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character in the original allopatric populations had a large degree of differentiation. As genetic admixture between the population increases with time however, the steepness of the cline is likely to decrease as the difference in character is eroded. However, if the character in the original allopatric populations was not very differentiated to begin with, the cline between the populations need not display a very steep gradient. Because both primary differentiation and secondary contact can therefore give rise to similar or identical clinal patterns (e.g. gently sloping clines), distinguishing which of these two processes is responsible for generating a cline is difficult and often impossible. However, in some circumstances a cline and a geographic variable (such as humidity) may be very tightly linked, with a change in one corresponding closely to a change in the other. In such cases it may be tentatively concluded that the cline is generated by primary differentiation and therefore moulded by environmental selective pressures.
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440:. For example, through environmental selection acting on populations and favouring particular allele frequencies, large genetic differences between populations may accumulate (this would be reflected in clinal structure by the presence of numerous very steep clines). If the local genetic differences are great enough, it may lead unfavourable combinations of genotypes and therefore to hybrids being at a decreased fitness relative to the parental lines. When this hybrid disadvantage is great enough, natural selection will select for
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232:, creating an intermediate zone. This secondary contact scenario may occur, for example, when climatic conditions change, allowing the ranges of populations to expand and meet. Because over time the effect of gene flow will tend to eventually swamp out any regional differences and cause one large homogenous population, for a stable cline to be maintained when two populations join there must usually be a selective pressure maintaining a degree of differentiation between the two populations.
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may be the result of two previously allopatric populations with a large degree of difference in the trait having only recently established gene flow, or where there is strong selection against hybrids. However, it may also reflect a sudden environmental change or boundary. Examples of rapidly changing environmental boundaries like this include abrupt changes in the heavy metal content of soils, and the consequent narrow clines produced between populations of
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421:). The possibility that clines may be a precursor to speciation was therefore ignored, as they were assumed to be evidence of the fact that in contiguous populations gene flow was too strong a force of homogenisation, and selection too weak a force of differentiation, for speciation to take place. However, the existence of particular types of clines, such as
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different geographical trajectory, being most vibrant where humidity is highest and becoming less vibrant further into the arid centre of the country. Because of this, Huxley described the notion of clines as an "auxiliary taxonomic principle,” meaning that clinal variation in a species is not awarded taxonomic recognition in the way
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character". In other words, areas on maps that demonstrate the same biological phenomenon or character will be connected by something that resembles a contour line. When mapping clines therefore, which follow a character gradation from one extreme to the other, isophenes will transect clinal lines at a right angle.
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In humans, clines in the frequency of blood types has allowed scientists to infer past population migrations. For example, the Type B blood group reaches its highest frequency in Asia, but become less frequent further west. From this, it has been possible to infer that some Asian populations migrated
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closer to the
Equator tend to be smaller than their more northerly or southerly conspecifics. One of the proposed reasons for this cline is that larger animals have a relatively smaller surface area to volume ratio and therefore improved heat conservancy – an important advantage in cold climates. The
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Continuous stepped clines consist of an overall shallow cline, interspersed by sections of much steeper slope. The shallow slope represents the populations, and the shorter, steeper sections the larger change in character between populations. Stepped clines can be further subdivided into horizontally
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The mechanism of selection maintaining the clines in this scenario is often intrinsic. This means that the fitness of individuals is independent of the external environment, and selection is instead dependent on the genome of the individual. Intrinsic, or endogenous, selection can give rise to clines
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Secondary contact could lead to a cline with a steep gradient if heterozygote disadvantage or frequency-dependent selection exists, as intermediates are heavily selected against. Alternatively, steep clines could exist because the populations have only recently established secondary contact, and the
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melanic morph of the peppered moth increased drastically in northern
England. This cline in morph colour, from a dominance of lighter morphs in the west of England (which did not suffer as heavily from pollution), to the higher frequency of melanic forms in the north, has slowly been degrading since
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Primary differentiation is demonstrated using the peppered moth as an example, with a change in an environmental variable such as sooty coverage of trees imposing a selective pressure on a previously uniformly coloured moth population. This causes the frequency of melanic morphs to increase the more
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recognised, these multiple independent clines may not act in concordance with each other. For example, it has been observed that in
Australia, birds generally become smaller the further towards the north of the country they are found. In contrast, the intensity of their plumage colouration follows a
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alone. It is unlikely that large-scale clines in genotype or phenotype frequency will be produced solely by drift. However, across smaller geographical scales and in smaller populations, drift could produce temporary clines. The fact that drift is a weak force upholding the cline however means that
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Clines produced through this way are generated by spatial heterogeneity in environmental conditions. The mechanism of selection acting upon organisms is therefore external. Species ranges frequently span environmental gradients (e.g. humidity, rainfall, temperature, or day length) and, according to
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Clines can be portrayed graphically on maps using lines that show the transition in character state from one end of the geographic range to the other. Character states can however additionally be represented using isophenes, defined by Ernst Mayr as "lines of equal expression of a clinally varying
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The steepness, or gradient, of a cline reflects the extent of the differentiation in the character across a geographic range. For example, a steep cline could indicate large variation in the colour of plumage between adjacent bird populations. It has been previously outlined that such steep clines
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that is created when these respective populations come into contact is then shaped by the opposing forces of selection and gene flow; even if selection against heterozygotes is great, if there is some degree of gene flow between the two populations, then a steep cline may be able to be maintained.
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Unlike in continuous clines, in discontinuous clines the populations of species are allopatric, meaning there is very little or no gene flow amongst populations. The genetic or phenotypic trait in question always shows a steeper gradient between groups than within groups, as in continuous clines.
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In obliquely stepped clines, conversely, each population also demonstrates a cline in the character, albeit of a shallower slope than the clines connecting the populations together. Huxley compared obliquely stepped clines to looking like a "stepped ramp", rather than taking on the formation of a
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can all produce patterns that deviate from those well-established signals which mark prey as being unpalatable. These individuals are then predated more heavily relative to their counterparts with "normal" markings (i.e. selected against), creating populations dominated by a particular pattern of
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that are either adapted to these soils with high metal content, or adapted to "normal" soil. Conversely, a shallow cline indicates little geographical variation in the character or trait across a given geographical distance. This may have arisen through weak differential environmental selective
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In continuous clines, all populations of the species are able to interbreed and there is gene flow throughout the entire range of the species. In this way, these clines are both biologically (no clear subgroups) and geographically (contiguous distribution) continuous. Continuous clines can be
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Horizontally stepped clines show no intra-population variation or gradation in the character, therefore displaying a horizontal gradient. These uniform populations are connected by steeper sections of the cline, characterised by larger changes in the form of the character. However, because in
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Continuous smooth clines are characterised by the lack of any abrupt changes or delineation in the genetic or phenotypic trait across the cline, instead displaying a smooth gradation throughout. Huxley recognised that this type of cline, with its uniform slope throughout, was unlikely to be
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According to Huxley, clines can be classified into two categories; continuous clines and discontinuous stepped clines. These types of clines characterise the way that a genetic or phenotypic trait transforms from one end of its geographical range of the species to the other.
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tend to covary with each other, such that birds found in arid areas near the
Equator tend to be much darker than those in less arid areas closer to the Poles. Since then, this rule has been extended to include many other animals, including flies, butterflies, and wolves.
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Such clines in characters can not be maintained through selection alone if much gene flow occurred between populations, as this would tend to swamp out the effects of local adaptation. However, because species usually tend to have a limited dispersal range (e.g. in an
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Although the term "cline" was first officially coined by Huxley in 1938, gradients and geographic variations in the character states of species have been observed for centuries. Indeed, some gradations have been considered so ubiquitous that they have been labelled
175:. In this way, when previously genetically or phenotypically uniform populations spread into novel environments, they will evolve to be uniquely adapted to the local environment, in the process potentially creating a gradient in a genotypic or phenotypic trait.
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Clines are generally thought to arise under one of two conditions: "primary differentiation" (also known as "primary contact" or "primary intergradation"), or "secondary contact" (also known as "secondary introgression", or "secondary intergradation").
516:, which was introduced in the early 1850s to the eastern United States, evolved a north-south gradient in size soon after its introduction. This gradient reflects the gradient that already existed in the house sparrow's native range in Europe.
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role of the environment in imposing a selective pressure and producing this cline has been heavily implicated due to the fact that
Bergmann's Rule has been observed across many independent lineages of species and continents. For example, the
240:, in which intermediate genotypes have a lower relative fitness than either homozygote genotypes. Because of this disadvantage, one allele will tend to become fixed in a given population, such that populations will consist largely of either
425:, in which populations did not differentiate in allopatry but the terminal ends of the cline nonetheless do not interbreed, cast into doubt whether complete geographical isolation of populations is an absolute requirement for speciation.
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Secondary contact between two previously isolated populations. Two previously isolated populations establish contact and therefore gene flow, creating an intermediate zone in the phenotypic or genotypic character between the two
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traits in the homozygous parental lines that reduce the likelihood of disadvantageous hybridisation - in other words, natural selection will favour traits that promote assortative mating in the parental lines. This is known as
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to the local environment, resulting in different genotypes or phenotypes being favoured in different environments. This diversifying force is countered by gene flow, which has a homogenising effect on populations and prevents
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gulls, the habitats of the end populations even overlap, which introduces questions as to what constitutes a species: nowhere along the cline can a line be drawn between the populations, but they are unable to interbreed.
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due to the cumulative effect of the many changes in phenotype along the cline. The populations elsewhere along the cline interbreed with their geographically adjacent populations as in a standard cline. In the case of
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clines produced this way are often random (i.e. uncorrelated with environmental variables) and subject to breakdown or reversal over time. Such clines are therefore unstable and sometimes called "transient clines".
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Because instrinsic selection is independent of the external environment, clines generated by selection against hybrids are not fixed to any given geographical area and can move around the geographic landscape. Such
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A cline is a spatial gradient in a single specific trait, rather than in a collection of traits; a single population can therefore have as many clines as it has traits, at least in principle. Additionally, as
61:), or phenotypic (e.g. body size, skin pigmentation) character. They can show either smooth, continuous gradation in a character, or more abrupt changes in the trait from one geographic region to the next.
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The gradient of a cline is related to another commonly referred to property, clinal width. A cline with a steep slope is said to have a small, or narrow, width, while shallower clines have larger widths.
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are a distinct type of cline where the geographical distribution in question is circular in shape, so that the two ends of the cline overlap with one another, giving two adjacent populations that rarely
196:, in the UK. During the 19th century, when the industrial sector gained traction, coal emissions blackened vegetation across northwest England and parts of northern Wales. As a result of this, lighter
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warning signal. As with heterozygote disadvantage, when these populations join, a narrow cline of intermediate individuals could be produced, maintained by gene flow counteracting selection.
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Bergmann's Rule demonstrated showing the difference in size between a larger northern fox, whose range spans colder regions, and a smaller desert fox, whose range is primarily in hot regions
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model), restricted gene flow can serve as a type of barrier which encourages geographic differentiation. However, some degree of migration is often required to maintain a cline; without it,
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of the moth were more visible to predators against the blackened tree trunks and were therefore more heavily predated relative to the darker morphs. Consequently, the frequency of the more
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where hybrids are a disadvantage relative to their parental lines (but which are nonetheless maintained through selection being counteracted by gene flow) are known as "tension zones".
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pressure, or where two populations established secondary contact a long time ago and gene flow has eroded the large character differentiation between the populations.
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refers to a population which differs from other populations in a number of characters, rather than the single character that varies amongst populations in a cline.
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Horizontally stepped clines, where intra-group variation is very small or non-existent and the geographic space separating groups shows a sharp change in character
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model. In contrast to this cline, blood Type A shows the reverse pattern, reaching its highest frequency in Europe and declining in frequency towards Asia.
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Because clines can exist in populations connected by some degree of gene flow, the generation of new species from a previously clinal population is termed
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Obliquely stepped clines, where there is some intra-group gradation, but this is less than the gradation in the character between populations
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Clinal characters change from one end of the geographic range to another. The extent of this change is reflected in the slope of the cline.
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continuous clines all populations exchange genetic material, the intergradation zone between the groups can never have a vertical slope.
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While selection can therefore clearly play a key role in creating clines, it is theoretically feasible that they might be generated by
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Clines generated through this mechanism have arisen through the joining of two formerly isolated populations which differentiated in
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On the Origin of
Species by means of natural selection, or the preservation of favoured races in the struggle for life
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A classic example of the role of environmental gradients in creating clines is that of the peppered moth,
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Saccheri, Illik J.; Rousset, Francois; Watts, Phillip C.; Brakefield, Paul M.; Cook, Laurence M. (2008).
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is likely to eventually occur, as local adaptation can cause reproductive isolation between populations.
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1334:"Evolutionary divergence among adjacent plant populations I. The evidence and its theoretical analysis"
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It was originally assumed that geographic isolation was a necessary precursor to speciation (
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butterflies sometimes display steep clines between populations, which are maintained through
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Discontinuous clines follow the same principles as continuous clines by displaying either
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Populations, species, and evolution: an abridgment of Animal species and evolution
432:. Both extrinsic and intrinsic selection can serve to generate varying degrees of
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491:, who observed in 1833 that environmental factors and the pigmentation of avian
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1285:"Genetic Drift Widens the Expected Cline but Narrows the Expected Cline Width"
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in characters through a variety of mechanisms. One way it may act is through
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Huxley, Julian (30 July 1938). "Clines: an
Auxiliary Taxonomic Principle".
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964:"Selection and gene flow on a diminishing cline of melanic peppered moths"
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in a single characteristic (or biological trait) of a species across its
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Gradual variation of the characteristics of a species along its territory
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Systematics and the Origin of
Species from the viewpoint of a zoologist
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towards Europe around 2,000 years ago, causing genetic admixture in an
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are sometimes used interchangeably, they do in fact differ in that
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The
Driving Forces of Evolution: Genetic Processes in Populations
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Clines are often cited to be the result of two opposing drivers:
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38:
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natural selection, different environments will favour different
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Another way in which selection can generate clines is through
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limitations to sooty emissions were introduced in the 1960s.
690:. Vol. 10. Princeton University Press. pp. 1–246.
483:. One commonly cited example of a gradient in morphology is
348:
Categories and subcategories of clines, as defined by Huxley
1199:"Strong natural selection in a warning-colour hybrid zone"
26:"Gene cline" redirects here. For the baseball player, see
772:
White, Timothy L.; Adams, W. T.; Neale, David B. (2007).
745:
If I Am To Be
Remembered: Correspondence Of Julian Huxley
298:
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staircase as in the case of horizontally stepped clines.
252:(homozygous recessive) individuals. The cline of hetero
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further sub-divided into smooth and stepped clines.
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1454:
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814:
737:
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1283:Polechová, Jitka; Barton, Nick (September 2011).
767:
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742:Dronamraju, Krishna R.; Needham, Joseph (1993).
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449:and plays an important role in parapatric and
371:stepped clines, and obliquely stepped clines.
140:and blurring any distinct genetic boundaries.
1904:
1521:
1171:
535:gulls interbreed in a ring around the arctic.
1331:
1196:
687:Geographic Variation, Speciation, and Clines
127:(also known as migration). Selection causes
1484:, Richard J. Huggett, 2nd ed. (2004), p. 20
1197:Mallet, James; Barton, Nicholas H. (1989).
1123:
1080:
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868:
821:. Belknap Press of Harvard University Press
93:, meaning "to lean.” While it and the term
1911:
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1332:Jain, S K; Bradshaw, A D (1 August 1966).
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1138:Hybrid Zones and the Evolutionary Process
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875:. Springer Science & Business Media.
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3145:Transgenerational epigenetic inheritance
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1383:Coyne, Jerry A.; Orr, H. Allen (2004).
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833:
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1083:"Alternative forms for genomic clines"
1081:Fitzpatrick, Benjamin M. (July 2013).
1026:"Hypothetical Mechanism of Speciation"
912:
895:
683:
619:
299:No selection (drift/migration balance)
272:). For example, aposematic signals in
85:was coined by Huxley in 1938 from the
53:. Clines usually have a genetic (e.g.
2908:Dialogues Concerning Natural Religion
1892:
1509:
1172:Jorgensen, Sven; Fath, Brian (2014).
857:
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465:
436:and thereby instigate the process of
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1457:Geoecology: An evolutionary approach
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499:Other ecogeographical rules include
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208:
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107:Drivers and the evolution of clines
30:. For people named Gene Klein, see
13:
2313:Evolutionary developmental biology
1216:10.1111/j.1558-5646.1989.tb04237.x
1043:10.1111/j.1558-5646.1969.tb03550.x
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282:. This is because heterozygosity,
14:
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3167:
3158:
3157:
1870:
1858:
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1372:. Jones & Bartlett Learning.
312:Clinal structure and terminology
2970:Extended evolutionary synthesis
2159:Gene-centered view of evolution
1487:
1475:
1446:
1401:
1376:
1325:
1276:
1247:
1190:
1074:
955:
3098:Hologenome theory of evolution
2965:History of molecular evolution
2191:Evolutionarily stable strategy
2080:Last universal common ancestor
1535:
906:
889:
563:Larus argentatus smithsonianus
456:
1:
2892:Renaissance and Enlightenment
1496:Encyclopaedia of Anthropology
1135:Harrison, Richard G. (1993).
599:
280:positive frequency dependence
266:frequency-dependent selection
32:Eugene Klein (disambiguation)
3219:Modern human genetic history
3103:Missing heritability problem
2730:Gamete differentiation/sexes
1482:Fundamentals of biogeography
1453:Huggett, Richard J. (1995).
1410:Fundamentals of Biogeography
1408:Huggett, Richard J. (2004).
869:Felsenstein, Joseph (2013).
853:. Columbia University Press.
386:Discontinuous stepped clines
162:soot there is on vegetation.
7:
1865:Evolutionary biology Portal
1301:10.1534/genetics.111.129817
1141:. Oxford University Press.
896:Darwin, Charles R. (1859).
539:Larus argentatus argentatus
507:in 1857, which states that
10:
3250:
2735:Life cycles/nuclear phases
2287:Trivers–Willard hypothesis
1432:. CRC Press. p. 163.
567:Larus argentatus argenteus
212:
25:
18:
3229:Biological classification
3153:
3053:
2978:
2882:
2809:
2765:
2620:
2524:
2341:
2300:
2233:Parent–offspring conflict
2169:
2038:Earliest known life forms
1959:
1926:
1830:
1777:
1726:
1669:
1617:
1571:
1543:
1254:Futuyma, Douglas (2005).
931:10.1093/genetics/28.2.114
238:heterozygote disadvantage
3086:Cultural group selection
2950:The eclipse of Darwinism
2922:On the Origin of Species
2897:Transmutation of species
1708:Nonecological speciation
1368:Hedrick, Philip (2011).
1174:Encyclopaedia of Ecology
684:Endler, John A. (1977).
555:Larus argentatus birulai
3091:Dual inheritance theory
2930:History of paleontology
1494:Birx, H. James (2006).
1370:Genetics of Populations
1024:Bazykin, A. D. (1969).
989:10.1073/pnas.0803785105
915:"Isolation by Distance"
913:Wright, Sewall (1943).
153:Primary differentiation
2779:Punctuated equilibrium
2100:Non-adaptive radiation
2048:Evolutionary arms race
1813:Punctuated equilibrium
1769:Character displacement
1589:Reproductive isolation
1558:Laboratory experiments
1262:. Sinauer Associates.
902:. London: John Murray.
569:
559:Larus argentatus vegae
524:
475:
434:reproductive isolation
414:
349:
321:
225:
163:
116:
3071:Evolutionary medicine
2945:Mendelian inheritance
2653:Biological complexity
2641:Programmed cell death
2333:Phenotypic plasticity
2053:Evolutionary pressure
2043:Evidence of evolution
1941:Timeline of evolution
1694:Ecological speciation
1609:Evidence of evolution
1087:Ecology and Evolution
594:isolation by distance
551:Larus fuscus heuglini
530:
522:
473:
430:parapatric speciation
419:allopatric speciation
412:Parapatric speciation
410:
403:Clines and speciation
347:
319:
222:
181:isolation by distance
160:
144:Development of clines
114:
3199:Evolutionary biology
3045:Teleology in biology
2940:Blending inheritance
2318:Genetic assimilation
2181:Artificial selection
1920:Evolutionary biology
1426:Wool, David (2006).
845:Mayr, Ernst (1942).
813:Mayr, Ernst (1963).
748:. World Scientific.
451:sympatric speciation
19:For other uses, see
3224:Population genetics
3209:Kinship and descent
3194:Ecology terminology
3108:Molecular evolution
3066:Ecological genetics
2935:Transitional fossil
2725:Sexual reproduction
2565:endomembrane system
2494:pollinator-mediated
2450:dolphins and whales
2228:Parental investment
1698:Parallel speciation
1412:. Psychology Press.
1351:10.1038/hdy.1966.42
980:2008PNAS..10516212S
974:(42): 16212–16217.
778:. CABI Publishing.
634:1938Natur.142..219H
3081:Cultural evolution
2196:Fisher's principle
2125:Handicap principle
2115:Parallel evolution
1979:Adaptive radiation
1778:Speciation in taxa
1713:Assortative mating
872:Numerical Taxonomy
570:
525:
481:ecological "rules"
476:
466:Examples of clines
415:
350:
322:
226:
164:
117:
51:geographical range
3214:Landscape ecology
3204:Genetic genealogy
3181:
3180:
2797:Uniformitarianism
2750:Sex-determination
2255:Sexual dimorphism
2250:Natural selection
2154:Unit of selection
2120:Signalling theory
1886:
1885:
1764:Secondary contact
1736:Hybrid speciation
1684:Natural selection
1671:Isolating factors
628:(3587): 219–220.
489:Constantin Gloger
357:Continuous clines
215:Secondary contact
209:Secondary contact
138:genetic admixture
3241:
3171:
3161:
3160:
2960:Modern synthesis
2720:Multicellularity
2715:Mosaic evolution
2600:auditory ossicle
2282:Social selection
2265:Flowering plants
2260:Sexual selection
1913:
1906:
1899:
1890:
1889:
1874:
1873:
1862:
1850:
1849:
1838:
1837:
1689:Sexual selection
1618:Geographic modes
1530:
1523:
1516:
1507:
1506:
1500:
1499:
1491:
1485:
1479:
1473:
1472:
1460:
1450:
1444:
1443:
1423:
1414:
1413:
1405:
1399:
1398:
1380:
1374:
1373:
1365:
1356:
1355:
1353:
1329:
1323:
1322:
1312:
1280:
1274:
1273:
1261:
1251:
1245:
1244:
1218:
1194:
1188:
1187:
1169:
1160:
1159:
1157:
1155:
1132:
1121:
1120:
1110:
1099:10.1002/ece3.609
1093:(7): 1951–1966.
1078:
1072:
1071:
1045:
1021:
1012:
1011:
1001:
991:
959:
953:
952:
942:
910:
904:
903:
893:
887:
886:
866:
855:
854:
852:
842:
831:
830:
828:
826:
820:
810:
797:
796:
794:
792:
769:
760:
759:
739:
730:
729:
723:
719:
717:
709:
681:
654:
653:
642:10.1038/142219a0
617:
193:Biston betularia
136:through causing
55:allele frequency
45:is a measurable
3249:
3248:
3244:
3243:
3242:
3240:
3239:
3238:
3184:
3183:
3182:
3177:
3149:
3076:Group selection
3049:
2974:
2878:
2805:
2767:Tempo and modes
2761:
2616:
2520:
2337:
2296:
2172:
2165:
2142:Species complex
1955:
1946:History of life
1922:
1917:
1887:
1882:
1826:
1809:Paleopolyploidy
1773:
1728:Hybrid concepts
1722:
1665:
1613:
1583:Species complex
1567:
1539:
1534:
1504:
1503:
1492:
1488:
1480:
1476:
1469:
1451:
1447:
1440:
1424:
1417:
1406:
1402:
1395:
1381:
1377:
1366:
1359:
1330:
1326:
1281:
1277:
1270:
1252:
1248:
1195:
1191:
1184:
1170:
1163:
1153:
1151:
1149:
1133:
1124:
1079:
1075:
1022:
1015:
960:
956:
911:
907:
894:
890:
883:
867:
858:
843:
834:
824:
822:
811:
800:
790:
788:
786:
775:Forest Genetics
770:
763:
756:
740:
733:
721:
720:
711:
710:
698:
682:
657:
618:
607:
602:
536:
501:Bergmann's Rule
468:
459:
405:
388:
359:
342:
340:Types of clines
314:
301:
217:
211:
155:
146:
109:
35:
24:
17:
12:
11:
5:
3247:
3237:
3236:
3231:
3226:
3221:
3216:
3211:
3206:
3201:
3196:
3179:
3178:
3176:
3175:
3165:
3154:
3151:
3150:
3148:
3147:
3142:
3137:
3132:
3127:
3126:
3125:
3115:
3110:
3105:
3100:
3095:
3094:
3093:
3088:
3083:
3073:
3068:
3063:
3057:
3055:
3051:
3050:
3048:
3047:
3042:
3041:
3040:
3035:
3030:
3029:
3028:
3018:
3013:
3008:
3003:
2998:
2988:
2982:
2980:
2976:
2975:
2973:
2972:
2967:
2962:
2957:
2952:
2947:
2942:
2937:
2932:
2927:
2926:
2925:
2916:Charles Darwin
2913:
2912:
2911:
2899:
2894:
2888:
2886:
2880:
2879:
2877:
2876:
2871:
2866:
2861:
2856:
2854:Non-ecological
2851:
2846:
2841:
2836:
2831:
2826:
2821:
2815:
2813:
2807:
2806:
2804:
2803:
2794:
2785:
2771:
2769:
2763:
2762:
2760:
2759:
2754:
2753:
2752:
2747:
2742:
2737:
2732:
2722:
2717:
2712:
2707:
2702:
2697:
2692:
2687:
2682:
2677:
2672:
2671:
2670:
2660:
2655:
2650:
2645:
2644:
2643:
2638:
2627:
2625:
2618:
2617:
2615:
2614:
2613:
2612:
2607:
2605:nervous system
2602:
2597:
2592:
2584:
2583:
2582:
2577:
2572:
2567:
2562:
2557:
2547:
2542:
2537:
2531:
2529:
2522:
2521:
2519:
2518:
2513:
2508:
2503:
2498:
2497:
2496:
2486:
2485:
2484:
2479:
2478:
2477:
2472:
2462:
2457:
2452:
2447:
2442:
2441:
2440:
2435:
2425:
2415:
2410:
2409:
2408:
2398:
2393:
2388:
2383:
2382:
2381:
2371:
2366:
2365:
2364:
2354:
2348:
2346:
2339:
2338:
2336:
2335:
2330:
2325:
2320:
2315:
2310:
2304:
2302:
2298:
2297:
2295:
2294:
2289:
2284:
2279:
2278:
2277:
2272:
2267:
2257:
2252:
2247:
2242:
2237:
2236:
2235:
2230:
2220:
2215:
2210:
2209:
2208:
2198:
2193:
2188:
2183:
2177:
2175:
2167:
2166:
2164:
2163:
2162:
2161:
2151:
2146:
2145:
2144:
2139:
2129:
2128:
2127:
2117:
2112:
2107:
2105:Origin of life
2102:
2097:
2092:
2090:Microevolution
2087:
2085:Macroevolution
2082:
2077:
2072:
2071:
2070:
2060:
2055:
2050:
2045:
2040:
2035:
2030:
2025:
2023:Common descent
2020:
2019:
2018:
2008:
2003:
2001:Baldwin effect
1998:
1997:
1996:
1991:
1981:
1976:
1971:
1965:
1963:
1957:
1956:
1954:
1953:
1948:
1943:
1938:
1933:
1927:
1924:
1923:
1916:
1915:
1908:
1901:
1893:
1884:
1883:
1881:
1880:
1868:
1856:
1844:
1831:
1828:
1827:
1825:
1824:
1817:Macroevolution
1802:
1797:
1792:
1787:
1781:
1779:
1775:
1774:
1772:
1771:
1766:
1761:
1751:
1732:
1730:
1724:
1723:
1721:
1720:
1718:Haldane's rule
1715:
1710:
1705:
1691:
1686:
1681:
1675:
1673:
1667:
1666:
1664:
1663:
1658:
1644:
1641:Founder effect
1621:
1619:
1615:
1614:
1612:
1611:
1606:
1601:
1596:
1591:
1586:
1575:
1573:
1572:Basic concepts
1569:
1568:
1566:
1565:
1560:
1555:
1550:
1544:
1541:
1540:
1533:
1532:
1525:
1518:
1510:
1502:
1501:
1486:
1474:
1467:
1445:
1438:
1415:
1400:
1393:
1375:
1357:
1344:(3): 407–441.
1324:
1295:(1): 227–235.
1275:
1268:
1246:
1209:(2): 421–431.
1189:
1182:
1161:
1147:
1122:
1073:
1036:(4): 685–687.
1013:
954:
925:(2): 114–138.
905:
888:
881:
856:
832:
798:
784:
761:
754:
731:
722:|journal=
697:978-0691081922
696:
655:
604:
603:
601:
598:
487:, named after
467:
464:
458:
455:
404:
401:
400:
399:
396:
387:
384:
383:
382:
381:
380:
376:
368:
358:
355:
341:
338:
313:
310:
300:
297:
213:Main article:
210:
207:
154:
151:
145:
142:
108:
105:
15:
9:
6:
4:
3:
2:
3246:
3235:
3232:
3230:
3227:
3225:
3222:
3220:
3217:
3215:
3212:
3210:
3207:
3205:
3202:
3200:
3197:
3195:
3192:
3191:
3189:
3174:
3170:
3166:
3164:
3156:
3155:
3152:
3146:
3143:
3141:
3138:
3136:
3133:
3131:
3128:
3124:
3121:
3120:
3119:
3118:Phylogenetics
3116:
3114:
3111:
3109:
3106:
3104:
3101:
3099:
3096:
3092:
3089:
3087:
3084:
3082:
3079:
3078:
3077:
3074:
3072:
3069:
3067:
3064:
3062:
3059:
3058:
3056:
3052:
3046:
3043:
3039:
3036:
3034:
3031:
3027:
3024:
3023:
3022:
3021:Structuralism
3019:
3017:
3014:
3012:
3009:
3007:
3004:
3002:
2999:
2997:
2996:Catastrophism
2994:
2993:
2992:
2989:
2987:
2984:
2983:
2981:
2977:
2971:
2968:
2966:
2963:
2961:
2958:
2956:
2955:Neo-Darwinism
2953:
2951:
2948:
2946:
2943:
2941:
2938:
2936:
2933:
2931:
2928:
2924:
2923:
2919:
2918:
2917:
2914:
2910:
2909:
2905:
2904:
2903:
2900:
2898:
2895:
2893:
2890:
2889:
2887:
2885:
2881:
2875:
2872:
2870:
2869:Reinforcement
2867:
2865:
2862:
2860:
2857:
2855:
2852:
2850:
2847:
2845:
2842:
2840:
2837:
2835:
2832:
2830:
2827:
2825:
2822:
2820:
2817:
2816:
2814:
2812:
2808:
2802:
2801:Catastrophism
2798:
2795:
2793:
2792:Macromutation
2789:
2788:Micromutation
2786:
2784:
2780:
2776:
2773:
2772:
2770:
2768:
2764:
2758:
2755:
2751:
2748:
2746:
2743:
2741:
2738:
2736:
2733:
2731:
2728:
2727:
2726:
2723:
2721:
2718:
2716:
2713:
2711:
2708:
2706:
2703:
2701:
2698:
2696:
2695:Immune system
2693:
2691:
2688:
2686:
2683:
2681:
2678:
2676:
2673:
2669:
2666:
2665:
2664:
2661:
2659:
2656:
2654:
2651:
2649:
2646:
2642:
2639:
2637:
2634:
2633:
2632:
2629:
2628:
2626:
2624:
2619:
2611:
2608:
2606:
2603:
2601:
2598:
2596:
2593:
2591:
2588:
2587:
2585:
2581:
2578:
2576:
2573:
2571:
2568:
2566:
2563:
2561:
2558:
2556:
2555:symbiogenesis
2553:
2552:
2551:
2548:
2546:
2543:
2541:
2538:
2536:
2533:
2532:
2530:
2528:
2523:
2517:
2514:
2512:
2509:
2507:
2504:
2502:
2499:
2495:
2492:
2491:
2490:
2487:
2483:
2480:
2476:
2473:
2471:
2468:
2467:
2466:
2463:
2461:
2458:
2456:
2453:
2451:
2448:
2446:
2443:
2439:
2436:
2434:
2431:
2430:
2429:
2426:
2424:
2421:
2420:
2419:
2416:
2414:
2411:
2407:
2404:
2403:
2402:
2399:
2397:
2394:
2392:
2389:
2387:
2384:
2380:
2377:
2376:
2375:
2372:
2370:
2367:
2363:
2360:
2359:
2358:
2355:
2353:
2350:
2349:
2347:
2345:
2340:
2334:
2331:
2329:
2326:
2324:
2321:
2319:
2316:
2314:
2311:
2309:
2306:
2305:
2303:
2299:
2293:
2290:
2288:
2285:
2283:
2280:
2276:
2273:
2271:
2268:
2266:
2263:
2262:
2261:
2258:
2256:
2253:
2251:
2248:
2246:
2243:
2241:
2238:
2234:
2231:
2229:
2226:
2225:
2224:
2223:Kin selection
2221:
2219:
2218:Genetic drift
2216:
2214:
2211:
2207:
2204:
2203:
2202:
2199:
2197:
2194:
2192:
2189:
2187:
2184:
2182:
2179:
2178:
2176:
2174:
2168:
2160:
2157:
2156:
2155:
2152:
2150:
2147:
2143:
2140:
2138:
2135:
2134:
2133:
2130:
2126:
2123:
2122:
2121:
2118:
2116:
2113:
2111:
2108:
2106:
2103:
2101:
2098:
2096:
2093:
2091:
2088:
2086:
2083:
2081:
2078:
2076:
2073:
2069:
2066:
2065:
2064:
2061:
2059:
2056:
2054:
2051:
2049:
2046:
2044:
2041:
2039:
2036:
2034:
2031:
2029:
2026:
2024:
2021:
2017:
2014:
2013:
2012:
2009:
2007:
2004:
2002:
1999:
1995:
1992:
1990:
1987:
1986:
1985:
1982:
1980:
1977:
1975:
1972:
1970:
1967:
1966:
1964:
1962:
1958:
1952:
1949:
1947:
1944:
1942:
1939:
1937:
1934:
1932:
1929:
1928:
1925:
1921:
1914:
1909:
1907:
1902:
1900:
1895:
1894:
1891:
1879:
1878:
1869:
1867:
1866:
1861:
1857:
1855:
1854:
1845:
1843:
1842:
1833:
1832:
1829:
1822:
1821:Chronospecies
1818:
1814:
1810:
1806:
1803:
1801:
1798:
1796:
1793:
1791:
1788:
1786:
1783:
1782:
1780:
1776:
1770:
1767:
1765:
1762:
1759:
1755:
1754:Reinforcement
1752:
1749:
1748:Recombination
1745:
1741:
1737:
1734:
1733:
1731:
1729:
1725:
1719:
1716:
1714:
1711:
1709:
1706:
1703:
1699:
1695:
1692:
1690:
1687:
1685:
1682:
1680:
1677:
1676:
1674:
1672:
1668:
1662:
1659:
1656:
1652:
1648:
1645:
1642:
1638:
1634:
1630:
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3061:Biogeography
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3006:Orthogenesis
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2839:Cospeciation
2834:Cladogenesis
2783:Saltationism
2740:Mating types
2663:Color vision
2648:Avian flight
2570:mitochondria
2308:Canalisation
2186:Biodiversity
1931:Introduction
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1599:Cladogenesis
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2757:Snake venom
2690:Eusociality
2668:in primates
2658:Cooperation
2586:In animals
2406:butterflies
2379:Cephalopods
2369:Brachiopods
2301:Development
2275:Mate choice
2028:Convergence
2011:Coevolution
1969:Abiogenesis
1877:WikiProject
1637:Centrifugal
1387:. Sinauer.
509:homeotherms
457:Clinal maps
442:pre-zygotic
270:aposematism
28:Gene Clines
3188:Categories
3001:Lamarckism
2979:Philosophy
2902:David Hume
2864:Peripatric
2859:Parapatric
2844:Ecological
2824:Anagenesis
2819:Allopatric
2811:Speciation
2775:Gradualism
2700:Metabolism
2560:chromosome
2550:Eukaryotes
2328:Modularity
2245:Population
2171:Population
2132:Speciation
2110:Panspermia
2063:Extinction
2058:Exaptation
2033:Divergence
2006:Cladistics
1994:Reciprocal
1974:Adaptation
1740:Polyploidy
1702:Allochrony
1679:Adaptation
1647:Parapatric
1629:Peripatric
1625:Allopatric
1594:Anagenesis
1537:Speciation
1385:Speciation
1176:. Newnes.
600:References
578:interbreed
438:speciation
275:Heliconius
246:homozygous
185:speciation
173:phenotypes
134:speciation
129:adaptation
72:subspecies
59:blood type
3135:Protocell
2986:Darwinism
2874:Sympatric
2623:processes
2511:Tetrapods
2460:Kangaroos
2386:Dinosaurs
2323:Inversion
2292:Variation
2213:Gene flow
2206:Inclusive
2016:Mutualism
1961:Evolution
1661:Sympatric
1258:Evolution
1203:Evolution
1030:Evolution
724:ignored (
714:cite book
284:mutations
230:allopatry
169:genotypes
125:gene flow
121:selection
81:The term
3163:Category
3038:Vitalism
3033:Theistic
3026:Spandrel
2710:Morality
2705:Monogamy
2580:plastids
2545:Flagella
2501:Reptiles
2482:sea cows
2465:primates
2374:Molluscs
2352:Bacteria
2240:Mutation
2173:genetics
2149:Taxonomy
2095:Mismatch
2075:Homology
1989:Cheating
1984:Altruism
1841:Category
1758:evidence
1563:Glossary
1338:Heredity
1319:21705747
1289:Genetics
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1233:28568556
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919:Genetics
791:28 March
328:Agrostis
89:κλίνειν
47:gradient
3234:Species
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2884:History
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2575:nucleus
2516:Viruses
2506:Spiders
2418:Mammals
2401:Insects
2201:Fitness
2137:Species
1936:Outline
1853:Commons
1805:Fossils
1795:Insects
1744:Klepton
1633:Quantum
1579:Species
1553:History
1498:. SAGE.
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976:Bibcode
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630:Bibcode
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254:zygotes
202:cryptic
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96:ecotype
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76:species
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2270:Fungi
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1951:Index
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1221:JSTOR
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