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Natural populations of widely-distributed animals often exhibit clinal variation in phenotypic traits or in allele frequencies of a particular gene over their geographical range. A planktotrophic intertidal snail, Littorina keenae is broadly distributed along the north-eastern Pacific coast through a large latitudinal range (24°50′N–43°18′N). We tested for latitudinal clines in two complex phenotypic traits – thermal tolerance and body size – and one single locus trait – heat shock cognate 70 (HSC70 )–i nL. keenae along almost its entire geographical range. We found only weak evidence for a latitudinal cline in the thermal tolerance and no evidence for a cline in allele frequencies at HSC70. However, as predicted by Bergmann’s rule, we detected a strong latitudinal cline that accounted for 60% of the variance in body size (R 2 = 0.598; P < 0.001). In contrast, body size did not significantly affect thermal tolerance. HSC70 showed no genetic differentiation among the populations, supporting our previous mitochondrial gene-based estimate of high gene flow during this snail’s free-swimming larval stage. Given that L. keenae experiences panmixia along its species range, the observed size cline may be partially or entirely caused by a phenotypically plastic response to local thermal environments rather than by genetic divergence in body size among populations in response to locally optimizing natural selection. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 100, 494–505.

Many gastropods have inherited conspicuous shell colour polymorphisms. A challenging question is, are colour frequencies under selection or is polymorphism owing to random evolutionary processes? The intertidal species Littorina saxatilis (a rock-dweller) and L. obtusata (confined to macroalgae) both have genetically determined shell colour variation. In Iceland, Littorina obtusata are mostly cryptic on brown macroalgae by having brown or yellow shells (~95% of the snails), while Littorina saxatilis often appears conspicuous to the background of dark rocks owing to non-cryptic colours (15–20%). This difference may be due to selective elimination of conspicuously coloured L. obtusata by visual predators, while L. saxatilis, largely living in another habitat, is not under a similarly intense colour selection. To test this hypothesis we increased the frequencies of conspicuous L. saxatilis in experimental populations (from

The marine snail Littorina saxatilis is highly polymorphic for shell colour. It lives in the heterogeneous intertidal zone, where there are sharp transitions in a number of abiotic factors that may influence the relative fitness of morphs. We investigated the hypothesis of selected variation by relating the colour distribution to five factors (wave exposure, substratum, shore level, sex, snail age), and to interactions between them. We compared patterns from geographical areas in Sweden, Iceland and Russia. Cryptic morphs (tessellated and different dark colours) generally dominated (80–98%) while conspicuous morphs (white, yellow, red and banded) were less common (2–20%). The colour frequencies were often related to wave exposure, substratum and shore level. Frequencies rarely varied with age and never with sex. In order to test the assumption that the different colours are genetically determined we cross-bred snails from Iceland in the laboratory. Both the presence of bands and the ground colours of the shell were inherited, and we have tentative support for a one-locus two-allele model for banding. Our results support a model of selected inherited colour variation, involving a number of different selective agents, the importance of which may vary between populations on local and geographical scales.

A planktotrophic larval development suggests a relatively high gene flow over long distances. Both Littorina littorea and Melarhaphe neritoides have an egg capsule and planktotrophic larvae which taken together are pelagic over a period of 4–8 weeks. In the absence of differential selection one would expect low levels of genetic differentiation over large distances in these two species. In this study of allozyme variation, low levels of differentiation were found over thousands of kilometres (Norway to Spain for L. littorea and Sweden to Greece for M. neritoides). This supports the hypothesis of two species with a high dispersal potential. A second expectation from neutral theory is that effective population size is positively correlated with average levels of genetic variation within species. In light of the generally high densities of local populations of these species and the high interpopulation migration rate, both L. littorea and M. neritoides may be considered as having high Ne s. Contrary to neutral expectation, L. littorea revealed very low levels of heterozygosity over its whole European range (mean Hexp= 0.015, 15 loci), while average heterozygosity of M. neritoides (Hexp= 0.084, 11 loci) was no more than in other littorinid species. This paper also reports the occurrence of M. neritoides on the Swedish west coast in 1988, 89 and 90, and two factors which may have promoted this unusually large invasion are discussed.

Twenty-three enzymes and five shell parameters were screened in 11 subpopulations of Littorina saxatilis Olivi (=L. rudis Maton) occupying different habitats over a 1 km stretch of coastline. Shell morphology varied considerably and consistently with respect to degree of exposure, and since there is evidence that such morphology is at least partly under genetic control, it is likely that natural selection selects particular genotypes at particular locations. There was significant allozyme heterogeneity between neighbouring subpopulations, sometimes only metres apart, but little of the allozyme variability could be related directly to environmental pressures. Thus, with the exception of the Odh locus, the considerable morphological differentiation between snails from exposed and sheltered sites was not reflected in differentiation of those genes coding for electrophoretically assayed enzymes. At the Odh locus, virtually all the genetic differentiation between subpopulations was attributable to differentiation between habitat types. Two loci, Sod-1 and Aat-1, showed highly significant genetic disequilibrium, and possible reasons for this are explored. The population structure as assessed electrophoretically accords well with the stepping-stone model which permits greater differentiation of neighbouring populations than the island model, and which seems realistic in the ovoviviparously reproducing L. saxatilis, where the greater part of gene flow is likely to occur through the occasional migration of adults between contiguous populations.

As adults Littorina littorea and L. saxatilis have overlapping distributions, both on the shore and on a geographical scale, this implies that they are affected by similar selective forces to a large extent. The two species have, however, different reproductive strategies. Littorina littorea has a planktonic larval stage lasting 4–6 weeks, whereas L. saxatilis has direct development. This influences dispersal rates of the two species, and gene flow and effective population size are both assumed to be much smaller in L. saxatilis than in L. littorea. Intraspeciflc variation in morphological and allozyme characters differs between the two species. Littorina littorea is homogeneous for both types of characters, both within and between populations, while populations of L. saxatilis may show pronounced differences in shell characters with micro-environment changes, and allozyme variation is markedly higher in this species too. The polymorphism in shell characters of L. saxatilis and the monomorphism of L. littorea, may be expected from theory as a consequence of the different dispersal strategies. Also the lack of divergence between geographically separate populations of L. littorea, but the high degree of between population differentiation in L. saxatilis, could be attributed to the high and low dispersal rates, respectively. However, the larger allozyme variation within subpopulations of L. saxatilis compared to that of L. littorea, is surprising as the former species has a smaller effective population size than the latter. Furthermore, there is no correlation between dispersal ability and overall allozyme heterozygosity, when 10 additional species of Littorina are included in the comparison. It is suggested that different evolutionary backgrounds of different species will possibly explain this, as neither selectionist nor neutralist models alone account for the observation. That is, an earlier passage through a population bottleneck may, for example, influence the genetic variation for a long period of time.

We tested in the laboratory three pairs of species belonging to three genera and families of marine gastropods, Monodonta turbinata, M. turbiformis (Trochidae), Littorina punctata, L. neritoides (Littorinidae), Cerithium scabridum, C. rupestre (Cerithiidae), for resistance to diverse inorganic (heavy metals and NaC1) and organic (detergents and crude oil) pollutants. Each pair consisted of one narrow-niche species with low genetic diversity and one broad-niche species with higher genetic diversity. Evidence is presented that in all three cases the species with a higher level of genetic diversity was more resistant to all pollutants than its counterpart. These results suggest that fitness is positively correlated with heterozygosity and support the niche-width-variation hypothesis in regard to pollutants. The results also have practical implications for the identification of optimum marine species as genetic monitors of pollution.

Six populations of Littorina rudis and ihree of L. arcana were screened for elcctrophorctically dctci table variation at 21 enzyme loci. These species reproduce by ovoviviparity and oviparily respectively, and both are members of the L. saxatilis species complex. The mean coefficient of genetic identity between the two species was high (0.957) and no diagnostic loci were observed. Allele frequency differences are maintained in sympatric populations of the two species. Intraspecific values of/ ranged from 0.944 to 0.995 and both species showed marked inter-population genetic heterogeneity, a feature consistent with their limited powers of dispersal. Heterozygosity is high in all populations of both species. Populations of I. rudis have higher mean heterozygosities (= 0.153) then L. arcane (H= 0.132).

Shell colour polymorphism was examined in populations of a mangrove snail Littorina sp. in Queensland, Australia. Three morphs were recognized, yellow, red and brown, and morph frequencies varied both between widely spaced populations and between islands less than 1 km apart. Morph frequencies also varied with time of year. There was a relationship between shell colour and position on the tree, with yellow snails more often occurring amongst the foliage and brown snails more often on trunks and branches. In some populations yellow snails appeared to survive better than the other morphs, while in other populations there was no difference. The evidence for the maintenance of the polymorphism by natural selection is discussed.

Snails in the genus Littorina (Gastropoda, Prosobranchia) were examined in an area of mangroves in Papua New Guinea. It is shown that these may be separated into three species belonging to the L. xabra complex. Two of the species live on the roots only while the third is restricted to the leaves. The root species are monomorphic and cryptic and co-exist with three other monomorphic cryptic species of snails. The leaf species is polymorphic. This is a particularly clear-cut case of the tendency seen in gastropods for species of dark, uniform habitats to be monomorphic, while those on bright, patterned foliage backgrounds are polymorphic. It is suggested that the polymorphism is maintained by resemblance of the morphs to different elements in the mangrove foliage background, which are present at rather constant frequencies over a wide area.