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In biology, a cline is a measurable gradient in a single characteristic (or biological trait) of a species across its geographical range. Clines usually have a genetic (e.g. allele frequency, blood type), 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.
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 Julian Huxley 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 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 subspecies or species are.
The term cline was coined by Huxley in 1938 from the Ancient Greek κλίνειν klinein, meaning "to lean.” While it and the term ecotype are sometimes used interchangeably, they do in fact differ in that ecotype 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.
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 isolation by distance 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, speciation is likely to eventually occur, as local adaptation can cause reproductive isolation between populations.
A classic example of the role of environmental gradients in creating clines is that of the peppered moth, Peppered moth, 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 morphs 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 Crypsis 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 limitations to sooty emissions were introduced in the 1960s.
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 in characters through a variety of mechanisms. One way it may act is through Underdominance, 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 AA (homozygote dominant) or aa (homozygous recessive) individuals. The cline of hetero 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. 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 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".
Another way in which selection can generate clines is through frequency-dependent selection. Characters that could be maintained by such frequency-dependent selective pressures include warning signals (aposematism). For example, aposematic signals in Heliconius butterflies sometimes display steep clines between populations, which are maintained through positive frequency dependence. This is because heterozygosity, and recombination 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 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.
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 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.
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.
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 parapatric speciation. Both extrinsic and intrinsic selection can serve to generate varying degrees of reproductive isolation and thereby instigate the process of speciation. 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 pre-zygotic 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. (2025). 9780080914565, Newnes. ISBN 9780080914565 This is known as reinforcement and plays an important role in parapatric and sympatric speciation. (2025). 9780878930890, Sinauer. ISBN 9780878930890
Other ecogeographical rules include Bergmann's Rule, coined by Carl Bergmann in 1857, which states that Homeothermy 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 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 house sparrow, 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.
Ring species Fundamentals of biogeography, Richard J. Huggett, 2nd ed. (2004), p. 20 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 interbreed 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 Larus 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.
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 towards Europe around 2,000 years ago, causing genetic admixture in an isolation by distance 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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