Gene regulation of shell banding in a land snail from Israel

The Israeli land snail, Xeropicta vestalis, offers a particularly clear example of gene regulation in relation to natural selection, in that within each population the appropriate phenotype is generated only at the correct part of the animals life cycle, and a contrasting phenotype develops when the forces of natural selection change. In the mountains of Jerusalem, where the winter is cold, the shells are dark. Westwards, towards the coastal plain where the winter is warmer, the shells gradually become paler. As dark shells absorb more radiation than pale ones, this clinal variation in morph frequencies can be explained in thermal terms. (Banded shells are also more cryptic than non-banded shells, so that in the mountains visual selection by predators may be an additional force which favours dark shells.) Xeropicta vestalis is an annual, semelparous species: the snails hatch in winter, become mature within one year, reproduce, and then die. In the coastal plain the snails are active throughout most of the year, and they have a long period in which to grow to reach adult size. In the mountains and hills, on the other hand, the snails are active for only a very short period. They spend most of their lives as small snails, in a state of aestivation. Xeropicta vestalis must be dark in mountains because when it finally awakens, it must very rapidly and hastily reach reproduction size. A dark shell, by speeding up temperature-dependent processes in this critical stage, assists the snail to mature rapidly. Shell darkness varies with age: in the mountains and hills the shells are moderately dark when they hatch, but become darker whilst growing in early winter. In the coastal plain also, the snails are moderately dark when they hatch; but here they become paler, whilst growing in winter and spring. In both cases, each snail is darker in the colder months and paler in the hot ones. A strategy of gene regulation of shell colour is thus favoured when the subsequent forces of the environment are very contrasting in their direction, very severe—yet also very predictable.     

Convergence in the shell-banding polymorphism of two sympatric landsnails

Xeropicta vestalis , a polymorphic helicelline as regards its shell-banding pattern, has shells that closely resemble those of Theba pisana , a helicine. These resemblances are found mainly in Israel's coastal plain, where distribution of the two species overlaps to a certain extent, and it hardly ever occurs in Israel's hilly region where T. pisana is missing. In the Jordan Valley however (where T. pisana is missing also), some T. pisana -like forms of X. vestalis may occur. Because X. vestalis is smaller than T. pisana , the resemblance refers mainly to juveniles and small adults of the latter, and less to large adults.
We examined whether there are any interspecific relations between the frequencies of visually analogous morphs, in mixed populations. No such relations was found. Further, mixed populations are so unstable in time that the chances of building up any such relation is highly unlikely.
It is difficult to accept that there are any interrelationships whatsoever (of apostatic selection as well as mimicry) between T. pisana and X. vestalis as regards their shell banding polymorphism. The convergence between the two species appears to represent parallel, but not interspecific forces of natural selection.     

Habitat and integration within indigenous communities of Xeropicta derbentina (Gastropoda: Hygromiidae) a recently introduced land snail in south-eastern France

Xeropicta derbentina is a recently introduced land snail in Provence, France. The present work aims at describing the habitats and the land snail communities that it has invaded. Also, its impact on the species richness and diversity of indigenous communities is assessed. Two areas have been studied in order to cover the range of conditions that X. derbentina is encountering within its current French distribution. Sixty‐five samples were analysed from the most Mediterranean site, Auriol, and 140 from a northern site, Craus de St‐Michel‐l’Observatoire. From these samples, X. derbentina is found to be a thermophilous open‐land species mostly inhabiting human‐impacted habitats. It occurs in large populations (more than 100 individuals collected in 15 min) in habitat characterizing the first stages of secondary successions, from cultivated crops to dense fallow lands. However, within a heterogeneous landscape and if enough time has allowed its dispersal, X. derbentina can be found in almost any kinds of habitat, such as garrigues and woodlands. Consequently, X. derbentina occurs in diverse land snail communities. At the local scale, its large populations could result in a significant decrease in species richness and diversity. At the regional scale, however, the introduction of X. derbentina seems to have increased land snail species richness.     

Appearance of morphological novelty in a hybrid zone between two species of land snail

On the small oceanic island of Chichijima, two endemic species of land snails, Mandarina mandarina and M. chichijimana, have discrete distributions separated by a hybrid zone. This study investigates the potential of hybridization as a source of morphological novelty in these snails. Mandarina mandarina possesses a shell with a higher whorl expansion rate and a smaller protoconch than M. chichijimana, relative to shell size. The number of whorls and shell size of M. mandarina do not differ from those of M. chichijimana, because the effect of higher expansion rate on number of whorls and size of the former is compensated for by its smaller protoconch. The whorl expansion rate and protoconch diameter of the individuals from the hybrid populations are intermediate or typical of either of the two species, and their average values show clinal changes along the hybrid zone. However, the hybrid populations include exceptionally high shells with many whorls and flat shells with few whorls, which are never found in the pure populations of either species. In addition, gradual increases in variance in shell height and number of whorls were found from the edges to the center of the hybrid zone. A combination of low expansion rate (typical of M. chichijimana) and a small protoconch (typical of M. mandarina) produces a shell with an extremely large number of whorls because of the geometry of shell coiling. However, the combination of high expansion rate and a large protoconch produces a shell with an extremely small number of whorls. Because of the correlation between the number of whorls and shell height, shells with an exceptional number of whorls possess an extraordinarily high or flat spire. Hybrids can inherit a mosaic of characters that, as they play out during growth, lead to novel adult morphologies. These findings emphasize the importance of hybridization as a source of morphological variation and evolutionary novelty in land snails.     

Shell colour polymorphism in a mangrove snail Littorina sp. (Prosobranchia: Littorinidae)

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.     

Epistasis, phenotypic disequilibrium and contrasting associations with climate in the land snail Theba pisana

Hotter conditions favour effectively unbanded (EUB) shells in the snail Theba pisana. T. pisana is also polymorphic for colour of the shell's apex, determined by a pair of alleles at a locus linked to the banding locus. Apex colour is epistatic to shell banding, such that banded snails with a dark apex have darker bands. Annual censuses over 22 years across an ecotone between a sheltered Acacia thicket and open dune vegetation showed a persistent association of both EUB shells and pale apex with the Open habitat. The parallel variation was due partly to strong phenotypic disequilibrium, as the combination of EUB with dark apex was rare. Nevertheless, in fully banded shells the frequency of pale apex was also higher in the Open habitat, confirming independent, parallel associations of the two contributors to paleness. Within the Acacia habitat, temporal variation of the frequencies of banding morphs was much greater than for apex colour, and EUB shells were associated with hotter summers. Consistent with its primary effect only on the very small snails, apex colour did not vary with summer conditions, but instead, higher frequencies of pale apices were associated with sunnier winters. The intensity of selection was lower on apex colour than shell banding, due partly to the constraint of phenotypic disequilibrium. The shell traits in T. pisana are an example of complex responses to climatic variation, in which phenotypic disequilibrium constrains evolution of apex colour, but separate mechanisms of selection are evident.     

Influence of shell colour on the mortality rate of the land snail Arianta arbustorum under different microclimatic regimes

The mortality of phenotypic shell colour morphs and age classes of the snail Arianta arbustorum in several microclimatic conditions was recorded. An analysis of variance was performed on five factors: adaptation temperature, relative humidity, shell colour, age class and test temperature. There were no significant differences in the mortality between different adaptation temperatures or relative humidities, but the interaction of these two factors was highly significant. There were significant differences in mortality rate between test temperature and age class. The mortality of the brown morph was higher than the yellow one at all adaptation temperatures (overall, P < 0.05). There were no significant differences between the mortality rates of the two morphs at different relative humidities. The mortality of the brown morph was higher than that of the yellow at six out of seven test temperatures. Juvenile snails survived significantly better than adults at all test temperatures.     

Thirty-four years of climatic selection in the land snail Theba pisana

Few continuous, long-term studies have measured the intensity and variability of natural selection within a framework of clear adaptive hypotheses. In the snail Theba pisana, the proportion of effectively unbanded shells is higher in exposed habitats than in adjacent acacia thickets, which has been explained by microclimatic selection. Comparisons across an ecotone for 34 consecutive years determined the combined effects on morph frequencies of habitat and changes in weather conditions during summer. The long-term average (±s.e.) frequency of effectively unbanded shells was 0.577±0.011 in the open habitat when compared with 0.353±0.005 in the acacia. The persistent association of shell banding with habitat accounted for 34% of the variation in morph frequencies. Differences among years were also large, representing 23% of the variation. Higher proportions of effectively unbanded snails were associated with hotter, sunnier summers. Thus, temporal variation supports the hypothesis of microclimatic selection, consistent with the spatial association with habitat. Based on observed rates of change, the mean annual selection on this polymorphism was about 0.13, but with a large variance: s was as high as 0.5, but ⩽0.05 in about 40% of the years. The large variance and frequent reversals in direction of selection indicate a potential for rapid genetic change, but with little net change in morph frequencies over three decades, highlighting the value of long-term continuous studies of populations facing natural environmental variation.     

Characterization of 17 microsatellite loci in the Japanese land snail genera Mandarina,Ainohelix, and Euhadra (Mollusca,Gastropoda, Pulmonata)

Land snails of the genera Mandarina, Euhadra and Ainohelix are useful for understanding the ecology and evolution of speciation and adaptation, so we have developed 17 microsatellite loci for these species. As in other land snails, most of the loci are highly polymorphic compound repeats, with a great size range between alleles. The loci should be useful in understanding gene‐flow, genetic structure and speciation in these 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.     

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.     

Effects of inundation and salt on the survival of ants in a sandy coastal plain

Abstract. 1. Survival of four species of ants, Myrmica rubra, Myrmica scabrinodis, Lasius niger and Lasius flavus , exposed to prolonged inundation and the drinking of brackish water, was experimentally determined.
2. In most of the experiments, survival of Lasius flavus workers was much worse than either Lasius niger or Myrmica scabrinodis .
3. After inundation with brackish water, and drinking of brackish water for more than 3 weeks, survival of workers of Myrmica rubra was also more affected than that of Lasius niger and Myrmica scabrinodis .
4. As a rule, survival of dealated queens after inundation appeared to be better in Lasius flavus and Lasius niger , but worse in Myrmica rubra , compared with worker survival.
5. After surviving inundation, the capacity to produce eggs and workers was only slightly affected in queens of both Lusius species.
6. The conclusions based on the experimental mortality rates seem to be consistent both with ant species distribution and with frequency of inundation and salt stress in different parts of the coastal plain and surrounding sand-dunes on the Dutch Wadden island Schiermonnikoog.     

An experimental test of the role of predators in the maintenance of a genetically based polymorphism

Polymorphisms provide one of the most useful tools for understanding the maintenance of genetic and phenotypic variation in nature. We have previously described a genetically based polymorphism in dorsal patterning that is expressed by female brown anole lizards, Anolis sagrei, which occur in Bar, Diamond and intermediate Diamond-Bar morphs. Previous studies of island populations in The Bahamas support a role for selection in maintaining the polymorphism, but the agents responsible for this selection remain unclear. We tested two main hypotheses regarding the importance of predation as a selective agent that maintains the polymorphism within populations. First, we tested whether correlational selection favours different combinations of morph, locomotor performance and escape behaviour by measuring morph-specific natural selection on sprint speed, running endurance and the propensity of females to either 'freeze' or 'run' in response to attempted capture. Morphs did not differ in any of these traits, nor did correlational selection consistently favour any particular combinations of morph and antipredator behaviour. Second, we experimentally excluded bird and snake predators from two entire island populations, allowed these predators access to two additional islands and then measured subsequent differences in natural selection on morphs in each population. Predators reduced the survival of Bar and Diamond females, but not of genetically intermediate Diamond-Bar females. These results provide limited evidence that predation may play a role in maintaining this polymorphism, although the functional traits that could account for differential susceptibility to predation remain unclear.     

The effects of temperature and humidity on the nocturnal activity of different shell colour morphs of the land snail Arianta arbustorum

The noctural activities of the phenotypic shell colour morphy and age classes (adults and juveniles) of Arianta arbustorum were recorded 1 day week^-1 for 4 weeks in several laboratory microclimatic conditions. Six constant temperatures between 3 and 18C and four levels of relative humidities between 34 and 98% were maintained. A light regime of 16 h light: 8 h dark was used. There are highly significant differences in activity at different levels of adaptation temperature and relative humidity. The interaction between these factors is significant. There are no significant differences in nocturnal activity between the two phenotypic shell colours nor between the two age classes, but the interaction between morph and relative humidity is significant. The interaction between age classes and relative humidity is also significant. Yellows are more active than browns at high humidities, but less active at low. They are therefore likely to be behaviourally more responsive than browns in an environment of fluctuating humidities. This result is discussed in relation to the maintenance of the polymorphism.     

On the gynodioecious polymorphism in Saxifraga granulata L. (Saxifragaceae)

Sexual and vegetative fitness components in hermaphrodite and female plants of the self-compatible, perennial herb Saxifraga granulata are compared using material derived from a gynodioecious population in northern England.
Females produced only 57% as many seeds as hermaphrodites, but their ovule offspring were 1.28 times as fit as those of hermaphrodites, and females were more vegetatively vigorous. The advantages to females in ovule offspring quality and in vegetative reproduction counteract their disadvantages in pollen and seed production and therefore probably play a role in the maintenance of the gynodioecious polymorphism. Pollination ecology, resource reallocation and inbreeding depression all appear to contribute to the observed sex differences in fitness.     

Ecological speciation in an island snail: evidence for the parallel evolution of a novel ecotype and maintenance by ecologically dependent postzygotic isolation

Speciation is the process by which reproductive isolation evolves between populations. Two general models of speciation have been proposed: ecological speciation, where reproductive barriers evolve due to ecologically based divergent selection, and mutation‐order speciation, where populations fix different mutations as they adapt to similar selection pressures. I evaluate these alternative models and determine the progress of speciation in a diverse group of land snails, genus Rhagada, inhabiting Rosemary Island. A recently derived keeled‐flat morphotype occupies two isolated rocky hills, while globose‐shelled snails inhabit the surrounding plains. The study of one hill reveals that they are separated by a narrow hybrid zone. As predicted by ecological speciation theory, there are local and landscape level associations between shell shape and habitat, and the morphological transition coincides with a narrow ecotone between the two distinct environments. Microsatellite DNA revealed a cline of hybrid index scores much wider than the morphological cline, further supporting the ecological maintenance of the morphotypes. The hybrid zone does not run through an area of low population density, as is expected for mutation‐order hybrid zones, and there is a unimodal distribution of phenotypes at the centre, suggesting that there is little or no prezygotic isolation. Instead, these data suggest that the ecotypes are maintained by ecologically dependent postzygotic isolation (i.e. ecological selection against hybrids). Mitochondrial and Microsatellite DNA indicate that the keeled‐flat form evolved recently, and without major historical disruptions to gene flow. The data also suggest that the two keeled‐flat populations, inhabiting similar rocky hills, have evolved in parallel. These snails provide a complex example of ecological speciation in its early stages.     

Radiation of the land snail genus Sagda (Pulmonata: Sagdidae): comparative morphology, biogeography and ecology of the species of north-central Jamaica

The radiation of the endemic Jamaican land snail genus Sagda is examined through analysis of the comparative shell morphology, anatomy, ecology and biogeography of the four species inhabiting north-central Jamaica: S. centralis sp. nov. , S. spei, S. bondi and S. montegoensis. The species are all rather similar in their morphology and ecology. The shells show some differences in the presence or absence of internal lamellae, the position of the lamellae, the form of the depression in the centre of the base, the regularity of sculpture (low ribs) and the pattern of microsculpture. Slight differences in the form of the radular teeth are found between some species. The structure of the secondary ureter is peculiar: in all species it is open shortly behind the mantle collar, then branches into two closed tubes which open separately on the outside of the mantle collar. Despite its great complexity, the reproductive system is rather uniform among species; differences are seen in the number of pockets that form spines on the spermatophore. The complex, three-branched spermatophore is formed within the entire epiphallar branch of the penis, whereas the appendicular branch apparently assumes the copulatory function. When inactive, all species are found associated with rocks or, less commonly, among leaf litter. Activity occurs primarily at night and mostly on leaf litter rather than rocks. The snails eat dead plant material, including a variety of plant species. All species (with the possible exception of S. montegoensis) are ovoviviparous. They overlap broadly in the range of climates occupied. The geographic ranges of the four species are overlapping, although Sagda bondi has a parapatric distribution in relation to the western species S. grandis. Fossil and subfossil evidence shows that shifts in the ranges of the species have occurred in the late Quaternary; Pleistocene-Holocene environmental changes, habitat destruction by man, and possibly other environmental changes during historical times are probably involved. Radiations of other land snail groups on tropical islands have generally produced greater morphological and ecological differentiation than in Sagda. Allopatric or parapatric species distributions appear to be more common in other land snail radiations.