Amphibian decline,pond loss and reduced population connectivity under agricultural intensification over a 38 year period

Habitat loss, together with less obvious land-use changes such as intensified farming practice, can have significant adverse impacts on biodiversity. An important factor in determining the ability of species to cope with such changes is their potential to sustain a populations network by dispersal across the landscape. Habitat quality and structure are particularly important for surface-dwelling species with low dispersal abilities, such as amphibians. To assess this ecological function, ponds in a coastal and typically rural area of northern France were surveyed for amphibians in 1974, 1992 and 2011. These repeated surveys yielded different outcomes for different species groups. Three rare species persisted in more or less specialized habitat types. Two moderately common species declined, but kept strongholds in coastal dunes and associated marshes. Five common species with broad ecological niches remained equally widespread. The Northern crested newt declined markedly and the Midwife toad declined dramatically, as did their breeding habitats in vegetated ponds and cattle drinking troughs. One species, the Moor frog, may have gone locally extinct. A model of relative resistance to amphibian dispersal was created for different landscape types, on a scale from 0 (low resistance) to 1 (high resistance). This generated values of 0.23 for pasture, 0.72 for arable and 0.98 for urban and transport. As pasture declined in the study area, while arable and urban/transport infrastructure increased, amphibian dispersal became more difficult. However, dispersal paths proved difficult to evaluate in a patchy landscape like the one surveyed, due to a paucity of spatial signal. Pond loss is a more tractable reason for the observed amphibian species decline than is the quality of intervening terrestrial habitat matrix. In 2011, 22 newly created ponds had species richness in line with pre-existing ponds and this will have counteracted a dwindling metapopulation structure, indicating that habitat creation/restoration can play a valuable role in conservation. The colonization of new ponds may also prove more informative for gauging the potential for amphibian dispersal in the landscape than the preceding decline.     

Size-structured effects of common carp on reproduction of pond-breeding amphibians

The role of fish in driving amphibian communities has been widely recognized. However, little is known about size-structured interactions between amphibian and fish populations. This study compared the taxonomic occurrence and densities of larval amphibians between unstocked ponds and ponds stocked with different age cohorts of common carp Cyprinus carpio differing in average body size. The average total densities of early and late breeding anurans known to be vulnerable to fish were by 1–2 orders of magnitude greater in the presence of young-of-the-year carp than that of older cohorts. The probabilities of occurrence of the most common taxa did not differ between ponds stocked with young-of-the-year fish and ponds free of carp, but were significantly larger in those ponds than in ponds stocked with large-size cohorts. No significant differences between pond categories were found for densities of unpalatable Bufo bufo larvae. In aquatic systems harbouring size-structured fish populations, a fish age/size gradient may explain differential habitat suitability for breeding amphibians better than the fish presence/absence dichotomy. When dominated by young cohorts incapable of predation or of adverse habitat alteration, fish-abundant waters are suitable for amphibian reproduction. Conversely, even a ‘non-predatory’ fish, after attaining large body size, may exert a detrimental impact on amphibian breeding success. These findings may be particularly important for amphibian conservation at pond fisheries characterized by spatial separation of age/size distributed stocks.     

Rainfall stochasticity controls the distribution of invasive crayfish and its impact on amphibian guilds in Mediterranean temporary waters

Invasive crayfish have severely impacted invaded aquatic ecosystems worldwide. We studied temporal and spatial variation in the range expansion of the red swamp crayfish at one of the first European localities to which it was introduced: Doñana National Park (SW Spain). In contrast to the rapid range expansion witnessed in other areas, this invasive crayfish has not spread across the entire park. Instead, its distribution has expanded during wet periods, but contracted during drought periods. The red swamp crayfish has caused steep amphibian declines in other invaded areas. However, after approximately 35 years of crayfish presence in Doñana National Park, we have yet to detect a reduction in the number or occurrence of amphibian species. Amphibians may thus be protected by the large abundance of temporary ponds in the area, which provides them with an effective refuge network. We show that natural fluctuations in annual rainfall and in the number of ponds filled can temporarily eliminate invasive crayfish from particular areas. This fact should be taken into account when attempting to reduce the impact of crayfish on aquatic communities, intensifying crayfish removal during particularly dry years, when it is most effective.     

Spatial genetic structure of <Emphasis Type="Italic">Lissotriton helveticus L</Emphasis>. following the restoration of a forest ponds network

Preserving amphibian genetic diversity through ecological restoration and conservation actions is a major challenge since their populations are declining worldwide. We studied the genetic diversity and spatial genetic structure of the palmate newt (Lissotriton helveticus) 2 years after the restoration of a pond network in northwestern France with the aim of reconstructing fine-scale genetic structure and patterns of colonization. We sampled newts from 29 forest ponds including both restored and non-degraded reference ponds, and genotyped 391 individuals at 12 microsatellite loci. We used two Bayesian clustering methods to spatially delineate genetic clusters, and we also detected potential recent migrants within the network. All ponds showed low levels of observed heterozygosity (Ho?=?0.534) and a mean F _IS of 0.251, possibly indicating a Wahlund or bottleneck effect. Pairwise F _ST suggested limited evidence of genetic differentiation among ponds. Within the pond network, we identified 3 to 4 genetic clusters. Combined with the detection of migrants, the results suggest an increase in gene flow within the restored pond network and that a high number of migrants came from the reference ponds. Our findings indicate an unexpected high dispersal ability for this small-bodied species. Overall, the absence of population structure represents a positive beginning for the restoration project. It also emphasizes the importance of spatial design in restoring a pond network and that such genetic data and methods should be used to monitor amphibians in restored habitats.     

Spatial scales for the management of amphibian populations

Conservation programs must be based on a detailed knowledge of the distribution of communities and populations, but this changes with the spatial scale of observation. In this context, we examined the multiscale patterns of spatial variation of a montane amphibian assemblage in Central Spain. The largest spatial scale examined (in a range of 1200 m) accounted for the maximum variability in both species richness and larval abundance, while the smallest scale examined (pond) was responsible for the maximum variation of larval abundances of many individual species. Habitat characteristics seemed to be more related to spatial variation of the amphibian assemblage at a particular spatial scale (in a range of 75 m), and distances among ponds were in part responsible for variation of larval abundance, but not species richness.     

The effect of fish and aquatic habitat complexity on amphibians

Fish introductions are considered one of the most widespread anthropogenic threats to aquatic ecosystems. Their negative impact on native amphibian communities has received increasing attention in recent years. We investigated the relationship between the introduced fish, emergent vegetation cover and native amphibians in man-made ponds generated by regulation and dam building along the Tarnava Mare Valley (Romania) during the last 40 years. We inventoried amphibians and fish inhabiting 85 permanent ponds and estimated habitat complexity focusing on emergent vegetation cover. Four amphibian species were found to be negatively associated with the presence of predatory fish. Species richness of ponds without fish and ponds without predatory fish did not differ significantly, whereas ponds containing only predatory fish had significantly lower amphibian richness. A significant positive relationship was found between the emergent vegetation cover and pond occupancy of six amphibian species and amphibian species richness. As a management recommendation, we suggest the restriction of fish introductions to non predatory fish and the maintenance of high emergent vegetation cover in the ponds. Handling editor: S. Declerck     

An expert-based landscape permeability model for assessing the impact of agricultural management on amphibian migration

The persistence of pond-breeding amphibian populations is influenced on different spatial scales ranging from the individual breeding pond to surrounding habitat patches to landscape clusters of breeding pond populations. The connectivity among breeding ponds as well as the availability of suitable terrestrial habitats surrounding the ponds plays a major role in long-term viability of amphibian species. Besides road traffic and urban structures agricultural land activity can disrupt landscape connectivity through the use of pesticides, fertilizers and physical activity such as tillage. We developed an expert-based model to assess the impact of agricultural management measures on the migration area and terrestrial habitat availability for seven amphibian species. The model is based on a Habitat Suitability Index to identify suitable terrestrial habitats and includes a landscape permeability approach. Size of migration areas, connectivity of breeding ponds and number of reachable terrestrial habitats were modeled considering species-specific migration ranges and habitat preferences. We consider how pesticides application might lead to fragmentation and isolation of amphibian breeding pond populations. Therefore the potential disrupting impact of pesticides was simulated by inflating landscape resistance to medium and high migration cost. One amphibian species showed a decrease of migration area by 48.3% and a decrease of reachable terrestrial habitats by 41.5% at high migration costs. Three additional species showed a decrease of their migration areas between 31.5 and 35.7%. At increased migration cost, some of the investigated populations were isolated at breeding pond level or restricted to pond clusters. Our model could be used to prioritize conservation efforts for pond-breeding amphibians with adequate consideration of agricultural land use and its impact on amphibian migration.     

Breeding site characteristics regulating life history traits of the brown tree frog, Litoria ewingii

Past research has determined the habitat requirements of amphibian species predominantly from presence/absence studies. This study tested the hypothesis that relationships between breeding site habitat components, life history traits and fitness may provide a higher resolution of biological data relating to the habitat requirements of amphibian species. We tested this novel approach by using Litoria ewingii as our model species. We correlated larval and metamorph life history traits with habitat variables at 28 small to medium sized ponds within a commercially logged forest in southern Tasmania, Australia. To avoid larval mortality due to pond desiccation, L. ewingii laid eggs and metamorphosed earlier in smaller ponds. Snout vent length at metamorphosis increased with elevation and metamorphosis was earlier in less shaded ponds. Breeding ponds that maximised the fitness of L. ewingii were higher elevation ponds with reduced shading, steeper bank slopes and reduced pond isolation. The findings of the study equip land managers with a greater ecological understanding of ecosystem function in relation to specific species. The methodological approach has broad application to conservation biology where an awareness of the specific habitat requirements of amphibians is critical to successful ecosystem management.     

Emerging conflicts for the environmental use of water in high-valuable rangelands. Can livestock water ponds be managed as artificial wetlands for amphibians?

Continental freshwater, irrespective of its origin, natural or artificial, may contribute significantly to biodiversity conservation. Because of the decline of natural aquatic habitats, an increasing concern exists about the role of water ponds as spots of biological richness. Amphibians are strongly at risk since the loss of aquatic habitats, among other factors, causes the isolation of their populations. The implementation of livestock ponds as artificial wetlands may be an effective measure for enhancing amphibian decaying communities. This policy assumes that managing ponds for wildlife conservation purposes joins livestock welfare requirements, but this hypothesis has not been specifically studied. The purpose of this research is to evaluate this premise in the Urbasa-Andia Natural Park, a high-valuable environmental area that holds a relevant amphibian community and has an extended grazing history. We analyse the relationship between the amphibian assemblages present and the design and attributes of a variety of drinking points previously chosen by embodying a high environmental heterogeneity of water resources. The results of this study indicate that the quality of the water stored varies largely along the season, degrading severely in summer because of the wading of animals (in unfenced ponds) and the low water recharge. The contamination, caused by increased enteric microorganisms and dissolved N, is likely to affect livestock more severely than amphibian populations, since the sensitive breeding stage of many amphibians occurs before the loss of water quality. Although the quality of the water is essential, and mammals (wild and domestic) have an influence on it, other factors that are less considered by environmental managers emerge as main drivers of amphibian assemblages, such as hydroperiod, predator occurrence and the environmental quality of the surrounding habitat.     

Metacommunity dynamics of amphibians in years with differing rainfall

Many studies investigated the habitat preference and behaviour ecology of individual amphibian species while we know less about how their community assembly reflects changes in environmental factors, including the role of climatic extremes. Community-level studies also allow us to apply trait-based analyses that are crucial for a better understanding of the functioning of amphibian communities and metacommunities. In two years with contrasting rainfall (2012 and 2013), we found amphibian species in 85 different waterbodies of a heterogeneous landscape in Central Europe (Hungary). Within the metacommunity framework, the contributions of local, landscape and spatial variables to community assembly were assessed. We also measured the local extinction–colonisation rates in the ponds for all species between the two years. To investigate the role of dispersal traits in explaining the spatial distribution of species, we studied the relationship between body size and the pure spatial fraction of variation. According to our results, the main drivers were the same in both the dry and wet year, but their relative contribution changed. Local variables played a predominant role in the assembly of the amphibian metacommunity. Spatial signals were more evident in the dry year. This implies not only the adverse effect of decreased connectivity due to the drying out of several habitats but also a loss of breeding sites for the studied amphibians. Local colonisation rates were higher in primarily terrestrial species (Hyla arborea, Pelobates fuscus, Bufo bufo) which only visit ponds during breeding. We found a negative relationship between the pure spatial effect and body size, suggesting an increased level of dispersal limitation in small-bodied species. Our results showed that while the strength and relative role of local and spatial processes changed between years, the role of dispersal traits in explaining the spatial distribution of species was similar. Specialisation to different habitats seems to be a major process in determining vertebrate metacommunities in landscapes. Dispersal traits of different species should be taken more into consideration in the practical conservation of amphibian habitats.     

Incorporating landscape connectivity into household pond configuration in a hilly agricultural landscape

Manmade ponds are common landscape features in rural areas and also important habitats for maintaining biodiversity. However, they are vulnerable to anthropogenic activities, land-use changes, and habitat degradation; many ponds being filled or (re)created arbitrarily. Little attention has been paid to quantifying the spatial structure of these manmade ponds at a landscape scale, nor to their potential functional benefits in promoting ecological flows and interactions between habitats for whole-ecosystem integrity. In this study, we investigated the patch-based landscape connectivity of household ponds, a particular type of domestic pond prevalent in hilly rural areas of China, by using least-cost path modelling and graph theory based network analysis. A hierarchical network was modelled consisting of 4606 individual ponds, 373 pond patches and 772 potential links within a 1.5-km threshold distance. Network importance analysis revealed that the largest pond patch contributes 24.5 % to network building and that patches with larger areas are generally more important. In contrast, the importance of the simulated links is only 2.3 % at most, indicating that the network has spatial redundancy which can strengthen resilience to uncertain disturbances. Our study moves beyond network simulation and importance assessment by directly relating the connectivity analysis to a real construction context through the incorporation of a spatially explicit land suitability analysis. This approach systematises the analysis of pond landscapes and guides integration with the wider landscape matrix. It provides operational spatial suggestions for holistic landscape planning across local to regional scales.     

Is the exotic red swamp crayfish (Procambarus clarkii) a current threat for the Mediterranean tree frog (Hyla meridionalis) in the Camargue (Southern France)?

The non-indigenous red swamp crayfish (Procambarus clarkii) has been shown to be a threat for amphibian conservation. Many amphibian species breed in temporary ponds to diminish predation risk as such ecosystems are free of large predators. However P. clarkii, occurring as an invasive species in the Camargue delta, can readily disperse on the ground and thus colonize isolated ponds. We studied the current impact of the exotic crayfish on the reproductive success of the Mediterranean tree frog (Hyla meridionalis). In a mesocosm experiment, we tested the effect of two crayfish densities (1 and 3 crayfish/m²) on tadpole abundance. We also tested in a field experiment, within a temporary pond, the crayfish’s predation on the tree frog’s eggs. Finally, we developed site occupancy models using data from 20 ponds to assess the effect of crayfish abundance on tadpole abundance. Neither the experiments, nor the site occupancy models showed a negative impact of the current crayfish abundance on the tree frog populations breeding in ponds. We found that recorded crayfish densities were lower than in other areas where crayfish has impacted amphibian populations, but we hypothesize that current crayfish abundance in the area may increase in the future, thus impacting tree frog populations.     

Seasonal patterns of activity and community structure in an amphibian assemblage at a pond network with variable hydrology

We studied community structure and seasonal activity patterns in a system of four ponds with seasonally-variable hydrology at a Mediterranean area in central Italy. We used a set of field methods to assess species presence and relative frequency of observation. The network of ponds was inhabited by six species of amphibians, two salamanders and four frogs. The breeding phenology of the six species did not vary remarkably among ponds, but there were significant differences among species in use of ponds. Factorial analysis of pond similarity drawn from percentage composition of the amphibian fauna, revealed that each of the four ponds was treatable as independent units, with no influence of relative inter-pond distance. PCA analysis allowed us to spatially arrange the amphibian species into three main groups: two were monospecific groups (i.e., Triturus vulgaris and Bufo bufo) and the third consisted of those species that selected not only the largest-deepest ponds, but also the ephemeral ones (i.e., Triturus carnifex, Hyla intermedia, the green frogs and Rana dalmatina). Our results suggest that the inter-pond differences in riparian vegetation, water depth, aquatic vegetation structure/abundance, and soil composition may produce differences among pond ecological characteristics (i.e., water turbidity and temperature, shelter availability, abundance of oviposition micro-sites), which may in turn influence different patterns of use by amphibians. To our knowledge, this is the first study emphasizing the potential role of heterochrony in the maintenance of a high species richness in Mediterranean amphibian communities. Preservation of freshwater vertebrate biodiversity requires management and protection not only of the main ponds and water bodies but also the temporary and ephemeral shallow ponds.     

Exotic Fish in Exotic Plantations: A Multi-Scale Approach to Understand Amphibian Occurrence in the Mediterranean Region

Globally, amphibian populations are threatened by a diverse range of factors including habitat destruction and alteration. Forestry practices have been linked with low diversity and abundance of amphibians. The effect of exotic Eucalyptus spp. plantations on amphibian communities has been studied in a number of biodiversity hotspots, but little is known of its impact in the Mediterranean region. Here, we identify the environmental factors influencing the presence of six species of amphibians (the Caudata Pleurodeles waltl, Salamandra salamandra, Lissotriton boscai, Triturus marmoratus and the anurans Pelobates cultripes and Hyla arborea/meridionalis) occupying 88 ponds. The study was conducted in a Mediterranean landscape dominated by eucalypt plantations alternated with traditional use (agricultural, montados and native forest) at three different scales: local (pond), intermediate (400 metres radius buffer) and broad (1000 metres radius buffer). Using the Akaike Information Criterion for small samples (AIC_c), we selected the top-ranked models for estimating the probability of occurrence of each species at each spatial scale separately and across all three spatial scales, using a combination of covariates from the different magnitudes. Models with a combination of covariates at the different spatial scales had a stronger support than those at individual scales. The presence of predatory fish in a pond had a strong effect on Caudata presence. Permanent ponds were selected by Hyla arborea/meridionalis over temporary ponds. Species occurrence was not increased by a higher density of streams, but the density of ponds impacted negatively on Lissotriton boscai. The proximity of ponds occupied by their conspecifics had a positive effect on the occurrence of Lissotriton boscai and Pleurodeles waltl. Eucalypt plantations had a negative effect on the occurrence of the newt Lissotriton boscai and anurans Hyla arborea/meridionalis, but had a positive effect on the presence of Salamandra salamandra, while no effect on any of the other species was detected. In conclusion, eucalypts had limited effects on the amphibian community at the intermediate and broad scales, but predatory fish had a major impact when considering all the scales combined. The over-riding importance of introduced fish as a negative impact suggests that forest managers should prevent new fish introductions and eradicate fish from already-occupied ponds whenever possible.     

The response of amphibian communities to fish and habitat features in Mediterranean permanent ponds

Generalized linear models were used to test the effect of fish, using ponds with and without fish and habitat features as covariates, on richness and abundance of amphibian species. Five fish species and six amphibian species were recorded in 60 permanent ponds located in central Italy. The choice of covariates (macrophyte cover and pond surface area) was made after studying the correlations. The richness of amphibian species was not significantly affected by fish presence or macrophyte cover, in line with previous studies, since almost all the fish species were non-predatory. However, abundance of urodeles (newts) was negatively affected by fish and positively affected by macrophyte cover. Although fish may strongly influence the abundance and composition of amphibian communities, the results indicate that the cover of aquatic macrophytes may increase the available habitat for amphibians and therefore their abundance. Anuran species preferred ponds where fish were present, since both groups preferred larger ponds. Concordance between fish and amphibian species composition was not found by the Mantel and Partial Mantel tests. This indicates that the fish assemblages do not predict which amphibian species occur in the pond.     

Acid sensitivity and water chemistry correlates of amphibian breeding ponds in northern Wisconsin,USA

Fifty-one amphibian breeding sites in northern Wisconsin were sampled in two surveys, in 1986 and 1987, to determine the chemical conditions and amphibian inhabitants present. All twenty-one ponds in 1986 and twenty-eight in 1987 were mildly acidic and sensitive to acidification. The pH levels encountered were generally not toxic to the resident amphibians, and may have been related to levels of phenolics in some cases. Soil type and forest canopy coverage appeared to influence the chemistry of these ponds and a direct relationship of amphibian species richness with pond size was found. Only larger ponds were long-lived enough for metamorphosis of amphibian larvae, suggesting an importance for recruitment. They were also the most sensitive ponds and the only ones to enter the chronically toxic pH range for ambystomatid salamanders, possibly placing them at risk to future acid inputs.     

Effects of terrestrial buffer zones on amphibians on golf courses

A major cause of amphibian declines worldwide is habitat destruction or alteration. Public green spaces, such as golf courses and parks, could serve as safe havens to curb the effects of habitat loss if managed in ways to bolster local amphibian communities. We reared larval Blanchard's cricket frogs (Acris blanchardi) and green frogs (Rana clamitans) in golf course ponds with and without 1 m terrestrial buffer zones, and released marked cricket frog metamorphs at the golf course ponds they were reared in. Larval survival of both species was affected by the presence of a buffer zone, with increased survival for cricket frogs and decreased survival for green frogs when reared in ponds with buffer zones. No marked cricket frog juveniles were recovered at any golf course pond in the following year, suggesting that most animals died or migrated. In a separate study, we released cricket frogs in a terrestrial pen and allowed them to choose between mown and unmown grass. Cricket frogs had a greater probability of using unmown versus mown grass. Our results suggest that incorporating buffer zones around ponds can offer suitable habitat for some amphibian species and can improve the quality of the aquatic environment for some sensitive local amphibians.     

The transferability of distribution models across regions: an amphibian case study

Aim Predicting species distribution is of fundamental importance for ecology and conservation. However, distribution models are usually established for only one region and it is unknown whether they can be transferred to other geographical regions. We studied the distribution of six amphibian species in five regions to address the question of whether the effect of landscape variables varied among regions. We analysed the effect of 10 variables extracted in six concentric buffers (from 100 m to 3 km) describing landscape composition around breeding ponds at different spatial scales. We used data on the occurrence of amphibian species in a total of 655 breeding ponds. We accounted for proximity to neighbouring populations by including a connectivity index to our models. We used logistic regression and information‐theoretic model selection to evaluate candidate models for each species. Location Switzerland. Results The explained deviance of each species’ best models varied between 5% and 32%. Models that included interactions between a region and a landscape variable were always included in the most parsimonious models. For all species, models including region‐by‐landscape interactions had similar support (Akaike weights) as models that did not include interaction terms. The spatial scale at which landscape variables affected species distribution varied from 100 m to 1000 m, which was in agreement with several recent studies suggesting that land use far away from the ponds can affect pond occupancy. Main conclusions Different species are affected by different landscape variables at different spatial scales and these effects may vary geographically, resulting in a generally low transferability of distribution models across regions. We also found that connectivity seems generally more important than landscape variables. This suggests that metapopulation processes may play a more important role in species distribution than habitat characteristics.     

Functional traits reveal environmental constraints on amphibian community assembly in a subtropical dry forest

The relationships between functional traits and environmental gradients are useful to identify different community assembly processes. In this work, we used an approach based on functional traits to analyse if changes in hydroperiod and tree covers of ponds are relevant for local amphibian community assembly processes. Ephemeral ponds with low vegetation cover are expected to impose constraints on different species with particular trait combinations and, therefore, to exhibit communities with lower functional diversity than more stable ponds with greater tree cover. Sampling was conducted in 39 temporary ponds located along vegetation and hydroperiod gradients in the most arid portion of the Chaco ecoregion. Seven functional traits were measured in each species present in the regional pool. Associations between these traits and environmental gradients were detected using multivariate ordination techniques and permutation test (RLQ and fourth‐corner analyses respectively). Functional diversity indices were then calculated and related to variations in the environmental gradients. The results obtained allowed us to identify different sets of traits associated with hydroperiod and tree cover, suggesting that these environmental variables are relevant for structuring amphibian communities according to interspecific variations in functional traits from both, larval and adult stages. Contrary to our expectations, communities associated with more stable ponds and with greater tree cover exhibited lower functional diversity than expected by chance (and were the ponds with highest species richness). This result indicates that the reduction in relative importance of environmental restrictions imposed by a very short hydroperiod and the lack of tree cover, favours different species of the regional pool that are similar in several functional traits. Accordingly, communities associated with stable ponds with high tree cover exhibited high functional redundancy.