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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.
520: 471: 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 408: 158: 112: 345: 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. 317: 220: 528: 325:
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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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
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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").
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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
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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
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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 291:
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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Interbreeding populations represented by a gradient of coloured circles around a geographic barrier
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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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Two populations with individuals moving between the populations to demonstrate gene flow
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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 407: 3075: 2699: 2526: 2515: 2469: 2412: 2361: 2141: 1945: 1808: 1582: 197: 50: 1300: 1256: 930: 491:, who observed in 1833 that environmental factors and the pigmentation of avian 157: 2915: 2787: 2729: 2089: 2084: 2022: 2000: 1816: 1640: 237: 1285:"Genetic Drift Widens the Expected Cline but Narrows the Expected Cline Width" 3187: 3117: 2995: 2954: 2800: 2554: 2222: 2217: 1820: 513: 344: 304: 236:
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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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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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).
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If I Am To Be Remembered: Correspondence Of Julian Huxley
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staircase as in the case of horizontally stepped clines.
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further sub-divided into smooth and stepped clines.
311: 1454: 1255: 846: 814: 737: 735: 1283:Polechová, Jitka; Barton, Nick (September 2011). 767: 765: 741: 3185: 742:Dronamraju, Krishna R.; Needham, Joseph (1993). 1282: 771: 732: 385: 1130: 1128: 1126: 1019: 1017: 762: 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: 1014: 868: 821:. Belknap Press of Harvard University Press 93:, meaning "to lean.” While it and the term 1911: 1897: 1528: 1514: 1363: 1361: 1332:Jain, S K; Bradshaw, A D (1 August 1966). 840: 838: 836: 808: 806: 804: 802: 152: 1349: 1308: 1214: 1138:Hybrid Zones and the Evolutionary Process 1106: 1041: 997: 987: 938: 875:. 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Clines usually have a genetic (e.g. 2908:Dialogues Concerning Natural Religion 1892: 1509: 1172:Jorgensen, Sven; Fath, Brian (2014). 857: 656: 606: 465: 436:and thereby instigate the process of 1847: 1493: 1457:Geoecology: An evolutionary approach 1425: 1416: 844: 812: 499:Other ecogeographical rules include 356: 208: 1871: 1162: 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 339: 282:. This is because heterozygosity, 14: 3245: 3167: 3158: 3157: 1870: 1858: 1846: 1835: 1834: 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:. 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(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: 1626: 1623: 1622: 1620: 1616: 1610: 1607: 1605: 1602: 1600: 1597: 1595: 1592: 1590: 1587: 1584: 1580: 1577: 1576: 1574: 1570: 1564: 1561: 1559: 1556: 1554: 1551: 1549: 1546: 1545: 1542: 1538: 1531: 1526: 1524: 1519: 1517: 1512: 1511: 1508: 1497: 1490: 1483: 1478: 1470: 1468:9780415086899 1464: 1461:. Routledge. 1459: 1458: 1449: 1441: 1439:9781578084456 1435: 1431: 1430: 1422: 1420: 1411: 1404: 1396: 1394:9780878930890 1390: 1386: 1379: 1371: 1364: 1362: 1352: 1347: 1343: 1339: 1335: 1328: 1320: 1316: 1311: 1306: 1302: 1298: 1294: 1290: 1286: 1279: 1271: 1269:9780878931873 1265: 1260: 1259: 1250: 1242: 1238: 1234: 1230: 1226: 1222: 1217: 1212: 1208: 1204: 1200: 1193: 1185: 1183:9780080914565 1179: 1175: 1168: 1166: 1150: 1148:9780195069174 1144: 1140: 1139: 1131: 1129: 1127: 1118: 1114: 1109: 1104: 1100: 1096: 1092: 1088: 1084: 1077: 1069: 1065: 1061: 1057: 1053: 1049: 1044: 1039: 1035: 1031: 1027: 1020: 1018: 1009: 1005: 1000: 995: 990: 985: 981: 977: 973: 969: 965: 958: 950: 946: 941: 936: 932: 928: 924: 920: 916: 909: 901: 900: 892: 884: 882:9783642690242 878: 874: 873: 865: 863: 861: 851: 850: 841: 839: 837: 819: 818: 809: 807: 805: 803: 787: 785:9781845932862 781: 777: 776: 768: 766: 757: 755:9789814505192 751: 747: 746: 738: 736: 727: 715: 707: 703: 699: 693: 689: 688: 680: 678: 676: 674: 672: 670: 668: 666: 664: 662: 660: 651: 647: 643: 639: 635: 631: 627: 623: 616: 614: 612: 610: 605: 597: 595: 589: 586: 585: 579: 574: 568: 564: 560: 556: 552: 548: 547:sensu stricto 544: 540: 534: 529: 521: 517: 515: 514:house sparrow 510: 506: 505:Carl Bergmann 502: 497: 494: 490: 486: 485:Gloger's Rule 482: 472: 463: 454: 452: 448: 447:reinforcement 443: 439: 435: 431: 426: 424: 420: 413: 409: 397: 394: 393: 392: 377: 373: 372: 369: 365: 364: 363: 354: 346: 337: 333: 330: 329: 318: 309: 306: 305:genetic drift 296: 292: 289: 288:recombination 285: 281: 277: 276: 271: 267: 262: 260: 255: 251: 248:dominant) or 247: 243: 239: 233: 231: 221: 216: 206: 203: 199: 195: 194: 188: 186: 182: 176: 174: 170: 159: 150: 141: 139: 135: 130: 126: 122: 113: 104: 102: 98: 97: 92: 88: 84: 79: 77: 73: 68: 67:Julian Huxley 62: 60: 56: 52: 48: 44: 40: 33: 29: 22: 3130:Polymorphism 3113:Astrobiology 3061:Biogeography 3016:Saltationism 3006:Orthogenesis 2991:Alternatives 2920: 2906: 2839:Cospeciation 2834:Cladogenesis 2783:Saltationism 2740:Mating types 2663:Color vision 2648:Avian flight 2570:mitochondria 2308:Canalisation 2186:Biodiversity 1931:Introduction 1875: 1863: 1851: 1839: 1655:Ring species 1650: 1604:Cospeciation 1599:Cladogenesis 1548:Introduction 1495: 1489: 1481: 1477: 1456: 1448: 1428: 1409: 1403: 1384: 1378: 1369: 1341: 1337: 1327: 1292: 1288: 1278: 1257: 1249: 1206: 1202: 1192: 1173: 1152:. Retrieved 1137: 1090: 1086: 1076: 1033: 1029: 971: 967: 957: 922: 918: 908: 898: 891: 871: 848: 823:. Retrieved 816: 789:. Retrieved 774: 744: 686: 625: 621: 590: 582: 573:Ring species 571: 566: 562: 558: 554: 550: 546: 543:Larus fuscus 542: 538: 532: 503:, coined by 498: 477: 460: 427: 423:ring species 416: 389: 360: 351: 334: 326: 323: 302: 293: 273: 263: 259:hybrid zones 249: 241: 234: 227: 224:populations. 192: 189: 177: 165: 147: 118: 100: 94: 90: 82: 80: 63: 42: 36: 3140:Systematics 3011:Mutationism 2829:Catagenesis 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 1241:15899619 1233:28568556 1154:17 April 1117:23919142 1068:33832575 1060:28562864 1008:18854412 949:17247074 919:Genetics 791:28 March 328:Agrostis 89:κλίνειν 47:gradient 3234:Species 3054:Related 2884:History 2745:Meiosis 2680:Empathy 2675:Emotion 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. 1310:3176109 1225:2409217 1108:3728937 1052:2406862 999:2571026 976:Bibcode 940:1209196 825:1 April 650:4124055 630:Bibcode 493:plumage 367:common. 254:zygotes 202:cryptic 101:ecotype 96:ecotype 91:klinein 76:species 39:biology 3173:Portal 2849:Hybrid 2685:Ethics 2527:organs 2489:Plants 2475:lemurs 2470:humans 2455:horses 2445:hyenas 2433:wolves 2428:canids 2362:origin 1800:Plants 1651:Clines 1465:  1436:  1391:  1317:  1307:  1266:  1239:  1231:  1223:  1180:  1145:  1115:  1105:  1066:  1058:  1050:  1006:  996:  947:  937:  879:  782:  752:  706:409931 704:  694:  648:  622:Nature 549:), 3: 198:morphs 2636:Death 2631:Aging 2610:brain 2396:Fungi 2357:Birds 2270:Fungi 2068:Event 1951:Index 1785:Birds 1237:S2CID 1221:JSTOR 1064:S2CID 1048:JSTOR 646:S2CID 584:Larus 565:, 7: 561:, 6: 557:, 5: 553:, 4: 541:, 2: 533:Larus 87:Greek 83:cline 78:are. 43:cline 21:Cline 3123:Tree 2595:hair 2535:Cell 2438:dogs 2423:cats 2413:Life 2391:Fish 2344:taxa 1790:Fish 1463:ISBN 1434:ISBN 1389:ISBN 1315:PMID 1264:ISBN 1229:PMID 1178:ISBN 1156:2018 1143:ISBN 1113:PMID 1056:PMID 1004:PMID 968:PNAS 945:PMID 877:ISBN 827:2018 793:2018 780:ISBN 750:ISBN 726:help 702:PMID 692:ISBN 531:The 286:and 123:and 41:, a 2621:Of 2590:eye 2540:DNA 2525:Of 2342:Of 1346:doi 1305:PMC 1297:doi 1293:189 1211:doi 1103:PMC 1095:doi 1038:doi 994:PMC 984:doi 972:105 935:PMC 927:doi 638:doi 626:142 537:1: 171:or 74:or 37:In 3190:: 1819:· 1815:· 1811:· 1746:· 1742:· 1700:· 1653:· 1639:· 1635:· 1631:· 1418:^ 1360:^ 1342:21 1340:. 1336:. 1313:. 1303:. 1291:. 1287:. 1235:. 1227:. 1219:. 1207:43 1205:. 1201:. 1164:^ 1125:^ 1111:. 1101:. 1089:. 1085:. 1062:. 1054:. 1046:. 1034:23 1032:. 1028:. 1016:^ 1002:. 992:. 982:. 970:. 966:. 943:. 933:. 923:28 921:. 917:. 859:^ 835:^ 801:^ 764:^ 734:^ 718:: 716:}} 712:{{ 700:. 658:^ 644:. 636:. 624:. 608:^ 453:. 250:aa 242:AA 57:, 2799:/ 2790:/ 2781:/ 2777:/ 1912:e 1905:t 1898:v 1823:) 1807:( 1760:) 1756:( 1750:) 1738:( 1704:) 1696:( 1657:) 1649:( 1643:) 1627:( 1585:) 1581:( 1529:e 1522:t 1515:v 1471:. 1442:. 1397:. 1354:. 1348:: 1321:. 1299:: 1272:. 1243:. 1213:: 1186:. 1158:. 1119:. 1097:: 1091:3 1070:. 1040:: 1010:. 986:: 978:: 951:. 929:: 885:. 829:. 795:. 758:. 728:) 708:. 652:. 640:: 632:: 545:( 244:( 34:. 23:.

Index

Cline
Gene Clines
Eugene Klein (disambiguation)
biology
gradient
geographical range
allele frequency
blood type
Julian Huxley
subspecies
species
Greek
ecotype

selection
gene flow
adaptation
speciation
genetic admixture

genotypes
phenotypes
isolation by distance
speciation
Biston betularia
morphs
cryptic
Secondary contact

allopatry

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