Seeking shelter from the storm: Conservation and management of imperiled species in a changing climate

Climate change is anticipated to exacerbate the extinction risk of species whose persistence is already compromised by habitat loss, invasive species, disease, or other stressors. In coastal areas of the southeastern United States (USA), many imperiled vertebrates are vulnerable to hurricanes, which climate models predict to become more severe in the 21st century. Despite this escalating threat, explicit adaptation strategies that address hurricane threats, in particular, and climate change more generally, are largely underrepresented in recovery planning and implementation. We provide a basis for stronger emphasis on strategic planning for imperiled species facing the increasing threat of catastrophic hurricanes. Our reasoning comes from observations of short‐term environmental and biological impacts of Hurricane Michael, which impacted the Gulf Coast of the southeastern USA in October 2018. During this storm, St. Marks National Wildlife Refuge, located along the northern Gulf of Mexico's coast in the panhandle region of Florida, received storm surge that was 3.0–3.6 m (NAVD88) above sea level. Storm surge pushed sea water into some ephemeral freshwater ponds used for breeding by the federally threatened frosted flatwoods salamander (Ambystoma cingulatum). After the storm, specific conductance across all ponds measured varied from 80 to 23,100 µS/cm, compared to 75 to 445 µS/cm in spring 2018. For 17 overwashed wetlands that were measured in both spring and fall 2018, posthurricane conductance observations were, on average, more than 90 times higher than in the previous spring, setting the stage for varying population responses across this coastal landscape. Importantly, we found live individual flatwoods salamanders at both overwashed and non‐overwashed sites, although we cannot yet assess the demographic consequences of this storm. We outline actions that could be incorporated into climate adaptation strategies and recovery planning for imperiled species, like A. cingulatum, that are associated with freshwater coastal wetlands in hurricane‐prone regions.     

Climate and coastal dune vegetation: disturbance, recovery, and succession

The sand dune habitats found on barrier islands and other coastal areas support a dynamic plant community while protecting areas further inland from waves and wind. Foredune, interdune, and backdune habitats common to most coastal dunes have very different vegetation, likely because of the interplay among plant succession, exposure, disturbance, and resource availability. However, surprisingly few long-term data are available describing dune vegetation patterns. A nine-year census of 294 plots on St. George Island, Florida suggests that the major climatic drivers of vegetation patterns vary with habitat. Community structure is correlated with the elevation, soil moisture, and percent soil ash of each 1 m² plot. Major storms reduce species richness in all three habitats. Principle coordinate analysis suggests that changes in the plant communities through time are caused by climatic events: changes in foredune vegetation are correlated with temperature and summer precipitation, interdune vegetation with storm surge, and backdune vegetation with precipitation and storm surge. We suggest that the plant communities in foredune, interdune, and backdune habitats tend to undergo succession toward particular compositions of species, with climatic disturbances pushing the communities away from these more deterministic trajectories.     

Canopy closure and amphibian diversity in forested wetlands

The crowns of trees and shrubs often overtop temporary wetlands in forested regions. By shading pond basins, canopy can dramatically change the conditions experienced by residents such as amphibians. In this study, we estimated the presence of 8 amphibian species across 17 temporary wetlands at the Yale–Myers Forest in northeastern Connecticut, USA. In addition, we quantified the light environment using a grid of hemispherical canopy photographs to calculate Global Site Factor (GSF) within each wetland. Amphibian richness was low in most wetlands, and most wetlands were relatively shaded. Amphibian richness increased in lighter wetlands. This result was not confounded by relationships with wetland size. Most amphibian species tended to be absent from heavily shaded wetlands (‘open canopy specialists’). However, three species were often found in the shadiest wetlands (‘canopy generalists’). Field transplant experiments using one canopy generalist and one open canopy specialist showed that development of the generalist was less affected by wetland light levels compared with performance of the specialist. These findings suggest that canopy may be an important determinant of amphibian diversity patterns across wetlands. Further, conservation strategies dependent on universally applied, inviolate shoreline vegetation buffers may inadvertently contribute to species loss. Because species differ in their sensitivity to changes in canopy, these losses may be predictable.     

Spatial variation in anuran richness,diversity, and abundance across montane wetland habitat in Volcanoes National Park,Rwanda

The spatial distribution of species has long sparked interest among ecologists and biogeographers, increasingly so in studies of species responses to climate change. However, field studies on spatial patterns of distribution, useful to inform conservation actions at local scales, are still lacking for many regions, especially the tropics. We studied elevational trends and species‐area relationships among anurans in wetland habitats within Volcanoes National Park (VNP) in Rwanda, part of the biodiverse Albertine Rift region. In VNP, wetlands are key sites for anuran reproduction, and anurans are likely threatened by wetland desiccation which has occurred for the last few decades. Between 2012 and 2017, we sampled anuran communities in ten VNP wetlands located along an elevational gradient of c. 600 m (from 2,546 to 3,188 m a.s.l.) and found at least eight species, including at least two Albertine Rift Endemics. We show that species richness, diversity, and abundance likely decline with a decrease in wetland size and with an increase in elevation, though additional sampling (e.g., at night) might be needed to derive definite conclusions. Larger wetlands at lower elevations contained most species and individuals, which indicates the potential threat of wetland size reduction (through desiccation) for anuran conservation. However, we also found that wetlands differed in species composition and that some species (e.g., Sclerophrys kisoloensis) were likely restricted in distribution to only a few of the smaller wetlands—suggesting that the conservation of each individual wetland should be prioritized, regardless of size. We propose that all wetlands in VNP require additional conservation measures, which should be based on knowledge gathered through long‐term monitoring of anuran communities and research on drivers of wetland decline. Only such extended research will allow us to understand the response of anurans in VNP to threats such as climate change and wetland desiccation.     

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.     

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.     

Amphibian species richness and distribution in relation to pond water chemistry in south-western Ontario, Canada

1. We assessed the patterns of amphibian species richness and distribution in relation to water chemistry over a large geographical area in 1992–94.
2. Thirteen amphibian species were observed at 180 ponds, with mean species richness 3.5 ± 0.13 species per pond (range zero to nine). Water samples were collected from 143 ponds, analysed for fifteen chemical variables, and further analysed by multivariate statistical techniques.
3. Water in the study area was hard, alkaline and well-buffered against pH change, and most ponds were eutrophic. Amphibian species richness was negatively correlated with five chemical variables (chloride, conductivity, magnesium, total hardness, turbidity).
4. Principle components analysis reduced the data set to four chemical components that explained 65.4% of the variance in the original variables. Principle component scores were retained for use in further multivariate tests. Multiple regression accounted for only 19.0% of the variance in amphibian species richness. Discriminant function analysis (DFA) was used to determine if water chemistry variables discriminated among species, but it was only able to classify 17.5% of cases correctly. DFA was also used to determine if water chemistry distinguished between used and unused sites for individual species. DFA was moderately successful, classifying 61–77% of cases correctly.
5. General water chemistry appears to play only a minor part in affecting amphibian species richness in south-western, Ontario. However, chemical variables may be helpful to distinguish between used and unused sites for some species.     

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.     

Fish reduce anuran abundance and decrease herpetofaunal species richness in wetlands

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Aquatic macroinvertebrate assemblages in mitigated and natural wetlands

Many wetlands have been constructed in West Virginia as mitigation for a variety of human disturbances, but no comprehensive evaluation on their success has been conducted. Macroinvertebrates are extremely valuable components of functioning wetland ecosystems. As such, benthic and water column invertebrate communities were chosen as surrogates for wetland function in the evaluation of 11 mitigation and 4 reference wetlands in West Virginia. Mitigation wetlands ranged in age from 4 to 21 years old. Overall familial richness, diversity, density and biomass were similar between mitigation and reference wetlands (p > 0.05). Within open water habitats, total benthic invertebrate density was higher in reference wetlands, but mass of common taxa from water column samples was higher in mitigation wetlands (p < 0.05) Planorbidae density from benthic samples in emergent habitats was higher in reference than mitigated wetlands. Benthic Oligochaeta density was higher across open water habitats in mitigation wetlands. All other benthic taxa were similar between wetland types. Among the most common water column orders, Isopoda density was higher in reference wetlands, but Physidae density was higher in mitigation wetlands. Within mitigation wetlands, emergent areas contained higher richness and diversity than open areas. These data indicate that mitigation and reference wetlands generally support similar invertebrate assemblages, especially among benthic populations. The few observed differences are likely attributable to differences in vegetative community composition and structure. Mitigation wetlands currently support abundant and productive invertebrate communities, and as such, provide quality habitat for wetland dependent wildlife species, especially waterbirds and anurans.     

Relation between Water Balance and Climatic Variables Associated with the Geographical Distribution of Anurans

Amphibian species richness increases toward the equator, particularly in humid tropical forests. This relation between amphibian species richness and environmental water availability has been proposed to be a consequence of their high rates of evaporative water loss. In this way, traits that estimate water balance are expected to covary with climate and constrain a species’ geographic distribution. Furthermore, we predicted that coexisting species of anurans would have traits that are adapted to local hydric conditions. We compared the traits that describe water balance in 17 species of anurans that occur in the mesic Atlantic Forest and xeric Cerrado (savannah) habitats of Brazil. We predicted that species found in the warmer and dryer areas would show a lower sensitivity of locomotor performance to dehydration (SLPD), increased resistance to evaporative water loss (REWL) and higher rates of water uptake (RWU) than species restricted to the more mesic areas. We estimated the allometric relations between the hydric traits and body mass using phylogenetic generalized least squares. These regressions showed that REWL scaled negatively with body mass, whereas RWU scaled positively with body mass. Additionally, species inhabiting areas characterized by higher and more seasonally uniform temperatures, and lower and more seasonally concentrated precipitation, such as the Cerrado, had higher RWU and SLPD than species with geographical distributions more restricted to mesic environments, such as the Atlantic Forest. These results support the hypothesis that the interspecific variation of physiological traits shows an adaptation pattern to abiotic environmental traits.     

Drivers of reptile and amphibian assemblages outside the protected areas of Western Ghats,India

Biodiversity conservation in forested landscapes outside protected areas is important to sustain populations of species with restricted ranges. However, such habitats face many anthropogenic threats, including logging, extraction of firewood and leaf-litter for mulch in plantations. In this study, we determined the effects of forest degradation on amphibians and reptiles in forests outside protected areas by measuring their species richness and community composition across a disturbance gradient from near pristine to highly degraded forests in Agumbe, Western Ghats, India. Twenty-one strip 15 m × 150 m transects were laid across the disturbance gradient and diurnal visual encounter surveys were conducted. Sampling was repeated three times per transect covering the dry, intermediate and wet seasons. Amphibian and reptile communities were affected by the decrease in canopy cover and leaf litter volume, respectively. Our results indicate that the collection of firewood and leaf-litter can severely affect amphibian and reptile populations. Structured conservation planning outside of protected areas is therefore imperative.     

Odonata,Amphibia and Environmental Characteristics in Motorway Stormwater Retention Ponds (Southern France)

Water and its protection against pollution is an urgent priority for all countries around the world. In that context, France, through its Water Law in 1992 obliged the motorway companies to build stormwater retention ponds along roads in order to protect the water resource from transport pollution and to control water flow during rainstorms. We propose to evaluate how much these ponds can be attractive for aquatic species and then evaluate their role in regional biodiversity. Six retention ponds, localized in the Mediterranean region, were investigated during one year (March 2002–March 2003) for their chemical and biological characteristics such as bottom type, trace metal composition, water quality and phytocenose composition. These variables were recorded and correlated with species richness of amphibian and dragonfly communities. Stormwater retention ponds showed a high concentration of copper and zinc in top sediment layer and herbicides in water column. Dragonfly richness was higher in ponds with a natural bottom than ones with an artificial bottom (PEHD membrane) while amphibian richness was more sensitive to the structure of the surrounding landscape. These habitats appeared to be very attractive for Odonata species and tend to favorize uncommon species present in the survey region such as Ischnura pumilio and Erythromma viridulum. Amphibian were found to be representative of anthropophilous guild.     

Amphibian species richness and distribution in relation to pond water chemistry in south-western Ontario, Canada

1. We assessed the patterns of amphibian species richness and distribution in relation to water chemistry over a large geographical area in 1992–94.
2. Thirteen amphibian species were observed at 180 ponds, with mean species richness 3.5 ± 0.13 species per pond (range zero to nine). Water samples were collected from 143 ponds, analysed for fifteen chemical variables, and further analysed by multivariate statistical techniques.
3. Water in the study area was hard, alkaline and well-buffered against pH change, and most ponds were eutrophic. Amphibian species richness was negatively correlated with five chemical variables (chloride, conductivity, magnesium, total hardness, turbidity).
4. Principle components analysis reduced the data set to four chemical components that explained 65.4% of the variance in the original variables. Principle component scores were retained for use in further multivariate tests. Multiple regression accounted for only 19.0% of the variance in amphibian species richness. Discriminant function analysis (DFA) was used to determine if water chemistry variables discriminated among species, but it was only able to classify 17.5% of cases correctly. DFA was also used to determine if water chemistry distinguished between used and unused sites for individual species. DFA was moderately successful, classifying 61–77% of cases correctly.
5. General water chemistry appears to play only a minor part in affecting amphibian species richness in south-western, Ontario. However, chemical variables may be helpful to distinguish between used and unused sites for some species.     

Amphibian and reptile communities of rainforest fragments: minimum patch size to support high richness and abundance

Information on animal communities inhabiting Neotropical fragmented landscapes is important for developing conservation strategies. The structure of amphibian and reptile communities in six tropical rainforest fragments (<20?ha) and two reference areas in continuous forest at Los Tuxtlas, Mexico was studied. A total of 3,481 individuals of 51 species of amphibians and reptiles were recorded across 12 bimonthly surveys during 2 years. Taxonomic composition was different between the smallest fragments and the reference areas. Six species were exclusive to large undisturbed forest and richness was significantly lower in the five smallest fragments (1.4–6.6?ha) compared with the largest patch, one or both of the reference areas. Amphibian abundance tended to be higher in large areas, while reptiles were more abundant in the five smallest fragments. Craugastor loki and Anolis uniformis were the dominant species in all sites, and particularly in the smaller fragments. Amphibian and reptile richness was positively related to larger patch sizes, deeper leaf litter, closed canopy cover, and higher relative humidity and negatively related to linear patch shape and high temperatures. Abundance of reptiles was positively associated to high temperatures, high density of woody debris, and closed canopy cover; it was negatively affected by linear patch shape, low humidity levels, and steeper slopes. While amphibian and reptile communities were strongly affected in vegetation fragments, these patches retained a considerable number of rainforest species. Fragments up to at least 17?ha have the potential for preserving communities with similar structure to those occurring in large tropical rainforests.     

Decline of some west-central Florida anuran populations in response to habitat degradation

Recent reports have suggested that a global decline in amphibian populations has taken place during the past few decades. Urban development is thought to affect the richness and abundances of species and, therefore, could be an important cause of decline. We estimated the richness and abundances of anurans in wetlands at a residential development and in similar wetlands at a nearby undeveloped park. The residential development originally was pine flatwoods habitat, as is the undeveloped park curiently. We also compared the anuran species' composition of the park in 1992 with the composition in 1974.Both richness and abundances of anurans in the residential development were different than those in the undeveloped park. Employing the same amount of sampling effort at both sites, we trapped or observed 11 species at the developement and 15 species at the park, and trapped 569 individuals at the development and 1224 individuals at the park. The anuran species richness at the undeveloped park in 1992 was nearly the same as in 1974; a single rare species apparently was not present in 1992. Of the 15 species present in both surveys, 14 showed higher abundances in 1992 than in 1974.We suggest that the current differences between the residential development and the park have resulted from degradation of both the uplands used by many species during the dry season and the temporary wetlands used by many species for reproduction. Four species especially sensitive to such degradation, Bufo quercicus, Scaphiopus h. holbrookii, Hyla femoralis, and H. gratiosa, were the species missing from the residential development.Not all species of anurans typical of pine flatwoods appeared to be affected adversely by development. Three species of ranids, Rana utricularia, R. grylio, and R. catesbeiana, were found in higher abundances at the residential development than at the park. These ranid species breed in a wide variety of aquatic systems, including the permanent bodies of water that are now abundant in the development, and probably use the uplands less than other anurans.If amphibian decline is international in scope, then the decline could be attributable either to global changes caused by humans, or to local, but widespread, environmental degradation, or to a combination of factors. While much recent popular focus has been on potential global causes of decline, we believe that this emphasis may have caused attention to be taken away from local causes that, as our study demonstrated, may be at least as important. We suggest that in many places, local environmental degradation is insidiously chipping away at amphibian diversity, and that more emphasis should be placed on these local causes than is now the case.     

Relationships among Amphibian Assemblage Structure,Wetland pH,and Forest Cover

A common mesofilter approach to conservation of biological diversity and ecosystem function used in agricultural and urban landscapes is maintenance of wetlands and an undisturbed terrestrial buffer surrounding wetlands. Although it is generally accepted that forest buffers protect wetland-associated biological diversity and ecosystem function, the effectiveness and optimal spatial extent of buffers is still an area of debate. During 2007 and 2008 we surveyed amphibians and environmental conditions associated with 54 depression wetlands on the Delmarva Peninsula of Maryland, USA, to examine the role of forest buffers and wetland characteristics in structuring amphibian communities. Forest cover within a 50-m buffer surrounding wetlands was correlated (r = −0.81) with wetland pH but no other wetland characteristics. Wetland pH, canopy cover, hydroperiod, and adjacent forest cover were important predictors of wetland use by individual amphibian species, with many species more likely to occur at wetlands that dried late in the hydrological year and with open canopies. At least one common species preferred circumneutral pH and several restricted-distribution species preferred lower pH (<5). Contrary to expectations, relationships between species occurrence and adjacent forest cover were negative. Our results suggest that current regulations that provide buffers of 30 m or less do not provide adequate protection of wetland water chemistry but that forest encroachment into wetlands may be a threat to the integrity of amphibian communities and should be the target of monitoring, future research, and management efforts. © 2021 The Wildlife Society.     

Influence of priority effects and pond location on invaded larval amphibian communities

In Florida, the Cuban Treefrog (Osteopilus septentrionalis) is a superb colonist and appears to be a significant driver of amphibian community dynamics. Decline of native anurans has been linked to possible competition with adult O. septentrionalis but interactions during the larval stage are largely unknown. Rearing O. septentrionalis tadpoles along with two native anurans, the Squirrel Treefrog (Hyla squirella) and the Southern Toad (Bufo terrestris) in both experimental artificial ponds and laboratory aquaria, the role of competition as the mechanism driving the dynamics of invaded amphibian communities in Florida was examined. Also examined was the role of priority effects and variation between pond locations in altering interactions between O. septentrionalis and native anuran larvae. Interspecific competition was strong during the larval stage; the presence of O. septentrionalis reduced larval performance and survival of native anurans. Pond location alone had little effect on interspecific interactions, but priority effects were strong. Pond location and priority effects acted together to influence species interactions. The selective influence of different interaction modifiers acted to increase or decrease the impacts of exotic species on native taxa.     

Species richness and composition patterns of clitellate (Annelida) assemblages in the treeless spring fens: the effect of water chemistry and substrate

Spring fens are isolated treeless wetlands of a high conservation value. Their environmental conditions are strongly related to their groundwater chemistry, which controls species distribution within various groups of organisms. Clitellates, a dominant group of non-insect aquatic fauna, however, have never been studied in these habitats. It is unclear from previous studies to what extent the distribution of aquatic non-insect taxa reflects water chemistry rather than the substrate structure. We studied 34 spring fens sampled in 17 isolated sites in the Western Carpathian Mountains to determine mainly the effects of water chemistry and substrate structure on variation in species richness and composition of clitellate assemblages as examples of the non-insect fauna. A total of 34 taxa were found, with 3–15 taxa collected per sample. Species richness was negatively correlated with water mineral concentration measured as water electric conductivity (r = −0.57, P < 0.001) and positively with TOC (r = 0.60, P < 0.001). Surprisingly, the lowest number of taxa was found in calcareous fens and richness increased towards Sphagnum-fens. There was a species turnover related to changes in mineral richness and substrate characters. The main change of species composition was promoted by changes in substrate structure. The second gradient of species composition was linked with the amount of nutrients, moisture, and dominance of sphagna, and was associated with an increase of eurytopic species in fens with high nutrient availability. It was difficult to separate the effects of water chemistry and substrate on clitellate species distributions owing to the fact that variation in tufa precipitation and vegetation was driven by water chemistry changes. This study presented the first quantitative data on fen clitellate assemblages, which appear to have an unusual pattern of species richness. In contrast to plants and molluscs, calcareous fens appeared to be a harsh environment for clitellate species. Only few specialized species, mainly Trichodrilus strandi, were able to establish viable populations. The significant effect of water chemistry on clitellate distribution patterns raises questions about the direct influence of water chemistry on non-insect aquatic taxa, which have previously been considered to be mostly determined by substrate characteristics.     

Broad-scale spatial patterns of canopy cover and pond morphology affect the structure of a Neotropical amphibian metacommunity

Spatial and environmental processes influence species composition at distinct scales. Previous studies suggested that the distribution of larval anurans at the landscape-scale is influenced by environmental gradients related to adult breeding site selection, such as pond canopy cover, but not by water chemistry. However, the combined effects of spatial, pond morphology, and water chemistry variables on metacommunity structure of larval anurans have not been analyzed yet. We used a partial redundancy analysis with variation partitioning to analyze the relative influence of pond morphology (e.g., depth, area, and aquatic vegetation), water chemistry, and spatial variables on a tadpole metacommunity from southeastern Brazil. We predict that pond morphology and canopy cover will influence the metacommunity at broad spatial scales, while water chemistry would play a larger role at finer scales. We found that broad-scale spatial patterns of pond canopy cover and pond morphology strongly influenced metacommunity structure, with water chemistry being not significant. Additionally, species composition was spatially autocorrelated at short distances. We suggest that the reproductive behavior of adult anurans is driving tadpole metacommunity dynamics, since pond morphology, but not water chemistry affects breeding site selection by adults. Our results contribute to the understanding of amphibian species diversity in tropical wetlands.