Restoring ponds for amphibians: a success story

Large-scale restoration of quality habitats is often considered essential for the recovery of threatened pond-breeding amphibians but only a few successful cases are documented, so far. We describe a landscape-scale restoration project targeted at two declining species—the crested newt (Triturus cristatus Laur.) and the common spadefoot toad (Pelobates fuscus Wagler)—in six protected areas in southern and southeastern Estonia. The ponds were restored or created in clusters to (i) increase the density and number of breeding sites at local and landscape levels; (ii) provide adjacent ponds with differing depths, hydroperiods and littoral zones and (iii) restore an array of wetlands connected to appropriate terrestrial habitat. In only 3 years, where 22 of the 405 existing ponds (5%) were restored and 208 new ponds (51%) created, the number of ponds occupied by the common spadefoot toad increased 6.5 times. Concerning the crested newt and the moor frog (Rana arvalis Nilsson), the increase was 2.3 and 2.5 times, respectively. The target species had breeding attempts in most of the colonised ponds—even more frequently than common species. Also, the amphibian species richness was higher in the restored than in the untreated ponds. The crested newt preferably colonised ponds that had some submerged vegetation and were surrounded by forest or a mosaic of forest and open habitats. The common spadefoot toad favoured ponds having clear and transparent water. Our study reveals that habitat restoration for threatened pond-breeding amphibians can rapidly increase their numbers if the restoration is implemented at the landscape scale, taking into account the habitat requirements of target species and the ecological connectivity of populations. When the remnant populations are strong enough, translocation of individuals may not be necessary. Guest editors: B. Oertli, R. Cereghino, A. Hull & R. Miracle Pond Conservation: From Science to Practice. 3rd Conference of the European Pond Conservation Network, Valencia, Spain, 14–16 May 2008     

Breeding phenology and larval distribution of amphibians in a Mediterranean pond network with unpredictable hydrology

Understanding spatial and temporal breeding patterns in Mediterranean amphibian communities is urgent considering the rate of habitat loss. Breeding phenology and breeding habitat selection by amphibians were analysed through the monthly occurrence of larvae in a mosaic of 198 Mediterranean temporary ponds during three years. A generalized linear model (GLM) coupled with principal component analysis showed that, for almost all species, occurrence was significantly positively correlated to pond depth. In addition, pond openness negatively affected the presence of some species. Temporal breeding patterns varied among species. Some species exhibited flexibility in their breeding date (Pelobates cultripes, Pelodytes punctatus, Hyla meridionalis, Rana perezi), while others did not (Triturus marmoratus, Triturus helveticus, Bufo calamita). When faced with inter-annual hydrological variability, the first group had a more constant breeding success than the second. Variable hydrological conditions caused differential larval occurrence of species between years. These fluctuations might favour long-term persistence of the whole amphibian community. We finally discuss the implications of our results for the management of amphibian habitats in the Mediterranean region.     

Micro-habitat use within a guild of newt larvae (Trituridae) in an Alpine lake

Population and ecological parameters such as numbers of larvae, microhabitat use, niche breadth and niche overlap of three species of syntopic larval newts (Alpine newt Triturus alpestris, Italian crested newt T. carnifex, and common newt T. vulgaris) were studied for two years in a small pond at 1160 m a.s.l. in NE Slovenia. Differences in microhabitat partitioning among larval newts were small. The largest niche breadth was estimated for larval T. alpestris, and the narrowest estimate was for larval T. carnifex in both years. Ecological differences seem to be very small and quite variable among sites and years. It appears that the developmental stage and size of newt larvae are more important in explaining resource partitioning than the characteristics of each species. Because of the absence of potential invertebrate predators and adult newts in the second half of the breeding season, the injuries could only be caused by intra-and interspecific predation attempts.     

Context-dependent dispersal,public information,and heterospecific attraction in newts

Dispersal is one of the main processes that determine community structure. Individuals make dispersal decisions according to environmental and/or social cues that reflect the fitness prospects in a given patch. The presence and abundance of heterospecifics within the same ecological guild, and/or their breeding success, may act as public information that influences movement decisions. To date, most studies investigating the role of heterospecific attraction have focused on habitat choice, using both experimental and correlational approaches. The present study is the first to examine how long-term variation in heterospecific density in breeding patches may affect dispersal patterns in spatially structured populations. We investigate how the dispersal decisions of the great crested newt (Triturus cristatus) are related to the variable density of two other newt species, the alpine newt (Ichthyosaura alpestris) and the palmate newt (Lissotriton helveticus). To examine this issue, we used capture–recapture data collected in an experimental pond network over a 20-year period. The results revealed that the great crested newt’s dispersal is context dependent and is affected by variation in heterospecific density: individuals were less likely to emigrate from ponds with high heterospecific density and were more likely to immigrate to ponds with high heterospecific density. These findings suggest that individuals adjust their dispersal decisions at least partly based on public information provided by heterospecifics. This mechanism may play a critical role in the dynamics of spatially structured populations and community functioning.

     

High aquatic niche overlap in the newts Triturus cristatus and T. marmoratus (Amphibia,Urodela)

We studied spatial niche metrics of large-bodied newts (Triturus cristatus and T. marmoratus) in three breeding ponds in western France. Adults and larvae were sampled with underwater funnel traps. Larvae were identified to the species with diagnostic microsatellite DNA markers. The distribution of adult T. cristatus and T. marmoratus across pond regions differed in one out of six cases, no differences were observed between larvae (two ponds studied). Niche overlap and niche breadth indices across resource states defined as pond regions or individual traps were high (Schoener's C: pond regions 0.60–0.98, traps 0.35–0.71; Levins' B: pond regions 0.71–0.98, traps 0.35–0.76). Adults of large-bodied newts significantly differed in resource use from small-bodied newts (T. helveticus). The results are discussed in view of the occurrence of interspecific breeding attempts, and the unpredictable ecological characteristics of newt breeding ponds.     

The application of eDNA for monitoring of the Great Crested Newt in the UK

Current ecological surveys for great crested newts are time‐consuming and expensive and can only be carried out within a short survey window. Additional survey methods which would facilitate the detection of rare or protected species such as the great crested newt (Triturus cristatus) would be extremely advantageous. Environmental DNA (eDNA) analysis has been utilized for the detection of great crested newts in Denmark. Here, the same methodology has been applied to water samples taken from UK ponds concurrently with conventional field surveying techniques. Our eDNA analysis exhibited an 84% success rate with a kappa coefficient of agreement between field and eDNA surveys of 0.86. One pond determined to be negative for great crested newt by field survey was positive by eDNA analysis, revealing the potential for improved detection rates using this methodology. Analysis of water samples collected in late summer indicates that eDNA analysis could be used to detect great crested newt after the optimal survey window for current field techniques had passed. Consequently, eDNA analysis could augment currently stipulated techniques for great crested newt surveying as a relatively quick and inexpensive tool for collecting great crested newt presence and distribution data within the UK instead of or prior to full field surveys.     

Limitation of similarity and feeding habits in three syntopic species of newts (Triturus, Amphibia)

Differences in body size between both species and sex were observed to be close to Hutchinsonian ratios in an assemblage of three species of newts (Triturus italicus, T alpestris and T carnifex) in southern Italy According to feeding, differences in body size were not related with a strict food niche segregation The diet of larger newt included indeed all the prey eaten by smaller newts but showed new prey types Differences between species were more marked than between sexes of the same species, despite a strong sexual dimorphism in body size Larger newts foraged in a wider range of microhabitats and exhibited a greater diversity of foraging tactics Triturus italicus fed essentially on plankton and chironomids in shallow water at the banks of the pond Triturus alpestris foraged also in deeper water, particularly on lsopods in vegetation Prey fallen on the water surface constituted the main part of T carnifex diet According to recent data on newt evolution, speciation consisted in the reduction of body size Our data show that this process has been related with a specialization of the diet and of habitat use     

Modeling spatial distribution of amphibian populations: a GIS approach based on habitat matrix permeability

Predictions of occurrence of two amphibian species, the common toad and the alpine newt, were made using information on land use surrounding breeding ponds. A geographical information system (GIS) was used to compile a landuse map, from which permeability estimates (friction) were derived. Potential migration zones based on friction and maximum migration distance were then modeled. Contacts between several migration zones suggest the potential for migration between ponds by adult individuals. The ability of the migration zones to enhance predictions of species occurrence was tested using generalized additive models (GAMs), and several landscape variables were selected as determinants of amphibian presence. The area of a migration zone and the number of ponds within that zone were positively related to both toad and newt presence, suggesting the importance of buffer habitats around each pond in amphibian conservation. Toad presence declined with cultivated field area and newt presence declined with vineyard area, suggesting the negative effect of agricultural activity on amphibians. The friction-based modeling approach improved the prediction of toad presence when compared to a more classical analysis of habitat composition within a circular zone centered on the focal pond. Prediction of newt presence was, however, less accurate than prediction of toad presence. Despite its exploratory nature and the subjectivity of permeability estimates, this study shows the usefulness of GIS in the functional analysis of a landscape, with potential applications in biological conservation. It also highlights the need for improving our knowledge of habitat use by animals.     

New polymorphic microsatellite loci for the Macedonian crested newt, Triturus macedonicus, and cross-priming testing in four other crested newt species

Eleven microsatellites have been characterized for Triturus macedonicus. Nine loci showed different variation patterns in a sample of 40 individuals from a single breeding pond in Zagori province (Greece), with an average number of 4.2 alleles per locus and an expected heterozygosity ranging from 0.1199 to 0.8079. Distinct cross-priming amplification rates were recovered on four additional crested newt species. Two monomorphic T. macedonicus loci were polymorphic in other Triturus species. The microsatellites developed herein could be a useful intraspecific genetic tool to undertake fine-scale population genetic analyses as well as in the study of contact zones between crested newt species.     

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.     

Spatial and temporal variability of aquatic habitat use by amphibians in a hydrologically modified landscape

1. Habitat loss is a major driver of biodiversity decline worldwide. Temporary waterbodies are especially vulnerable because they are sensitive both to human impact and to climatic variations. Pond‐breeding amphibians are often dependent on temporary waterbodies for their reproduction, and hence are sensitive to loss of temporary ponds. 2. Here we present the results of a 5‐year study regarding the use of temporary aquatic habitats by amphibians in a hydrologically modified area of Eastern Europe (Romania). The annual number of aquatic habitats varied between 30 and ～120. Each aquatic habitat was characterised by a number of variables such as: ‘type’ (pond, drainage ditch and archaeological ditch), ‘hydroperiod’ (number of weeks the ponds were filled in a given year), ‘depth’ (cm), ‘area’ (m²) and the density of predatory insects (‘predation’). The turnover rate for each amphibian species for each wetland was calculated based on the pond occupancy. 3. Eight amphibian species were recorded from the aquatic habitats. Hydroperiod was the most important variable, positively influencing wetland use by amphibians and their reproductive success. Most species preferred drainage ditches for reproduction, and the reproductive success was highest in this habitat type every year. For most of the species, the local extinction rate was higher than the colonisation rate in the first 4 years, but the situation reversed in the last year of the study when wetland use by amphibians sharply increased because of high rainfall. 4. This study confirms the importance for amphibians of maintaining and managing aquatic habitat diversity at small spatial scales. Man‐made aquatic habitats such as drainage ditches may be important habitats for amphibians, and this should be considered in restoration activities.     

Effects of forest degradation on butterfly communities in the Gwangneung Forest

This study was carried out to clarify the response of butterfly communities on forest degradation in the Gwangneung Forest, Korea. We monitored butterfly communities with varying degrees of human activities by conducting a line transect twice a month in 2011. A total of 70 species and 4676 individuals of butterflies were observed in four sites: natural forest (NR), two plantation forests, and the Korean National Arboretum (AR). The result on niche breadth, habitat breadth and habitat type of butterfly was not consistent with our predictions. Species richness of habitat type was only significantly different between NF and AR. Species diversity was significantly different among sites. Butterfly diversity associated with landscape patterns based on aerial photographs supported a mosaic concept. A forest management plan to conserve butterfly diversity in forests is necessary to maintain various habitats and to ensure that grasslands are protected.     

Abundance–range size relationships in the herbaceous flora of central England

1 Using data from a survey of over 10 000 1-m² quadrats in a 3000-km² area, we examined the relationship between abundance and range for the vascular plant flora of central England.
2 At the level of the whole landscape, abundance was not related to local, regional or national range. Local, regional and national range were closely related to each other.
3 At the level of the whole landscape, range was significantly and positively related to both niche breadth (expressed as the range of habitats exploited) and to habitat availability, although niche breadth appeared to be more important. Abundance was not related to niche breadth or habitat availability. Since specialist species are mainly confined to uncommon habitats (especially wetlands), we conclude that the relationship between range and niche breadth is not an artefact of widespread species passively sampling more habitats.
4 At the level of individual habitat types, significant positive relationships between range and abundance were common. These relationships remained after controlling for the effects of phylogeny. For predominantly annual weed communities, the relationship was linear, but for perennial communities it was markedly 'upper triangular', i.e. all combinations of range and abundance were found except wide range/low abundance. The evidence suggests that this difference can be attributed to the greater mobility of annual weeds.     

若尔盖不同生境植物多样性与植物竞争强度和生态位重叠度的关系

植物多样性与植物竞争强度和生态位重叠度的关系会随环境发生变化。为探究上述关系在若尔盖地区的表现形式,于2021年8月对若尔盖地区典型的水生、湿生、湿生-中生和中生植物群落进行调查,构建了新的植物竞争强度（Competition intensity,CI）公式并进行测算,计算了植物群落的植物多样性指数（包括物种丰富度、Shannon-Weiner指数、Simpson指数和Pielou均匀度指数）和生态位重叠度（Niche overlap of species,NOS）,分析了植物群落物种多样性指数与CI和NOS的关系。结果表明：1）从水生到中生生境,植物多样性指数均呈增加趋势（P<0.05）;2）湿生-中生生境的CI显著高于湿生生境（P<0.05）,湿生生境的NOS高于水生生境（P<0.05）;CI与NOS无显著相关性,但在湿生生境中两者呈倒抛物线关系（P<0.05）。3）整体来看,植物群落的物种丰富度与CI呈抛物线关系（P<0.05）,与NOS无显著关系（P>0.05）;Shannon-Weiner指数、Simpson指数和Pielou均匀度指数均与NOS呈线性正相关（P<0.05）,与CI无显著关系（P>0.05）;从单个生境看,湿生-中生生境的Simpson指数、Shannon-Weiner指数和Pielou均匀度指数与CI呈线性负相关（P<0.05）,其余生境的上述多样性指数与CI无显著关系（P>0.05）;各生境的植物多样性指数均与NOS无显著相关性（P>0.05）。本研究表明,从水生到中生生境,若尔盖地区的植物多样性呈增加趋势,但植物多样性与物种竞争强度和生态位重叠度的关系较复杂。本研究结果有助于理解若尔盖高原植物多样性的形成机制。     

Hydroperiod,predators and the distribution of physid snails across the freshwater habitat gradient

1. Studies of species distributions across environmental gradients further our understanding of mechanisms regulating species diversity at the landscape scale. For some freshwater taxa the habitat gradient from small, shallow and temporary ponds to large, deep and permanent lakes has been shown to be an important environmental axis. Freshwater snails are key players in freshwater ecosystems, but there are no comprehensive studies of their distributions across the entire freshwater habitat gradient. Here we test the hypothesis that snail species in the family Physidae are distributed in a non‐random manner across the habitat gradient. We sampled the snails, their predators and the abiotic environment of 61 ponds and lakes, spanning a wide range in depth and hydroperiod. 2. Temporary habitats had the lowest biomass of predators. Shallow permanent ponds had the highest biomass of invertebrate predators but an intermediate fish biomass. Deep ponds and lakes had the highest fish biomass and intermediate invertebrate biomass. Five species of physids occurred in the regional species pool and 60 of the 61 ponds and lakes surveyed contained physid snails. Each pond and lake contained an average of just 1.2 physid species, illustrating limited membership in local communities and substantial among‐site heterogeneity in species composition. 3. Physids showed strong sorting along the habitat gradient, with Physa vernalis found in the shortest hydroperiod ponds and Aplexa elongata, P. gyrina, P. acuta and P. ancillaria found in habitats of successively greater permanence. When organised into a site‐by‐species incidence matrix with sites ordered according to their hydroperiods, we found the pattern of incidence to be highly coherent, showing that much of the heterogeneity in species composition from one pond to another is explained by hydroperiod. We also found that the number of species replacements along this gradient was higher than random, showing that replacement is more important than nesting in describing species composition in ponds of different hydroperiod. 4. Discriminant analysis showed that pond depth, invertebrate biomass and fish biomass were the best predictors of species composition. Analysis of these niche dimensions showed that P. vernalis and A. elongata were most successful in shallow, temporary ponds with few predators. P. gyrina and P. acuta were typically found in ponds of intermediate depth and high predator abundance. P. ancillaria was found in the deepest lakes, which had abundant fish predators but few invertebrate predators. Of the five species considered, P. ancillaria, P. vernalis and A. elongata were relatively specialised with regard to key habitat characteristics, P. gyrina was moderately generalised and P. acuta was remarkably generalised, since it alone occurred across the entire freshwater habitat gradient. The exceptional habitat breadth of P. acuta stands in contrast to distributional studies of other freshwater taxa and deserves further attention.     

Habitat structure determines spider diversity in highland ponds

Wetlands (e.g. ponds, meadows) can be found in many landscapes, playing an important role in maintaining regional biodiversity and supporting heterogeneous communities. Spiders are diversified predators that are highly influenced by changes in plant community structure, heterogeneous habitats sustain high spider diversity and abundance. We investigated the characteristics of spider biodiversity in ponds with different habitat structures, by examining patterns across habitats of ponds with different vegetation levels. Sampling took place in four occasions over a year. We compared spider abundance, species richness and composition among ponds including distinct vegetation variables, related to life form, type of leaves, coverage and height. Overall 1174 individuals (194 adults) of 11 families and 37 morphospecies were sampled. We found mostly expected differences in the manner that communities were structured between different habitats. Thus, higher variability of abundance was explained for higher habitat structure of ponds. We also found differences in species composition between ponds with low emergent vegetation and higher habitat structures. Additionaly, spiders were consistently structured more by turnover than nestedness components, with a greater beta diversity of web-builders. Our results suggest varying levels of habitat structures and species substitution shape pond spider communities, depending on habitat heterogeneity and spider guild. Those findings demonstrate the clear role of spatial habitat structure, with more spider species preferring to build webs or actively hunt at vegetated environments on ponds.     

Waterbird assemblages and associated habitat characteristics of farm ponds in the Western Cape, South Africa

Natural freshwater wetlands are among the most threatened habitats on Earth. Effective wetland biodiversity conservation can not, however, be evaluated without fully understanding the roles of artificial waterbodies as refuges for water-dependent plants and animals. Waterbird assemblages were examined on 59 farm ponds in the Elgin and Caledon districts of the Western Cape, South Africa. This study examines the relationship between waterbird use and habitat characteristics of farm ponds. Patterns of temporal and spatial variation of waterbird species richness and abundance were quantified in relation to the habitat characteristics of each pond. Cluster analysis and multiple regression analyses identified surface area of the farm ponds as an important variable determining the presence and abundance of many waterbird species. Structural diversity in terms of vegetation in and around the ponds was especially important in determining their usage by waterbirds. These variables were evaluated in terms of creating a mosaic of habitat types (by varying vegetation structure and pond topography), which may be a useful way to enhance waterbird diversity at farm ponds. This study concludes that the high number of farm ponds in the transformed habitat matrix of the Western Cape plays an important role in conserving waterbirds.     

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.     

Population structure and genetic diversity of the threatened pygmy newt Triturus pygmaeus in a network of natural and artificial ponds

Pond physical characteristics (connectivity, hydroperiod) have shown to be highly relevant in explaining species presence, reproductive success, and survival in breeding-pond amphibians. However, few studies have addressed the influence that these factors may have on the genetic variability of pond populations. We examined genetic variation at 11 microsatellite loci in Iberian endemic, the pygmy newt (Triturus pygmaeus), from 58 breeding ponds in the Doñana National Park (Andalusia), including both temporary ponds and artificially deepened ponds that remain wet during the whole year. Temporary ponds are located in the North part of the region where the surrounding habitat-wet meadows-facilitates the connectivity among populations, whereas the deepest ponds (‘zacallones’) are located in the southern edge embedded in a matrix of unsuitable habitat (thickets and dry underbrush). We investigated genetic diversity and structure within and among ponds. Our results show that both regions (Doñana-North and Doñana-South) are well-differentiated and form two main clusters. We found higher genetic diversity within ponds from the North region, which also exhibited a higher degree of genetic admixture in comparison with populations from the southern edge. Although we found an isolation-by-distance pattern within each cluster, it arose due to the effect of a few isolated ponds located on the edge of each zone, suggesting the existence of substantial gene flow between ponds in the core area. According to our findings, landscape’s permeability to movement (pond connectivity) may constitute a more important factor than hydroperiod length in determining the genetic diversity and viability of pygmy newt populations in this area. Although Doñana populations show a good state, more peripheral and isolated populations present a more worrisome condition as a result of fragmentation and thus, require conservation efforts. Our study provides key insights that could help guide management practices of this threatened and poorly-studied salamander.