Variation in populations of Cepaea nemoralis (L.) in North Spain

In a survey of 105 populations of Cepaea nemoralis sampled in North Spain, the distribution of morph polymorphism seems to have a climatic component. The area sampled corresponds with the two main climatic regions in the Iberian Peninsula: Green and Brown Iberia, which have an Atlantic and Continental climate, respectively. Paler colour and banding phenotypes are found in the Atlantic zone, while the dark ones are commoner in Brown Iberia. This may arise from climatic selection related to behaviour patterns, which may be particularly important in limiting situations with harsh climates.     

The effect of artificial photoperiod on growth and reproduction in the land snail Cepaea nemoralis

Summary

Specimens of Cepaea nemoralis were raised from egg to adult in the laboratory for 15.5 months in two artificial photoperiods: short-day (LD 8:16) and long-day (LD 16:8). Over 20% of the snails in each photoperiod were functional adults by the end of the experiment and had fully developed male and female reproductive systems as determined both by dissection and by oviposition. A total of 925 eggs were laid by animals reared in a long-day photoperiod, whereas animals in a short-day photoperiod laid 677 eggs. Animals reared in a long-day photoperiod grew slightly faster (2.04 mg/day compared to 1.73 mg/day), began laying eggs about 18 days sooner, and suffered higher mortality (19% compared to 11%) than animals reared in a short-day photoperiod. These results clearly establish for C. nemoralis that gametogenesis and the development of a functional hermaphroditic reproductive system are neither prevented nor significantly retarded by exposing the animals to a short-day photoperiod for their entire pre-reproductive life.     

Food overlap in co-existing populations of the land snails Cepaea nemoralis (L.) and Cepaeahortensis (Müll)

Faecal analysis of adult Cepaea nemoralis and Cepaea hortensis from a mixed population on chalk grassland shows that the two snail species select the same plant material as food. Herbs are selected in preference to grasses and Urtica dioica is particularly favoured. C. hortensis has the more pronounced preference for senescent material. These results are discussed in relation to competition between the two species.     

Patterns of morphological and molecular polymorphism in the land snail Cepaea nemoralis

In the land snail Cepaea nemoralis , allele frequencies at loci controlling shell polymorphisms often show large areas of remarkable constancy which are separated by steep clines from neighbouring areas with strikingly different allele frequencies. It has recently been claimed that these 'area effects' exemplify a general tendency for population differentiation without geographic isolation in a variety of organisms of relatively low mobility. As such they could represent an early phase in the process of speciation. If this is true, population differentiation of shell polymorphisms in Cepaea would be expected to be accompanied by parallel differentiation at other gene loci, such as those detected by gel electrophoresis.
We have studied populations of C. nemoralis in North Wales and in the Valle de Aran of the Pyrenees. Levels of molecular heterogeneity are comparable to those found in related animals which show much less visible polymorphism. In spite of some statistical problems inherent in the analysis ol overlapping geographic patterns, there is no clear association between the patterns of geographic variation at the visible and molecular levels. Claims that genetic differentiation in the visible polymorphisms between C. nemoralis populations are a special case of the formation of geographic races are therefore probably not justified.     

The habitats of the land snail Cepaea nemoralis (L.) on downland and their ecogenetie significance

Population densities of adult Cepaea nemoralis (L.) were estimated for nine chalk grassland sites on the South Downs. Estimates of cover, species composition and height of vegetation were also obtained. The density of C. nemoralis was positively associated with Bromus erectus cover, and negatively with Festuca spp. cover. This finding, confirmed by an earlier survey, suggests that the density of C. nemoralis in chalk grassland is directly or indirectly governed by mammalian grazing pressure, and not by the availability of suitable food. Intensely grazed downland is an unsuitable, or at best a marginal habitat for C. nemoralis. This conclusion is discussed with particular reference to 'area effects' in the shell character polymorphism, often found in downland populations of C. nemoralis.     

Variation in body colour in western Irish populations of Cepaea nemoralis (L.)

Samples of Cepaea nemoralis (L.) taken from sand-dune, lowland and mountain populations show differences in body colour. Overall, sand-dune populations are much paler than those in the other two areas. On sand-dunes, darker and more variable populations are found in densely vegetated habitats; in the lowland, darker forms occur further inland but there is no correlation with vegetation; mountain populations are darker and less variable on north- than south-facing slopes. Overall differences between areas can be ascribed to climatic selection and differences in variation within areas to niche width.     

The karyotype of Cepaea sylvatica (Pulmonata: Helicidae) and its relationship to those of C. hortensis and C. nemoralis

The karyotypes of Cepaea nemoralis (L.) and C. hortensis (Müller), with 2n=44 and a conspicuously large pair of chromosomes, are described and compared with that of C. sylvatica (Draparnaud) which has 2n=50. The karyotype of C. sylvatica also has a conspicuously large pair of chromosomes but the comparison suggests that these have an independent origin from those in the 2n = 44 species. There is no evidence that the large chromosomes in C. nemoralis and C. hortensis have originated from simple fusion of chromosomes from a 2n=50 karyotype with chromosomes all sub-equal such as is reported for C. vindobonensis. It may be that such a karyotype with little size differentiation amongst the chromosomes is not a primitive feature in the Helicinae. The relationship of shell colour and banding polymorphism to the chromosome architecture is discussed.     

Shell polymorphism in a southern peripheral population of Cepaea nemoralis (L.) (Pulmonata: Helicidae) in Spain

Populations of the land snail Cepaea nemoralis have been investigated at the southern limits of its range in Spain, the Upper Tajo region, in order to analyse the distribution of the species and the variation of polymorphism in relation to environmental conditions. For this purpose two natural subareas, which differ in topography, soil nature, climate and vegetation have been considered. Cepaea nemoralis is confined to the river banks of the area; it is found in open and shaded biotopes of subarea 1 where it is relatively common, and it is almost restricted to shaded biotopes of subarea 2 where it becomes progressively rarer towards the south. Relative abundance of the species may be related to the amount of calcareous soils and climate, the restriction in its southern distribution being probably the result of extreme solar radiation, high temperature and low humidity. The two subareas differ in the frequency of the main polymorphic characters. These differences may be related to the kind of biotope. Subarea 1 richer in more open and exposed biotopes have higher frequencies of yellow, 00000, 00300, and unfused bands, whereas the restriction to poplar groves because of the lack of suitable exposed biotopes in subarea 2 may account for the higher proportion of pink five-bandeds with the bands fused together. This kind of variation with biotope is consistent with an explanation in terms of differences in the temperature absorbance of different morphs and a metabolic response to differences in temperature amplitude of the biotopes. Other characters (lip colour, punctate bands, hyalozonate bands, etc.) are not polymorphic in this region, thus, suggesting that at the periphery environmental conditions are important not only in determining the distribution of the species but exerting a marked effect on the genetic structure of the populations.     

Opioid systems,behavioral thermoregulation and shell polymorphism in the land snail,Cepaea nemoralis

Summary Numerous ecological studies have dealt with the shell color and banding polymorphism of the land snail Cepaea nemoralis. The present field and laboratory investigations focus on the roles of opioid systems in modulating the thermal preferences and behavioral thermoregulation of various morphological types of Cepaea. Evidence is presented that differences in opioid modulation of the thermal responses of Cepaea are associated with shell polymorphism. It is shown that the effects of the prototypic opiate agonist, morphine, and antagonist, naloxone, on behavioral thermoregulation in Cepaea vary with the shell banding pattern and thermal microhabitat. In both the field and laboratory, morphine (0.10, 1.0 and 10 g per snail) caused significant dose- and time-dependent increases in the temperatures selected by various morphological types of Cepaea. The palest shell type (yellow, unbanded) with the highest basal preferred temperature displayed the greatest response to morphine, the shell type (yellow, 2-banded) with an intermediate basal preferred temperature showed an intermediate response to morphine, and the darkest shell type (yellow, 5-banded) with lowest basal preferred temperature showed the least increase in preferred temperature after administration of morphine. These effects of morphine were blocked and reversed by naloxone (1.0 g), with the opiate antagonist by itself (1.0 and 10 g) causing a significant decrease in behaviorally selected temperatures. The unbanded and 2-banded morphs displayed significantly greater decreases in preferred temperatures after treatment with naloxone than did the 5-banded morph, which showed minimal responses. It is suggested that these differences in opioid modulation of thermal preferences and behavioral thermoregulation may contribute to the polymorphic thermal preferences of natural populations of Cepaea.Abbreviations B yellow five-banded shell type - I yellow two-banded shell type - U yellow unbanded shell type     

Selection and disequilibrium in Cepaea nemoralis

The snail Cepaea nemoralis (L.) is usually polymorphic for linked loci conferring pink or yellow shell ground colour and banding or lack of bands. An unlinked locus, mid‐banded, modifies the appearance of banded shells. These characters have a very wide range of frequencies across the species range and even within limited areas. In Britain, frequencies differ between populations from shaded woodland habitats and those from closely adjacent open habitats. Using data from the Evolution MegaLab Project, it is shown that the greater frequency of pink unbanded in shaded habitats is associated with a greater excess of the favoured allele combinations, measured as linkage disequilibrium. Examination of the relationship of disequilibrium to allele frequency in samples from the full geographical range and all habitats and climates shows that, in general, the allele combinations at high frequencies are in gametic excess. This suggests that selection tends to diversify rather than move frequencies towards stable equilibria. The result is important in relation to the as yet unresolved question how the polymorphism is maintained. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Temporal changes of gene frequencies in Cepaea hortensis

There have been few studies analysing long-term changes of gene frequencies in natural populations. This work is the first report of such changes in the land snail Cepaea hortensis (Mull).
Collections of C. hortensis were made at Silbury Hill, Wiltshire in 1957, 1963 and 1978. The banded phenotype significantly declined in frequency between 1963 and 1978. It is argued that the decline, which was statistically homogeneous at 15 separate sites within the area, was a consequence of natural selection. The average selection coefficient against the banded homozygote is estimated to have been greater than 1096.     

Persistence and extinction in some Cepaea populations

Eight artificial populations of Cepaea nemoralis kept in enclosures excluding visual predators, built up to peak numbers and then crashed in periods of between 10 and 20 years. Fluctuations in numbers were caused by variation in recruitment, not in adult survival. One population survived through a very low population size (14 individuals or less), and shows no significant change in morph frequencies. The very slight fluctuation in morph frequency has a probabilty of chance occurrence of 7%. Studies of 250 years or more are probably needed to distinguish systematic effects on gene frequency from random variation. The lack of any major alteration in morph frequencies, however, suggests that there has been no genetic revolution in this population.     

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.     

Variation in the distribution and fitness of the brown morph of Cepaea nemoralis (L.)

The distribution of the brown rnorph of the landsnail Cepaea nemoralis in part of the Brendon Hills, Somerset is described. The morph is at a high frequency on north-facing slopes, valley bottoms, and in a broad partly-enclosed basin. It is argued that low temperature is the selective agent. Alternative explanations are considered. In two population cages adult brown-shelled snails survived less well than pink five-banded ones during the summer months, this trend being reversed in the autumn. It is concluded that the experimental results support the 'climatic' interpretation of distributional data.     

Genetic variation in two land snails, Cepaea nemoralis and Succinea putris (Gastropoda, Pulmonata), from sites differing in heavy metal content

Allozyme variation was determined in two land snail species (Cepaea nemoralis and Succinea putris) from four localities in northern Belgium. In each locality we selected a polluted and a nearby, less-polluted, reference plot. We examined whether (i) genetic variability differed between the polluted and reference plots, (ii) populations from polluted plots experienced recent bottlenecks, and (iii) certain allele or genotype frequencies were associated with the pollution. Our results suggest that (i) about 13% of the genetic differentiation in C. nemoralis and 5% in S. putris was due to differences among polluted and reference plots, (ii) polluted and reference plots had comparable levels of genetic variation, but in C. nemoralis observed heterozygosities were higher in polluted plots, (iii) most plots showed significant evidence for recent bottlenecks, irrespective of the degree of pollution, so that bottlenecks seem poor indicators of pollution-induced stress in land snails, and (iv) mutagenic or pollution-induced modifications did not seem to account for new allozyme variants in polluted sites. The observed patterns of genetic variation may be explained by the action of genetic drift, pollution-mediated selection, restricted gene flow, or a combination of these processes.