Gene frequency changes in Cepaea snails on the Marlborough Downs over 25 years

In 1985 we resurveyed the sites on the Marlborough Downs in southern England at which Cain and Currey in 1960/61 sampled Cepaea snails and thence introduced the term 'area effects' to describe large areas of uniform morph frequency. Some sites no longer harboured Cepaea and at others the species composition had changed, with a general spread of Cepaea hortensis at the expense of Cepaea nemoralis. The majority, however, permitted comparison of morph frequencies between the two surveys. In C. nemoralis, we detected a significant overall decrease in the frequency of the brown morph and estimate selection as 5–9% per generation. There was no apparent change in frequencies of banded morphs. In C. hortensis we detected a significant overall increase in the frequency of unbanded shells (1–3% selection per generation) and an almost significant decrease in the frequency of fusions within the banded class. There was insufficient colour polymorphism in C. hortensis to allow analysis of colour morph frequencies. These changes—all in the direction of reduced absorption of solar energy—resemble others detected in both species at other localities in southern England. Possible explanations include large-scale climatic effects and changes in vegetation.     

Anti-apostatic selection by wild birds on quasi-natural morphs of the land snail Cepaea hortensis: a generalised linear mixed models approach

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.     

Distribution patterns of morph frequencies in the snail Cepaea hortensis in Iceland

In Iceland Cepaea hortensis Müll. reaches its northern limit of distribution occurring in restricted areas along the south coast. Samples of snails collected along steep cliffs in two such restricted areas were effectively monomorphic for shell colour (yellow), but were polymorphic for banding pattern and revealed strong area effects. In both localities samples from sites facing east contained varying frequencies of unbanded snails whereas in ecologically similar sites facing south unbanded were absent. In an experimental population released at a south-facing site unbanded snails failed to become established whereas banded individuals survived and bred. It is suggested that the area effects are maintained through non-visual (most likely climatic) selection though other possibilities cannot be ruled out.     

The distribution and variation of three species of land snail near Rickmansworth, Hertfordshire

The distribution of Arianta arbustorum, Cepaea nemoralis and C. hortensis , and the variations in the shell polymorphism of the Cepaea species have been studied in an area of low chalk downland and alluvial river valleys near Rickmansworth, Hertfordshire. The distribution of A. arbustorum is restricted, the species being largely confined to the bottoms of river valleys, especially in hedgerows. Variations in dampness seem the most likely limiting factor. C. hortensis is widespread and found in many habitats, but C. nemoralis is largely confined to woods, and is especially common in beechwoods. The morph-frequencies of C. nemoralis and C. hortensis vary in such a way as to suggest the operation of climatic and visual selection. In C. hortensis there are area effects for the presence or absence of pink five-banded shells, and there is an overall linkage disequilibrium resulting in an excess of unbandeds in non-yellow shells. Reasonable evidence for the operation of random factors in determining the morph-frequencies of populations of Cepaea in the area has been found only for three very disturbed populations. It is suggested that the large number of selective forces known to act on Cepaea populations in this area will account for the variation in morph-frequencies in populations from the same area or habitat.     

Lake Erie water snakes revisited: Morph- and age-specific variation in relative crypsis

Summary Populations of water snakes (Nerodia sipedon insularum) on islands in western Lake Erie are variable in colour pattern, consisting of unbanded, intermediate, and banded morphs. In contrast, mainland populations (N. s. sipedon) consist solely of banded morphs. Previous investigators hypothesized that natural selection favoured unbanded morphs on exposed island shorelines and banded morphs in overgrown mainland habitats and that gene flow from mainland populations was responsible for the persistence of banded morphs on islands. To clarify the potential role of natural selection, I quantified relative crypsis among morphs and age classes of water snakes by comparing the size of patches making up their colour patterns with the size of patches in island and mainland backgrounds. This analysis reveals that if unbanded morphs are more cryptic than intermediate and banded morphs on islands, it is only in the young-of-the-year age class. For older snakes on islands and for all snakes on the mainland, unbanded morphs are consistently less cryptic than intermediate and banded morphs. Given these results, the net direction of selection in island populations should depend on the intensity of predation on different age classes of snakes. Overall, selection may favour unbanded morphs (e.g. if predation occurs primarily on young-of-the-year), intermediate and banded morphs (e.g. if predation occurs primarily on older snakes), or be weak or absent (e.g. certain combinations of predation on young-of-the-year and older snakes). Using estimates of relative crypsis to guide reanalysis of morph frequency data, I find support for the hypothesis that unbanded morphs are favoured by natural selection in island populations.     

Opportunity for natural selection and gene flow in an isolated Zapotec-speaking community in southern Mexico in the throes of a secular increase in size

Our object in this paper is to analyze the opportunity for natural selection and gene flow in an isolated Zapotec-speaking community in the valley of Oaxaca, southern Mexico, that is undergoing a secular increase in body size. Surveys were conducted in the community in 1968, 1978, and 2000, including anthropometric and census data. No secular change was found in the growth status of schoolchildren and adult height between 1968 and 1978; subsequently, major secular gains in height occurred among children and adolescents between 1978 and 2000. The 1978 household data were used to compute gene flow (3.3%) and opportunity for selection intensity (I = 1.312). Migration and other demographic information was obtained from household census data for 1978 and 2000, and mortality information was extracted from community records and archives. These data were used to compute gene flow and opportunity for natural selection. Gene flow increased from 3.3% to 4.7% and intensity of natural selection decreased from 1.312 to 0.272 from 1978 to 2000. Variance in fertility increased slightly over time (12.25 to 13.69). Opportunity for selection was dominant during the prereproductive period in 1978, but approached 0 for the mortality component in 2000, resulting in a marked decrease in the mortality component (Im) of selection (0.626 and 0.019, respectively) and total opportunity for selection (I = 1.312 and 0.272, respectively). Secular increase in height and markedly decreased opportunity for natural selection (1) were associated with better health and nutritional conditions. Genotype-environment interaction and environmental influences are apparently the predominant causes of the secular trend. If natural selection plays a role in causing the secular trend, it is a small one.     

Effects of experimental stress on the glycogen content of Cepaea hortensis (Mollusca)

A method for measuring the glycogen content of the hepatopancreas of Cepaea hortensis is described. Glycogen content was determined for snails from field population cages and from a nearby natural population. Results showed that caged snails possessed lower levels of glycogen than did the natural population, while the lowest levels of glycogen were associated with higher snail densities in the cages. There is evidence for an increase in glycogen content of C. hortensis during the late summer in nature and in experimental populations. The results are discussed with regard to the selective effect that glycogen may have on Cepaea and to the interpretation of events occurring in populations of Cepaea that glycogen assay could provide.     

Environmental effects on adult shell size in Cepaea hortensis

In natural populations of the land snail Cepaea hortensis there is often a negative association between shell size and population density. The usual interpretation of such associations, both in C. hortensis and other species, is that they represent an ecophenotypic effect that is part of a system of density-dependent natality, rather than a genetically-based effect that is produced through density-related selection. However, this interpretation is usually based on supposition rather than evidence. Here we present the results of an experiment in which small juveniles collected from two sampling sites within a colony known to exhibit a negative size-density association were grown to maturity under constant laboratory conditions. The highly significant inter-site difference observed in the wild disappeared in the laboratory-reared snails, indicating the direct influence of environmental factors on size. Because of the nature of the data-sets from which density-dependent natality is inferred, this result has a clearer connection with such density dependence than conventional within-population heritability estimates.     

Trill consistency is an age-related assessment signal in banded wrens

Older males tend to have a competitive advantage over younger males in sexual selection. Therefore, it is expected that signals used in sexual selection change with age. Although song repertoire size in songbirds is often mentioned as an age-related trait, many species, including the banded wren (Thryothorus pleurostictus), do not increase their repertoires after the first year. Here, we show that banded wrens reproduce the trill notes in their songs with less variability between them (i.e. more consistently) when they grow older. In a playback experiment, we also show that banded wrens discriminate between younger and older birds based on structural aspects of their song. In a second experiment, banded wrens also respond differentially to natural songs versus songs with artificially enhanced consistency. We argue that consistency in trill note reproduction may be achieved through practice. Sexual selection in the form of male–male competition may therefore operate on a phenotypic trait, the expression of which is enhanced by practice.     

Analysis of polymorphic variation in different types of habitat

A correlation between the distribution of an organism and features of its environment can be taken as indirect evidence of natural selection. Biologists may therefore collect samples from polymorphic populations at a number of locations, classify the locations into habitat types, and consider whether the distribution of morphs varies with the habitat. Statistical aspects of this type of study are discussed in this paper. A randomization test for habitat effects is proposed and a negative binomial model is suggested for the distribution of morphs from random locations within one type of habitat. Data on the distribution of Cepaea hortensis and C. nemoralis snails in southern England provide an example. For both species there is clear evidence of differences between habitats, although the morph distributions are rather variable within habitats. The negative binomial model suggests that, for the snail data, variation in morph proportions is mainly due to location differences. The binomial sampling error is relatively unimportant unless the sample size at a location is very small. Therefore it is reasonable to analyse morph proportions by standard methods without giving different weights to data from different locations. The snail data are analysed in this way. Discriminant function analyses are used to test for habitat effects. The relationships between C. hortensis and C. nemoralis morph frequencies within one habitat are examined by a canonical correlation analysis.     

Selection on laying date is connected to breeding density in the pied flycatcher

Timing of reproduction and clutch size are important determinants of breeding success, especially in seasonal environments. Several recent bird population studies have shown changes in breeding time and in natural selection on it. These changes have often been linked with climate change, but few studies have investigated how the traits or natural selection are actually connected with climatic factors. Furthermore, the effect of population density on selection has been rarely considered, despite the potential importance of density in demographic processes. We studied variation in natural selection on laying date and on clutch size in relation to measures of spring phenology and population density in a long-term study of pied flycatchers in SW Finland. The phenological stage of the environment at mean egg-laying did not affect the direction of selection on either laying date or on clutch size. There was, however, stronger selection for earlier laying date when the breeding density of the population was high, suggesting that early breeding is not necessarily beneficial as such, but that its importance is emphasized when high population density increases competition. In addition, early breeding was favoured when the pre-breeding period was cool, which may indicate an increased advantage for the fittest individuals in harsher conditions. In the middle of the twentieth century, there was selection for large clutch size, which subsequently ceased, along with an overall decrease in recruit production. Our results indicate that attention should be paid to demographic factors such as breeding density when studying natural selection and temporal changes in it.     

DIFFERENTIAL AVOIDANCE OF CORAL SNAKE BANDED PATTERNS BY FREE-RANGING AVIAN PREDATORS IN COSTA RICA

Empirical studies of mimicry have rarely been conducted under natural conditions. Field investigations of some lepidopteran systems have provided a bridge between experiments examining artificial situations and the mimicry process in nature, but these systems do not include all types of mimicry. The presence of dangerous or deadly models is thought to alter the usual rules for mimicry complexes. In particular, a deadly model is expected to protect a wide variety of mimics. Avoidance of different types of mimics should vary according to how closely they resemble the model. Coral snake mimicry complexes in the neotropics may provide natural systems in which these ideas can be examined, but there is no direct evidence that the patterns of venomous coral snakes or potential mimics are avoided in the wild. Plasticine replicas of snakes were used to assess the frequency of avian predation attempts as a function of color pattern. Avian predators left identifiable marks on the replicas, the position of which indicated that replicas were perceived as potentially dangerous prey items by birds. The number of attacks on unmarked brown replicas was greater than that on tricolor coral snake banded replicas. This result was true whether replicas were placed on natural or plain white backgrounds, suggesting that coral snake banded patterns function aposematically. In a separate experiment, replicas representing all six patterns of proposed coral mimics at the study site were attacked less often than unmarked brown replicas. Within these six banded patterns, some were attacked significantly more often than others. This study provides direct field evidence that coral snake banded patterns are avoided by free-ranging avian predators and supports theoretical predictions about mimicry systems involving deadly models.     

COLOR PATTERN POLYMORPHISM IN THE LAKE ERIE WATER SNAKE,NERODIA SIPEDON INSULARUM

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.     

Cepaea on Salisbury Plain: patterns of variation, landscape history and habitat stability

Patterns of variation in Cepaea nemoralis and C. hortensis on Salisbury Plain are related to its landscape history. Areas of arable or mixed farming, with hedgerows and woods, show patterns of variation with habitat and topography. Areas of downland, until recently heavily grazed by sheep, show area effects. Cepaea populations in these downland areas would have been much rarer, and more isolated when grazing was intense. This pattern is repeated in other parts of southern England, showing that area effects in Cepaea are strongly associated with areas of habitat instability, where Cepaea survived as small and isolated populations until recently. This evidence enhances the plausibility of explanations of area effects involving isolation, founder effect or local selection, and co-adaptation.     

Spouse selection in Canada, 1921-78: an examination by age, sex and religion

This paper presents time series data on spouse selection in Canada for the period 1921-1978. Homogamous and endogamous trends are examined by age, sex, and religion. In the case of religious endogamy, most groups have shown decreasing proportions of marriages. There are also considerable differences in rates of endogamy among religious groups, and these differences have largely persisted over time. Until 1971, the proportions of brides marrying older and younger grooms decreased, while the proportion marrying the same age increased, after which these trends reversed. Younger males increasingly tended to select brides of their own age group until the 1960s, while older males have been increasingly heterogamous since World War II. While these changes in spouse selection have been examined with respect to changes in the unmarried population, the relation between the 2 trends is not clear, and it is difficult to account fully for the observed trends in spouse selection with respect to age.     

Difference in habitat selection by different morphs of the land snail Arianta arbustorum

The behaviour of morphs of Arianta arbustorum has been investigated using a new technique involving a paint which fades in daylight. The technique was used to study the response of different morphs to exposure to sunlight at a homogeneous woodland habitat and a heterogeneous open habitat. The yellow morph showed a significantly higher rate of fading than the brown morph. A mark, release and recapture programme was used to analyse the association between frequencies of different morphs recaptured and not recaptured. In both habitats, there was a general tendency for the frequency of recapture to increase with increasing age in both morphs. Habitat selection resulting from different behaviour of different morphs of A. arbustorum is discussed. Morph frequencies can be related to habitat, the brown morphs of A. arbustorum are significantly less frequent in the open habitat. The behavioural responses of Cepaea hortensis , present with A. arbustorum in two other populations, to the exposure of sunshine were also studied. Climatic selection by solar energy associated with different morphs, species and habitat are also discussed.     

Disequilibrium in some Cepaea populations

Phenotypic disequilibrium is a measure available, along with morph frequency, in many published sets of sample data recording the polymorphism of shell colour and pattern in the snail Cepaea nemoralis. The relation of disequilibrium to morph frequency for the colour (pink/yellow) and banding (unbanded/banded) loci has been examined for a large and widespread set of data. The direction of disequilibrium is a function of frequency at the two loci in a way that suggests that selection favours combinations of common morphs, whichever they are. This could indicate that such combinations are common because they have been selected. The data are therefore consistent with the proposal that populations are generally subject to selection of varying directions at different places and times, acting on the phenotype. In combination with migration, such selection could lead to prolonged polymorphism.     

Apostatic selection by song thrushes (Turdus philomelos) feeding on the snail Cepaea hortensis

There is no direct evidence that predators exert apostatic selection (the systematic overpredation of commoner forms) on live, naturally polymorphic prey. This study tested whether captive song thrushes ( Turdus philomelos ) select apostatically when presented with dimorphic populations of yellow five-banded and yellow unhanded morphs of the snail Cepaea hortensis. Four thrushes were used. Two were presented with a 9: 1 ratio of five-bandeds to unbandeds and two were presented with 1:9 ratios. Each thrush was given four trials in succession. In each trial 30 snails were presented and the trial was stopped when 15 had been eaten. There were no differences in shell size between morphs or between eaten and uneaten snails of each morph. Three thrushes selected apostatically and one thrush exerted virtually no selection. Overall, there was statistically significant apostatic selection.     

Interisland gene flow among populations of the buff‐banded rail (Aves: Rallidae) and its implications for insular endemism in Oceania

Studies of fragmented habitat, such as island archipelagos, provide insights into the microevolutionary processes that drive early stages of diversification. Here, we examined genetic variation and gene flow among populations of the widespread buff‐banded rail Gallirallus philippensis in Oceania to understand the factors that promote speciation associated within this bird lineage. We analysed mtDNA Control Region sequences and six microsatellite loci from a total of 152 individuals of buff‐banded rail on islands and continental areas. We used a phylogeographic model‐testing approach and a structured spatial design ranging from within to among archipelagoes in the south Pacific. Buff‐banded rail populations in the Philippines archipelago and nearby Palau and Wallacea had high genetic diversity while those in geographically distant Australia showed lower variation. Other archipelagos sampled were found to have less genetic diversity and included haplotypes closely related to Wallacea (Bismarck, Vanuatu, New Caledonia) or Australia (New Zealand, Samoa, Fiji, Cocos Islands). Nucleotide diversity and allele frequency declined with degree of geographic isolation but haplotype diversity remained more even. However, both nucleotide and haplotype diversities were positively correlated with land area. Microsatellite data for a subset of locations showed moderate to high genetic differentiation and significant pairwise F_ST despite a relatively high migration rate. Our results are mostly consistent with a model of abrupt genetic changes due to founder events with multiple dispersals into Australia from Wallacea and Bismarck. Australia has probably been the source of birds for islands in the Pacific. This is shown by decreasing genetic diversity and growing genetic differentiation when distances separating populations increased from Australia. A history of range expansion and divergent natural selection may help explain the existence of numerous sympatric Gallirallus island endemics.