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.     

Temporal morph frequency changes in sand‐dune populations of Cepaea nemoralis (L.)

Changes in morph frequency over time in sand‐dune populations of Cepaea nemoralis in the British Isles have been examined using the Evolution Megalab database. Frequencies in colonies on a sand dune at Berrow, Somerset were estimated in 2008–9. This extends a survey started in 1926 by A. E. Boycott and C. Diver and continued in the mid‐20th Century by B. C. Clarke and J. J. Murray. An increase in the frequency of the mid‐banded morph, noted in earlier work, has continued. The apparent decrease in brown was not confirmed but the yellow frequency has increased. At a range of dunes in the British Isles, comparisons between data from the mid‐20th Century and the early 21st Century indicate an increase in yellow and mid‐banded morphs and a decrease in banding and brown morphs. These results differ from the overall trends derived from Europe‐wide comparison of early data with Evolution Megalab data, in which banded and mid‐banded increased in frequency, whereas there was no overall change in yellow. The general pattern of regional variation has been retained, although there is also high heterogeneity between samples, suggesting that a variety of factors are involved in explaining the changes. The mean shifts in frequency are consistent with climatic change over the period. Dunes are probably the most likely habitat in which to detect such a change. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 108 , 315–322.     

Change and stability in a steep morph‐frequency cline in the snail Cepaea nemoralis (L.) over 43 years

Populations of the polymorphic land snail Cepaea nemoralis (L.) from Deepdale, Derbyshire, UK, sampled in 1965–67, showed a pattern of area effects, with steep clines among groups of populations differing in shell colour and banding morph frequencies. In 2010, most of these populations were resampled. In particular, a continuous transect made in 1967 of 42 quadrats (18.34 × 18.34 m) across a steep cline in several morph frequencies was completely resampled. In the dale as a whole, yellow shells had increased in frequency. In the transect, the frequencies of banding morphs showed no significant changes, although colour morphs showed some changes. Pink shells had increased in frequency in a section in which scrub had developed, and brown shells had increased in frequency in the area in which they had originally been at the highest frequency. In each case, the selection coefficients were of the order of 4%. Yellow had increased elsewhere. Nevertheless, both in the dale as a whole and in the transect, the pattern of geographical change in morph frequencies had remained essentially the same. Estimates of migration based on previous studies of marked snails and on modelling of the effect of drift and migration suggest that, regardless of whether the cline is a product of differential selection or of the gradual merging of previously separate founding populations, it has been in existence for a long time, and that migration occurs over greater distances than estimated from direct observation on marked snails. Although we can demonstrate that selection has occurred, the origin and maintenance of the cline and others similar to it remain in doubt; the development and maintenance of polymorphism in this species may require consideration of several processes operating on different time scales. © 2012 The Linnean Society of London     

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.     

Evolutionary change in Cepaea nemoralis shell colour over 43 years

We compared shell colour forms in the land snail Cepaea nemoralis at 16 sites in a 7 × 8 km section of the Province of Groningen, the Netherlands, between 1967 and 2010. To do so, we used stored samples in a natural history collection and resampled the exact collection localities. We found that almost all populations had experienced considerable evolutionary change in various phenotypes, possibly due to population bottlenecks and habitat change after repeated land consolidation schemes in the area. More importantly, we found a consistent increase in yellow effectively unbanded snails at the expense of brown snails. This is one of the expected adaptations to climate change (this area of the Netherlands has warmed by 1.5–2.0 °C over the time period spanned by the two sampling years), and the first clear demonstration of this in C. nemoralis.     

Morph frequencies in the snail Cepaea nemoralis: changes with time and their interpretation

Colonies of the polymorphic snail Cepaea nemoralis on the sand dune at Point of Air, north Wales, first sampled in 1962 by A.J. Cain and P.M. Sheppard, have been re-sampled after 35 years. The frequency of the mid-banded morph has increased. Overall, yellow frequency has decreased at the expense of pink, but the change is in different directions in different parts of the dune. Yellow mid-banded and pink five-banded, phenotype combinations controlled by alleles at unlinked loci, are in excess within colonies, which suggests an effect of selection. It is argued that changes in morph frequency are due, at least in part, to movement between colonies. Cepaea populations are slow to respond to environmental changes and displacement of individuals may be greater than is often assumed. It is therefore difficult to interpret frequency changes over time. Correlated changes in groups of colonies may not always be interpretable as indicating selection.     

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.     

Altitudinal and seasonal variation in microsatellite allele frequencies of Drosophila buzzatii

Variation in climate, particularly temperature, is known to affect the genetic composition of populations. Although there have been many studies of latitudinal variation, comparisons of populations across altitudes or seasons, particularly for animal species, are less common. Here, we study genetic variation (microsatellite markers) in populations of Drosophila buzzatii collected along altitudinal gradients and in different seasons. We found no differences in genetic variation between 2 years or between seasons within years. However, there were numerous cases of significant associations between allele frequencies or expected heterozygosities and altitude, with more than half showing nonlinear relationships. While these associations indicate possible selection and local altitudinal adaptation, direct tests gave strong evidence for selection affecting two loci and weaker evidence for five other loci. Two loci that are located within an inversion (including the one with strongest evidence for selection) show a linear increase in genetic diversity with altitude, likely due to thermal selection. Parallel associations with altitude here and with latitude in Australian populations indicate that selection is operating on chromosomal regions marked by some of the loci.     

Rapid evolution in unstable habitats: a success story of the polymorphic land snail Cepaea nemoralis (Gastropoda: Pulmonata)

The present study reports on a natural experiment with twelve replicates in which rapid, predictable and consistent divergence of Cepaea nemoralis populations occurred in response to repeated selection gradient of adjacent open and shaded habitats. Because the frequencies of various genetically‐based phenotypes varied widely among surveyed populations, and there was a large overlap between habitat types, no overall association with habitat was apparent. In paired comparisons, however, significant changes were consistently towards higher frequencies of light morphs in the open than corresponding shaded habitats, and this result is attributable to natural selection. This shows that the knowledge of the genetic composition of reference populations is often essential for discerning selection from random processes. At each site, a different morph combination contributed to the divergence of populations, indicating that there are many genetic solutions to similar ecological problems; this likely enhances the maintenance of high levels of polymorphism. Adaptation of populations occurred in contemporary time and was fast. In one case where it was possible to follow changes, significant shifts in morph frequencies occurred within just two snail generations (selection coefficients of 0.404 and 0.518). High evolvability may be one of the factors contributing to the ecological success of Cepaea nemoralis. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 251–262.     

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.     

Influences of extreme weather,climate and pesticide use on invertebrates in cereal fields over 42 years

Cereal fields are central to balancing food production and environmental health in the face of climate change. Within them, invertebrates provide key ecosystem services. Using 42 years of monitoring data collected in southern England, we investigated the sensitivity and resilience of invertebrates in cereal fields to extreme weather events and examined the effect of long‐term changes in temperature, rainfall and pesticide use on invertebrate abundance. Of the 26 invertebrate groups examined, eleven proved sensitive to extreme weather events. Average abundance increased in hot/dry years and decreased in cold/wet years for Araneae, Cicadellidae, adult Heteroptera, Thysanoptera, Braconidae, Enicmus and Lathridiidae. The average abundance of Delphacidae, Cryptophagidae and Mycetophilidae increased in both hot/dry and cold/wet years relative to other years. The abundance of all 10 groups usually returned to their long‐term trend within a year after the extreme event. For five of them, sensitivity to cold/wet events was lowest (translating into higher abundances) at locations with a westerly aspect. Some long‐term trends in invertebrate abundance correlated with temperature and rainfall, indicating that climate change may affect them. However, pesticide use was more important in explaining the trends, suggesting that reduced pesticide use would mitigate the effects of climate change.     

Salamander morph frequencies do not evolve as predicted in response to 40 years of climate change

The global climate is changing rapidly, yet biotic responses remain uncertain. Most studies focus on changes in species ranges or plastic responses like phenology, but adaptive evolution could be equally important. Studying evolutionary responses is challenging given limited historical data and a poor understanding of genetically variable traits under selection. We take advantage of a historical dataset to test for an adaptive response to climate change in a widespread, polymorphic amphibian, the eastern red‐backed salamander Plethodon cinereus. We resurveyed color morph frequencies across New England to test for an adaptive shift in response to climate change. We modeled historical and present‐day morph proportions as a function of climate and tested the accuracy of predictions both within and across different time periods. Our models showed moderate accuracy when predicting morph frequencies within time periods, but poor accuracy across time periods. Despite substantial changes in climate and significant relationships between morph frequency and climate variables within periods, we found no evidence for the predicted shift in morph frequencies across New England. The relationship between climate and color morph frequencies is likely more complex than originally suggested, potentially involving the interplay of additional factors such as microclimate variation, land use changes, and frequency‐dependent selection. Model extrapolation and changes in the correlation structure of climate variables also likely contributed to poor predictive ability. Evolution could provide a means to moderate the effects of climate change on many species. However, we often do not understand the direct links between climate variation, traits, and fitness. Therefore, forecasting climate‐mediated evolution remains an ongoing and important challenge for understanding climate change threats to species.     

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.     

近56年来科尔沁沙地气候变化特征

根据1951-2006年的气温和降水资料,研究了干旱少雨、生态环境脆弱、土地荒漠化严重的科尔沁沙地近56年的气候变化特征.结果表明:1951-2006年,科尔沁沙地的气温以0.28℃·10 a~(-1)的速度上升,远大于全球近50年来0.13℃·10 a~(-1)的平均增温速率;各季节气温都呈上升趋势,冬、春季增温速率极显著(P<0.001),增温速率分别为0.46和0.39℃·10 a~(-1);年最高气温(0.17℃·10 a~(-1))与年最低气温(0.42℃·10 a~(-1))均呈极显著地增加趋势(P<0.01);降水量年际间波动较大,无明显的变化趋势,各季节降水量也没有明显的变化规律;年降水日数呈显著减少趋势(1.3 d·10 a~(-1)),各季的降水日数虽都有逐年减少趋势,但没有达到显著水平;年降水强度和各季降水强度都没有明显的变化规律;年总小雨日数呈显著的减少趋势(P<0.05),减少速率为1.0 d·10 a~(-1).

Abstract：

Based on the 1951-2006 air temperature and precipitation data from seven meteoro-logical stations in Horqin Sandy Land, the past 56-year climate change characteristics in this semi-arid area with vulnerable eco-environment and severe land desertification were analyzed. From 1951 to 2006, the mean annual air temperature in this area increased by 0. 28℃·10 a~(-1), much higher than the global average value of 0.13℃·10 a~(-1) in recent 50 years. The increasing trend of the temperature appeared in all four seasons, but was only significant in winter and spring (P<0.01), with an increasing rate of 0. 46℃·10 a~(-1) and 0.39℃·10 a~(-1), respec-tively. Both the maximum and the minimum mean annual temperatures increased significantly (P <0. 01), and the increasing rate was about 0.17℃·10 a~(-1) and 0.42℃·10 a~(-1), respec-tively. The annual precipitation fluctuated among years, with no obvious change trend at yearly and seasonal scales, while the precipitation days decreased significantly at yearly scale (1.3 d·10 a~(-1)) but not at seasonal scale. No obvious change patterns were observed in the precipitation intensity both in yearly and seasonal scales. The annual light-rain days decreased significant (P <0.05) in a rate of 1.0 d·10 a~(-1).     

Variation in sexual size dimorphism among populations: testing the differential‐plasticity hypothesis

Sexual size dimorphism (SSD) is a common phenomenon in animals and varies widely among species and among populations within species. Much of this variation is likely due to variance in selection on females vs. males. However, environmental variables could have different effects on females vs. males, causing variation in dimorphism. In this study, we test the differential‐plasticity hypothesis, stating that sex‐differential plasticity to environmental variables generates among‐population variation in the degree of sexual dimorphism. We examined the effect of temperature (22, 25, 28, and 31 °C) on sexual dimorphism in four populations of the cockroach Eupolyphaga sinensis Walker (Blattaria: Polyphagidae), collected at various latitudes. We found that females were larger than males at all temperatures and the degree of this dimorphism was largest at the highest temperature (31 °C) and smallest at the lowest temperature (22 °C). There is variation in the degree of SSD among populations (sex*population interaction), but differences between the sexes in their plastic responses (sex*temperature interaction) were not observed for body size. Our results indicated that sex‐differential plasticity to temperature was not the cause of differences among populations in the degree of sexual dimorphism in body size.     

Reduced variability in range‐edge butterfly populations over three decades of climate warming

Populations at the high latitude edge of species’ geographical ranges are thought to show larger interannual population fluctuations, with subsequent higher local extinction risk, than those within the ‘core’ climatic range. As climate envelopes shift northward under climate warming, however, we would expect populations to show dampened variability. We test this hypothesis using annual abundance indices from 19 butterfly species across 79 British monitoring sites between 1976 and 2009, a period of climatic warming. We found that populations in the latter (warmer) half of the recording period show reduced interannual population variability. Species with more southerly European distributions showed the greatest dampening in population variability over time. Our results suggest that increases in population variability occur towards climatic range boundaries. British sites, previously existing at the margins of suitable climate space, now appear to fall closer to the core climatic range for many butterfly species.     

Lifetime fitness components in female colour morphs of a damselfly: density‐ or frequency‐dependent selection?

In many damselfly species mature females exhibit colour polymorphism: one female morph resembles the conspecific male (androchrome) while the others do not (gynochromes). Hypotheses for the maintenance of such polymorphisms differ mainly as to whether they are based on density- and/or frequency-dependent selection and on the nature of the frequency dependence. We collected lifetime fitness data (individual lifespan, number of copulations and number of ovipositions) for female morphs of the damselfly Ischnura elegans from 15 insectaries differing in population parameters (density, sex ratio and ratio of andro- to gynochromes). Both density and frequency affected a specific set of the studied fitness components. While morph frequency influenced lifespan, sex ratio influenced the number of copulations, and density affected lifespan and the number of ovipositions. Clearly, discrepancies among studies may be generated if the studied fitness components differ. Our final fitness estimate, the number of ovipositions, was only influenced by density, thereby not supporting frequency-based hypotheses. Contrary to expectation under the current density-based hypothesis, androchromes compared to gynochromes had a lower number of ovipositions at high density. We discuss our findings in the light of mechanisms maintaining the female polymorphism.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 86 , 515–523.