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.
133-421: There are four geographic modes of speciation in nature, based on the extent to which speciating populations are isolated from one another: allopatric , peripatric , parapatric , and sympatric . Whether genetic drift is a minor or major contributor to speciation is the subject of much ongoing discussion. Rapid sympatric speciation can take place through polyploidy , such as by doubling of chromosome number;
266-572: A census to quantify the size of a resident population within a given jurisdiction. The term is also applied to non-human animals , microorganisms , and plants , and has specific uses within such fields as ecology and genetics . The word population is derived from the Late Latin populatio (a people, a multitude), which itself is derived from the Latin word populus (a people). In sociology and population geography , population refers to
399-437: A population is often defined as a set of organisms in which any pair of members can breed together. They can thus routinely exchange gametes in order to have usually fertile progeny, and such a breeding group is also known therefore as a gamodeme. This also implies that all members belong to the same species. If the gamodeme is very large (theoretically, approaching infinity), and all gene alleles are uniformly distributed by
532-477: A ( non-silent ) mutation , which is almost certain to be deleterious. It therefore behooves sexual creatures to avoid mates sporting rare or unusual features ( koinophilia ). Sexual populations therefore rapidly shed rare or peripheral phenotypic features, thus canalizing the entire external appearance, as illustrated in the accompanying image of the African pygmy kingfisher , Ispidina picta . This uniformity of all
665-501: A bird's lifetime. Activational hormones occur during puberty and adulthood and serve to 'activate' certain behaviors when appropriate, such as territoriality during breeding season. Organizational hormones occur only during a critical period early in development, either just before or just after hatching in most birds, and determine patterns of behavior for the rest of the bird's life. Such behavioral differences can cause disproportionate sensitivities to anthropogenic pressures. Females of
798-668: A common ancestor may be unclear or unknown; a collection of three such species is referred to as a "trichotomy". Polyploidy is a mechanism that has caused many rapid speciation events in sympatry 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 ). It has been suggested that many of
931-409: A concept discredited by (for example) George C. Williams , John Maynard Smith and Richard Dawkins 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
1064-423: A distinct phenotype . 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
1197-459: A diverse array of sexually dimorphic traits. Aggressive utility traits such as "battle" teeth and blunt heads reinforced as battering rams are used as weapons in aggressive interactions between rivals. Passive displays such as ornamental feathering or song-calling have also evolved mainly through sexual selection. These differences may be subtle or exaggerated and may be subjected to sexual selection and natural selection . The opposite of dimorphism
1330-476: A few programs, most notably the Chinese government's one-child per family policy, have resorted to coercive measures. In the 1970s, tension grew between population control advocates and women's health activists who advanced women's reproductive rights as part of a human rights -based approach. Growing opposition to the narrow population control focus led to a significant change in population control policies in
1463-408: A group of human beings with some predefined feature in common, such as location, race , ethnicity , nationality , or religion . In ecology , a population is a group of organisms of the same species which inhabit the same geographical area and are capable of interbreeding . The area of a sexual population is the area where interbreeding is possible between any opposite-sex pair within
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#17327900653401596-553: A large role in the changing of sex by the fish. It is often seen that a fish will change its sex when there is a lack of a dominant male within the social hierarchy. The females that change sex are often those who attain and preserve an initial size advantage early in life. In either case, females which change sex to males are larger and often prove to be a good example of dimorphism. In other cases with fish, males will go through noticeable changes in body size, and females will go through morphological changes that can only be seen inside of
1729-1011: A less bright or less exaggerated color during the off-breeding season. This occurs because the species is more focused on survival than on reproduction, causing a shift into a less ornate state. Consequently, sexual dimorphism has important ramifications for conservation. However, sexual dimorphism is not only found in birds and is thus important to the conservation of many animals. Such differences in form and behavior can lead to sexual segregation , defined as sex differences in space and resource use. Most sexual segregation research has been done on ungulates, but such research extends to bats , kangaroos , and birds. Sex-specific conservation plans have even been suggested for species with pronounced sexual segregation. The term sesquimorphism (the Latin numeral prefix sesqui - means one-and-one-half, so halfway between mono - (one) and di - (two)) has been proposed for bird species in which "both sexes have basically
1862-425: A naturally occurring part of development, for example plumage. In addition, the strong hormonal influence on phenotypic differences suggests that the genetic mechanism and genetic basis of these sexually dimorphic traits may involve transcription factors or cofactors rather than regulatory sequences. Sexual dimorphism may also influence differences in parental investment during times of food scarcity. For example, in
1995-640: 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 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
2128-417: 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 loci are different, that hawthorn flies mature later in the season and take longer to mature than apple flies; and that there
2261-414: A relic of infection by Wolbachia 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,
2394-425: 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
2527-414: A significant role in male reproductive success. Males have a propensity to be larger than females of a comparable age but it is unclear whether the size increase is due to a growth spurt at the time of the sexual transition or due to the history of faster growth in sex changing individuals. Larger males are able to stifle the growth of females and control environmental resources. Social organization plays
2660-542: A speciation phenomenon if the variation becomes strongly drastic and favorable towards two different outcomes. Sexual dimorphism is maintained by the counteracting pressures of natural selection and sexual selection. For example, sexual dimorphism in coloration increases the vulnerability of bird species to predation by European sparrowhawks in Denmark. Presumably, increased sexual dimorphism means males are brighter and more conspicuous, leading to increased predation. Moreover,
2793-434: 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
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#17327900653402926-467: A view known as punctuated equilibrium . (See diagram, and Darwin's dilemma .) 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 ( Zea mays ), for instance, was created in Mexico in only a few thousand years, starting about 7,000 to 12,000 years ago. This raises
3059-474: Is Lamprologus callipterus , a type of cichlid fish. In this fish, the males are characterized as being up to 60 times larger than the females. The male's increased size is believed to be advantageous because males collect and defend empty snail shells in each of which a female breeds. Males must be larger and more powerful in order to collect the largest shells. The female's body size must remain small because in order for her to breed, she must lay her eggs inside
3192-543: Is Lasioglossum hemichalceum , which is a species of sweat bee that shows drastic physical dimorphisms between male offspring. Not all dimorphism has to have a drastic difference between the sexes. Andrena agilissima is a mining bee where the females only have a slightly larger head than the males. Weaponry leads to increased fitness by increasing success in male–male competition in many insect species. The beetle horns in Onthophagus taurus are enlarged growths of
3325-627: Is monomorphism , when both biological sexes are phenotypically indistinguishable from each other. Common and easily identified types of dimorphism consist of ornamentation and coloration, though not always apparent. A difference in the coloration of sexes within a given species is called sexual dichromatism, commonly seen in many species of birds and reptiles. Sexual selection leads to exaggerated dimorphic traits that are used predominantly in competition over mates. The increased fitness resulting from ornamentation offsets its cost to produce or maintain, suggesting complex evolutionary implications, but
3458-414: Is a gene flow 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 rudis , D. valentini and D. portschinskii all hybridize with each other in their hybrid zone ; however, hybridization
3591-423: Is a good indicator for females because it shows that they are good at obtaining a food supply from which the carotenoid is obtained. There is a positive correlation between the chromas of the tail and breast feathers and body condition. Carotenoids play an important role in immune function for many animals, so carotenoid dependent signals might indicate health. Frogs constitute another conspicuous illustration of
3724-407: Is a sexually dimorphic trait. Theropoda It has been hypothesized that male theropods possessed a retractable penis, a feature similar to modern day crocodilians . Crocodilian skeletons were examined to determine whether there is a skeletal component that is distinctive between both sexes, to help provide an insight on the physical disparities between male and female theropods. Findings revealed
3857-423: Is advantageous to both parties because it avoids damaging the developing fruit and wasting the pollinator's effort on unrewarding visits. In effect, the strategy ensures that pollinators can expect a reward every time they visit an appropriately advertising flower. Females of the aquatic plant Vallisneria americana have floating flowers attached by a long flower stalk that are fertilized if they contact one of
3990-587: Is also displayed by dichromatism. In butterfly genera Bicyclus and Junonia , dimorphic wing patterns evolved due to sex-limited expression, which mediates the intralocus sexual conflict and leads to increased fitness in males. The sexual dichromatic nature of Bicyclus anynana is reflected by female selection on the basis of dorsal UV-reflective eyespot pupils. The common brimstone also displays sexual dichromatism; males have yellow and iridescent wings, while female wings are white and non-iridescent. Naturally selected deviation in protective female coloration
4123-673: Is also seen in frog species like P. bibroni i . Male painted dragon lizards, Ctenophorus pictus . are brightly conspicuous in their breeding coloration, but male colour declines with aging . Male coloration appears to reflect innate anti-oxidation capacity that protects against oxidative DNA damage . Male breeding coloration is likely an indicator to females of the underlying level of oxidative DNA damage (a significant component of aging) in potential mates. Possible mechanisms have been proposed to explain macroevolution of sexual size dimorphism in birds. These include sexual selection, selection for fecundity in females, niche divergence between
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4256-470: Is believed that this is obtained by the ingestion of green Lepidopteran larvae, which contain large amounts of the carotenoids lutein and zeaxanthin . This diet also affects the sexually dimorphic colours in the human-invisible ultraviolet spectrum. Hence, the male birds, although appearing yellow to humans, actually have a violet-tinted plumage that is seen by females. This plumage is thought to be an indicator of male parental abilities. Perhaps this
4389-506: Is called homoploid hybrid speciation. It is considered very rare but has been shown in Heliconius butterflies and sunflowers . Polyploid speciation, which involves changes in chromosome number, is a more common phenomenon, especially in plant species. Theodosius Dobzhansky , 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
4522-487: 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. 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
4655-410: Is displayed in mimetic butterflies. Many arachnid groups exhibit sexual dimorphism, but it is most widely studied in the spiders. In the orb-weaving spider Zygiella x-notata , for example, adult females have a larger body size than adult males. Size dimorphism shows a correlation with sexual cannibalism , which is prominent in spiders (it is also found in insects such as praying mantises ). In
4788-401: Is generally impossible to determine. This is illustrated in the diagram depicting the distribution of hominin species through time since the hominins separated from the line that led to the evolution of their closest living primate relatives, the chimpanzees. Population Population is the term typically used to refer to the number of people in a single area. Governments conduct
4921-687: Is known in ceratopsids, although there is evidence that the more primitive ceratopsian Protoceratops andrewsi possessed sexes that were distinguishable based on frill and nasal prominence size. This is consistent with other known tetrapod groups where midsized animals tend to exhibit markedly more sexual dimorphism than larger ones. However, it has been proposed that these differences can be better explained by intraspecific and ontogenic variation rather than sexual dimorphism. In addition, many sexually dimorphic traits that may have existed in ceratopsians include soft tissue variations such as coloration or dewlaps , which would be unlikely to have been preserved in
5054-455: Is little evidence of interbreeding (researchers have documented a 4–6% hybridization rate) suggests that sympatric speciation is occurring. Reinforcement, also called the Wallace effect , is the process by which natural selection increases reproductive isolation. It may occur after two populations of the same species are separated and then come back into contact. If their reproductive isolation
5187-678: Is more prominently selected for in less dimorphic species of spiders, which often selects for larger male size. In the species Maratus volans , the males are known for their characteristic colorful fan which attracts the females during mating. Ray-finned fish are an ancient and diverse class, with the widest degree of sexual dimorphism of any animal class. Fairbairn notes that "females are generally larger than males but males are often larger in species with male–male combat or male paternal care ... [sizes range] from dwarf males to males more than 12 times heavier than females." There are cases where males are substantially larger than females. An example
5320-477: Is not all nature in confusion instead of the species being, as we see them, well defined? 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
5453-609: Is not even known to the nearest million, so there is a considerable margin of error in such estimates. Researcher Carl Haub calculated that a total of over 100 billion people have probably been born in the last 2000 years. Population growth increased significantly as the Industrial Revolution gathered pace from 1700 onwards. The last 50 years have seen a yet more rapid increase in the rate of population growth due to medical advances and substantial increases in agricultural productivity, particularly beginning in
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5586-402: Is of a subdued brown coloration. The plumage of the peacock increases its vulnerability to predators because it is a hindrance in flight, and it renders the bird conspicuous in general. Similar examples are manifold, such as in birds of paradise and argus pheasants . Another example of sexual dichromatism is that of nestling blue tits . Males are chromatically more yellow than females. It
5719-564: 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 subject to population bottlenecking. 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
5852-611: Is often referred to as the demographic transition . Human population planning is the practice of altering the rate of growth of a human population. Historically, human population control has been implemented with the goal of limiting the rate of population growth. In the period from the 1950s to the 1980s, concerns about global population growth and its effects on poverty, environmental degradation , and political stability led to efforts to reduce population growth rates. While population control can involve measures that improve people's lives by giving them greater control of their reproduction,
5985-595: 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
6118-469: 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 . Without reinforcement,
6251-434: Is seen in the bee species Macrotera portalis in which there is a small-headed morph, capable of flight, and large-headed morph, incapable of flight, for males. Anthidium manicatum also displays male-biased sexual dimorphism. The selection for larger size in males rather than females in this species may have resulted due to their aggressive territorial behavior and subsequent differential mating success. Another example
6384-480: Is selected for, which is seen in the family Araneidae . All Argiope species, including Argiope bruennichi , use this method. Some males evolved ornamentation including binding the female with silk, having proportionally longer legs, modifying the female's web, mating while the female is feeding, or providing a nuptial gift in response to sexual cannibalism. Male body size is not under selection due to cannibalism in all spider species such as Nephila pilipes , but
6517-407: Is strong when the factor of environmental selection is also introduced. Environmental selection may support a smaller chick size if those chicks were born in an area that allowed them to grow to a larger size, even though under normal conditions they would not be able to reach this optimal size for migration. When the environment gives advantages and disadvantages of this sort, the strength of selection
6650-490: 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 niches . A niche must be available in order for a new species to be successful. Ring species such as Larus gulls have been claimed to illustrate speciation in progress, though
6783-409: 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 , which would control hybridization. This is sometimes called
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#17327900653406916-504: Is the dragonet , in which males are considerably larger than females and possess longer fins. Sexual dimorphism also occurs in hermaphroditic fish. These species are known as sequential hermaphrodites . In fish, reproductive histories often include the sex-change from female to male where there is a strong connection between growth, the sex of an individual, and the mating system within which it operates. In protogynous mating systems where males dominate mating with many females, size plays
7049-448: 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
7182-576: Is thought to have arisen from hybrid speciation. Hybridization is an important means of speciation in plants, since polyploidy (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 in small populations. Hybridization without change in chromosome number
7315-715: Is very likely that the world's population will stop growing before the end of the 21st century. Further, there is some likelihood that population will actually decline before 2100. Population has already declined in the last decade or two in Eastern Europe, the Baltics and in the former Commonwealth of Independent States. The population pattern of less-developed regions of the world in recent years has been marked by gradually declining birth rates. These followed an earlier sharp reduction in death rates. This transition from high birth and death rates to low birth and death rates
7448-411: Is weakened and the environmental forces are given greater morphological weight. The sexual dimorphism could also produce a change in timing of migration leading to differences in mating success within the bird population. When the dimorphism produces that large of a variation between the sexes and between the members of the sexes, multiple evolutionary effects can take place. This timing could even lead to
7581-619: 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
7714-604: The blue-footed booby , the female chicks grow faster than the males, resulting in booby parents producing the smaller sex, the males, during times of food shortage. This then results in the maximization of parental lifetime reproductive success. In Black-tailed Godwits Limosa limosa limosa females are also the larger sex, and the growth rates of female chicks are more susceptible to limited environmental conditions. Sexual dimorphism may also only appear during mating season; some species of birds only show dimorphic traits in seasonal variation. The males of these species will molt into
7847-443: The heterozygote leads to selection for behaviours or mechanisms that prevent their interbreeding . 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 . Even if there
7980-489: The ovules ). Each pollen grain accordingly may be seen as a male plant in its own right; it produces a sperm cell and is dramatically different from the female plant, the megagametophyte that produces the female gamete. Insects display a wide variety of sexual dimorphism between taxa including size, ornamentation and coloration. The female-biased sexual size dimorphism observed in many taxa evolved despite intense male-male competition for mates. In Osmia rufa , for example,
8113-438: 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 . Under these circumstances, not only
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#17327900653408246-407: The 'fittest' available male. Sexual dimorphism is a product of both genetics and environmental factors. An example of sexual polymorphism determined by environmental conditions exists in the red-backed fairywren . Red-backed fairywren males can be classified into three categories during breeding season : black breeders, brown breeders, and brown auxiliaries. These differences arise in response to
8379-569: The 1960s, made by the Green Revolution . In 2017 the United Nations Population Division projected that the world's population would reach about 9.8 billion in 2050 and 11.2 billion in 2100. In the future, the world's population is expected to peak at some point, after which it will decline due to economic reasons, health concerns, land exhaustion and environmental hazards. According to one report, it
8512-496: The United States Census Bureau, the world population hit 6.5 billion on 24 February 2006. The United Nations Population Fund designated 12 October 1999 as the approximate day on which world population reached 6 billion. This was about 12 years after the world population reached 5 billion in 1987, and six years after the world population reached 5.5 billion in 1993. The population of countries such as Nigeria
8645-496: The Wallace effect after the evolutionary biologist Alfred Russel Wallace 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
8778-580: The adult members of a sexual species has stimulated the proliferation of field guides on birds, mammals, reptiles, insects, and many other taxa , in which a species can be described with a single illustration (or two, in the case of sexual dimorphism ). 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,
8911-482: The area and more probable than cross-breeding with individuals from other areas. In humans , interbreeding is unrestricted by racial differences, as all humans belong to the same species of Homo sapiens. In ecology, the population of a certain species in a certain area can be estimated using the Lincoln index to calculate the total population of an area based on the number of individuals observed. In genetics,
9044-616: 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. All forms of natural speciation have taken place over
9177-533: The bird's body condition: if they are healthy they will produce more androgens thus becoming black breeders, while less healthy birds produce less androgens and become brown auxiliaries. The reproductive success of the male is thus determined by his success during each year's non-breeding season, causing reproductive success to vary with each year's environmental conditions. Migratory patterns and behaviors also influence sexual dimorphisms. This aspect also stems back to size dimorphism in species. It has been shown that
9310-415: The birds now known as Darwin's finches to be a classic case of adaptive evolutionary radiation. 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 , since small populations often undergo bottlenecks . Genetic drift
9443-1001: The body. For example, in sockeye salmon , males develop larger body size at maturity, including an increase in body depth, hump height, and snout length. Females experience minor changes in snout length, but the most noticeable difference is the huge increase in gonad size, which accounts for about 25% of body mass. Sexual selection was observed for female ornamentation in Gobiusculus flavescens , known as two-spotted gobies. Traditional hypotheses suggest that male–male competition drives selection. However, selection for ornamentation within this species suggests that showy female traits can be selected through either female–female competition or male mate choice. Since carotenoid-based ornamentation suggests mate quality, female two-spotted guppies that develop colorful orange bellies during breeding season are considered favorable to males. The males invest heavily in offspring during incubation, which leads to
9576-480: The breaking up of a large sexual population (panmictic) into smaller overlapping sexual populations. This failure of panmixia leads to two important changes in overall population structure: (1) the component gamodemes vary (through gamete sampling) in their allele frequencies when compared with each other and with the theoretical panmictic original (this is known as dispersion, and its details can be estimated using expansion of an appropriate binomial equation ); and (2)
9709-436: 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 phenotypes both in time and space – which would be
9842-470: The caudal chevrons of male crocodiles, used to anchor the penis muscles, were significantly larger than those of females. There have been criticisms of these findings, but it remains a subject of debate among advocates and adversaries. Ornithopoda Studies of sexual dimorphism in hadrosaurs have generally centered on the distinctive cranial crests , which likely provided a function in sexual display. A biometric study of 36 skulls found sexual dimorphism
9975-400: The costs and evolutionary implications vary from species to species. The peafowl constitute conspicuous illustrations of the principle. The ornate plumage of peacocks, as used in the courting display, attracts peahens . At first sight, one might mistake peacocks and peahens for completely different species because of the vibrant colours and the sheer size of the male's plumage; the peahen
10108-548: The course of evolution ; however, debate persists as to the relative importance of each mechanism in driving biodiversity . One example of natural speciation is the diversity of the three-spined stickleback , a marine fish that, after the last glacial period , has undergone speciation into new freshwater 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 of fish including variations in fins, changes in
10241-410: 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. It has been argued that the resolution of Darwin's first dilemma lies in the fact that out-crossing sexual reproduction has an intrinsic cost of rarity. The cost of rarity arises as follows. If, on
10374-429: 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 , which can be considered the same species as several varieties of wild ox , gaur , and yak ; and with domestic sheep that can interbreed with the mouflon . The best-documented creations of new species in
10507-654: The degree of preservation. The availability of well-preserved remains is not a probable outcome as a consequence of decomposition and fossilization . Some paleontologists have looked for sexual dimorphism among dinosaurs using statistics and comparison to ecologically or phylogenetically related modern animals. Apatosaurus and Diplodocus Female Apatosaurus and Diplodocus had interconnected caudal vertebrae that allowed them to keep their tails elevated to aid in copulation. Discovering that this fusion occurred in only 50% of Apatosaurus and Diplodocus skeletons and 25% of Camarasaurus skeletons indicated that this
10640-508: The early 1980s. Sexual dimorphism Sexual dimorphism is the condition where sexes of the same species exhibit different morphological characteristics, including characteristics not directly involved in reproduction . The condition occurs in most dioecious species, which consist of most animals and some plants. Differences may include secondary sex characteristics , size, weight, color, markings, or behavioral or cognitive traits. Male-male reproductive competition has evolved
10773-448: The empty shells. If she grows too large, she will not fit in the shells and will be unable to breed. The female's small body size is also likely beneficial to her chances of finding an unoccupied shell. Larger shells, although preferred by females, are often limited in availability. Hence, the female is limited to the growth of the size of the shell and may actually change her growth rate according to shell size availability. In other words,
10906-516: The estrogen pathway. The sexual dimorphism in lizards is generally attributed to the effects of sexual selection, but other mechanisms including ecological divergence and fecundity selection provide alternative explanations. The development of color dimorphism in lizards is induced by hormonal changes at the onset of sexual maturity, as seen in Psamodromus algirus , Sceloporus gadoviae , and S. undulates erythrocheilus . Sexual dimorphism in size
11039-457: The evidence, "...accumulated from top-down studies of adaptation and reproductive isolation". Sexual selection can drive speciation in a clade, independently of natural selection . 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
11172-443: The exhausted anthers after a day or two and perhaps change their colours as well while the pistil matures; specialist pollinators are very much inclined to concentrate on the exact appearance of the flowers they serve, which saves their time and effort and serves the interests of the plant accordingly. Some such plants go even further and change their appearance once fertilized, thereby discouraging further visits from pollinators. This
11305-435: 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 or apomixis , as for unknown reasons many asexual organisms are polyploid. Rare instances of polyploid mammals are known, but most often result in prenatal death. Hybridization between two different species sometimes leads to
11438-418: 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
11571-485: The expense of the smaller, rarer species, eventually driving them to extinction . Eventually, only a few species remain, each distinctly different from the other. Rarity not only imposes the risk of failure to find a mate, but it may also incur indirect costs, such as the resources expended or risks taken to seek 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
11704-465: The expression of advantageous recessive phenotypes, reducing the recombination load, and reducing the cost of sex. The hawthorn fly ( Rhagoletis pomonella ), 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 ,
11837-526: 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 mutations ", which can be, and are, used to trace
11970-420: The extraordinary diversity of crustaceans living in the depths of Siberia's Lake Baikal . 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
12103-417: The female is larger/broader than males, with males being 8–10 mm in size and females being 10–12 mm in size. In the hackberry emperor females are similarly larger than males. The reason for the sexual dimorphism is due to provision size mass, in which females consume more pollen than males. In some species, there is evidence of male dimorphism, but it appears to be for distinctions of roles. This
12236-564: The female, which looks different from the males. Various other dioecious exceptions, such as Loxostylis alata have visibly different sexes, with the effect of eliciting the most efficient behavior from pollinators, who then use the most efficient strategy in visiting each gender of flower instead of searching, say, for pollen in a nectar-bearing female flower. Some plants, such as some species of Geranium have what amounts to serial sexual dimorphism. The flowers of such species might, for example, present their anthers on opening, then shed
12369-678: The first aspect, and it is now widely agreed that the critical factor behind the origin of new species is reproductive isolation. In On the Origin of Species (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 Firstly, why, if species have descended from other species by insensibly fine gradations, do we not everywhere see innumerable transitional forms? Why
12502-651: The form of reduced survival. This means that even if the trait causes males to die earlier, the trait is still beneficial so long as males with the trait produce more offspring than males lacking the trait. This balance keeps dimorphism alive in these species and ensures that the next generation of successful males will also display these traits that are attractive to females. Such differences in form and reproductive roles often cause differences in behavior. As previously stated, males and females often have different roles in reproduction. The courtship and mating behavior of males and females are regulated largely by hormones throughout
12635-734: The fossil record. Stegosaurians A 2015 study on specimens of Hesperosaurus mjosi found evidence of sexual dimorphism in the shape of the dermal plates. Two plate morphs were described: one was short, wide, and oval-shaped, the other taller and narrower. In a large proportion of mammal species, males are larger than females. Both genes and hormones affect the formation of many animal brains before " birth " (or hatching ), and also behaviour of adult individuals. Hormones significantly affect human brain formation, and also brain development at puberty. A 2004 review in Nature Reviews Neuroscience observed that "because it
12768-417: 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 ,
12901-691: The gametes within it, the gamodeme is said to be panmictic. Under this state, allele ( gamete ) frequencies can be converted to genotype ( zygote ) frequencies by expanding an appropriate quadratic equation , as shown by Sir Ronald Fisher in his establishment of quantitative genetics. This seldom occurs in nature: localization of gamete exchange – through dispersal limitations, preferential mating, cataclysm, or other cause – may lead to small actual gamodemes which exchange gametes reasonably uniformly within themselves but are virtually separated from their neighboring gamodemes. However, there may be low frequencies of exchange with these neighbors. This may be viewed as
13034-705: The head or thorax expressed only in the males. Copris ochus also has distinct sexual and male dimorphism in head horns. Another beetle with a distinct horn-related sexual dimorphism is Allomyrina dichotoma, also known as the Japanese rhinoceros beetle . These structures are impressive because of the exaggerated sizes. There is a direct correlation between male horn lengths and body size and higher access to mates and fitness. In other beetle species, both males and females may have ornamentation such as horns. Generally, insect sexual size dimorphism (SSD) within species increases with body size. Sexual dimorphism within insects
13167-420: The laboratory were performed in the late 1980s. William R. Rice and George W. Salt bred Drosophila melanogaster fruit flies 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,
13300-444: The larger males are better at coping with the difficulties of migration and thus are more successful in reproducing when reaching the breeding destination. When viewing this from an evolutionary standpoint, many theories and explanations come into consideration. If these are the result for every migration and breeding season, the expected results should be a shift towards a larger male population through sexual selection. Sexual selection
13433-463: The level of homozygosity rises in the entire collection of gamodemes. The overall rise in homozygosity is quantified by the inbreeding coefficient (f or φ). All homozygotes are increased in frequency – both the deleterious and the desirable. The mean phenotype of the gamodemes collection is lower than that of the panmictic original – which is known as inbreeding depression. It is most important to note, however, that some dispersion lines will be superior to
13566-508: The male's ability to collect large shells depends on his size. The larger the male, the larger the shells he is able to collect. This then allows for females to be larger in his brooding nest which makes the difference between the sizes of the sexes less substantial. Male–male competition in this fish species also selects for large size in males. There is aggressive competition by males over territory and access to larger shells. Large males win fights and steal shells from competitors. Another example
13699-517: 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 (1859), under the heading "Difficulties with the Theory". There are several suggestions as to how mate choice might play a significant role in resolving Darwin's dilemma . If speciation takes place in the absence of natural selection, it might be referred to as nonecological speciation . New species have been created by animal husbandry , but
13832-593: The more ornamented or brightly colored sex. Such differences have been attributed to the unequal reproductive contributions of the sexes. This difference produces a stronger female choice since they have more risk in producing offspring. In some species, the male's contribution to reproduction ends at copulation, while in other species the male becomes the main (or only) caregiver. Plumage polymorphisms have evolved to reflect these differences and other measures of reproductive fitness, such as body condition or survival. The male phenotype sends signals to females who then choose
13965-692: The most common causes for mortality in young fish. Most flowering plants are hermaphroditic but approximately 6% of species have separate males and females ( dioecy ). Sexual dimorphism is common in dioecious plants and dioicous species. Males and females in insect-pollinated species generally look similar to one another because plants provide rewards (e.g. nectar ) that encourage pollinators to visit another similar flower , completing pollination . Catasetum orchids are one interesting exception to this rule. Male Catasetum orchids violently attach pollinia to euglossine bee pollinators. The bees will then avoid other male flowers but may visit
14098-526: The number or size of their bony plates, variable jaw structure, and color differences. During allopatric (from the ancient Greek allos , "other" + patrā , "fatherland") speciation, a population splits into two geographically isolated populations (for example, by habitat fragmentation due to geographical change such as mountain formation ). The isolated populations then undergo genotypic or phenotypic divergence as: (a) they become subjected to dissimilar selective pressures; (b) different mutations arise in
14231-418: The panmictic original, while some will be about the same, and some will be inferior. The probabilities of each can be estimated from those binomial equations. In plant and animal breeding , procedures have been developed which deliberately utilize the effects of dispersion (such as line breeding, pure-line breeding, backcrossing). Dispersion-assisted selection leads to the greatest genetic advance (ΔG=change in
14364-771: The phenotypic mean), and is much more powerful than selection acting without attendant dispersion. This is so for both allogamous (random fertilization) and autogamous (self-fertilization) gamodemes. According to the UN, the world's population surpassed 8 billion on 15 November 2022, an increase of 1 billion since 12 March 2012. According to a separate estimate by the United Nations, Earth's population exceeded seven billion in October 2011. According to UNFPA , growth to such an extent offers unprecedented challenges and opportunities to all of humanity. According to papers published by
14497-436: The plants become sexually mature. Every sexually reproducing extant species of the vascular plant has an alternation of generations; the plants we see about us generally are diploid sporophytes , but their offspring are not the seeds that people commonly recognize as the new generation. The seed actually is the offspring of the haploid generation of microgametophytes ( pollen ) and megagametophytes (the embryo sacs in
14630-418: The presence of specific sex-related behaviour is common to many lizards; and vocal qualities which are frequently observed in frogs . Anole lizards show prominent size dimorphism with males typically being significantly larger than females. For instance, the average male Anolis sagrei was 53.4 mm vs. 40 mm in females. Different sizes of the heads in anoles have been explained by differences in
14763-410: The principle. There are two types of dichromatism for frog species: ontogenetic and dynamic. Ontogenetic frogs are more common and have permanent color changes in males or females. Ranoidea lesueuri is an example of a dynamic frog with temporary color changes in males during the breeding season. Hyperolius ocellatus is an ontogenetic frog with dramatic differences in both color and pattern between
14896-408: The production of more exaggerated ornaments in males may come at the cost of suppressed immune function. So long as the reproductive benefits of the trait due to sexual selection are greater than the costs imposed by natural selection, then the trait will propagate throughout the population. Reproductive benefits arise in the form of a larger number of offspring, while natural selection imposes costs in
15029-450: 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 mutations " which do not affect the functionality or appearance of the carrier, are thus usually disadvantageous, and their chance of proving to be useful in
15162-404: The result is progeny which are immediately reproductively isolated from the parent population. New species can also be created through hybridization , followed by reproductive isolation, if the hybrid is favoured by natural selection. In addressing the origin of species, there are two key issues: Since Charles Darwin's time, efforts to understand the nature of species have primarily focused on
15295-690: The same area. The best known example of sympatric speciation is that of the cichlids of East Africa inhabiting the Rift Valley lakes , particularly Lake Victoria , Lake Malawi and Lake Tanganyika . 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 and sexual selection . A 2008 study suggests that sympatric speciation has occurred in Tennessee cave salamanders . Sympatric speciation driven by ecological factors may also account for
15428-585: The same plumage pattern, though the female is clearly distinguishable by reason of her paler or washed-out colour". Examples include Cape sparrow ( Passer melanurus ), rufous sparrow (subspecies P. motinensis motinensis ), and saxaul sparrow ( P. ammodendri ). Examining fossils of non-avian dinosaurs in search of sexually dimorphic characteristics requires the supply of complete and articulated skeletal and tissue remains. As terrestrial organisms, dinosaur carcasses are subject to ecological and geographical influence that inevitably constitutes
15561-479: The sexes, and allometry, but their relative importance is still not fully understood . Sexual dimorphism in birds can be manifested in size or plumage differences between the sexes. Sexual size dimorphism varies among taxa, with males typically being larger, though this is not always the case, e.g. birds of prey , hummingbirds , and some species of flightless birds. Plumage dimorphism, in the form of ornamentation or coloration, also varies, though males are typically
15694-526: The sexes. At sexual maturity, the males display a bright green with white dorsolateral lines. In contrast, the females are rusty red to silver with small spots. The bright coloration in the male population attracts females and is an aposematic sign to potential predators. Females often show a preference for exaggerated male secondary sexual characteristics in mate selection. The sexy son hypothesis explains that females prefer more elaborate males and select against males that are dull in color, independent of
15827-511: The sexual preference in colorful females due to higher egg quality. In amphibians and reptiles, the degree of sexual dimorphism varies widely among taxonomic groups . The sexual dimorphism in amphibians and reptiles may be reflected in any of the following: anatomy; relative length of tail; relative size of head; overall size as in many species of vipers and lizards ; coloration as in many amphibians , snakes , and lizards, as well as in some turtles ; an ornament as in many newts and lizards;
15960-410: The situation may be more complex. The grass Anthoxanthum odoratum may be starting parapatric speciation in areas of mine contamination. 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 plants in
16093-414: The size dimorphic wolf spider Tigrosa helluo , food-limited females cannibalize more frequently. Therefore, there is a high risk of low fitness for males due to pre-copulatory cannibalism, which led to male selection of larger females for two reasons: higher fecundity and lower rates of cannibalism. In addition, female fecundity is positively correlated with female body size and large female body size
16226-567: The species' vision. Similar sexual dimorphism and mating choice are also observed in many fish species. For example, male guppies have colorful spots and ornamentations, while females are generally grey. Female guppies prefer brightly colored males to duller males. In redlip blennies , only the male fish develops an organ at the anal-urogenital region that produces antimicrobial substances. During parental care, males rub their anal-urogenital regions over their nests' internal surfaces, thereby protecting their eggs from microbial infections, one of
16359-542: The thousands of free-floating flowers released by a male. Sexual dimorphism is most often associated with wind-pollination in plants due to selection for efficient pollen dispersal in males vs pollen capture in females, e.g. Leucadendron rubrum . Sexual dimorphism in plants can also be dependent on reproductive development. This can be seen in Cannabis sativa , a type of hemp, which have higher photosynthesis rates in males while growing but higher rates in females once
16492-443: 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
16625-532: 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
16758-769: 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 fruit flies after several generations by placing them in different media, starch- and maltose-based media. [REDACTED] 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
16891-668: 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 genes . Island genetics is the term associated with the tendency of small, isolated genetic pools to produce unusual traits. Examples include insular dwarfism and the radical changes among certain famous island chains, for example on Komodo . The Galápagos Islands are particularly famous for their influence on Charles Darwin. During his five weeks there he heard that Galápagos tortoises could be identified by island, and noticed that finches differed from one island to another, but it
17024-501: The two species would have uncontrollable inbreeding . Reinforcement may be induced in artificial selection experiments as described below. 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
17157-460: The whinchat in Switzerland breed in intensely managed grasslands. Earlier harvesting of the grasses during the breeding season lead to more female deaths. Populations of many birds are often male-skewed and when sexual differences in behavior increase this ratio, populations decline at a more rapid rate. Also not all male dimorphic traits are due to hormones like testosterone, instead they are
17290-462: 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 horses and donkeys . One reasoning behind this
17423-470: Was exhibited in the crest of 3 species of hadrosaurids. The crests could be categorized as full (male) or narrow (female) and may have given some advantage in intrasexual mating-competition. Ceratopsians According to Scott D. Sampson, if ceratopsids were to exhibit sexual dimorphism, modern ecological analogues suggest it would be found in display structures, such as horns and frills. No convincing evidence for sexual dimorphism in body size or mating signals
17556-413: Was only nine months later that he speculated 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
17689-782: 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. 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 palaeontologists such as Niles Eldredge and Stephen Jay Gould have argued that species usually remain unchanged over long stretches of time, and that speciation occurs only over relatively brief intervals,
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