation gene causing reproductive isolation was reported. It causes hybrid sterility between related subspecies. The order of speciation of three groups from a common ancestor may be unclear or unknown; a collection of three such species is referred to as a "trichotomy".


Speciation via polyploidy

Polyploid Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
y is a mechanism that has caused many rapid speciation events in
sympatry 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 sh ...
because offspring of, for example, tetraploid x diploid matings often result in triploid sterile progeny. However, among plants, not all polyploids are reproductively isolated from their parents, and gene flow may still occur, such as through triploid hybrid x diploid matings that produce tetraploids, or matings between meiotically unreduced gametes from diploids and gametes from tetraploids (see also
hybrid speciation Hybrid speciation is a form of speciation where hybridization between two different species leads to a new species, reproductively isolated from the parent species. Previously, reproductive isolation between two species and their parents was tho ...
). It has been suggested that many of the existing plant and most animal species have undergone an event of polyploidization in their evolutionary history. Reproduction of successful polyploid species is sometimes asexual, by
parthenogenesis Parthenogenesis (; from the Greek grc, παρθένος, translit=parthénos, lit=virgin, label=none + grc, γένεσις, translit=génesis, lit=creation, label=none) is a natural form of asexual reproduction in which growth and developmen ...
or
apomixis In botany, apomixis is asexual reproduction without fertilization. Its etymology is Greek for "away from" + "mixing". This definition notably does not mention meiosis. Thus "normal asexual reproduction" of plants, such as propagation from cuttin ...
, as for unknown reasons many asexual organisms are polyploid. Rare instances of polyploid mammals are known, but most often result in prenatal death.


Hybrid speciation

Hybridization between two different species sometimes leads to a distinct
phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology or physical form and structure, its developmental processes, its biochemical and physiological prope ...
. This phenotype can also be fitter than the parental lineage and as such natural selection may then favor these individuals. Eventually, if reproductive isolation is achieved, it may lead to a separate species. However, reproductive isolation between hybrids and their parents is particularly difficult to achieve and thus hybrid speciation is considered an extremely rare event. The
Mariana mallard The Mariana mallard or Oustalet's duck (''Anas oustaleti'') is an extinct species of duck of the genus ''Anas'' that was endemic to the Mariana Islands. Its taxonomic status is debated, and it has variously been treated as a full species, a s ...
is thought to have arisen from hybrid speciation. Hybridization is an important means of speciation in plants, since
polyploid Polyploidy is a condition in which the cells of an organism have more than one pair of ( homologous) chromosomes. Most species whose cells have nuclei (eukaryotes) are diploid, meaning they have two sets of chromosomes, where each set contains ...
y (having more than two copies of each chromosome) is tolerated in plants more readily than in animals. Polyploidy is important in hybrids as it allows reproduction, with the two different sets of chromosomes each being able to pair with an identical partner during meiosis. Polyploids also have more genetic diversity, which allows them to avoid
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. ...
in small populations. Hybridization without change in chromosome number is called homoploid hybrid speciation. It is considered very rare but has been shown in ''
Heliconius ''Heliconius'' comprises a colorful and widespread genus of brush-footed butterflies commonly known as the longwings or heliconians. This genus is distributed throughout the tropical and subtropical regions of the New World, from South America a ...
''
butterflies Butterflies are insects in the macrolepidopteran clade Rhopalocera from the order Lepidoptera, which also includes moths. Adult butterflies have large, often brightly coloured wings, and conspicuous, fluttering flight. The group comprises ...
and
sunflower The common sunflower (''Helianthus annuus'') is a large annual forb of the genus ''Helianthus'' grown as a crop for its edible oily seeds. Apart from cooking oil production, it is also used as livestock forage (as a meal or a silage plant), as ...
s. Polyploid speciation, which involves changes in chromosome number, is a more common phenomenon, especially in plant species.


Gene transposition

Theodosius Dobzhansky Theodosius Grigorievich Dobzhansky (russian: Феодо́сий Григо́рьевич Добржа́нский; uk, Теодо́сій Григо́рович Добржа́нський; January 25, 1900 – December 18, 1975) was a prominent ...
, who studied fruit flies in the early days of genetic research in 1930s, speculated that parts of chromosomes that switch from one location to another might cause a species to split into two different species. He mapped out how it might be possible for sections of chromosomes to relocate themselves in a genome. Those mobile sections can cause sterility in inter-species hybrids, which can act as a speciation pressure. In theory, his idea was sound, but scientists long debated whether it actually happened in nature. Eventually a competing theory involving the gradual accumulation of mutations was shown to occur in nature so often that geneticists largely dismissed the moving gene hypothesis. However, 2006 research shows that jumping of a gene from one chromosome to another can contribute to the birth of new species. This validates the reproductive isolation mechanism, a key component of speciation.


Rates

There is debate as to the rate at which speciation events occur over geologic time. While some evolutionary biologists claim that speciation events have remained relatively constant and gradual over time (known as "Phyletic gradualism" – see diagram), some
palaeontologist Paleontology (), also spelled palaeontology or palæontology, is the scientific study of life that existed prior to, and sometimes including, the start of the Holocene epoch (roughly 11,700 years before present). It includes the study of foss ...
s such as
Niles Eldredge Niles Eldredge (; born August 25, 1943) is an American biologist and paleontologist, who, along with Stephen Jay Gould, proposed the theory of punctuated equilibrium in 1972. Education Eldredge began his undergraduate studies in Latin at Colum ...
and
Stephen Jay Gould Stephen Jay Gould (; September 10, 1941 – May 20, 2002) was an American paleontologist, evolutionary biologist, and historian of science. He was one of the most influential and widely read authors of popular science of his generation. Gould sp ...
have argued that species usually remain unchanged over long stretches of time, and that speciation occurs only over relatively brief intervals, a view known as ''
punctuated equilibrium In evolutionary biology, punctuated equilibrium (also called punctuated equilibria) is a theory that proposes that once a species appears in the fossil record, the population will become stable, showing little evolutionary change for most of i ...
''. (See diagram, and Darwin's dilemma.)


Punctuated evolution

Evolution can be extremely rapid, as shown in the creation of domesticated animals and plants in a very short geological space of time, spanning only a few tens of thousands of years.
Maize Maize ( ; ''Zea mays'' subsp. ''mays'', from es, maíz after tnq, mahiz), also known as corn (North American and Australian English), is a cereal grain first domesticated by indigenous peoples in southern Mexico about 10,000 years ago. The ...
(''Zea mays''), for instance, was created in
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
in only a few thousand years, starting about 7,000 to 12,000 years ago. This raises the question of why the long term rate of evolution is far slower than is theoretically possible. Evolution is imposed on species or groups. It is not planned or striven for in some Lamarckist way. The mutations on which the process depends are random events, and, except for the "
silent mutation Silent mutations are mutations in DNA that do not have an observable effect on the organism's phenotype. They are a specific type of neutral mutation. The phrase ''silent mutation'' is often used interchangeably with the phrase '' synonymous mutat ...
s" which do not affect the functionality or appearance of the carrier, are thus usually disadvantageous, and their chance of proving to be useful in the future is vanishingly small. Therefore, while a species or group might benefit from being able to adapt to a new environment by accumulating a wide range of genetic variation, this is to the detriment of the ''individuals'' who have to carry these mutations until a small, unpredictable minority of them ultimately contributes to such an adaptation. Thus, the ''capability'' to evolve would require
group selection Group selection is a proposed mechanism of evolution in which natural selection acts at the level of the group, instead of at the level of the individual or gene. Early authors such as V. C. Wynne-Edwards and Konrad Lorenz argued that the behav ...
, a concept discredited by (for example) George C. Williams,
John Maynard Smith John Maynard Smith (6 January 1920 – 19 April 2004) was a British theoretical and mathematical evolutionary biologist and geneticist. Originally an aeronautical engineer during the Second World War, he took a second degree in genetics und ...
and
Richard Dawkins Richard Dawkins (born 26 March 1941) is a British evolutionary biologist and author. He is an emeritus fellow of New College, Oxford and was Professor for Public Understanding of Science in the University of Oxford from 1995 to 2008. An ath ...
"Remarks on an earlier article by [Elliot] Sober icand
David Sloan Wilson David Sloan Wilson (born 1949) is an American evolutionary biologist and a Distinguished Professor Emeritus of Biological Sciences and Anthropology at Binghamton University. He is a son of author Sloan Wilson, and co-founder of the Evolution ...
, who made a more extended argument in their recent book ''Unto Others : The Evolution and Psychology of Unselfish Behavior''"
"Commentary on Wilson & Sober: Group Selection." as selectively disadvantageous to the individual. The resolution to Darwin's second dilemma might thus come about as follows: If sexual individuals are disadvantaged by passing mutations on to their offspring, they will avoid mutant mates with strange or unusual characteristics. Mutations that affect the external appearance of their carriers will then rarely be passed on to the next and subsequent generations. They would therefore seldom be tested by natural selection. Evolution is, therefore, effectively halted or slowed down considerably. The only mutations that can accumulate in a population, on this punctuated equilibrium view, are ones that have no noticeable effect on the outward appearance and functionality of their bearers (i.e., they are "silent" or "
neutral mutation Neutral mutations are changes in DNA sequence that are neither beneficial nor detrimental to the ability of an organism to survive and reproduce. In population genetics, mutations in which natural selection does not affect the spread of the mutat ...
s", which can be, and are, used to trace the relatedness and age of populations and species.) This argument implies that evolution can only occur if mutant mates cannot be avoided, as a result of a severe scarcity of potential mates. This is most likely to occur in small, isolated communities. These occur most commonly on small islands, in remote valleys, lakes, river systems, or caves, or during the aftermath of a
mass extinction An extinction event (also known as a mass extinction or biotic crisis) is a widespread and rapid decrease in the biodiversity on Earth. Such an event is identified by a sharp change in the diversity and abundance of multicellular organisms. It ...
. Under these circumstances, not only is the choice of mates severely restricted but population bottlenecks, founder effects, genetic drift and inbreeding cause rapid, random changes in the isolated population's genetic composition. Furthermore, hybridization with a related species trapped in the same isolate might introduce additional genetic changes. If an isolated population such as this survives its genetic upheavals, and subsequently expands into an unoccupied niche, or into a niche in which it has an advantage over its competitors, a new species, or subspecies, will have come into being. In geological terms, this will be an abrupt event. A resumption of avoiding mutant mates will thereafter result, once again, in evolutionary stagnation. In apparent confirmation of this punctuated equilibrium view of evolution, the
fossil record A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved in ...
of an evolutionary progression typically consists of species that suddenly appear, and ultimately disappear, hundreds of thousands or millions of years later, without any change in external appearance. Graphically, these fossil species are represented by lines parallel with the time axis, whose lengths depict how long each of them existed. The fact that the lines remain parallel with the time axis illustrates the unchanging appearance of each of the fossil species depicted on the graph. During each species' existence new species appear at random intervals, each also lasting many hundreds of thousands of years before disappearing without a change in appearance. The exact relatedness of these concurrent species is generally impossible to determine. This is illustrated in the diagram depicting the distribution of hominin species through time since the
hominins The Hominini form a taxonomic tribe of the subfamily Homininae ("hominines"). Hominini includes the extant genera ''Homo'' (humans) and '' Pan'' (chimpanzees and bonobos) and in standard usage excludes the genus ''Gorilla'' (gorillas). The t ...
separated from the line that led to the evolution of their closest living primate relatives, the chimpanzees. For similar evolutionary time lines see, for instance, the paleontological list of African dinosaurs, Asian dinosaurs, the
Lampriformes Lampriformes is an order of ray-finned fish. Members are collectively called lamprids (which is more properly used for the Lampridae) or lampriforms, and unite such open-ocean and partially deep-sea Teleostei as the crestfishes, oarfish, op ...
and
Amiiformes The Amiiformes order of fish has only one extant species, the bowfin (''Amia calva''). These Amiiformes are found in the freshwater systems of North America, in the United States and parts of southern Canada. They live in freshwater streams, ri ...
.


See also

* Bateson–Dobzhansky–Muller model * Chronospecies * Court jester hypothesis *
Macroevolution Macroevolution usually means the evolution of large-scale structures and traits that go significantly beyond the intraspecific variation found in microevolution (including speciation). In other words, macroevolution is the evolution of taxa abov ...
*
Selection (genetic algorithm) Selection is the stage of a genetic algorithm in which individual genomes are chosen from a population for later breeding (using the crossover operator). A generic selection procedure may be implemented as follows: #The fitness function {{no foot ...
*
Species problem The species problem is the set of questions that arises when biologists attempt to define what a species is. Such a definition is called a species concept; there are at least 26 recognized species concepts. A species concept that works well for se ...


References


Bibliography

* * * * * * * The book is available from . * * Reprinted in * * *
1982 edition
via Archive.org. * * * * * * * * * * * * * * * *


Further reading

* * * * * * *


External links

* * * {{Authority control Ecology Evolutionary biology Sexual selection