Designing Wetlands for Amphibians: The Importance of Predatory Fish and Shallow Littoral Zones in Structuring of Amphibian Communities

Under section 401 and section 404 of the Clean Water Act, permission to degrade existing natural wetlands in the USA may be conditional on restoring or creating ‘replacement’ wetlands. Success of wetland mitigation efforts in adequately replacing lost wildlife habitats depends on our good understanding of key ecological attributes that affect the structure of wetland faunal communities. We examined the effects of the presence of predatory fish, shallow vegetated littoral zone, emergent vegetation cover, wetland age and size on amphibian diversity in 42 replacement wetlands located in the Ohio’s North Central Tillplain ecoregion. We recorded 13 species of pond-breeding amphibians, and the average local species richness (α-richness) was 4.2 ± 1.7 species per site (range 1–7). There is strong evidence for the positive association between amphibian species richness and presence of a shallow littoral zone, and the negative association with presence of predatory fish. There was no evidence for the association between species richness and age, size, amount of forest cover within 200 m, nor the amount of emergent vegetation cover at the study sites. It is estimated that local species richness in wetlands with shallows was 1.76 species higher on average than in wetlands without shallows (95% CI from 0.75 to 2.76). The presence of predatory fish was associated with an average reduction in species richness by an estimated 1.21 species (95% CI from 0.29 to 2.11). Replacement wetlands were placed in areas with little or no existing forest cover, and amphibian species associated with forested wetlands were either rare (eastern newt, spotted salamander) or not present at all (marbled salamander, wood frog). In addition, we surveyed all replacement wetlands constructed under section 401 in Ohio since 1990, and found that predatory fish were present in 52.4% of the sites and that shallows were absent from 42.7% of the sites. Our results indicate that current wetland replacement practices could have a negative effect on the amphibian diversity within our region.     

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.     

Amphibian species richness across environmental gradients

Large‐scale field patterns are a fundamental source of inferences on processes responsible for variation in species richness among habitats. We examined species richness of larval amphibian communities in 37 ponds over seven years on the Univ. of Michigan's E. S. George Reserve. Ordination of the community incidence matrix indicated a strong major axis of variation in species associations that was correlated with pond hydroperiod, surface area and forest canopy cover. Communities were significantly nested with those species found in ponds with high canopy cover, small area and short hydroperiod being nested subsets of those found in ponds with contrasting characteristics. Presence of fish had strong negative effects on species richness; relaxation of this effect also was apparent when fish were extirpated from ponds by drought. We employed a model selection analysis to identify the most appropriate statistical model for predicting the long‐term average species richness of these ponds from local abiotic and biotic (predator and competitor density) factors. A model including only the abiotic factors was overwhelmingly superior for the anurans; hierarchical partitioning indicated that area and canopy cover alone accounted for over 70% of the independent effects of predictor variables. The global model including both abiotic and biotic factors was the best supported model for the caudates, and correspondingly hierarchical partitioning suggested that area, hydroperiod, invertebrate predators and caudate biomass all accounted for 9–16% of the independent effects. Overall, biotic factors accounted for much less of the variation in species richness than abiotic factors. The patterns in larger, open‐canopy ponds provided little evidence of competitive effects on species richness, though there were patterns consistent with competitive effects in small, closed‐canopy ponds. The unusual temporal and spatial extent of these data enabled us to critically evaluate ideas regarding patterns in larval amphibian communities, and the effects of area, disturbance (hydroperiod) and productivity (canopy cover) on species richness of these communities. These results have important implications to the conservation of amphibian species richness in freshwater wetlands, which are among the most threatened ecosystems worldwide.     

Intermediate Pond Sizes Contain the Highest Density,Richness, and Diversity of Pond-Breeding Amphibians

We present data on amphibian density, species richness, and diversity from a 7140-ha area consisting of 200 ponds in the Midwestern U.S. that represents most of the possible lentic aquatic breeding habitats common in this region. Our study includes all possible breeding sites with natural and anthropogenic disturbance processes that can be missing from studies where sampling intensity is low, sample area is small, or partial disturbance gradients are sampled. We tested whether pond area was a significant predictor of density, species richness, and diversity of amphibians and if values peaked at intermediate pond areas. We found that in all cases a quadratic model fit our data significantly better than a linear model. Because small ponds have a high probability of pond drying and large ponds have a high probability of fish colonization and accumulation of invertebrate predators, drying and predation may be two mechanisms driving the peak of density and diversity towards intermediate values of pond size. We also found that not all intermediate sized ponds produced many larvae; in fact, some had low amphibian density, richness, and diversity. Further analyses of the subset of ponds represented in the peak of the area distribution showed that fish, hydroperiod, invertebrate density, and canopy are additional factors that drive density, richness and diversity of ponds up or down, when extremely small or large ponds are eliminated. Our results indicate that fishless ponds at intermediate sizes are more diverse, produce more larvae, and have greater potential to recruit juveniles into adult populations of most species sampled. Further, hylid and chorus frogs are found predictably more often in ephemeral ponds whereas bullfrogs, green frogs, and cricket frogs are found most often in permanent ponds with fish. Our data increase understanding of what factors structure and maintain amphibian diversity across large landscapes.     

Effects of an introduced crayfish, Procambarus clarkii, on the distribution of south-western Iberian amphibians in their breeding habitats

The introduction of exotic species in aquatic habitats is one of the causes for the amphibian declines observed worldwide. In the 1970s, the red swamp crayfish Procambarus clarkii was introduced in the southwest Iberian Peninsula, where no native crayfish occur. In this study we assess the effect of P. clarkii presence in the breeding site distribution of each of the 13 southwest Iberian amphibians, while simultaneously accounting for the effects of potentially confounding habitat variables, as well as for the effects of the other large aquatic predators in the study area – predatory fish. Amphibian species richness was lower in places where P. clarkii was present than in places without P. clarkii , regardless of fish presence. After accounting for habitat variables and fish, crayfish presence was a negative predictor of the breeding probability for all urodeles ( Pleurodeles waltl , Salamandra salamandra , Triturus boscai and T. marmoratus ) and for two anurans ( Pelobates cultripes and Bufo bufo ). The majority of the species affected usually breed in temporary ponds without fish, but that may be colonized by the crayfish. The ongoing expansion of P. clarkii may eventually lead to a growing isolation of amphibian populations and ultimately to local extinctions and a permanent alteration of the amphibian communities in southwest Iberian Peninsula.     

Habitat associations of pool-breeding amphibians in eastern Massachusetts,USA

There is a pressing need to develop a sound conservation strategy for pool-breeding amphibians, which includes gaining a better understanding of the habitat and landscape-scale characteristics associated with populations. To investigate relationships between amphibian species richness and characteristics of breeding pools and surrounding landscapes, we surveyed 85 pools in eastern Massachusetts (USA) in 1996 and 1997. A total of 11 species was detected, with most pools having 2–5 species. Pools were typically small, 77.6% were <0.2 ha, but most pools (72%) had hydroperiods that persisted at least into August in most years. Based on linear regression analyses, species richness was positively associated with three within-pool variables (pool surface area, hydroperiod, and the amount of emergent vegetation), and a landscape-level variable (presence of another breeding pool within 1 km), while one within-pool variable (tree canopy cover) exhibited a significant negative association with species richness. These within-pool habitat variables and connectivity to other breeding pools are important characteristics to consider when attempting to identify breeding sites that could provide core habitat in conservation reserves designed for the conservation of pool-breeding amphibian species richness. Conservation reserves for pool-breeding amphibian populations should include pool complexes functioning as habitat for metapopulations. Core pool habitats within such reserves should be large (0.5–1.0 ha), with seasonal hydroperiods that persist into August or that dry in some years, and with sufficient emergent vegetation to provide diverse microhabitats and refugia.     

The relationship between wetland hydroperiod and nestedness patterns in assemblages of larval amphibians and predatory macroinvertebrates

Assemblages exhibit nested distributional patterns if the species found in species-poor locations also occur in progressively richer locations. We investigated patterns of nestedness in assemblages of larval amphibians and predatory macroinvertebrates in 42 isolated freshwater wetlands in southern New Hampshire, USA. These wetlands varied markedly in hydroperiod and we predicted that nestedness would be relatively weak because changes in disturbance processes (the relative threat of desiccation and predation) along the hydroperiod gradient often generate distinct assemblages. Contrary to expectations we found that both amphibian and macroinvertebrate assemblages were strongly nested not only with respect to species richness but also with respect to hydroperiod and wetland size, which were positively correlated. We attribute our results to the increased colonization rates and decreased extinction rates associated with increasing hydroperiod, and to concomitant increases in wetland size, habitat heterogeneity/complexity, and possibly water temperature. Moreover, the impact of predatory fishes on species richness and composition of amphibians and macroinvertebrates was relatively minor. We found that amphibians had a significantly lower degree of nestedness than macroinvertebrates, suggesting that a higher proportion of amphibian species found in species-poor assemblages was unlikely to occur in species-rich assemblages of amphibians (e.g. wood frogs and spotted salamanders). The degree of nestedness appeared to be influenced primarily by hydroperiod and wetland size for amphibians, whereas nestedness of macroinvertebrates was influenced by unknown factors (possibly water temperature) in addition to hydroperiod and wetland size. The high degrees of nestedness observed in amphibian and macroinvertebrate assemblages imply that protection of larger, more permanent wetlands may be more important for conserving native biological diversity than protection of smaller, non-permanent wetlands. However, non-permanent wetlands are used by several species of conservation concern that often do not occur in larger and more permanent wetlands.     

Impact of three aquatic invasive species on native plants and macroinvertebrates in temperate ponds

Biological plant invasions pose a serious threat to native biodiversity and have received much attention, especially in terrestrial habitats. In freshwater ecosystems impacts of invasive plant species are less studied. We hypothesized an impact on organisms from the water column and from the sediment. We then assessed the impact of three aquatic invasive species on the plants and macroinvertebrates: Hydrocotyle ranunculoides, Ludwigia grandiflora and Myriophyllum aquaticum. Our research on 32 ponds in Belgium indicated that the reduction in the native plant species richness was a common pattern to invasion. However, the magnitude of impacts were species specific. A strong negative relationship to invasive species cover was found, with submerged vegetation the most vulnerable to the invasion. Invertebrate richness, diversity and abundance were measured in sediments of invaded and uninvaded ponds along a gradient of H. ranunculoides, L. grandiflora, and M. aquaticum species cover. We found a strong negative relationship between invasive species cover and invertebrate abundance, probably due to unsuitable conditions of the detritus for invertebrate colonization. Taxonomic compositions of aquatic invertebrate assemblages in invaded ponds differed from uninvaded ponds. Sensitive benthos, such as mayflies were completely absent in invaded ponds. The introduction of H. ranunculoides, L. grandiflora, and M. aquaticum in Belgian ponds has caused significant ecological alterations in the aquatic vegetation and the detritus community of ponds.     

Is livestock grazing compatible with amphibian diversity in the High Mountains of Córdoba, Argentina?

The High Mountains of Córdoba, Argentina have a long evolutionary history of grazing by large herbivores. However, about 400?years ago, European livestock were introduced and gradually replaced native herbivores. Since the 1920s, domestic herbivores have been the only large herbivores present in the area, causing severe soil erosion and a threat to the system diversity. The endemic fauna of the region includes four amphibian species. We evaluated the effect of livestock rearing on amphibian diversity and water bodies in woodlands and grasslands of the High Mountains of Córdoba. The work was conducted on stream stretches and ponds in two contrasting grazing situations: an area with livestock presence and another area where livestock was excluded 14?years ago. The following variables were recorded at each sampling site: amphibian richness and abundance, percentage of emergent, submerged and peripheral vegetation in areas surrounding the water bodies, water pH, and water dissolved O₂. No significant differences were detected in amphibian diversity between streams of both grazing situations, whereas a greater diversity (p?<?0.01) was observed in ponds in grazed grasslands. Our results suggest that livestock rearing, qualitatively measured as grazing and 14?years of livestock exclusion, in the study area would not have negative effects on amphibian diversity. This finding might be due to the long evolutionary grazing history of the area, large-scale livestock exclusion exhibiting a novel scenario. The ongoing reintroduction of native grazers may provide the benefits of grazing without the consequent soil erosion and habitat degradation associated with domestic livestock.     

The introduced fish, rotan (Perccottus glenii), depresses populations of aquatic animals (macroinvertebrates, amphibians, and a fish)

The fish rotan (Perccottus gleniiDybowski) was accidentally introduced into European Russia from the Amur River basin. Rotan is capable of colonising small waterbodies – favourable breeding sites of native amphibians. To reveal its influence on the native aquatic fauna, monitoring of small waterbodies has been carried out since 1994 in the region of Lake Glubokoe Reserve (Moscow Province, Russia). The fish's diet includes a wide range of animal species of all trophic levels. Rotan considerably decreases the species richness of aquatic macroinvertebrates and larval amphibians. As a rule, most amphibian species (Triturus cristatus, T. vulgaris, Rana temporaria, R. arvalis, R. lessonae) and the fish Carassius carassius failed to breed successfully in ponds inhabited by rotan. In contrast, the toad Bufo bufo bred successfully in such sites because its larvae are distasteful to rotan. Rotan–amphibian interactions are discussed.     

Pathways of ornamental and aquarium fish introductions into urban ponds of Epping Forest (London,England): the human vector*

To examine the role of humans in the non‐native fish introductions, we measured the frequency of occurrence and density of non‐native fishes in ponds (Epping Forest, Essex, England) that had been restored (drained of water and voided of fish or treated with rotenone) on a known date and into which no piscivorous or non‐native fishes had subsequently been stocked intentionally. For each pond, the period of time since pond restoration, pond area, distance to nearest residential housing, distance to nearest footpath, distance to nearest water body or stream, and the proportion of pond vegetated were measured. The occurrence of both non‐native and unexpected native fish species was non‐random, and the number of ornamental varieties was found to increase as pond distance from the nearest road decreased. Variety richness of each of three categories of fish (non‐native, goldfish Carassius auratus and native) was significantly correlated with at least two of the following variables: distance from nearest road, nearest footpath and nearest pond. The rate of non‐native fish introductions (adjusted variety richness per year) could also be estimated from pond distance to the nearest road, being about 3.5 ornamental varieties introduced per year in ponds adjacent to roads, but the rate appears to be much greater in ponds that had recently (<1.5 years) undergone restoration. Implications for conservation and management, as well as the potential role of societal issues such as recreational activities, cultural and religious practices, are discussed.     

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.     

农业景观中环境因素对两栖类生物分布的影响

以黄河下游地区封丘县为研究区,在地理信息系统和R软件的支持下,在栖息地水平和景观水平上,分析了两栖类生物分布与栖息地变量(池塘水体溶解氧DO、电导率COND、浊度TURB、水体氨氮NH₃-N、池塘内植被盖度VEG_interior、池塘内挺水植物盖度EMER、池塘边缘植被盖度VEG_edge、两栖类幼虫捕食者PRE和池塘面积AREA)和采样点池塘250、500、1000、2000 m半径缓冲区内景观变量(池塘面积AP、与最近道路的距离DR、耕地面积AF、林地面积AW、Shannon多样性指数SHDI和蔓延度指数CONTAG)的关系.结果表明: 中华大蟾蜍的分布主要与栖息地变量中的PRE和较大缓冲区半径内的景观变量(AW2000、DR2000、SHDI1000、CONTAG1000)相关;黑斑侧褶蛙的分布与栖息地变量中的EMER和较小缓冲区半径内的景观变量(SHDI500)关系密切;金线侧褶蛙的分布主要与栖息地变量中的VEG_interior和PRE相关;泽陆蛙的分布主要与较大缓冲区半径内的景观变量(AF2000、SHDI2000)相关;两栖类生物总体上主要与栖息地变量中的EMER和较小缓冲区半径内的景观变量(SHDI500)相关.在农业景观中,保护两栖类生物应该保护永久性和半永久性池塘、提高池塘内部植被盖度、增强栖息地池塘之间的功能连通性.     

Species richness patterns of amphibians in southwestern Ontario ponds

Abstract. In southwestern Ontario amphibian species richness (α-diversity) was investigated at 180 ponds from 1992 to 1994. Patterns of species richness were compared among regions and the relationship between species richness and local habitat and regional landscape variables was investigated. Patterns of incidence differed among regions, with species that use woodlands being rare in one of the regions. Repeated measures analysis of variance indicated that species richness differed significantly among regions but not among sub-regions nested within regions. Species richness did not change significantly over time and there was no region by year effect. Species richness was highly correlated with local variables related to fish predation and to regional variables related to forest cover. Multiple regression indicated that a combination of local and regional variables best accounted for the variance in species richness, but the amount of regional woodlands was the single most important variable. The pattern of species richness can be explained by historical deforestation as the primary process.     

Environmental influences on fish assemblages in irrigation ponds

Ponds are common features of the landscape and are considered important for freshwater biodiversity conservation. Although fish have a significant impact on the lentic ecosystems, the environmental factors that regulate fish assemblages in human-created water bodies, such as irrigation ponds, remain unclear. We evaluated the relationship between environmental factors and the fish assemblage structure in 31 ponds located in northern Japan. Species richness (range: 1–9) was positively correlated with the size of the inflow channel. Multivariate analyses revealed that the size of the inflow channel was a better predictor for species richness than lake morphology (surface area and maximum depth), vegetation coverage, water quality (turbidity, pH, DO, and EC), distance to the main channel, and distance to an adjacent pond. Species richness was significantly different between ponds with and without an inflow channel. Furthermore, three of the four most commonly observed species are thought to be relatively tolerant to low oxygen. Given that ponds have a relatively high local extinction rate resulting from exposure to stressful conditions, such as low oxygen and/or small population sizes, our results suggest that immigration from surrounding water bodies plays an important role in maintaining species richness of pond-dwelling fish.     

Dragonfly species richness on man-made ponds: effects of pond size and pond age on newly established assemblages

We studied the abundance and species richness of adult dragonflies in 11 artificial ponds which were recently established (within 2years). We found that the adult dragonfly assemblage patterns were influenced by pond size as well as pond age. The species richness was positively correlated with the pond size, which was because the distributional patterns of species were significantly nested according to pond area. The species richness was highly correlated with pond age in association with the vegetation cover within ponds. It was suggested that the species richness was enhanced by the increasing immigration rate of species which favor well-vegetated ponds.     

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.     

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.     

Direct and indirect effects of predation on tadpole community structure in the Amazon rainforest

Abstract The relationship between the distribution of predators (fish, odonates and water beetles) and prey assemblages (amphibian larvae) was investigated in the tropical rainforest of central Amazonas, Brasil. The anuran community uses a variety of waterbodies for reproduction, ranging from streams and streamside ponds to isolated forest ponds. Predators in this system include fish in streams and streamside ponds, and invertebrates (primarily odonate naiads and beetles) in forest ponds. Tadpole species richness and assemblage structure were related to fish density and species richness. No relationships between tadpole assemblages and abiotic pond characteristics were detected. The presence offish explained much of the variation in both species composition and species richness within and among ponds. Some species of tadpole were consistently found to coexist with high densities of fish. Path analyses suggest that while fish have a strong direct effect on tadpole associations and species richness, they also have an indirect effect through invertebrate predators (odonate larvae and coleopteran beetles). Prey survival-strategies such as palatability and behaviour may explain how tadpole species composition is affected by predators at the community level. These findings suggest that the observed patterns of habitat use by larval anurans may be structured in response to the distribution of key predators (fish) in this system.