Habitat-related dispersal in the rock-dwelling land snail Chondrina clienta

Habitat type may influence dispersal in a variety of animal species We examined dispersal in the rock-dwelling land snail Chondrina clienta m a limestone pavement, on vertical rock walls, a pile of stones and a stone wall on the Baltic island of Oland, Sweden Dispersal was estimated by recording movements of marked C clienta in natural populations over 3 yr Dispersal differed significantly between habitat types The largest distances dispersed were recorded in the limestone pavement (264 cm yr¹, median distance) and on vertical rock walls (96 cm yr¹), simple habitats for travelling snails Dispersal was less in the stone pile (68 cm yr¹) and on the stone wall (88 cm yr¹) that are more complex habitats with multiple layers of pieces of stones Distances dispersed also varied among vertical rock walls, indicating that other factors such as exposure of the rock surface and size of the habitat may be important in determining snail dispersal The results of two experiments indicated that grassland vegetation may inhibit dispersal in C clienta , and that isolated stones covered with lichens might serve as stepping stones for dispersing snails in otherwise unsuitable grassland Snail size (age) influenced distances moved, but might only be important in determining daily movements, not dispersal over longer periods Dispersal in C clienta is habitat-specific and cannot be characterized by a single parameter     

Associations between shell morphology and land crab predation in the land snail Cerion

1. The land crab Gecarcinus lateralis is a significant predator of the abundant Bahamian land snails of the genus Cerion . The crabs typically 'scissor' the cylindrical shells in half or break the lip and peel back the shell to reach the animal which withdraws two or three whorls into the shell. Scars on shells of live adults at 73 sites in the Bahamas and Florida Keys show that about 8% (range: 0–44%) of the snails have survived attacks of this type.
2. An artificial crab claw was used to investigate the compressive force required to break Cerion shells of different morphotypes. Defining shell strength as the ability to withstand compressive forces, 10 morphotypes were found that exhibited mean relative strengths of between 30 and 300 newtons. Feeding trials with one adult crab showed that snails whose shells could withstand compressive forces of > 95 N were safe from this individual predator.
3. Multiple linear regression analyses showed that both shell size (length and width) and shell wall thickness were the ultimate determinants of shell strength. Ribs strengthen the shell by contributing to wall thickness and also by increasing overall shell width. The thickened adult shell lip and collabral ribs provide effective protection from attack by peeling.     

Effects of food availability and intra-and interspecific interaction on the dispersal tendency in the land snail Chondrina clienta

The land snails Chondrina clienta and Baleaperversa compete most probably for a limited food resource (calcicolous lichens) on rock-faces on the Baltic Island of Öland (Sweden). Two laboratory and three field experiments were conducted to determine whether food availability and intra- and interspecific interactions affect the dispersal tendency in C. clienta . Under laboratory conditions, individuals of C. clienta showed a higher tendency to disperse from previously grazed than from ungrazed pieces of limestone (their natural substratum), whereas conspecific density or presence of B. perversa had no effect. However, when snails had been kept on the pieces of limestone for 40 days prior to testing, dispersal tended to increase with increasing density of conspecifics. In the field, marked individuals of C. clienta were released at different densities on vertical rock walls. Density of conspecifics at the release points did not influence the distances travelled. Similarly, intraspecific density did not affect dispersal when the snails' food resource (lichens) had been experimentally reduced on the quarry walls. Neither was the dispersal tendency influenced by different crowding conditions experienced for 40 days prior to release. The discrepancy between the results of laboratory and field experiments suggest that additional factors acting on dispersal are of importance in natural populations of C. clienta .     

Genital dimorphism in natural populations of the land snail Chondrina clienta and the influence of the environment on its expression

Several species of simultaneously hermaphroditic land snails show a genital dimorphism: aphallic individuals differ from euphallic ones by a lack of male copulatory organs (penis plus genital retractor muscle). Aphallic individuals can self-fertilize or outcross as females but not as males. Thus, the mating system of a population may be significantly influenced by the proportion of aphallic individuals.
We present data on the frequency of aphally in 23 natural populations of the rock-dwelling land snail Chondrina clienta on the Baltic island of Öland, Sweden. The populations varied greatly in percentage of aphallic individuals, ranging from 52.2 to 99.1%, (grand mean 77.7%)). This variation did not follow any geographical pattern.
In a laboratory experiment, we examined whether food supply (high or low) and/or population density (high or low) experienced during ontogeny affected the expression of genital dimorphism. Snails derived from a population with 99.1% aphallic individuals and raised under different food and density conditions did not differ from the original population in frequency of aphally. By contrast, when snails from a population with 66.7% aphallic individuals were raised on a low food supply, more individuals became euphallic than expected under complete genetic determination. These results suggest that, in addition to a genetic component. the expression of the genital dimorphism can be influenced by environmental conditions.     

Microgeographic evolution of snail shell shape and predator behavior

Genetic divergence in geographically isolated populations is a prerequisite for allopatric speciation, one of the most common modes of speciation. In ecologically equivalent populations existing within a small, environmentally homogeneous area, an important role for environmentally neutral divergence is often found or inferred. We studied a species complex of conspicuously shaped Opisthostoma land snails on scattered limestone outcrops within a small area of lowland rainforest in Borneo. We used shell morphometrics, mitochondrial and nuclear DNA sequences, and marks of predation to study the factors involved in allopatric divergence. We found that a striking geographic divergence exists in shell morphology, which is partly associated with neutral genetic divergence. We also found geographic differentiation in the behavior of the snails' invertebrate predator and evidence of an evolutionary interaction between aspects of shell shape and predator behavior. Our study shows that adaptation to biotic aspects of the environment may play a more important role in allopatric speciation than previously suspected, even on a geographically very small scale.     

Population regulation in the land snail Arianta arbustorum: density effects on adult size,clutch size and incidence of egg cannibalism

Summary The relationships between local population density and adult size, clutch size and spatial distribution of egg batches were investigated in 11 natural populations of the land snail Arianta arbustorum in a forest near Uppsala, Sweden. Shell size of adults decreased with increasing population density as did clutch size. Within populations, clutch size scaled allometrically with shell size indicating size-specific fecundity. It is hypothesized that food unpalatability caused by mucus deposition slows down juvenile growth rate in high density populations, resulting in small adults and thus reducing their fecundity in subsequent years. The influence of the distance between batches on the incidence of egg cannibalism by hatchlings was examined in a laboratory experiment. In this experiment the number of eggs cannibalized increased with decreasing distance to the batch of hatching snails. Thus, in the field, eggs of highly aggregated batches suffer a high risk of cannibalism. In the 3 populations with the highest snail density, 21–39% of all batches were deposited close to each other (nearest neighbour distance 5 cm, i.e. less than hatchlings more within 1 day). These findings indicate that egg cannibalism can act as a population regulating factor.     

Density and environmental effects on shell size in some sand dune snail populations

Populations ol three species of land snail, Helicella itala. Candidula intersecta and Cochlicella acuta are studied on sand dunes on Coll in the Inner Hebrides.
Population density and mean shell size were estimated on 18 sample sites. The sites were ranked for six environmental factors. Environmental factors which are related to the dune vegetational succession account for much of the variation in the densities of C. intersecta and C. acuta . The density of H. itala , however, shows no strong association with these factors. Both snail density and shell size are reactively independent of total soil calcium levels. For each of the three species, mean shell size is negatively associated with population density; the environmental factors account for little of the variation in shell size.
Four possible mechanisms are suggested to account for the variation of shell size with density. It is argued that a direct influence of density on shell size, possibly mediated by mucus conditioning of the environment is the most probable mechanism. There is some evidence to suggest an inter-specific effect whereby C. intersecta density affects H. itala shell size.     

Density and environmental effects on shell size in some sand dune snail populations

Populations ol three species of land snail, Helicella itala. Candidula intersecta and Cochlicella acuta are studied on sand dunes on Coll in the Inner Hebrides. Population density and mean shell size were estimated on 18 sample sites. The sites were ranked for six environmental factors. Environmental factors which are related to the dune vegetational succession account for much of the variation in the densities of C. intersecta and C. acuta. The density of H. itala, however, shows no strong association with these factors. Both snail density and shell size are reactively independent of total soil calcium levels. For each of the three species, mean shell size is negatively associated with population density; the environmental factors account for little of the variation in shell size. Four possible mechanisms are suggested to account for the variation of shell size with density. It is argued that a direct influence of density on shell size, possibly mediated by mucus conditioning of the environment is the most probable mechanism. There is some evidence to suggest an inter-specific effect whereby C. intersecta density affects H. itala shell size.     

Adaptive responses and invasion: the role of plasticity and evolution in snail shell morphology

Invasive species often exhibit either evolved or plastic adaptations in response to spatially varying environmental conditions. We investigated whether evolved or plastic adaptation was driving variation in shell morphology among invasive populations of the New Zealand mud snail (Potamopyrgus antipodarum) in the western United States. We found that invasive populations exhibit considerable shell shape variation and inhabit a variety of flow velocity habitats. We investigated the importance of evolution and plasticity by examining variation in shell morphological traits 1) between the parental and F₁ generations for each population and 2) among populations of the first lab generation (F₁) in a common garden, full‐sib design using Canonical Variate Analyses (CVA). We compared the F₁ generation to the parental lineages and found significant differences in overall shell shape indicating a plastic response. However, when examining differences among the F₁ populations, we found that they maintained among‐population shell shape differences, indicating a genetic response. The F₁ generation exhibited a smaller shell morph more suited to the low‐flow common garden environment within a single generation. Our results suggest that phenotypic plasticity in conjunction with evolution may be driving variation in shell morphology of this widespread invasive snail.     

Polygyrid land snail phylogeny: external sperm exchange, early North American biogeography, iterative shell evolution

A fresh assessment of reproductive behaviour and anatomy, combined with new allozyme data, results in a phylogenetic hypothesis for the 23 polygyrid genera that differs substantially from previous attempts. Analyses based on this phylogeny suggest that (a) evolutionary transition from internal to external sperm exchange occurred only once in polygyrids, with unusual functional intermediates still extant; (b) polygyrid biogeographic history paralleled that of plethodontid salamanders, with hypothesized vicariance events at about 145, 120, 65 and 40 MA, and, dispersal events at about 55 and 40 MA; and (c) iterative shell evolution occurred despite a phylogenetic constraint on ontogenetic whorl-expansion rate.     

Regeneration of excised shell by the invasive apple snail Pomacea canaliculata

After excision of shell from Pomacea canaliculata, survival rates, repair process, haemocyte response, and calcium content were observed and quantified for three weeks. The following experimental treatments were used: black-colored individuals with large (BL-group) or small (BS-group) shell heights, yellow-colored individuals with large (YL-group) or small (YS-group) shell heights. The survival rates in BL-, YL-, BS-, and YS-groups after making partial excisions of snail shells were, respectively, 93.59, 94.87, 92.31, and 96.15%. Freshly regenerated shell could be seen at 1 day among the four groups after induction of shell regeneration. Regeneration shell filled the wound after 5–10 days. The area of regenerated shell in yellow-colored individuals was larger than that of black-colored individuals, but the thickness of regenerated shell was not significantly different. Multiple (four times) inductions of shell regeneration significantly enhanced the thickness of newly formed shells. Two types of circulating haemocytes (hyalinocytes and granulocytes) were identified under light microscopy based on Wright’s staining. Artificial induction of shell regeneration rendered a significant increase in the number of total circulating haemocytes which was followed by a decrease to pre-wounding levels. This peak value of total circulating haemocytes was 2.86, 2.43, 1.69, and 1.71-fold higher than pre-wounding levels in group BL, YL, BS, and YS, respectively. The number of total circulating haemocytes in yellow-colored individuals was significantly higher than in black-colored. Calcium concentrations in the mantles of snails are favorable for calcium nucleation. It is postulated that calcium content increased in the mantle increase at the start of the experiment and then returned to pre-wounding levels at the end of the experiment. This study shows that apple snails have effective shell regeneration abilities and that haemocytes and calcium transportation appear to play an important role in shell growth and regeneration.     

Variation in multiple paternity and sperm utilization patterns in natural populations of a simultaneous hermaphrodite land snail

Mating frequency has important implications for patterns of sexual selection and sexual conflict, and hence for issues such as the maintenance of genetic diversity and speciation. We assessed the level of multiple paternity and sperm utilization patterns in four natural populations of the simultaneous hermaphrodite land snail Arianta arbustorum using four polymorphic microsatellite loci. A total of 1088 offspring from 26 wild‐caught snails were genotyped to determine the number of fathers siring each brood and paternity skew in succeeding clutches. Multiple paternity was detected in the offspring of all 26 mother snails examined with the contribution of two to six fathers. The four populations examined differed in the level of multiple paternity. Snails in the population with the highest density of adults showed the highest level of multiple paternity, whereas snails in the population with the lowest density exhibited the lowest value of multiple paternity. Highly skewed paternity patterns were found in the progeny of 15 (57.7%) of the 26 mother snails. The number and identity of fathers siring the offspring of single mothers also varied among successive clutches. Furthermore, genetic analyses indicate a low level of self‐fertilization in one of the four populations. © 2010 The Linnean Society of London, Biological Journal of the Linnean Society, 2010, 99 , 350–361.     

Microgeographical variation in shell strength in the flat periwinkles Littorina obtusata and Littorina mariae

The strength of molluscan shells has been shown to vary in adaptive ways in a number of species and one of the main factors thought to be involved is shell-crushing by predators. A recent study found that the sibling species of flat periwinkle Littorina obtusata and Littorina mariae showed significant differences in the rates at which shell strength increased with shell length in specimens which had been collected from the same location, where the species were sympatric. This paper describes differences between the shells of the two species from a number of localities around Milford Haven in Dyfed, Wales, and local geographical variation in the shells. Littorina mariae, which is normally found at lower tidal levels than L. obtusata, matures at a smaller shell length. Both species reinforce the shell as they grow since shell strength, determined as the maximum force applied by a hydraulic tensile testing machine before the shell cracked, is strongly positively allometric; it increases at a rate close to the cube of shell length whilst isometric growth would result in strength increasing in proportion to the square of shell length. Because L. mariae matures earlier and reinforces the shell at a smaller size, the mature shell of L. mariae is substantially stronger on average than that of a similar sized but immature L. obtusata. At maturity the shell strengths of the two species are not very different despite the substantial difference in mean shell length. Strength varies significantly from shore to shore, and with the level of the shore from which the animals were collected. Strength increases down the shore in both species. Shell strength decreases with exposure to wave action in L. mariae but increases with exposure in L. obtusata; there is also substantial shore-to-shore variation which is not explained by exposure. Path analysis was used to explore the relationship between shell strength and other measured shell parameters (mass, length, height, thickness). The best predictor of shell strength in both species is a parameter which is heavily positively loaded on LN (shell mass) and strongly offset by negative loadings on LN (shell length) and LN (shell height). This is logical because for a given shell length a heavier shell will be thicker and stronger, whilst for a given shell mass a bigger shell will be thinner and therefore weaker. Such differential variation of shell mass and shell length explains most of the geographical variation observed in shell strength; shells are stronger in snails collected from one place than from another because, for the same shell length they are heavier or, to put it the other way, because at the same shell mass, they are smaller.     

Latitudinal and altitudinal body size variation among north-west European land snail species

Aim To investigate latitudinal and altitudinal patterns in body size variation among north‐west European land snail species, as well as factors influencing such patterns. Location Latitudinal patterns were investigated in north‐west Europe from the British Isles and France in the west to Finland, Poland, Slovakia and Hungary in the east and from Norway in the north to France (with the exception of the Mediterranean part of the country), Switzerland, Austria and Hungary in the south. Altitudinal patterns were examined in the Alps in Austria and Switzerland. Methods Data on latitudinal ranges of 366 north‐west European land snail species, on altitudinal ranges of 175 species from Austria and 150 species from Switzerland, and on their body sizes were used to test for the presence of interspecific latitudinal or altitudinal body size patterns. Four methods, Stevens’ method, the midpoint method, the across‐species method, and a phylogenetically controlled analysis (CAIC) were applied. Results As a result of the predominance of some small bodied clades at higher latitudes and some large bodied clades at lower latitudes, body size of north‐west European land snails decreases significantly with increasing latitude. However, little of the body size variation across species is explained by latitude and the phylogenetically controlled analysis showed that the decrease of body size with increasing latitude is not a result of repeated and independent evolution of an association between body size and latitude in many clades. There is no significant correlation between body size of land snail species and altitude in the Alps although a negative correlation of body size and altitude is frequent within species. Main conclusion If phylogenetic effects are controlled for, neither latitudinal nor altitudinal patterns in body size could be found. Bergmann's rule, which predicts a positive correlation between body size of species and latitude, could not be confirmed for north‐west European land snails.     

Macroevolutionary pattern of sexual size dimorphism in geckos corresponds to intraspecific temperature‐induced variation

Many animal lineages exhibit allometry in sexual size dimorphism (SSD), known as ‘Rensch’s rule’. When applied to the interspecific level, this rule states that males are more evolutionary plastic in body size than females and that male‐biased SSD increases with body size. One of the explanations for the occurrence of Rensch’s rule is the differential‐plasticity hypothesis assuming that higher evolutionary plasticity in males is a consequence of larger sensitivity of male growth to environmental cues. We have confirmed the pattern consistent with Rensch’s rule among species of the gecko genus Paroedura and followed the ontogeny of SSD at three constant temperatures in a male‐larger species (Paroedura picta). In this species, males exhibited larger temperature‐induced phenotypic plasticity in final body size than females, and body size and SSD correlated across temperatures. This result supports the differential‐plasticity hypothesis and points to the role phenotypic plasticity plays in generating of evolutionary novelties.     

Population density, body size, and phenotypic plasticity of brood size in a burying beetle

Theory predicts that organisms living in heterogeneous environmentswill exhibit phenotypic plasticity. One trait that may be particularlyimportant in this context is the clutch or brood size becauseit is simultaneously a maternal and offspring characteristic.In this paper, I test the hypothesis that the burying beetle,Nicrophorus orbicollis, adjusts brood size, in part, in anticipationof the reproductive environment of its adult offspring. N. orbicollisuse a small vertebrate carcass as a food resource for theiryoung. Both parents provide parental care and actively regulatebrood size through filial cannibalism. The result is a positivecorrelation between brood size and carcass size. Adult bodysize is an important determinant of reproductive success forboth sexes, but only at higher population densities. I testthree predictions generated by the hypothesis that beetles adjustbrood size in response to population density. First, averageadult body size should vary positively with population density.Second, brood size on a given-sized carcass should be larger(producing more but smaller young) in low-density populationsthan in high-density populations. Third, females should respondadaptively to changes in local population density by producinglarger broods when population density is low and small broodswhen population density is high. All three predictions weresupported using a combination of field and laboratory experiments.These results (1) show that brood size is a phenotypically plastictrait and (2) support the idea that brood size decisions arean intergenerational phenomenon that varies with the anticipatedcompetitive environment of the offspring.     

Dynamic gastropods: stable shell polymorphism despite gene flow in the land snail Arianta arbustorum

Based on a sequence fragment of mtDNA we analysed the genetic differentiation of the land snail Arianta arbustorum in the Alpine massif Gesäuse, where four morphotypes occur in close vicinity. We aimed at understanding actual and historical evolutionary processes among the morphotypes. With 135 haplotypes diverging up to 12.5%, genetic diversity was extremely high. Phylogenetic analyses indicated that A. arbustorum has colonized the Gesäuse several times in consecutive waves and that introgression of neutral markers across morphotypes and selection on shell shape have played an important role. Gene flow was more likely between locally close, different morphotypes than among distant populations of the same morphotype again indicating that cohesion of morphotypes was mainly due to selection. The actual distribution of morphotypes and haplotypes as well as the high genetic diversity also suggested that the area provided refugia during the Pleistocene glaciations.