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1.
Populations of the water snake, Nerodia sipedon, on islands in western Lake Erie are polymorphic for color pattern. These populations include banded, intermediate, and unbanded morphs while surrounding mainland populations consist solely of the banded morph. The hypothesis that this polymorphism is maintained by strong selection and migration pressures is widely accepted. Unbanded morphs are apparently more cryptic along island shorelines while banded morphs are more cryptic on the mainland. Migration of banded morphs from the mainland explains their persistence in island populations. Data collected in a capture-mark-recapture program on six islands provide no evidence of differential selection among morphs; morph frequencies do not differ among age classes, between once-captured and multiply-captured snakes, or between scarred and unscarred snakes. Furthermore, herring gulls, the most common snake predators in the island area, appear to detect banded and unbanded model snakes with equal ease. High site fidelity of water snakes and the distribution of morphs among islands suggest that migration from the mainland is not common. However, islands close to each other are similar in morph frequency, and water snakes have colonized islands elsewhere in the Great Lakes, indicating that some migration does occur. Recently, the frequency of banded morphs has increased in island populations while adult population sizes have declined. This increase in banded morphs is interpreted as reflecting an increased impact of migration from the mainland into these reduced populations. One scenario for the evolution and maintenance of this polymorphism is that selection was important in establishing unbanded morphs in island populations as they became isolated from the mainland. As populations declined to their present size, the impact of migration from the mainland increased and is now swamping the effect of selection. Further declines in island population size may result in fixation of the banded morph.  相似文献   

2.
A classic example of natural selection, that of color-pattern variation in Lake Erie island populations of water snakes, was reexamined to overcome shortcomings resulting from classification of snakes into discrete color-pattern categories and use of cross-sectional data. Four continuously varying color-pattern components (DB, the number of dorsal blotches; LB, the number of lateral blotches; ROWS, the height of lateral blotches measured in scale rows; and VEXT, the extent of ventral pigmentation) were analyzed. Patterns of natural selection were predicted from the relationship between color-pattern scores and independent measures of relative crypsis. Tests for natural selection were carried out using longitudinal data on neonate to juvenile-aged snakes and cross-sectional data on juvenile to adult-aged snakes. As predicted, the form of selection differed between younger and older age classes of snakes: selection resulted in a reduction in DB and LB among neonate and juvenile snakes but had little influence on color-pattern components in older snakes. The correspondence between observed patterns of natural selection and predictions based on the relationship between color-pattern scores and relative crypsis supports the hypothesis that differential predation by visual predators on younger age classes of snakes is the mechanism of selection. Gene flow from mainland populations or the initial lack of an allele necessary for reduced pattern may explain why selection has not resulted in greater differentiation between island and mainland populations.  相似文献   

3.
Local populations of the adder, Vipera berus, are polymorphic for dorsal colour pattern, containing both melanistic (black) and zig-zag patterned individuals. Colour patterns in snakes influence crypsis and thermoregulatory capacity and therefore may be subjected to natural selection. To find an explanation for the maintenance of this polymorphism I examined temporal and spatial variation in morph frequency, and tested for differential selection among morphs using data from a six year capture-mark-recapture study. The data derive from six groups of islands in the Baltic Sea off the Swedish east coast, two mainland localities near the coast, and one inland locality. Morph frequency did not change over time within a population but varied among populations: melanistic individuals were not found at the inland locality, but comprised from 17 to 62% of the coastal and island populations. Adders frequently moved between islands within a group, but the tendency to disperse was independent of morph. These results suggest that the polymorphism is stable and maintained by a deterministic process. Scar frequency was twice as high among melanistic as among zig-zag snakes, and melanistic individuals were easier to capture, indicating that predation may be higher on the melanistic morph. Colour morphs did not differ in body size, but analysis of recapture data shows evidence for differential survival among morphs. Zig-zag males survived better than melanistic males, but the relative survival rates of morphs were reversed in females. This difference was consistent through time and may be due to sexual differences in behaviour, with melanism increasing predation intensity when associated with male but not with female behaviour. Opposing fitness consequences of colour pattern in the two sexes may help maintain colour polymorphism within populations of Vipera berus.  相似文献   

4.
Polymorphisms provide one of the most useful tools for understanding the maintenance of genetic and phenotypic variation in nature. We have previously described a genetically based polymorphism in dorsal patterning that is expressed by female brown anole lizards, Anolis sagrei, which occur in Bar, Diamond and intermediate Diamond-Bar morphs. Previous studies of island populations in The Bahamas support a role for selection in maintaining the polymorphism, but the agents responsible for this selection remain unclear. We tested two main hypotheses regarding the importance of predation as a selective agent that maintains the polymorphism within populations. First, we tested whether correlational selection favours different combinations of morph, locomotor performance and escape behaviour by measuring morph-specific natural selection on sprint speed, running endurance and the propensity of females to either 'freeze' or 'run' in response to attempted capture. Morphs did not differ in any of these traits, nor did correlational selection consistently favour any particular combinations of morph and antipredator behaviour. Second, we experimentally excluded bird and snake predators from two entire island populations, allowed these predators access to two additional islands and then measured subsequent differences in natural selection on morphs in each population. Predators reduced the survival of Bar and Diamond females, but not of genetically intermediate Diamond-Bar females. These results provide limited evidence that predation may play a role in maintaining this polymorphism, although the functional traits that could account for differential susceptibility to predation remain unclear.  相似文献   

5.
The evolution of striking phenotypes on islands is a well‐known phenomenon, and there has been a long‐standing debate on the patterns of body size evolution on islands. The ecological causes driving divergence in insular populations are, however, poorly understood. Reduced predator fauna is expected to lower escape propensity, increase body size and relax selection for crypsis in small‐bodied, insular prey species. Here, we investigated whether escape behaviour, body size and dorsal coloration have diverged as predicted under predation release in spatially replicated islet and mainland populations of the lizard species Podarcis gaigeae. We show that islet lizards escape approaching observers at shorter distances and are larger than mainland lizards. Additionally, we found evidence for larger between‐population variation in body size among the islet populations than mainland populations. Moreover, islet populations are significantly more divergent in dorsal coloration and match their respective habitats poorer than mainland lizards. These results strongly suggest that predation release on islets has driven population divergence in phenotypic and behavioural traits and that selective release has affected both trait means and variances. Relaxed predation pressure is therefore likely to be one of the major ecological factors driving body size divergence on these islands.  相似文献   

6.
Gene flow and melanism in Lake Erie garter snake populations   总被引:3,自引:0,他引:3  
Melanistic garter snakes ( Thamnophis sirtalis ) are unusually common near Lake Erie, apparently because selection for thermoregulatory ability in cool lake-shore habitats (which favours melanistic morphs) outweighs selection for crypsis (which favours striped morphs). However, morph frequencies are highly variable among sites, suggesting that random genetic drift also influences colour pattern. In an effort to better understand the evolutionary processes influencing garter snake colour patterns, we estimated Fx and Nm (the number of migrants per generation) among island and mainland populations from patterns of allozymic variation detected using electrophoresis. Estimates of Nm were high, ranging from 2.7 to 37.6 between pairs of study sites and making it unlikely that differences in morph frequencies among sites were solely the result of random genetic drift. Furthermore, differences in F st estimates between colour pattern (a one-locus two-allele trait) and allozyme loci suggest that colour pattern alleles are not in Hardy-Weinberg equilibrium, most likely as a result of natural selection. Comparison of allozymic data from Lake Erie with those from more distant sites suggests that gene flow occurs over long distances in T. sirtalis.  相似文献   

7.
Introduced species have contributed significantly to the extinction of endemic species on islands. They also create new selection pressures on their prey that may result in modified life history strategies. Introduced viperine snakes (Natrix maura) have been implicated in the decline of the endemic midwife toad of Mallorca (Alytes muletensis). A comparison of A. muletensis tadpoles in natural pools with and without snakes showed that those populations subject to snake predation possessed longer tails with narrower tail fins but deeper tail muscles. Field and laboratory experiments showed that these changes in tail morphology could be induced by chemical and tactile cues from snakes. Populations of tadpoles that were subject to snake predation also displayed clear bimodal size-frequency distributions, with intermediate-sized tadpoles missing from the pools completely. Tadpoles in pools frequented by snakes developed faster in relation to their body size than those in pools without snakes. Variation in morphology between toad populations may therefore be caused by a combination of size-selective predation and tadpole plasticity. The results of this study indicate that the introduction of alien species can result in selection for induced defences, which may facilitate coexistence between predator and prey under certain conditions.  相似文献   

8.
It is a well-known phenomenon that islands can support populations of gigantic or dwarf forms of mainland conspecifics, but the variety of explanatory hypotheses for this phenomenon have been difficult to disentangle. The highly venomous Australian tiger snakes (genus Notechis) represent a well-known and extreme example of insular body size variation. They are of special interest because there are multiple populations of dwarfs and giants and the age of the islands and thus the age of the tiger snake populations are known from detailed sea level studies. Most are 5000-7000 years old and all are less than 10,000 years old. Here we discriminate between two competing hypotheses with a molecular phylogeography dataset comprising approximately 4800 bp of mtDNA and demonstrate that populations of island dwarfs and giants have evolved five times independently. In each case the closest relatives of the giant or dwarf populations are mainland tiger snakes, and in four of the five cases, the closest relatives are also the most geographically proximate mainland tiger snakes. Moreover, these body size shifts have evolved extremely rapidly and this is reflected in the genetic divergence between island body size variants and mainland snakes. Within south eastern Australia, where populations of island giants, populations of island dwarfs, and mainland tiger snakes all occur, the maximum genetic divergence is only 0.38%. Dwarf tiger snakes are restricted to prey items that are much smaller than the prey items of mainland tiger snakes and giant tiger snakes are restricted to seasonally available prey items that are up three times larger than the prey items of mainland tiger snakes. We support the hypotheses that these body size shifts are due to strong selection imposed by the size of available prey items, rather than shared evolutionary history, and our results are consistent with the notion that adaptive plasticity also has played an important role in body size shifts. We suggest that plasticity displayed early on in the occupation of these new islands provided the flexibility necessary as the island's available prey items became more depauperate, but once the size range of available prey items was reduced, strong natural selection followed by genetic assimilation worked to optimize snake body size. The rate of body size divergence in haldanes is similar for dwarfs (h(g) = 0.0010) and giants (h(g) = 0.0020-0.0025) and is in line with other studies of rapid evolution. Our data provide strong evidence for rapid and repeated morphological divergence in the wild due to similar selective pressures acting in different directions.  相似文献   

9.
For conservation purposes islands are considered safe refuges for many species, particularly in regions where introduced predators form a major threat to the native fauna, but island populations are also known to possess low levels of genetic diversity. The New Zealand archipelago provides an ideal system to compare genetic diversity of large mainland populations where introduced predators are common, to that of smaller offshore islands, which serve as predator-free refuges. We assessed microsatellite variation in South Island robins (Petroica australis australis), and compared large mainland, small mainland, natural island and translocated island populations. Large mainland populations exhibited more polymorphic loci and higher number of alleles than small mainland and natural island populations. Genetic variation did not differ between natural and translocated island populations, even though one of the translocated populations was established with five individuals. Hatching failure was recorded in a subset of the populations and found to be significantly higher in translocated populations than in a large mainland population. Significant population differentiation was largely based on heterogeneity in allele frequencies (including fixation of alleles), as few unique alleles were observed. This study shows that large mainland populations retain higher levels of genetic diversity than natural and translocated island populations. It highlights the importance of protecting these mainland populations and using them as a source for new translocations. In the future, these populations may become extremely valuable for species conservation if existing island populations become adversely affected by low levels of genetic variation and do not persist.  相似文献   

10.
In an effort to clarify the evolutionary processes influencing color-pattern variation in Lake Erie island water snake (Nerodia sipedon) populations, rates of gene flow among island and mainland populations were estimated from patterns of allozymic variation detected using electrophoresis. Rates of gene flow were high with Nm, the number of migrants per generation, averaging 25.5 among island sites, 9.2 between the Ontario mainland and the islands, and 3.6 between the Ohio mainland and the islands. Based on estimates of current population size from mark-recapture work and of past population size extrapolated from the extent of shoreline habitat, values of m between island and mainland populations ranged from 0.0008–0.01. Synthesis of estimates of the rate of gene flow with information on inheritance of color pattern, the strength of natural selection, and population history supports the hypothesis that color-pattern variation in island populations results from a balance between gene flow and natural selection. However, depending on the mode of inheritance of color pattern, stochastic processes such as drift may have been important in the initial stages of differentiation between island and mainland populations.  相似文献   

11.
Abstract Many animal species display striking color differences with respect to geographic location, sex, and body region. Traditional adaptive explanations for such complex patterns invoke an interaction between selection for conspicuous signals and natural selection for crypsis. Although there is now a substantial body of evidence supporting the role of sexual selection for signaling functions, quantitative studies of crypsis remain comparatively rare. Here, we combine objective measures of coloration with information on predator visual sensitivities to study the role of crypsis in the evolution of color variation in an Australian lizard species complex (Ctenophorus decresii). We apply a model that allows us to quantify crypsis in terms of the visual contrast of the lizards against their natural backgrounds, as perceived by potential avian predators. We then use these quantitative estimates of crypsis to answer the following questions. Are there significant differences in crypsis conspicuousness among populations? Are there significant differences in crypsis conspicuousness between the sexes? Are body regions “exposed” to visual predators more cryptic than “hidden” body regions? Is there evidence for local adaptation with respect to crypsis against different substrates? In general, our results confirmed that there are real differences in crypsis conspicuousness both between populations and between sexes; that exposed body regions were significantly more cryptic than hidden ones, particularly in females; and that females, but not males, are more cryptic against their own local background than against the background of other populations. Body regions that varied most in contrast between the sexes and between populations were also most conspicuous and are emphasized by males during social and sexual signaling. However, results varied with respect to the aspect of coloration studied. Results based on chromatic contrast (“hue’ of color) provided better support for the crypsis hypothesis than did results based on achromatic contrast (“brightness’ of color). Taken together, these results support the view that crypsis plays a substantial role in the evolution of color variation and that color patterns represent a balance between the need for conspicuousness for signaling and the need for crypsis to avoid predation.  相似文献   

12.
Variations in visible genetic polymorphisms are assumed to decrease in populations on small islands because of intense founder effects, genetic drift and inbreeding. However, we have found evidence of a marked enhancement of colour polymorphisms within populations on small oceanic islands that were colonized from the mainland. The source populations on the mainland of the land snail Euhadra peliomphala in four oceanic islands were estimated by phylogenetic analysis of mitochondrial DNA sequences. Diversity of shell colour was higher in the island populations than in the source populations on the mainland. In addition, the shell colour morphs differed not only among populations from different islands but also between the island populations and the source populations on the mainland. By contrast, no mtDNA variations were found in any of the island populations, even though the source populations possessed high mtDNA diversity. Thus, components of colour morphs changed in the island populations after their colonization, and colour polymorphisms are enhanced in these islands despite the loss of genetic variation. The above findings suggest that ecological mechanisms such as morphological release owing to a release from competition may overcome the tendency toward reduced genetic variation in islands to enhance the colour polymorphism.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 81 , 417–425.  相似文献   

13.
The mechanisms behind the evolution and maintenance of conspicuous visible polymorphisms comprising tens of morphs present a challenge to evolutionary theory. However, for cryptic forms Endler (Evol. Biol., 11, 1978, 319) conjectured that complex backgrounds facilitate polymorphism because in such habitats there are several ways to resemble the resting surface. We use computer simulation to explore the evolution of cryptic morphs on increasingly complex backgrounds under regimes that include selection for crypsis, apostatic predation and dietary wariness. We show that there is a monotonic increase in the number of morphs evolving in a population as the potential number of cryptic morphs increases. The relationship is very weak with selection for crypsis alone, but much stronger with the addition of apostatic selection. In contrast, when dietary wariness is added to the model the plot of number of morphs maintained, as a function of the potential number of cryptic forms available, is minimized at an intermediate number of cryptic forms, i.e. is V-shaped. These counter-intuitive patterns are robust to varying strengths of apostatic selection and different implementations of dietary wariness, and are more pronounced when predator and prey generation lengths are similar.  相似文献   

14.
G. M. Livshits 《Oecologia》1981,51(2):220-226
Summary Microspatial variation of banded and unbanded shell morphs frequencies as well as number of individuals per m2, mortality, migration and burrowing into the ground were examined in a population of snail Brephulopsis bidens found in South Crimea mountains (USSR). Differential values of relative survival of morphs were determined by their thermotolerance. The relative survival of the banded morph was lower at the west sites of population area (W=0.273), and increased gradually up to 1 at the east sites. Survival of the banded morph was dependent on its burrowing activity. Differences in relative survival of morphs decreased parallel with increasing general mortality of snails.Burrowing activity and intensity of migration of the banded morph were significantly higher than that in unbanded. In experiments with artificial shaded sections, the banded morph preferred shaded sections, whereas unbanded chose illuminated sites. All these differences in behaviour probably form the main factors for microspatial variation of morph frequencies.  相似文献   

15.
In Batesian mimicry, a harmless species (the ‘mimic’) resembles a dangerous species (the ‘model’) and is thus protected from predators. It is often assumed that the mimetic phenotype evolves from a cryptic phenotype, but it is unclear how a population can transition through intermediate phenotypes; such intermediates may receive neither the benefits of crypsis nor mimicry. Here, we ask if selection against intermediates weakens with increasing model abundance. We also ask if mimicry has evolved from cryptic phenotypes in a mimetic clade. We first present an ancestral character-state reconstruction showing that mimicry of a coral snake (Micrurus fulvius) by the scarlet kingsnake (Lampropeltis elapsoides) evolved from a cryptic phenotype. We then evaluate predation rates on intermediate phenotypes relative to cryptic and mimetic phenotypes under conditions of both high- and low-model abundances. Our results indicate that where coral snakes are rare, intermediate phenotypes are attacked more often than cryptic and mimetic phenotypes, indicating the presence of an adaptive valley. However, where coral snakes are abundant, intermediate phenotypes are not attacked more frequently, resulting in an adaptive landscape without a valley. Thus, high-model abundance may facilitate the evolution of Batesian mimicry.  相似文献   

16.
Anti-apostatic selection occurs when predators feed disproportionately on rare prey, a process that leads to a decrease in variation within prey populations. Most of the evidence for this phenomenon comes from experiments using artificial prey items distinguished by simple artificial colour differences. We describe an experiment performed at 10 different sites in which we tested whether selection by wild birds is anti-apostatic when presented with high density populations of shells of the polymorphic land snail, Cepaea hortensis , stuffed with pastry. At each site we presented two sorts of populations in sequence: 45 yellow unbanded with 5 yellow banded, and 5 yellow unbanded with 45 yellow banded. Selection was measured using an analysis based on generalised linear mixed models (GLMMs) that has broad applicability to all resource selection studies where extraneous variance is a problem. Using this analytic method, we found that birds altered their behaviour to produce significant anti-apostatic selection in direct response to changes in frequency of the quasi-natural Cepaea morphs.  相似文献   

17.
Galician exposed shore populations of the direct developing periwinkle Littorina saxatilis are strikingly polymorphic, with an ornamented and banded upper shore form and a smooth and unbanded lower shore form. Intermediates between the two pure forms occur in a narrow mid shore zone together with the parental forms. We have previously shown that the two pure forms share the same gene pool but that mating between them is non-random. This is due to a non-random microdistribution in the zone of overlap, and also to assortative mating. In this study we present data which show that intermediate (hybrid) females mate less often than pure females in micropatches dominated by either of the pure forms, but not in micropatches in which the two pure forms are equally common. Thus, sexual fitness in intermediate females depends on the frequency of both pure morphs. Furthermore, sexual selection against intermediate females also varies with the densities of snails within each micro patch. The biological mechanisms which may explain this particular reduction of female hybrid fitness are discussed. Assortative mating between the pure morphs is sometimes almost complete, while both morphs do not mate the intermediates assortatively. In the light of this, sexual selection against intermediate females may contribute considerably to restrict gene flow between the pure forms.  相似文献   

18.
Aposematic signals represent one of the most accessible traits to evaluate the interaction of natural and sexual selection on signal evolution. Here we investigate the contributions of these two selective forces on the aposematic signal evolution of the highly polytypic strawberry poison frog, Oophaga pumilio, of Bocas del Toro, Panama. Previous research has shown that the brightness of O. pumilio warning coloration can inform predators of the toxicity levels associated with different populations of the archipelago. Other studies suggest that sexual selection may be influencing warning signal brightness within populations via female mate choice (Isla Solarte, Isla Bastimentos, and Aquacate Peninsula populations) and male–male competition (Isla Solarte). Here we present two non-exclusive scenarios for how natural and sexual selection interact to drive phenotypic variation across this archipelago: (1) predators impose a selective regime whereby populations above a toxicity-brightness threshold are at liberty to diversify via sexual selection and below which populations are constrained to maintain a stricter resemblance to a more cryptic population mean, and (2) synergistic/additive effects of inter- and intrasexual selection drive the evolution of brighter males within populations above this toxicity threshold. We investigate whether aposematic patterns of divergence across the archipelago relative to the common mainland phenotype meet these predictions using existing data on O. pumilio morph toxicity measures and overall conspicuousness estimates to an avian predator. Using standardized z-scores to evaluate the range of trait values, we find that indeed the population representative of the common mainland phenotype (Almirante) represents an intermediate level of both toxicity and conspicuousness, and that derived Bocas del Toro populations vary in each of those components in directions predicted by the proposed scenarios. Furthermore, we find greater divergence towards conspicuousness than crypsis, a pattern suggestive of sexual and natural selection acting synergistically in morphs with high toxicity.  相似文献   

19.
Population divergence in sexual signals may lead to speciation through prezygotic isolation. Sexual signals can change solely due to variation in the level of natural selection acting against conspicuousness. However, directional mate choice (i.e., favoring conspicuousness) across different environments may lead to gene flow between populations, thereby delaying or even preventing the evolution of reproductive barriers and speciation. In this study, we test whether natural selection through predation upon mate‐choosing females can favor corresponding changes in mate preferences. Our study system, Oophaga pumilio, is an extremely color polymorphic neotropical frog with two distinctive antipredator strategies: aposematism and crypsis. The conspicuous coloration and calling behavior of aposematic males may attract both cryptic and aposematic females, but predation may select against cryptic females choosing aposematic males. We used an experimental approach where domestic fowl were encouraged to find digitized images of cryptic frogs at different distances from aposematic partners. We found that the estimated survival time of a cryptic frog was reduced when associating with an aposematic partner. Hence, predation may act as a direct selective force on female choice, favoring evolution of color assortative mating that, in turn, may strengthen the divergence in coloration that natural selection has generated.  相似文献   

20.
Variations at both the genetic and phenotypic levels play animportant role in responses to food and food-related stimuli.Knowledge of such variations is crucial to understanding howpopulations adapt to changing environments. We investigatedthe dietary preferences of 2 tiger snake populations and comparedthe responses of diet-naive animals (laboratory-born neonates),diet-controlled animals (laboratory-reared juveniles), and naturaldiet–experienced animals (wild-caught adults) to visualand chemical cues from 6 prey types (mouse, skink, silver gull,chicken, shearwater, and frog). The mainland population inhabitsa swamp, feeds mostly on frogs, and suffers heavy predation.The second population inhabits a small nearby offshore islandwith no standing water (no frogs); feeds mostly on skinks, mice,and, as adults, silver gull chicks; and suffers no known predation.Although different prey are eaten in the 2 populations, adultwild-caught snakes from both populations showed a significantpreference for 3 types of prey (frog, mouse, and chick), irrespectiveof their natural diet. Neonates responded to all prey cues morethan they did to control stimuli in both populations. However,the island neonates showed significantly higher interest insilver gull chick stimuli (the main prey of the island adultsnakes) than did their mainland conspecifics. Laboratory-bredjuveniles displayed behavioral plasticity by significantly increasingtheir response to mice after being fed baby mice for 7 months.We conclude that genetic-based differences in food-related cuesare important in tiger snakes but that they are also capableof behavioral plasticity. Island adult and neonate snakes exhibitedresponses to prey types no longer consumed naturally (frog),suggesting that behavioral characters may have been retainedfor long periods under relaxed selection. Island neonates showeda strong interest in a novel prey item (silver gull). This resultcomplements previous work describing how island snakes havedeveloped the ability to swallow larger prey than usual, aswell as seemingly developing a taste for them.  相似文献   

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