The survival, weight-loss and behaviour of three species of land snail in conditions of low humidity

Cepaea nemoralis L., C. hortensis Mull, and Arianta arbustorum L. show differences in survival, rate of water-loss and behaviour under conditions of low humidity in laboratory studies. C. nemoralis survives better, ceases activity more rapidly, and loses proportionally less weight in low humidity than C. hortensis , which in turn shows the same properties with respect to A. arbustorum . The differences between these three species are reflected in differences in distribution in parts of Britain.     

The effect of temperature on the activity of three species of Helicid snail (Mollusca: Gastropoda)

The activity of the land snails Cepaea nemoralis (L.), C. hortensis (Müll.) and Arianta arbustorum (L.) was measured over 24 hours at the constant temperatures of 0, 3, 8, 17, 22 and 26°C at 100% relative humidity and with 16 hours light and 8 hours dark. The activity of all three species varies significantly with temperature and light, and there are differences in the activities of the species at each temperature. A. arbustorum is relatively more active than Cepaea at low rather than high temperatures, as is C. hortensis with respect to C. nemoralis , though to a lesser extent. There is little diurnal rhythm in activity at 0°C, but all species become more nocturnal as temperature increases; C. nemoralis is most, and A. arbustorum least nocturnal. The results obtained assist in explaining the observed geographical and local distributions of the species concerned.     

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.     

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.     

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.     

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.     

Variation with altitude and habitat in Cepaea hortensis (Müll.)

Variation in morph, chromosome and gene frequencies in Cepaea hortensis are described for a valley system in the north slopes of the Brendon Hills, Somerset. Climatic data from three sites show that there are considerable differences in temperature regime at different altitudes and also that there are major climatic differences between the valley system and a more maritime district further west. Pronounced intercolony variation that can be related to altitude and geographical position suggests that temperature and/or rainfall may cause frequency differences. There is some variation with habitat suggesting the action of visual selection, but there are also consistent differences between habitats which are not in accordance with this hypothesis.
The linkage states between the alleles of the colour and banding genes vary with altitude.
Variation in the distribution of Arianta arbustorum and Cepaea nemoralis suggest that climatic differences are limiting factors.     

Gene frequencies, genetic background and environment in Pyrenean populations of Cepaea nemoralis (L.)

Microgeographical genetic differentiation of populations of the polymorphic land snail Cepaea nemoralis (L.) has been the subject of intensive investigation. The importance of environmental selection acting at the loci which control the shell polymorphism in determining the distribution of genes in this species remains a matter of considerable controversy. Some populations of C. nemoralis in the central Pyrenees show striking local associations of morph frequency (and particularly of the frequency of unhanded shells) with topography which have been ascribed to the action of natural selection.
In the Ter valley (eastern Spanish Pyrenees) populations described here, however, the pattern of association of the frequencies of the shell morphs with topography is quite different from that found in the central Pyrenees. In particular, the distribution of unhanded shells is almost the inverse of that found in some previously studied populations. The general picture of variation in relation to topography in Pyrenean C. nemoralis populations is now seen to be one of great local consistency within valley systems, but of considerable differences between valley systems. This variation in population structure may be due either to cryptic and undetected environmental differences between valley systems or to evolutionary divergence between their C. nemoralis populations. The difficulty of deciding between these two conflicting hypotheses emphasizes the limitations of the technique of simply examining the distribution of individual genes in natural populations as a method of analysing their evolutionary history.     

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.     

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.     

The chromosomes of Cepaea vindobonensis (Pulmonata: Helicidae) and the relationship between this species and the rest of the genus

The karyotype of Cepaea vindobonensis (Pfeiffer) (2 n = 50) is defined and compared with those of C. sylvatica (Draparnaud), C. nemoralis (L.) and C. hortensis (Müller). Cepaea vindobonesis shares its chromosome number with C. sylvatica. The genus Cepaea thus consists of two pairs of species, one with 2 n = 50 the other with 2 n = 44. The karyotype of C. vindobonensis is distinct from that of C. sylvatica as it lacks any discontinuity in its range of chromosome length and has all three of its longest chromosome pairs nearly identical. The two 2 n = 50 species both have an unusual meiotic bivalent showing a 'satellite-type' structure. It is apparent that the 2 n = 50 species are more closely related to each other than to the 2 n = 44 species. No precise definition of the evolutionary history can be given but a working hypothesis is suggested.     

The effect of interspecific interaction on spatial distribution patterns in experimental populations of Cepaea nemoralis (L.) and C. hortensis (Müll.)

'Choice-chamber' population cages were used to investigate distribution patterns in mixed and unmixed populations of closely related species of landsnails, C. nemoralis (L.) and C. hortensis (Müll.). Numbers of snails migrating into cut-grass cages were affected by mixing the two species. The direction of response was not the same for both siblings. The results indicate that interspecific interaction could affect species' distribution patterns in free-living populations.     

The effects of density and interspecific interaction on mortality in experimental populations of adult Cepaea (Held.)

Field population cages were used to investigate the effects of density and interspecific interactions on adult mortality in two closely related species of landsnail, C. nemoralis (L.) and C. hortensis (Mull). High densities were associated with increased mortality in both species. Increased mortalities were also observed in populations in which the two species were placed together, compared to those in which each was kept separate. However, this trend was not consistent for populations which contrasted in density levels. The effect of C. hortensis on C. nemoralis was greater at low densities than at high densities. The results suggest that coexistence in nature may be unstable and could result in spatial displacement.     

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.     

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.     

Contrasting patterns of variation in urban populations of Cepaea (Gastropoda: Pulmonata): a tale of two cities

The shell colour and banding polymorphisms in urban populations of Cepaea snails were studied in Sheffield (England) and Wrocław (Poland), which are two cities of similar size. Both Cepaea nemoralis and Cepaea hortensis were found in Sheffield, but only C. nemoralis was found in Wrocław. In neither city was variation with habitat apparent, nor was there evidence of large-scale geographical patterns. Spatial autocorrelation analyses revealed only vaguely defined local clusters. There was a greater degree of among-population variation in Sheffield, and more cases of extreme frequencies and low levels of polymorphism. Wrocław populations were more uniform and highly polymorphic. These differences can be related to the history of these snails in each city. Sheffield has been colonized mainly in the last 20 years, and some parts of the city with apparently suitable habitat remain unoccupied. Leptokurtic dispersal of small propagules followed by local spread appears likely. Values of F _ST resemble those seen in districts elsewhere with 'Area Effects'. Although there are ancient woodlands within the city, they have been colonized too recently for selection to achieve a new balance. Wrocław has held Cepaea populations for at least 100 years, and they are denser and more continuous. Shaded habitats are recent and temporary; the overall pattern of variation matches that seen in open habitats in districts elsewhere in which variation with habitat is apparent, and values of F _ST also correspond. Population history and the character of dispersal clearly affect the patterns of variation observed.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 27–39.     

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.     

Cepaea nemoralis in a hostile environment: continuity, colonizations and morph-frequencies over time

In the 1930s Cyril Diver sampled and scored populations of the polymorphic snail Cepaea nemoralis on the South Haven Peninsula, Dorset. His recording methods make it possible to be confident about the absence of the species from parts of the peninsula, which has acid soils and changing habitats, and is generally unfavourable for the species. A resurvey in 1999 revealed a pattern of retreats and extensions of distribution related to local environmental change. Where populations have persisted over the period, the pattern of morph-frequency distributions has remained the same; there is equivocal evidence for a small increase in the frequency of midbanded shells. While most wetland populations have disappeared, new populations have become established along a roadside, and on newly-stabilized foredune ridges. Morph-frequencies in these new populations relate to those of the nearest established populations, and are sometimes monomorphic. The results are discussed in relation to the history of the peninsula, and to the role of migration in determining observed morph-frequency distributions.     

Citizen science reveals unexpected continental-scale evolutionary change in a model organism

Organisms provide some of the most sensitive indicators of climate change and evolutionary responses are becoming apparent in species with short generation times. Large datasets on genetic polymorphism that can provide an historical benchmark against which to test for recent evolutionary responses are very rare, but an exception is found in the brown-lipped banded snail (Cepaea nemoralis). This species is sensitive to its thermal environment and exhibits several polymorphisms of shell colour and banding pattern affecting shell albedo in the majority of populations within its native range in Europe. We tested for evolutionary changes in shell albedo that might have been driven by the warming of the climate in Europe over the last half century by compiling an historical dataset for 6,515 native populations of C. nemoralis and comparing this with new data on nearly 3,000 populations. The new data were sampled mainly in 2009 through the Evolution MegaLab, a citizen science project that engaged thousands of volunteers in 15 countries throughout Europe in the biggest such exercise ever undertaken. A known geographic cline in the frequency of the colour phenotype with the highest albedo (yellow) was shown to have persisted and a difference in colour frequency between woodland and more open habitats was confirmed, but there was no general increase in the frequency of yellow shells. This may have been because snails adapted to a warming climate through behavioural thermoregulation. By contrast, we detected an unexpected decrease in the frequency of Unbanded shells and an increase in the Mid-banded morph. Neither of these evolutionary changes appears to be a direct response to climate change, indicating that the influence of other selective agents, possibly related to changing predation pressure and habitat change with effects on micro-climate.     

At the north-eastern extremity: variation in <Emphasis Type="Italic">Cepaea nemoralis</Emphasis> around Gdańsk,northern Poland

Variation in the shell colour and banding polymorphism in the land snail Cepaea nemoralis was studied in 260 populations in the region of Gdańsk, northern Poland. Unlike in other regions of Poland, many populations contain brown shells. Populations from shaded habitats have higher frequencies of brown than those from open and intermediate habitats, largely at the expense of yellow shells. Nearly all brown shells are also unbanded. Apart from this disequilibrium, banding morphs among yellow and pink shells show no relationship to habitat. There are no broad geographical trends in morph-frequencies, but there are very strong correlations among populations very close together, revealed both by pairwise analysis and Moran’s I. Principal Component Analyses show that these correlations relate to overall genetic similarity at the loci involved. The populations are at the north-eastern limits of the species’ range; habitats are mostly anthropogenic, and comparisons with studies in two urban areas (Wrocław, SW Poland, and Sheffield, central England) suggest that the patterns of variation seen are a product of human transport of propagules followed by local dispersal. The effect of habitat here is much less marked than in regions much further west, but it indicates that natural selection has occurred.