Application of harmonic radar technology to monitor tree snail dispersal

Abstract. Planned conservation efforts for tree snails of the endangered genus Achatinella, endemic to the island of O'ahu, Hawai'i, will include translocations among the remaining wild and captive‐bred populations. In order to establish optimal levels of artificial migration among neighboring groups of snails within fragmented populations, efforts to determine natural dispersal rates through direct observation were initiated. Capture–mark–recapture (CMR) efforts have proved inadequate for obtaining the requisite dispersal estimates, due to low recapture probabilities. In addition, snail dispersal beyond the boundaries of a finite CMR study site was indistinguishable from mortality. In the preliminary study reported here, both the low recapture probability and dispersal detection problems of past CMR efforts were addressed by using harmonic radar tracking. This approach yielded rough dispersal estimates that were unattainable using CMR alone by providing 100% recapture rates even beyond the normal survey area boundaries. Extensive snail movements within clusters of connected trees were frequently observed after tracking for merely a few hours, although movements between unconnected trees were rare and recorded only after monthly survey intervals. Just 11 out of 40 tracked snails made between‐tree movements (average distance of 4.94±1.52 m) during the entire 7‐month study, and provided the only data utilizable for inferring gene flow in and out of subpopulations. Meteorological data loggers were deployed when tracking began to look for an association between such snail movement and weather fluctuations. The resultant data indicate that increases in both wind gusts and humidity facilitate dispersal (R²=0.77, p‐value <0.001), and that passive wind dispersal alone may be responsible for many snail movements (R²=0.59, p‐value=0.0014). Despite having provided coarse estimates of short‐term dispersal and corresponding wind influences, the limitations of the radar method can be substantial.     

Extinction,turnover and species diversity in an experimentally fragmented California annual grassland

Summary Many natural populations are subdivided among partially isolated habitat patches, but the influence of habitat patchiness per se on species immigration, extinction, and the resulting patterns of species diversity, has received virtually no experimental study. In an experiment designed to test the effects of habitat subdivision on local community structure, we compare the diversity and annual turnover of flowering plant species in 3 treatments of the same total area, but subdivided to different degrees. We experimentally fragmented a California winter annual grassland into isolated plots, two of 32 m², eight of 8 m², and 32 of 2 m², each treatment representing a combined area of 64 m². Insularization of the experimental habitat fragments is provided by grazing sheep. The effects of plot area on species diversity, extinction, and turnover are consistent with the MacArthur-Wilson model. Species richness increases with the degree of habitat subdivision. Extinction, immigration, and turnover, however, are relatively independent of the degree of subdivision. These experimental results contrast with predictions that habitat subdivision necessarily results in greater rates of extinction accompanied by reduced species diversity.     

The dynamics of species richness in an experimentally restored calcareous grassland

Abstract. This paper concerns the hypothesis that shoot (light) competition is the main interaction determining the community change during succession from a relatively species-poor deciduous forest (an overgrown former grassland) to a species-rich grassland, while root (nutrient) competition is of little importance. In a 4-yr restoration experiment, clearcutting, mowing and root trenching were used as treatments. The results did not reject the hypothesis. However, the significance of separating two kinds of shoot competition - ‘coarse-scale (between different growth forms) and ‘fine-scale’ (between similar growth forms) became evident. Release from the ‘coarse-scale’ shoot competition (between different growth forms) increased species richness at the beginning of the experiment. This change was interpreted as the replacement of one species pool (shade-tolerant herbaceous perennials) by another (light-demanding herbaceous perennials), the second pool containing considerably more species. The importance of ‘fine-scale’ shoot competition increased gradually - the levelling of competition by mowing resulted in a more pronounced increase in species richness during successive years. The elimination of ‘coarse-scale’ root competition seemed to be important to some extent only in combination with another treatment - mowing. Initial colonization of the cleared area by individual species was a stochastic process which had little relevance to life-history traits. True grassland species were able to colonize quickly. On the community scale, the developing community still remained relatively poor in species. In all plots which were cleared but not mown, succession already started to reverse towards woody vegetation in the third year.     

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.     

Population dynamics in a long-lived seabird: I. Impact of breeding activity on survival and breeding probability in unbanded king penguins

Understanding the trade-off between current reproductive effort, future survival and future breeding attempts is crucial for demographic analyses and life history studies. We investigated this trade-off in a population of king penguins (Aptenodytes patagonicus) marked individually with transponders using multistate capture-recapture models. This colonial seabird species has a low annual proportion of non-breeders (13%), despite a breeding cycle which lasts over 1 year. To draw inferences about the consequences of non-breeding, we tested for an effect of reproductive activity on survival and on the probability of subsequent breeding. We found that birds non-breeding in year t show the same survival rate as breeders (two-states analysis: breeding and non-breeding). However, breeders had a lower probability of breeding again the following year. This negative phenotypic correlation suggests the existence of reproductive costs affecting future breeding probability, but it might also be strengthened by late arrival for courtship in year t. A three-state analysis including breeding success revealed that failed breeders in year t have a lower probability to reproduce successfully in year t + 1 than non-breeders in year t, providing some evidence for the existence of reproductive costs. Moreover, successful breeders showed higher survival probability. This positive phenotypic correlation between current reproduction and subsequent survival supports the hypothesis of an heterogeneity in individual quality. Males breeding in year t had a lower probability to breed again in year t + 1 than females, suggesting higher reproductive costs for this sex. Such additional costs might be due to higher male parental investment in the final phase of chick-rearing, which also delays the arrival of males in year t + 1, and decreases their breeding probability. Our study is the first to explore the breeding biology and the demography of penguins without the disturbance of flipper-bands.     

Intraspecific range dynamics and niche evolution in Candidula land snail species

The range dynamics of a species can either be governed by the spatial tracing of the fundamental environmental niche or by adaptation that allows to occupy new niches. Therefore, the investigation of spatial variation in the realized environmental niche is central to the understanding of species range limit dynamics. However, the study of intraspecific niche variation has been neglected in most phylogeographical studies. We studied the spatial distribution of the realized environmental niche in three land snail species of the genus Candidula , integrating phylogeographical methods, morphometrics, and spatial biodiversity informatics . The phylogeographical analyses showed significant range expansions in all species. These expansions were accompanied in Candidula gigaxii by a shift in the realized environmental niche, the species Candidula unifasciata followed its ancestral niche during expansion while the climate changed in the area of origin and Candidula rugosiuscula tracked the ancestral environmental conditions. The significant niche shifts were associated with potentially adaptive changes of shell morphology. We propose our presented approach as a practicable framework to test hypotheses on intraspecific niche evolution. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 90 , 303–317.     

Effective population sizes and migration rates in fragmented populations of an endangered insect (Coenagrion mercuriale: Odonata)

1. Effective population sizes (N(e)) and migration rates (m) are critical evolutionary parameters that impact on population survival and determine the relative influence of selection and genetic drift. While the parameter m is well-studied in animal populations, N(e) remains challenging to measure and consequently is only rarely estimated, particularly in insect taxa. 2. We used demographic and genetic methods to estimate N(e) and m in a fragmented population of the endangered damselfly Coenagrion mercuriale to better understand the contrast between genetic and field estimates of these parameters and also to identify the spatial scale over which populations may become locally adapted. 3. We found a contrast between demographic- and genetic-based estimates of these parameters, with the former apparently providing overestimates of N(e), owing to substantial underestimation of the variance in reproductive success, and the latter overestimating m, because spatial genetic structure is weak. 4. The overall N(e) of sites within the population network at Beaulieu Heath, the largest C. mercuriale site in the UK, was estimated to vary between approximately 60 and 2700. 5. While N(e) was not correlated with either the total numbers of adults (N) or the area of habitat, this parameter was always less than N, because of substantial variance in reproductive success. The ratio N(e)/N varied between 0.006 and 0.42 and was generally larger in smaller populations, possibly representing some 'genetic compensation'. 6. From a simple genetic model and these data on N(e) and m, it seems that populations of C. mercuriale have the potential to respond to localized spatial variation in selection and this would need to be considered for future genetic management of this endangered species.     

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.     

Population dynamics and production of the freshwater snail Chilina gibbosa Sowerby 1841 (Chilinidae,Pulmonata) in a North-Patagonian reservoir

Chilina gibbosa is an endemic snail widely distributed in Patagonia, Argentina. Due to its importance in the benthic fauna and in the diet of some fish in the oligo-mesotrophic reservoir Ezequiel Ramos Mexía (39° 30′ S, 69° 00′ W), special attention has been given to its life cycle, growth patterns and annual production. Samples were taken monthly at five littoral stations between June 1983 and July 1984. Mean abundance and biomass of C. gibbosa were much higher in vegetated stations dominated by Potamogeton berteroanus (Station 1 : 583 ind. m⁻², 5.95 g AFDM m⁻²) or by Nitella clavata (Station 5 : 275 ind. m⁻², 4.18 g AFDM m⁻²) than bare stations with low transparency or stations with other macrophytes. The snails presented a clustered spatial pattern and their abundance was significantly correlated with macrophyte wet biomass only when this was above 250 g m⁻². Analysis of size distributions showed an annual life cycle with a reproductive period in the summer. However, differences in recruitment and growth occurred probably due to differences in water temperature and food availability. Growth was maximum in summer and almost absent during winter. Hence, shell growth data fit a sigmoid curve well, and growth was somewhat higher at Station 1. Annual production at Stations 1 and 5, estimated by the ‘growth increment summation’ method (28.8 g AFDM m⁻² and 14.18 g AFDM m⁻² respectively), was among the highest recorded for pulmonate gastropods, possibly due to a low interspecific competition. The P : B ratio values were within the range for univoltine gastropods (4.84 and 3.39). The high productivity and turnover rate of these snails grant a high availability of food for the abundant molluscivore, the silverside Patagonina hatcheri.     

Small-scale spatial dynamics of plant species in a grassland community over six years

Abstract. In a species-rich mountain grassland in the Krkonosse Mts., Czechoslovakia, data from four permanent plots of 50 cm x 50 cm were recorded annually from 1985 to 1990 to study the spatial dynamics of the species. Plots were divided into 15 x 15 subplots and the number of vegetative units of all plants within each subplot was determined. There was not much net change at the plot level, but the subplots were very dynamic. Two aspects of the spatial dynamics of the species were followed: (1) persistence, i.e. the tendency of the species to remain in the same subplot, and (2) long-distance spreading, i.e. movement to subplots beyond the immediate neighbourhood. Species differed widely in their persistence and longdistance spreading and were classified into mobility types: long-range guerrilla, short-range guerrilla, phalanx and 'sitting’. The mobility types were, to a certain extent, correlated with the growth form of plants, but some species of one growth form showed different types of small-scale dynamics and some species with different growth forms had the same spatial dynamics.     

A land snail's view of a fragmented landscape

Habitat fragmentation may influence the genetic structure of populations, especially of species with low mobility. So far, these effects have been mainly studied by surveying neutral markers, and much less by looking at ecologically relevant characters. Therefore, we aimed to explore eventual patterns of covariation between population structuring in neutral markers and variation in shell morphometrics in the forest‐associated snail Discus rotundatus in relation to habitat fragment characteristics. To this end, we screened shell morphometric variability and sequence variation in a fragment of the mitochondrial 16S rDNA gene in D. rotundatus from the fragmented landscape of the Lower Rhine Embayment, Germany. The 16S rDNA of D. rotundatus was highly variable, with a total of 118 haplotypes (384 individuals) forming four clades and one unresolved group. There was a geographic pattern in the distribution of the clades with the river Rhine apparently separating two groups. Yet, at the geographic scale considered, there was no obvious effect of fragmentation on shell morphometrics and 16S rDNA variation because G_ST often was as high within, as between forests. Instead, the age of the habitat and (re‐)afforestation events appeared to affect shell shape and 16S rDNA in terms of the number of clades per site. The ecologically relevant characters thus supported the presumably neutral mitochondrial DNA markers by indicating that populations of not strictly stenecious species may be (relatively) stable in fragments. However, afforestation after large clearcuts and habitat gain after the amendment of deforestation are accompanied by several, seemingly persistent peculiarities, such as altered genetic composition and shell characters (e.g. aperture size). © 2009 The Linnean Society of London, Biological Journal of the Linnean Society, 2009, 98 , 839–850.     

Multiscale patterns of movement in fragmented landscapes and consequences on demography of the snail kite in Florida

1. Habitat loss and fragmentation are major factors affecting vertebrate populations. A major effect of these habitat alterations is that they reduce movement of organisms. Despite the accepted importance of movement in driving the dynamics of many natural populations, movement of vertebrates in fragmented landscapes have seldom been estimated with robust statistical methods. 2. We estimated movement probabilities of snail kites Rosthramus sociabilis within the remaining wetlands in Florida. Using both radio-telemetry and banding information, we used a multistate modelling approach to estimate transition probabilities at two temporal scales (month; year) and multiple spatial scales. We examined kite movement among wetlands altered by three different levels of fragmentation: among wetlands separated by small physical barriers (e.g. road); among wetlands separated by moderate amount of matrix (< 5 km); and among wetlands separated by extensive matrix areas (> 15 km). 3. Kites moved extensively among contiguous wetlands (movement probability 0.29 per month), but significantly less among isolated wetlands (movement probability 0.10 per month). 4. Kites showed high levels of annual site fidelity to most isolated wetlands (probability ranged from 0.72 to 0.95 per year). 5. We tested the effects of patch size and interpatch distance on movement. Our modelling indicated an effect of both distance and patch size on juveniles' movement (but not adult) when examining movements among fragments. 6. Only a small proportion of kites escaped a regional drought by moving to refugia (wetlands less affected by drought). Many individuals died after the drought. During drought adult survival dropped by 16% while juvenile survival dropped by 86% (possibly because juveniles were less likely to reach refugia). 7. We hypothesize that fragmentation may decrease kite's resistance to drought by restricting exploratory behaviour.     

Heterozygosity-fitness correlations in a bottlenecked island species: a case study on the Seychelles warbler

We used capture-mark-recapture models to investigate the effects of both individual and parental heterozygosity, measured at microsatellite loci on the survival of Seychelles warblers (Acrocephalus sechellensis), an endemic island species which went through a severe population bottleneck in the middle of the last century. We found that an individual's survival was not correlated with multilocus heterozygosity, or with heterozygosity at any specific locus. However, maternal, but not paternal, multilocus heterozygosity was positively associated with offspring survival, but only in years with low survival probabilities. A nestling cross-fostering experiment showed that this was a direct maternal effect as there was an effect of the genetic mother's, but not of the social mother's, heterozygosity. Heterozygosity-fitness correlations at microsatellite markers were generally assumed to reflect genome-wide effects. Although this might be true in partially inbred populations, such correlations may also arise as a result of local effects with specific markers being closely linked to genes which determine fitness. However, heterozygosity at the individual microsatellite loci was not correlated and therefore does not seem to reflect genome-wide heterozygosity. This suggests that even in a small bottlenecked population, heterozygosity-fitness correlations may not be caused by genome-wide effects. Support for the local effects hypothesis was also equivocal; although three specific loci were associated with offspring survival, including all single-locus heterozygosities as independent predictors for the variation in survival was not supported by the data. Furthermore, in contrast to the local effects hypothesis, the loci which contributed most to the heterozygosity-survival relationship were not more polymorphic than the other loci. This study highlights the difficulties in distinguishing between the two hypotheses.     

Characterization of 13 polymorphic microsatellite loci for two land snail species, Trochulus villosus and T. sericeus (Gastropoda: Pulmonata: Hygromiidae)

Thirteen new microsatellite loci were isolated and tested on two land snail species, Trochulus villosus and T. sericeus (Pulmonata: Hygromiidae), resulting in a set of eight polymorphic markers for each species. The expected heterozygosity was high for all loci and species (between 0.616 and 0.944). Such levels of variability will allow detailed insights into the population genetic structure of some Trochulus species.     

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.     

破碎栖息地中物种灭绝机制

栖息地毁坏既会直接降低物种多度,又会间接地降低物种迁移繁殖力,同时还会改变原有的种间平衡.尽管已有研究表明栖息地毁坏是物种灭绝的主要原因之一,但是尚未揭示破碎的栖息地中物种灭绝的驱动机制.通过元胞自动机模拟了物种灭绝对栖息地毁坏空间异质性响应的基础上,进一步研究了栖息地毁坏和种间竞争对物种灭绝的影响.结果发现:强物种的灭绝主要来自栖息地毁坏,而弱物种的灭绝,在随机毁坏下,主要由栖息地毁坏与种间竞争共同决定,而在边缘毁坏下则主要由种间竞争所引起的.栖息地毁坏与种间竞争共同引起的物种灭绝的时间非常短,而栖息地毁坏或种间竞争所引起的物种灭绝时间则较长.