SPATIAL POPULATION STRUCTURE IN THE WHIRLIGIG BEETLE DINEUTUS ASSIMILIS: EVOLUTIONARY INFERENCES BASED ON MITOCHONDRIAL DNA AND FIELD DATA

The spatial population structure of the pond-living water beetle Dineutus assimilis (Coleoptera: Gyrinidae) was investigated through a field study of population dynamics and dispersal, with a concurrent assessment of the spatial distribution of mitochondrial DNA (mtDNA) restriction-fragment-length polymorphism (RFLP). A comprehensive 2-yr survey within a 60-km² study area revealed pronounced fluctuations in local abundances, including extinctions and colonizations. The recapture of marked individuals showed that dispersal among ponds is frequent in both males and females and connects populations on a large geographic scale (maximum observed flight distance: 20 km). The population structure of D. assimilis is thus characterized by both pronounced genetic drift and frequent gene flow. Together, these two forces generate a pattern of very local and transient genetic differentiation. Mitochondrial DNA samples collected within a few kilometers indicate highly significant spatial structure, if newly founded demes or those that experienced recent bottlenecks are included. These results based on four demes within the study area were placed into a regional context by further samples collected at distances of 100 km and 200 km. F_st estimates computed on increasing spatial scales were variable but showed no increasing trend. Thus, gene flow exerts a strong homogenizing force over a wide geographic range but is counteracted locally by genetic drift. These findings highlight the need to supplement estimates of F_st with additional data to arrive at valid interpretations of the genetic information. More generally, this study raises questions about how to capture the relevant features of dynamic, subdivided populations to understand their evolutionary dynamics.     

Zooplankton on the move: first results on the quantification of dispersal of zooplankton in a set of interconnected ponds

In systems of interconnected ponds or lakes, the dispersal of zooplankton may be mediated by the active population component, with rivulets and overflows functioning as dispersal pathways and the dispersal being unidirectional. Such systems offer the possibility to study the impact of dispersal rates on local population dynamics and community structure, and provide opportunities to quantify dispersal in the field in a straightforward manner. In this study, dispersal of active zooplankton populations among interconnected ponds was quantified directly in the field by sampling the small waterways connecting the ponds. The number of dispersing zooplankton sampled in connecting elements was on average high (almost 7000 ind h^–1). However, the contribution of dispersing individuals to total population size in the target ponds was very limited (< 1% 24 h^–1.). Only a weak diel pattern in dispersal rates was observed.     

Relative importance of different dispersal vectors for small aquatic invertebrates in a rock pool metacommunity

The extent and frequency of passive overland dispersal of freshwater invertebrates as well as the relative importance of different dispersal vectors is not well documented. Although anecdotal evidence subscribing the feasibility of individual vectors in various aquatic systems is abundant, dispersal rates have rarely been quantified for different vectors in one study system. Earlier studies also usually investigated dispersal potential rather than actual dispersal rates. In this study we have estimated passive dispersal rates of invertebrate propagules within a cluster of temporary rock pools via water, wind and amphibians in a direct way. Overflows after heavy rains mediated dispersal of a large number of propagules through eroded channels between pools, which were collected in overflow traps. Taking into account model based predictions of overflow frequency, this corresponds with average dispersal rates of 4088 propagules/channel yr^?1. Wind dispersal rates as measured by numbers of propagules collected on sticky traps mounted between pool basins were very high (average dispersal rate: 649 propagules m^?2 in one month) and were positively related to the proximity of source populations. Finally, invertebrate propagules were also isolated from the faeces of African clawed frogs Xenopus laevis caught from the pools (on average 368 propagules/frog). The combination of short distance wind and overflow dispersal rates likely explain the dominant species sorting and mass effect patterns observed in the metacommunity in a previous study. Amphibian mediated dispersal was much less important as the Xenopus laevis population was small and migrations very rare. Based on our own results and available literature we conclude that both vector and propagule properties determine local passive dispersal dynamics of freshwater invertebrates. Accurate knowledge on rates and vectors of dispersal in natural systems are a prerequisite to increase our understanding of the impact of dispersal on ecology (colonisation, community assembly, coexistence) and evolution (gene flow, local adaptation) in fragmented environments.     

Freshwater snail diversity: effects of pond area,habitat heterogeneity and isolation

Summary A large number of eutrophic ponds were surveyed for the presence of freshwater gastropods. Factors thought to influence the distribution of the snails were evaluated. As the investigated area has a homogeneous geological background physicochemical factors probably have a low effect on the local distribution of snails. There was a significant, positive regression between pond area and the number of gastropod species in the pond, but the regression only explained a minor part of the variation in species numbers. Multiple regression of an extended number of variables, associated with habitat complexity and dispersal, indicated that, in addition to area, macrophyte diversity and the mean number of gastropods in the 5 closest ponds (S₅) were important in explaining the distribution of gastropods. An increase in the number of macrophyte species increases the number of available microhabitats and refuges from predators. S₅ probably influences the dispersal rates between ponds. The gastropods in this area are thought to have additional dispersal modes, besides aerial dispersal with birds, and this probably increases the immigration rates and/or decreases the extinction rate.     

The spatio-temporal distribution of adult Ceutorhynchus assimilis in a crop of winter oilseed rape in relation to the distribution of their larvae and that of the parasitoid Trichomalus perfectus

The spatio-temporal distribution of Ceutorhynchus assimilis Payk. (Coleoptera: Curculionidae) adults caught in a rectangular grid of flight traps in a crop of winter oilseed rape (Brassica napus L.) was mapped and was analysed using Spatial Analysis by Distance IndicEs (SADIE). Their distribution was compared to that of their larvae and that of their parasitoid Trichomalus perfectus (Walker) (Hymenoptera: Pteromalidae) in pods. The distribution of immigrating C. assimilis adults was consistent with their arrival at the crop boundaries and movement within the crop towards its centre. Adult C. assimilis were aggregated at all times, invasion being on two fronts, leading to the formation of two major clusters within the crop. Large areas of the crop remained relatively unpopulated. During the emigration phase, numbers declined simultaneously in all parts of the crop. The distributions of adult and larval C. assimilis and of larval T. perfectus were spatially associated. The distribution of the parasitoid did not show a density dependent relationship with that of its host. We discuss the movements of insects which underlie their population distributions, the value of integrating spatial information into improved management strategies for C. assimilis and the potential for the spatial targeting of insecticides to reduce the amount applied and to conserve T. perfectus.     

Time‐restricted flight ability influences dispersal and colonization rates in a group of freshwater beetles

Variation in the ability to fly or not is a key mechanism for differences in local species occurrences. It is increasingly acknowledged that physiological or behavioral mechanisms rather than morphological differences may drive flight abilities. However, our knowledge on the seasonal variability and stressors creating nonmorphological differences in flight abilities and how it scales to local and regional occurrences is very limited particularly for small, short‐lived species such as insects. Here, we examine how flight ability might vary across seasons and between two closely related genera of freshwater beetles with similar geographical ranges, life histories, and dispersal‐related morphology. By combining flight experiments of >1,100 specimens with colonization rates in a metacommunity of 54 ponds in northern and eastern Europe, we have analyzed the relationship between flight ability and spatio‐environmental distribution of the study genera. We find profound differences in flight ability between the two study genera across seasons. High flight ability for Acilius (97% of the tested individuals flew during the experiments) and low for Graphoderus (14%) corresponded to the different colonization rates of newly created ponds. Within a 5‐year period, 81 and 31% of the study ponds were colonized by Acilius and Graphoderus, respectively. While Acilius dispersed throughout the season, flight activity in Graphoderus was restricted to stressed situations immediately after the emergence of adults. Regional colonization ability of Acilius was independent of spatial connectivity and mass effect from propagule sources. In contrast, Graphoderus species were closely related to high connectivity between ponds in the landscape. Our data suggest that different dispersal potential can account for different local occurrences of Acilius and Graphoderus. In general, our findings provide some of the first insights into the understanding of seasonal restrictions in flight patterns of aquatic beetles and their consequences for species distributions.     

The influence of predator regime on the behaviour and mortality of a freshwater amphipod, <Emphasis Type="Italic">Gammarus pulex</Emphasis>

In species with restricted dispersal, traits may become genetically fixed leading to local adaptations. Therefore, predator avoidance in a prey species may differ between populations experiencing different predator regimes, but also between sexes within a population due to different vulnerability to predators. In this study we used male and female Gammarus pulex from two different predator regimes: fishless ponds, where invertebrates are the dominant predators and ponds with predatory fish. In the laboratory we examined refuge use, mortality, leaf decomposition rate and pair-formation in G. pulex when exposed to predator cues from either invertebrate predators or fish. Individuals from fish ponds spent more time in refuge and had a higher mortality than those from fishless ponds independent of predator cues. There was no effect of pond predator regime or predator cues on leaf decomposition rates. Further, fewer individuals formed pairs in G. pulex from fish ponds than from fishless ponds. Male G. pulex had a higher mortality and a higher decomposition rate than females independent of predator cues. However, there was no difference in refuge use between sexes. Our study shows that there are general differences in behaviour traits, both between predator regimes and sexes in G. pulex.     

Limited influence of local and landscape factors on finescale gene flow in two pond‐breeding amphibians

Dispersal and gene flow within animal populations are influenced by the composition and configuration of the landscape. In this study, we evaluated hypotheses about the impact of natural and anthropogenic factors on genetic differentiation in two amphibian species, the spotted salamander (Ambystoma maculatum) and the wood frog (Lithobates sylvaticus) in a commercial forest in central Maine. We conducted this analysis at two scales: a local level, focused on factors measured at each breeding pond, and a landscape level, focused on factors measured between ponds. We investigated the effects of a number of environmental factors in six categories including Productivity, Physical, Land Composition, Land Configuration, Isolation and Location. Embryos were sampled from 56 spotted salamander breeding ponds and 39 wood frog breeding ponds. We used a hierarchical Bayesian approach in the program GESTE at each breeding pond and a random forest algorithm in conjunction with a network analysis between the ponds. We found overall high genetic connectivity across distances up to 17 km for both species and a limited effect of natural and anthropogenic factors on gene flow. We found the null models best explained patterns of genetic differentiation at a local level and found several factors at the landscape level that weakly influenced gene flow. This research indicates multiscale investigations that incorporate local and landscape factors are valuable for understanding patterns of gene flow. Our findings suggest that dispersal rates in this system are high enough to minimize genetic structuring and that current forestry practices do not significantly impede dispersal.     

Genotypic diversity and differentiation among populations of two benthic freshwater diatoms as revealed by microsatellites

Given their large population sizes and presumed high dispersal capacity, protists are expected to exhibit homogeneous population structure over large spatial scales. On the other hand, the fragmented and short‐lived nature of the lentic freshwater habitats that many protists inhabit promotes strong population differentiation. We used microsatellites in two benthic freshwater diatoms, Eunotia bilunaris ‘robust’ and Sellaphora capitata, sampled from within a pond and connected ponds, through isolated ponds from the same region to western Europe to determine the spatial scale at which differentiation appears. Because periods of low genotypic diversity contribute to population differentiation, we also assessed genotypic diversity. While genotypic diversity was very high to maximal in most samples of both species, some had a markedly lower diversity, with up to half (Eunotia) and over 90% (Sellaphora) of the strains having the same multilocus genotype. Population differentiation showed an isolation‐by‐distance pattern with very low standardized F_ST values between samples from the same or connected ponds but high values between isolated ponds, even when situated in the same region. Partial rbcL sequences in Eunotia were consistent with this pattern as isolated ponds in the same region could differ widely in haplotype composition. Populations identified by Structure corresponded to the source ponds, confirming that ‘pond’ is the main factor structuring these populations. We conclude that freshwater benthic diatom populations are highly fragmented on a regional scale, reflecting either less dispersal than is often assumed or reduced establishment success of immigrants, so that dispersal does not translate into gene flow.     

Genetic differentiation of submerged plant populations and taxa between habitats

Ceratophyllum spp., Callitriche spp., Zannichellia spp. and Potamogeton pectinatus L. are widespread submerged macrophyte species, often occurring at high abundance and forming an integral part of the vegetation of many types of shallow aquatic systems. Several species occur in both freshwater and brackish water habitats. Most have a mixed reproduction system and can reproduce sexually by seeds and propagate asexually by rhizomes, turions, root tubers or axillary tubers. It is hypothesized that sexual propagules are more important than vegetative fragments to ensure long-distance-dispersal, which in case of frequent bird or water flow-mediated dispersal should lead to lowered genetic differentiation. At a regional level, we used dominant ISSR markers in a multi-species approach and observed the largest clonal differentiation between brackish water and freshwater populations of the western European lowland (Belgium). Differentiation was pronounced at taxon level (e.g. Zannichellia), as a salinity gradient (P. pectinatus) or as a coastal-to-inland conductivity gradient (Callitriche obtusangula). These differences and trends suggested a very limited dispersal at regional level across both habitats and regions. To test the hypothesis whether vegetative reproduction and dispersal may have an important function in maintenance of the species at local scale, we investigated the microsatellite diversity and clonal distribution within and between populations of P. pectinatus from a single catchment, representing upstream forest ponds and downstream river sites along the Woluwe (Brussels, Belgium). Clonal diversity was low on average, however, with a higher number of multilocus genotypes in upstream forest ponds than in downstream river sites. A few but abundant clones were present along various stretches of the river indicating clonal spread and establishment over larger distances within the river. Clonal dispersal at a local scale was more pronounced in river than in pond habitats, indicating a higher relative importance of water flow than bird-mediated dispersal for establishment of P. pectinatus in river sites. Dispersal of seeds and establishment of seedlings were assumed more effective within ponds than in river habitats. Upstream forest ponds can be regarded as source populations and refuges of clonal diversity for recolonization of the more stressful downstream river habitat.     

Fine‐ and Broad‐Scale Approaches to Understanding the Evolution of Aggression in Crickets

Male field crickets frequently engage in agonistic contests to establish dominance in social interactions and gain access to mate attraction territories. Crickets (Orthoptera: Gryllidae) are often used as a model taxon to study aggression, but limited documentation of aggression in some cricket species hinders our understanding of its evolutionary costs and benefits. Our study investigated cricket aggression at two scales: the within‐species scale for two cricket species, Gryllus assimilis and G. veletis, whose aggression had not been adequately documented and the among‐species scale to detect evolutionary patterns in species’ levels of aggression. In both G. veletis and G. assimilis, winners spent more time being aggressive than losers, but they were not larger or heavier. Collectively, our results reveal that G. veletis males are more aggressive than G. assimilis. Male G. veletis had higher aggression scores that male G. assimilis. The majority of G. veletis contests escalated to grappling (a highly aggressive behavior), while less than one quarter of G. assimilis contests escalated to grappling. Further, G. veletis males transitioned between two of the most aggressive behaviors most often while G. assimilis transitioned between two of the least aggressive behaviors most often. We integrate this new information on aggression for G. assimilis and G. veletis with previously documented aggression data for many cricket species to investigate aggression in a broader evolutionary context than previously possible. Within a phylogenetic context, we test the hypothesis that species whose males use burrows from which to call and attract females are more aggressive than species with non‐burrowing males. We found evidence consistent with this hypothesis; species with burrowing males tended to be more aggressive than species with non‐burrowing males. Together, our study provides fine‐scale understanding of aggression in two cricket species and broad‐scale evolutionary context for aggression across cricket species.     

How did this snail get here? Several dispersal vectors inferred for an aquatic invasive species

1. How species reach and persist in isolated habitats remains an open question in many cases, especially for rapidly spreading invasive species. This is particularly true for temporary freshwater ponds, which can be remote and may dry out annually, but may still harbour high biodiversity. Persistence in such habitats depends on recurrent colonisation or species survival capacity, and ponds therefore provide an ideal system to investigate dispersal and connectivity. 2. Here, we test the hypothesis that the wide distributions and invasive potential of aquatic snails is due to their ability to exploit several dispersal vectors in different landscapes. We explored the population structure of Physa acuta (recent synonyms: Haitia acuta, Physella acuta, Pulmonata: Gastropoda), an invasive aquatic snail originating from North America, but established in temporary ponds in Doñana National Park, southern Spain. In this area, snails face land barriers when attempting to colonise other suitable habitat. 3. Genetic analyses using six microsatellite loci from 271 snails in 21 sites indicated that (i) geographically and hydrologically isolated snail populations in the park were genetically similar to a large snail population in rice fields more than 15 km away; (ii) these isolated ponds showed an isolation‐by‐distance pattern. This pattern broke down, however, for those ponds visited frequently by large mammals such as cattle, deer and wild boar; (iii) snail populations were panmictic in flooded and hydrologically connected rice fields. 4. These results support the notion that aquatic snails disperse readily by direct water connections in the flooded rice fields, can be carried by waterbirds flying between the rice fields and the park and may disperse between ponds within the park by attaching to large mammals. 5. The potential for aquatic snails such as Physa acuta to exploit several dispersal vectors may contribute to their wide distribution on various continents and their success as invasive species. We suggest that the interaction between different dispersal vectors, their relation to specific habitats and consequences at different geographic scales should be considered both when attempting to control invasive freshwater species and when protecting endangered species.     

Disturbance from pond management obscures local and regional drivers of assemblages of primary producers

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Self‐recruitment in a Caribbean reef fish: a method for approximating dispersal kernels accounting for seascape

Characterizing patterns of larval dispersal is essential to understanding the ecological and evolutionary dynamics of marine metapopulations. Recent research has measured local dispersal within populations, but the development of marine dispersal kernels from empirical data remains a challenge. We propose a framework to move beyond point estimates of dispersal towards the approximation of a simple dispersal kernel, based on the hypothesis that the structure of the seascape is a primary predictor of realized dispersal patterns. Using the coral reef fish Elacatinus lori as a study organism, we use genetic parentage analysis to estimate self‐recruitment at a small spatial scale (<1 km). Next, we determine which simple kernel explains the observed self‐recruitment, given the influx of larvae from reef habitat patches in the seascape at a large spatial scale (up to 35 km). Finally, we complete parentage analyses at six additional sites to test for export from the focal site and compare these observed dispersal data within the metapopulation to the predicted dispersal kernel. We find 4.6% self‐recruitment (CI_95%: ±3.0%) in the focal population, which is explained by the exponential kernel y = 0.915^x (CI_95%: y = 0.865^x, y = 0.965^x), given the seascape. Additional parentage analyses showed low levels of export to nearby sites, and the best‐fit line through the observed dispersal proportions also revealed a declining function y = 0.77^x. This study lends direct support to the hypothesis that the probability of larval dispersal declines rapidly with distance in Atlantic gobies in continuously distributed habitat, just as it does in the Indo‐Pacific damselfishes in patchily distributed habitat.     

Strong spatial influence on colonization rates in a pioneer zooplankton metacommunity

The magnitude of community-wide dispersal is central to metacommunity models, yet dispersal is notoriously difficult to quantify in passive and cryptic dispersers such as many freshwater invertebrates. By overcoming the problem of quantifying dispersal rates, colonization rates into new habitats can provide a useful estimate of the magnitude of effective dispersal. Here we study the influence of spatial and local processes on colonization rates into new ponds that indicate differential dispersal limitation of major zooplankton taxa, with important implications for metacommunity dynamics. We identify regional and local factors that affect zooplankton colonization rates and spatial patterns in a large-scale experimental system. Our study differs from others in the unique setup of the experimental pond area by which we were able to test spatial and environmental variables at a large spatial scale. We quantified colonization rates separately for the Copepoda, Cladocera and Rotifera from samples collected over a period of 21 months in 48 newly constructed temporary ponds of 0.18–2.95 ha distributed in a restored wetland area of 2,700 ha in Doñana National Park, Southern Spain. Species richness upon initial sampling of new ponds was about one third of that in reference ponds, although the rate of detection of new species from thereon were not significantly different, probably owing to high turnover in the dynamic, temporary reference ponds. Environmental heterogeneity had no detectable effect on colonization rates in new ponds. In contrast, connectivity, space (based on latitude and longitude) and surface area were key determinants of colonization rates for copepods and cladocerans. This suggests dispersal limitation in cladocerans and copepods, but not in rotifers, possibly due to differences in propagule size and abundance.     

Passive external transport of freshwater invertebrates by elephant and other mud‐wallowing mammals in an African savannah habitat

1. Recent findings hint at the potential importance of mammals affecting the spatial dynamics of aquatic organisms in areas where mammals live in close association with water. Perhaps the most iconic example of such an environment is the African savannah. 2. We investigated dispersal patterns of freshwater organisms among a set of temporary ponds in SE Zimbabwe to test the hypothesis that large mammals, and particularly African elephants (Loxodonta africana), can be important vectors of aquatic organisms. Dispersal kernels were reconstructed by hatching mud collected from ‘rubbing’ trees located at increasing distances from a set of isolated ponds. To assess the relative importance of other mammalian vectors, the vertical distribution of mud on rubbing trees was mapped and related to the body size of candidate vector species. 3. Laboratory hatching of mud samples revealed large numbers of propagules of 22 invertebrate taxa as well as some aquatic macrophytes. Dispersing communities reflected source communities and diverged with increasing distance from the source. Both dispersal rates and richness of transported taxa decreased significantly with dispersal distance. No indications for differences in dispersal capacity among propagule types were detected. Instead, common propagules were more likely to travel greater distances. Most mud was attached to trees at heights >1.5 m, implicating elephants as the dominant vector. Vertical distributions of tree mud, however, also revealed clustering at heights up to 50 cm and 90–120 cm corresponding to the height of warthog, rhinoceros and buffalo, respectively. Finally, variation in the vertical distribution of mud on trees in combination with differences in vector vagility suggests that local differences in vector species composition may affect passive dispersal dynamics of aquatic organisms. 4. Based on vagility and vector load, mud‐wallowing mammals emerge as highly effective vectors that, in some areas, may be more important in transporting aquatic organisms than traditionally recognised vectors such as waterbirds. Since most large‐ and medium‐sized mammals currently have restricted geographic distributions, it is likely that mammal‐mediated dispersal was more important in the past.     

Vision-based ability of an ant-mimicking jumping spider to discriminate between models,conspecific individuals and prey

Myrmarachne assimilis, an ant-like (myrmecomorphic) jumping spider (Araneae, Salticidae) from the Philippines, is a Batesian mimic of Oecophylla smaragdina, the Asian weaver ant. Salticids are well known for their acute eyesight and the elaborate vision-based display behaviour they adopt during encounters with conspecific individuals, but most salticids are not myrmecomorphic. Despite its unusual morphology, M. assimilis adopts display behaviour during intraspecific interactions that is similar to the display behaviour of more typical salticids. The specificity with which M. assimilis deploys display behaviour is investigated and provides insights into this mimic’s ability to differentiate, by sight alone, between models, conspecific individuals and prey. During each standardized test, an adult M. assimilis female was in a large cage along with a small transparent glass vial, a stimulus animal being enclosed in the vial such that potential optical cues, but not potential chemical cues, were available to the tested M. assimilis individual. Depending on the test, the stimulus animal was another adult M. assimilis female, a house fly (prey) or an ant (Camponotus sp. or O. smaragdina). Only the conspecific female consistently elicited display from M. assimilis, implying that M. assimilis is a Batesian mimic that can, when relying on vision alone, discriminate between conspecific individuals, models and prey. Received 12 June 2006; revised 22 September 2006; accepted 26 September 2006.     

Habitat patch size alters the importance of dispersal for species diversity in an experimental freshwater community

Increased dispersal of individuals among discrete habitat patches should increase the average number of species present in each local habitat patch. However, experimental studies have found variable effects of dispersal on local species richness. Priority effects, predators, and habitat heterogeneity have been proposed as mechanisms that limit the effect of dispersal on species richness. However, the size of a habitat patch could affect how dispersal regulates the number of species able to persist. We investigated whether habitat size interacted with dispersal rate to affect the number of species present in local habitats. We hypothesized that increased dispersal rates would positively affect local species richness more in small habitats than in large habitats, because rare species would be protected from demographic extinction. To test the interaction between dispersal rate and habitat size, we factorially manipulated the size of experimental ponds and dispersal rates, using a model community of freshwater zooplankton. We found that high‐dispersal rates enhanced local species richness in small experimental ponds, but had no effect in large experimental ponds. Our results suggest that there is a trade‐off between patch connectivity (a mediator of dispersal rates) and patch size, providing context for understanding the variability observed in dispersal effects among natural communities, as well as for developing conservation and management plans in an increasingly fragmented world.     

Microsatellite variation and population structure of a recovering Tree frog (Hyla arborea L.) metapopulation

Numbers and sizes of populations of the European tree frog in The Netherlands have dramatically decreased in the second half of the last century due to extensive habitat destruction and fragmentation. We have studied the genetic structure of a slowly recovering meta-population. Strong genetic differentiation, estimated at eight microsatellite loci, was found between clusters of populations (F _st-values above 0.2). Within clusters, consisting of ponds within a radius of about 5 km, European tree frog populations were less differentiated (F _st<0.08) and exact tests showed that most of the ponds within clusters were not significantly differentiated. Although local population sizes have been increasing since 1989, and some new ponds have been colonised in the direct vicinity of ponds that have been populated continuously, little evidence for gene flow between clusters of ponds was found (only one exception). Furthermore, levels of genetic diversity were low compared to populations in comparable areas elsewhere in Europe. Therefore, a continuous conservation effort is needed to prevent any further loss of genetic diversity. The alleviation of important barriers to dispersal between the clusters should be given a high priority for the restoration of the meta-population as a whole.     

Habitat requirements and local persistence of three damselfly species (odonata: coenagrionidae)

Habitat requirements and population persistence were investigated in three damselfly species, all coastal plain pond specialists: Enallagma recurvatum, E. laterale, and E. pictum. Because of geographic restriction, two are of special concern to conservation, E. recurvatum and E. laterale. We surveyed more than 70 ponds on Cape Cod, Massachusetts, and collected adult presence-absence data during the summers of 1999 and 2000. We achieved a detection rate approaching 100% for each species by visiting each pond up to three times. We looked for relationships between the presence of each damselfly species and presence of specific aquatic vegetation, the presence of the other Enallagma species, and the number of ponds within various distances of the 72 surveyed ponds. Using stepwise logistic regression, we found the following significant associations: E. recurvatum with the rush Juncus militaris; E. laterale with water lilies (Nuphar variegatum and Brasenia schreberi) the damselfly E. pictum, and the number of ponds within 2 km; and E. pictum with the water lily Nymphaea odorata, the damselfly E. laterale, and the number of ponds within 1.5 km and 2.5 km. Presence-absence data were used to calculate turnover and local extinction rates for each species between the two years. E. recurvatum's turnover and local extinction rates (33.3% and 41% respectively) were much higher than either E. laterale (9.8%, 11.5%) or E. pictum (7.7%, 10%). These results suggest that E. recurvatum occurs in a metapopulation, and that patch colonization rates might be important to local population persistence.