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     

Ultrastructure of snail grazing damage to calcicolous lichens

The identification of damaged lichens is often difficult due to changes in the morphology of regenerating specimens. We examined the Ultrastructure of grazing damages to four species of calcicolous lichens ( Aspicilia calcarea, Physcia adscendens, Tephromela atra and Xanthoria parietina ) and free-living cyanobacteria (family Chroococcaceae) caused by individuals of four species of land snails ( Chondrina clienta, Balea perversa, Clausilia bidentata and Helicigona lapicida ). We also investigated the radular structure of the four lichen-feeding snails to examine whether differences in radular morphology result in species-specific grazing damages. Individuals of all four snail species removed the cyanobacteria layer covering the limestone or lichen surfaces. The four lichen species were grazed to a different extent by the different snail species. SEM-images showed that B. perversa left distinct depressions on the thalli of A. calcarea , whereas H. lapicida grazed off the thalli of this lichen rather evenly. Both snail species left visible radular traces on the lichen surface. In contrast, Ch. clienta left shallow depressions without radular traces on the thalli of A. calcarea. In Tephromela atra , grazing damages were observed on both thallus and ascocarp. Ascocarps of T. atra were partly grazed by B. perversa. Helicigona lapicida grazing on T. atra removed more or less evenly the entire lichen tissue including the ascocarps. In foliose lichens, grazing by Ch. clienta, B. perversa and Cl. bidentata resulted in depressions of different depths, while H. lapicida removed entire pieces of the thalli. In general, radular traces were less distinct in foliose lichens than in crustose lichens.     

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.     

Microgeographical variation in shell size of the land snail Chondrina clienta

Microgeographical variation in shell morphology of the rock-dwelling land snail Chondrina clienta , collected from 30 sites within an area of 0.5 km² on the island of Öland, Sweden, was examined in relation to its own population density and that of a potential competitor (the land snail Balea perversa ) and to environmental factors. Dispersal of marked individuals averaged 83 cm per year within a stone pile and 291 cm in an area of exposed bedrock. Local population density of active C. clienta ranged from 5 to 794 individuals per m². Shell characters were highly intercorrelated, both within and between populations. Principal component analysis revealed that most of the interpopulational variation could be expressed by the single character of shell height, which ranged from 5.54 to 6.94 mm. In all populations, snails of a given size had the same whorl number. Shell size was not influenced by habitat type (exposed rock surface, stone pile or stone wall) or proportion of calcareous stones within habitat. It was, however, negatively correlated with local population density, indicating intraspecific competition, and positively correlated with the degree of plant cover within the habitat. Analysis of variance revealed additionally a density effect of B. perversa on shell size in C. clienta , probably as a result of interspecific competition. Breeding experiments using C. clienta from different sites and carried out under unifrom conditions caused most of the phenotypic variation to disappear, demonstrating the high phenotypic plasticity of the species.     

Differential herbivore damage to calcicolous lichens by snails

Abstract: A laboratory experiment was conducted to determine the damage caused by snail grazing to 35 calcicolous lichen species and cyanobacteria from the Baltic island of Öland, Sweden. Individuals of four species of land snails (Balea perversa, Chondrina clienta, Clausilia bidentata and Helicigona lapicida) were allowed to graze on identified lichen species growing on pieces of limestone. Snail grazing on thalli and ascocarps was classified into four categories ranging from no damage to completely eaten. In general, some lichen species were heavily grazed, whereas others were only slightly damaged or not injured. Aspicilia calcarea, Tephromela atra and Verrucaria nigrescens were preferred by all snail species except Chondrina clienta. In general, lichen thalli that were not immersed in the calcareous rock and cyanobacteria were preferred, whereas ascocarps were avoided by one of the snail species (C. clienta). Immersed perithecia with a carbonized outer layer were avoided by all snail species except C. bidentata. Possibilities of chemical and mechanical defence properties in calcicolous lichens are discussed.     

Herbivory on calcicolous lichens: different food preferences and growth rates in two co-existing land snails

A total of 32 calcicolous lichen species, one alga and one bryophyte were recorded on a limestone wall in the grassland Great Alvar on the Baltic island of Öland, Sweden. Fourteen (41%) of these 34 species and free-living cyanobacteria showed herbivore damage, most probably due to grazing by the land snails Chondrina clienta and Balea perversa which inhabited the limestone wall. Three laboratory experiments were conducted to examine the food preferences of individuals of C. clienta and B. perversa collected at this site and to evaluate any association between their preference and the net food quality of the lichens to the snails. Chondrina clienta and B. perversa exhibited food preferences, which differed significantly between species. Within species, variation in food choice was similar among individuals. This indicates that snail populations may be composed of polyphagous individuals with similar food preferences. Different lichen species were of different net food quality to the snails as indicated by growth rate differences. In both snail species the most preferred lichen species of the choice experiment caused the largest weight increase in juveniles, viz. Caloplaca flavovirescens for C. clienta and Aspicilia calcarea for B. perversa. This suggest that the snail species studied differ in their abilities to deal with secondary compounds and physical characteristics of certain lichens or that they can utilize the energy and nutrients of these lichens to a different extent. It is suggested that differential food preferences might reduce the intensity of interspecific competition for resources (lichens) between C. clienta and B. perversa.     

Trade-off between mating and predation risk in the marine snail, Littorina plena

Abstract. We investigated the hypothesis that predation risk affects mating decisions in the intertidal snail Littorina plena in Bamfield Inlet, Northeast Pacific. First, we conducted a field tethering experiment to test the assumption that mating pairs of snails are more susceptible to predation than solitary individuals, and then performed a laboratory experiment to quantify the effect of predation threat on the propensity of snails to form mating pairs. Our results support the hypothesis, in that "mating pairs" were more frequently killed than single snails in the field, and snails were less likely to form mating pairs in the laboratory when simulated predation risk was high (chemical cues from crushed conspecifics were added to the water) than when it was low (no risk cues were added to the water). In contrast to several earlier studies, we found no effect of individual size on snail susceptibility to predation, perhaps because our two size classes were contiguous and snails within them were not dissimilar enough. The results of the behavioral experiment were consistent with this lack of individual size effect on snail vulnerability; both size classes of snails showed a significant and similar tendency to decrease mating when predation risk was high. Taken together, the results of this and recent studies indicate that predators can considerably affect the behavior of littorinid snails, including their movement patterns, feeding, and reproduction. We argue that greater consideration should be given to how marine invertebrates trade off predation risk and activities related to reproduction.     

Pathogens as a factor limiting the spread of cannibalism in tiger salamanders

Summary Intraspecific predation is taxonomically widespread, but few species routinely prey on conspecifics. This is surprising as conspecifics could be a valuable resource for animals limited by food. A potential cost of cannibalism that has been largely unexplored is that it may enhance the risk of acquiring debilitating pathogens or toxins from conspecifics. We examined how pathogens affect variation in the incidence of cannibalism in tiger salamander larvae (Ambystoma tigrinum nebulosum), which occur as two environmentally-induced morphs, typicals and cannibals. Salamanders from one population were more likely than those in another to develop into cannibals, even when reared under identical conditions. Variation in the propensity to become a cannibal may be caused by variation in pathogen density. In the population with cannibals at low frequency, bacterial blooms in late summer correlated with massive die-offs of salamanders. The frequency of cannibals correlated significantly negatively with bacterial density in ten different natural lakes. In the laboratory, cannibals exposed to a diseased conspecific always preyed on the sick animal. As a result, cannibals wre more likely to acquire and die from disease than were typicals that were similarly exposed, or cannibals that were exposed to healthy conspecifics. Since conspecifics often share lethal pathogens, enhanced risk of disease may explain why cannibalism is generally infrequent. Pathogens may constrain not only the tendency to be behaviorally cannibalistic, but also the propensity to develop specialized cannibal morphologies.     

The role of intraspecific competition in the dispersal of an invasive fish

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Effects of body size and resource availability on dispersal in a native and a non-native estuarine snail

I manipulated snail densities of two coexisting species of salt marsh snail, Cerithidea californica Haldeman (native) and Batillaria attramentaria Sowerby (non-indigenous) to investigate how resource levels set by intraspecific competition may influence dispersal rates. I used two distinct size classes of the snails (mature and immature) to determine if the effects of competition on dispersal differed between developmental stages of a consumer. Dispersal attempts were measured within enclosure pens by counting snails climbing the sides of the enclosure. The influence of snail density per se and resource levels (which were set by snail densities) on dispersal rates were separated by comparing responses of snails to density before and after resources became depleted. For large snails of both species, dispersal increased as resource levels decreased, supporting the hypothesis that competition influences dispersal rates. Small snails of both species, in contrast, always dispersed at relatively higher rates than larger individuals, but were not influenced by variation in resource levels. This result corroborates other studies that have shown reduced competition in these species at smaller size, and suggests that another mechanism, such as genetically coded behavior to disperse when young, influences their behavior. Previous experiments demonstrated Batillaria's superior resource conversion efficiency; therefore, I had hypothesized that for any given resource level, Cerithidea would disperse more, because it was more affected by resource availability. Adult Batillaria, however, responded more sensitively to resource levels (i.e., dispersed more at any given resource level) than Cerithidea. This counter-intuitive result illustrates the potential importance of genetic limitations on behavioral responses available to a species. Constraints on behavioral responses may have been accentuated since Batillaria is a non-indigenous species whose evolved behavioral responses are not necessarily adapted to its present, non-native environment.     

Field observations of the alarm response to crushed conspecifics in the freshwater snail Pomacea canaliculata: effects of habitat,vegetation, and body size

The freshwater snail Pomacea canaliculata shows alarm responses to chemical cues released from injured conspecifics, but its behavioural responses in the field are unknown. We investigated effects of habitat (canals or paddy fields), vegetation, and body size on alarm responses in the field. Snails responded to crushed conspecifics within 4 min by burying themselves, but the proportions of self-buried snails were generally lower (0–28% depending on experiments) than those reported in the laboratory. Snails not only showed the alarm response, but also frequently fed on crushed conspecifics. There were no influences of habitat or body size on the proportions of individuals showing the alarm response. Nevertheless, in paddy fields with high-density vegetation a higher proportion of snails showed the alarm response than in paddy fields with low-density vegetation.     

Cannibalism in Arctic charr: do all individuals have the same propensity to be cannibals?

By comparing large Arctic charr Salvelinus alpinus , which had shown a persistent cannibalistic response varying from zero to very high in succeeding laboratory trials, with their individual cannibalistic behaviour after release into a natural lake inhabited by small Arctic charr, it was found that all Arctic charr had the potential to become cannibalistic, irrespective of their laboratory behaviour. More specifically, Arctic charr that never fed on prey fishes when offered them in the tank experiments turned to cannibalism when released in the lake, highlighting the potential difficulties in extrapolating laboratory results to natural settings. This was also true for naive fish that had no prior experience of eating live food. Since no significant increase in the number of prey consumed during each of the succeeding laboratory trials was found, and naive fish showed a response under natural conditions similar to that of their counterparts, the training of the Arctic charr (or experience or learning) probably had no effect upon the piscivorous or cannibalistic response after stocking. Thus, the study appeared to demonstrate that most variations in cannibalism in Arctic charr was simply a function of environmental conditions, depending on the density of conspecifics v . alternative prey, and the relative size difference between predator and prey, rather than any genetic influence.     

Beetle and plant density as cues initiating dispersal in two species of adult predaceous diving beetles

Dispersal can influence population dynamics, species distributions, and community assembly, but few studies have attempted to determine the factors that affect dispersal of insects in natural populations. Consequently, little is known about how proximate factors affect the dispersal behavior of individuals or populations, or how an organism’s behavior may change in light of such factors. Adult predaceous diving beetles are active dispersers and are important predators in isolated aquatic habitats. We conducted interrelated studies to determine how several factors affected dispersal in two common pond-inhabiting species in southern Alberta, Canada: Graphoderus occidentalis and Rhantus sericans. Specifically, we (1) experimentally tested the effect of plant and beetle densities on dispersal probabilities in ponds; (2) surveyed ponds and determined the relationships among beetle densities and plant densities and water depth; and (3) conducted laboratory trials to determine how beetle behavior changed in response to variation in plant densities, conspecific densities, food, and water depth. Our field experiment determined that both species exhibited density dependence, with higher beetle densities leading to higher dispersal probabilities. Low plant density also appeared to increase beetle dispersal. Consistent with our experimental results, densities of R. sericans in ponds were significantly related to plant density and varied also with water depth; G. occidentalis densities did not vary with either factor. In the laboratory, behavior varied with plant density only for R. sericans, which swam at low density but were sedentary at high density. Both species responded to depth, with high beetle densities eliciting beetles to spend more time in deeper water. The presence of food caused opposite responses for G. occidentalis between experiments. Behavioral changes in response to patch-level heterogeneity likely influence dispersal in natural populations and are expected to be important for observed patterns of individuals in nature. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Density-dependence across dispersal stages in a hermaphrodite land snail: insights from discrete choice models

Dispersal movements, i.e. movements leading to gene flow, are key behaviours with important, but only partially understood, consequences for the dynamics and evolution of populations. In particular, density-dependent dispersal has been widely described, yet how it is determined by the interaction with individual traits, and whether density effects differ between the three steps of dispersal (departure, transience, and settlement), remains largely unknown. Using a semi-natural landscape, we studied dispersal choices of Cornu aspersum land snails, a species in which negative effects of crowding are well documented, and analysed them using dispersal discrete choice models, a new method allowing the analysis of dispersal decisions by explicitly considering the characteristics of all available alternatives and their interaction with individual traits. Subadults were more dispersive than adults, confirming existing results. In addition, departure and settlement were both density dependent: snails avoided crowded patches at both ends of the dispersal process, and subadults were more reluctant to settle into crowded patches than adults. Moreover, we found support for carry-over effects of release density on subsequent settlement decisions: snails from crowded contexts were more sensitive to density in their subsequent immigration choices. The fact that settlement decisions were informed indicates that costs of prospecting are not as important as previously thought in snails, and/or that snails use alternative ways to collect information, such as indirect social information (e.g. trail following). The observed density-dependent dispersal dynamics may play an important role in the ability of C. aspersum to successfully colonise frequently human-disturbed habitats around the world.     

Effects of rearing density, age, sex, and food deprivation on flight initiation of the red flour beetle (Coleoptera: Tenebrionidae)

Effects of rearing density, adult density and sex ratio in the flight chamber, adult age, sex, presence or absence of food, and duration of food deprivation on rate of and time to flight initiation of the red flour beetle, Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae), were studied in the laboratory. Rates of flight initiation were slightly lower at lower rearing densities, but they did not differ with age or sex of adults, presence or absence of food in the flight chamber, or duration of food deprivation. Focal adults were less likely to fly when individuals of the opposite sex were present in flight chambers. Presence of the same sex or mixed sexes and numbers of individuals in flight chambers did not affect tendency to fly of focal individuals. Mean time to flight of older beetles (7-20 d old) was less than that of younger beetles (1-4 d old). No young beetles flew during the first 24 h of flight tests. Mean time to flight did not differ with rearing density, sex, presence or absence of food, or duration of food deprivation. Our results indicate that this species is highly dispersive during the adult stage and that flight does not seem to be associated with just prereproductive or postreproductive dispersal phases, high rearing density, or short to medium periods of food deprivation. Therefore, T. castaneum level of flight activity does not seem to be associated with factors that have been shown in the literature to increase dispersal by walking for this species and to increase flight initiation in other stored-product species.     

Effects of Familiarity and Population Density on Competitive Interactions and Growth: An Experimental Study on a Territorial Salmonid Fish

The deleterious effect of competition for space and food in animals increases with increasing population density. In contrast, familiarity towards conspecifics can relax the intensity of interference competition. Here, we hypothesized that familiarity towards conspecifics mitigates the effect of density‐dependent growth and dispersal behaviour in territorial animals. To test this, wild‐captured juvenile brown trout were subjected to two consecutive laboratory experiments. First, growth and fin erosion were measured for 40 d in a 2 × 2 factorial design manipulating density and familiarity. The density was manipulated via size of experimental tanks, while per capita food abundance and fish number was constant. All fish were subsequently exposed to an emergence test, giving them the option to leave their group and disperse to a novel unoccupied environment. The results show that familiarity increases growth and decreases the level of fin erosion (i.e. proxy of intensity of aggressive interactions). We found no significant effect of population density on growth rate. However, there was a tendency towards higher fin erosion in fish kept under high density. The growth of individuals was also affected by their size rank within the group, with the largest individuals in each group growing disproportionally faster than the rest of the group, probably due to their high social rank. However, the second and third fish in the size rank did not grow significantly faster and tended to suffer higher mortality than the rest of the group. During the emergence test, the largest individuals in the familiar groups left the shelter either as the first (six of 12 groups) or last (five of 12 groups) individual in the group, while no such pattern was observed in unfamiliar groups. Our results suggest that individuals in familiar groups receive less aggression and stress (i.e. fin damage) and grow faster than fish in unfamiliar groups. The mechanisms indicated in this laboratory study may be especially important in highly fecund organisms like fish which undergo density‐dependent bottlenecks during early life.     

High-quality habitat and facilitation ameliorate competitive effects of prior residents on new settlers

Many species disperse during their lifetime. Two factors that can affect the performance of individuals following dispersal are the presence of conspecifics and intrinsic habitat quality at the settlement site. Detecting the influence of these factors can be difficult for at least two reasons: (1) the outcomes of interactions with conspecifics are often variable including both competition and facilitation, and (2) selection of high quality habitats often leads to positive covariance between habitat quality and density. In this study, I investigate positive and negative effects of resident blue streak cleaner wrasse (Labroides dimidiatus) on the growth and survival of recently settled conspecifics while accounting for habitat quality. Juvenile L. dimidiatus settle near adult conspecifics, but likely have to compete with resident adults for access to food. However, field experiments indicate that settlers have access to more resources at occupied sites, and as a result, grow faster despite evidence for competition with residents. This result is a direct consequence of two factors: (1) resident conspecifics facilitate settlers by attracting client fish, and (2) resident conspecifics are strongly associated with high quality habitat. These results highlight the need to simultaneously consider habitat quality and competitive and facilitative interactions between conspecifics when making inferences about ecological processes from spatial patterns of individual performance.     

Modelling density-dependent fish shoal distributions in the laboratory and field

Density-dependent variables have long been established as an important area of ecological research, but the effects of the local density of conspecifics on grouping behaviour are less well-studied. We compared the influence of the density of conspecifics on the shoal size distribution of killifish, Fundulus diaphanus , in the laboratory and the field. In both environments we observed an increase in shoal size and shoal number with the density of individuals present. The increase in shoal size was markedly steeper in the field than in the laboratory, but direct comparison of the two was complicated by the fact that the absolute numbers of fish present at the field site were considerably higher than those used in the laboratory trials. We developed an individual-based model that was first used as a null model of shoal formation (defined by proximity to others) in fish with no shoaling tendency over the same range of densities used in the laboratory. Group size increased much more rapidly with increasing density in the laboratory than predicted by the null model. When we incorporated shoaling behaviour into our model, the laboratory results could be reproduced with high accuracy. However, when extrapolated to match conditions in the field, the model predicted smaller, more numerous shoals than were actually observed. We suggest this is due to heterogeneity of the field environment because fish were found to be highly aggregated in certain areas of our field site. The predictive power of laboratory studies for the field is discussed with regards to using individual-based modelling as a tool for deriving such predictions.     

To eat or not to eat—relationship of lichen herbivory by snails with secondary compounds and field frequency of lichens

Aims The biochemical defense of lichens against herbivores and its relationship to lichen frequency are poorly understood. Therefore, we tested whether chemical compounds in lichens act as feeding defense or rather as stimulus for snail herbivory among lichens and whether experimental feeding by snails is related to lichen frequency in the field.Methods In a no-choice feeding experiment, we fed 24 lichen species to snails of two taxa from the Clausilidae and Enidae families and compared untreated lichens and lichens with compounds removed by acetone rinsing. Then, we related experimental lichen consumption with the frequency of lichen species among 158 forest plots in the field (Schw?bische Alb, Germany), where we had also sampled snail and lichen species.Important findings In five lichen species, snails preferred treated samples over untreated controls, indicating chemical feeding defense, and vice versa in two species, indicating chemical feeding stimulus. Interestingly, compared with less frequent lichen species, snails consumed more of untreated and less of treated samples of more frequent lichen species. Removing one outlier species resulted in the loss of a significant positive relationship when untreated samples were analyzed separately. However, the interaction between treatment and lichen frequency remained significant when excluding single species or including snail genus instead of taxa, indicating that our results were robust and that lumping the species to two taxa was justified. Our results imply lichen-feeding snails to prefer frequent lichens and avoid less frequent ones because of secondary compound recognition. This supports the idea that consumers adapt to the most abundant food source.     

Costs of cannibalism in the presence of an iridovirus pathogen of Spodoptera frugiperda

Abstract.  1. The costs of cannibalism were examined in larvae of Spodoptera frugiperda (J. E. Smith) (Lepidoptera: Noctuidae) in the presence of conspecifics infected by a lethal invertebrate iridescent virus (IIV). The hypothesis of a positive correlation between insect density and the likelihood of disease transmission by cannibalism was examined in laboratory microcosms and a field experiment.
2. Transmission was negligible following peroral infection of early instars with purified virus suspensions or following coprophagy of virus-contaminated faeces excreted by infected insects. In contrast, 92% of the insects that predated infected conspecifics acquired the infection and died prior to adult emergence in the laboratory. Diseased larvae were more likely to be victims of cannibalism than healthy larvae.
3. The prevalence of cannibalism was density dependent in laboratory microcosms with a low density (10 healthy insects + one infected insect) or high density (30 healthy insects + one infected insect) of insects, and field experiments performed on maize plants infested with one or four healthy insects + one infected insect.
4. Cannibalism in the presence of virus-infected conspecifics was highly costly to S. frugiperda ; in all cases, insect survival was reduced by between ≈ 50% (laboratory) and ≈ 30% (field) in the presence of the pathogen. Contrary to expectations, the prevalence of disease was not sensitive to density because cannibalism resulted in self-thinning. As infected individuals are consumed and disappear from the population, the prevalence of disease will be determined by the timescale over which transmission can be achieved, and the rate at which individuals that have acquired an infection become themselves infectious to conspecific predators.