Structure and dynamics of an amphibian metacommunity in two regions

1. The concept of metacommunity is based on the hypothesis that species occurrence depends on species dynamics and interactions on local and regional scales via the movements of individuals between localities. Metacommunity approaches are currently being applied to pond breeding taxa such as amphibians. 2. Given that animal movement is also influenced by the physical quality of the matrix to be crossed to reach a breeding habitat and by the affinity of the species for specific terrestrial habitats, matrix characteristics may enhance or hinder dispersal success. These characteristics would, in turn, affect the composition of larval assemblages at local level and, consequently, determine metacommunity structure and dynamics. 3. Here we compared the structures and dynamics of two metacommunities with the same pool of anurans along similar freshwater gradients in two regions that are well differentiated in terms of their respective terrestrial matrix. 4. Abundance of tadpole species and species assemblage in the two regions were determined principally by local processes (at pond level); however, the structure and dynamics of the communities differed. In one region species abundance was explained in part by landscape factors and consequently showed lower co-occurrence and lower colonization rates (species sorting models) indicating that terrestrial habitat could restrict animal movements, whereas in the other region higher co-occurrence and higher colonization rates (mass effect models) indicated low dispersal limitations.     

Evidence of stochasticity driving anuran metacommunity structure in the Pantanal wetlands

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Pond canopy cover: a resource gradient for anuran larvae

1. The gradient in pond canopy cover strongly influences freshwater species distributions. This study tested the effects of canopy cover on the performance of two species of larval anurans, a canopy cover generalist (Rana sylvatica, the wood frog) and an open‐canopy specialist (R. pipiens, the leopard frog), and tested which factors co‐varying with canopy cover mediate these effects. 2. A field transplant experiment demonstrated that canopy cover had negative performance effects on both species. However, leopard frogs, which grow faster than wood frogs in open‐canopy ponds, were more strongly affected by closed‐canopy pond conditions. 3. Closed‐canopy ponds had lower temperature, dissolved oxygen (DO), and food nutritional quality as indicated by carbon‐to‐nitrogen ratio (C : N) analysis of field‐sampled food types, and of gut contents of transplanted larvae. 4. Laboratory experiments demonstrated that higher temperature and food quality but not DO substantially increased larval growth. However, only food quality increased growth rates of leopard frogs more than wood frogs. 5. The strong correlation of growth rates to gut content C : N in the field, and the similarity of growth curves as a function of resource quality in the field and laboratory, strongly suggest that resources are of primary importance in mediating intraspecific, and especially interspecific differences in performance across the canopy cover gradient.     

Anuran tadpole assemblages in riparian areas of the Middle Paraná River, Argentina

We investigated habitat variables influencing the species richness and composition of anuran tadpole assemblages in ponds from riparian areas of the Middle Paraná River. Tadpoles were collected at seven ponds that differed in hydroperiod and 10 habitat variables (water temperature, pH, conductivity, dissolved oxygen, turbidity, depth, size, time of isolation, macrophyte richness, and level of natural or anthropic disturbances) during a 5 year period. The presence/absence of potential tadpole predators was also recorded. Although we did not examine tadpole diets, we categorized anurans into ecomorphological guilds. We found that dissolved oxygen, maximum depth, pond size, macrophyte richness, and disturbances were significantly related to tadpole species richness. We also found that the richness of potential tadpole predator increased with longer pond hydroperiod. Thus, the number of tadpole species was higher in temporary and semi-permanent ponds without potential tadpole predators. The results of our study provide the first quantitative data on the tadpole assemblages in ponds associated with riparian areas of the Middle Paraná River Floodplain based on multi-year intensive sampling, a finding of both ecological interest and practical significance for future conservation management of anurans of Paraná River ecosystems.     

Body shape, burst speed and escape behavior of larval anurans

Variation in behavior, morphology and life history traits of larval anurans across predator gradients, and consequences of that variation, have been abundantly studied. Yet the functional link between morphology and burst-swimming speed is largely unknown. We conducted experiments with two divergent species of anurans, Scaphiopus holbrookii and Rana sphenocephala , to examine how behavior and morphology influence predator vulnerability, and whether tadpole shape is related to burst-swimming performance. Scaphiopus holbrookii , a species that typically uses ephemeral pools, was more active, exhibited slower burst speeds, and was more susceptible to predation than R. sphenocephala , a species associated with more permanent aquatic sites. Our analysis of morphology and burst speed defined a shared axis of shape variation associated with burst-swimming speed regardless of species. Tadpoles with a deeper tail fin and muscle and a smaller body produced faster speeds. The nature and breadth of the morphology–speed relationship suggests it may represent a generalized ecomorphological paradigm for larval anurans.     

Revisiting the contribution of larval characters to an analysis of phylogenetic relationships of basal anurans

The higher-level relationships of anurans have been explored by numerous studies, producing a variety of hypotheses. The relationships of the basal anurans ('archaeobatrachians') are, however, poorly known . In part, this may be because the adult morphology of basal anurans is derived and therefore may not provide suitable phylogenetic signal. Recently, several authors have shown the phylogenetic utility of information derived from anuran larvae. In this paper we conduct separate and combined analyses of anuran relationships based on adult and larval morphology. Our combined results suggest that anurans form two major clades − the pipoids and all other frogs. Evidence also suggests that, taken together, Neobatrachia and Pelobatoidea form a monophyletic group. We discuss support for various groupings as shown by the different data sets. We also comment on the consequences of our phylogenetic hypothesis for the interpretation of reduction of vertebral elements and evolution of Orton's tadpole types in anurans.  © 2003 The Linnean Society of London, Zoological Journal of the Linnean Society, 2003, 139 , 129−155.     

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.     

Spatial dependence of phenotype-environment associations for tadpoles in natural ponds

Within natural habitats, phenotypes are shaped by many environmental factors. Consequently, heterogeneity of these factors can promote phenotypic divergence. However, because environments exhibit heterogeneity at different spatial scales, phenotypic divergence should also exhibit such scale-dependence. Using hierarchical linear models, I determined how multiple environmental factors at two spatial scales affected the morphology of wood frog (Rana (Lithobates) sylvatica) tadpoles collected from natural ponds. Among ponds, predation risk intensity and tadpole density were strong predictors of tadpole morphology, while within ponds, other environmental variables such as water depth and leaf litter were more important. Spatial analyses revealed that water depth and leaf litter, but not predation risk intensity or tadpole density, exhibited heterogeneous spatial distributions within ponds, suggesting that spatial properties of environmental variables influenced the scale at which they shaped phenotypes. Furthermore, patterns of phenotypic variation with respect to predation risk intensity and tadpole density among ponds largely matched observations from previous laboratory studies. These results emphasize the importance of considering phenotype-environment associations across multiple spatial scales.     

Biodiversity–productivity relationship in ponds: Community and metacommunity patterns along time and environmental gradients

Primary production correlates with diversity in various ways. These patterns may result from the interaction of various mechanisms related to the environmental context and the spatial and temporal scale of analysis. However, empirical evidence on diversity‐productivity patterns typically considers single temporal and spatial scales, and does not include the effect of environmental variables. In a metacommunity of macrophytes in ephemeral ponds, we analysed the diversity‐productivity relationship patterns in the field, the importance of the environmental variables of pond size and heterogeneity on such relationship, and the variation of these patterns at local (community level) and landscape scales (metacommunity level) across 52 ponds on twelve occasions, over five years (2005–2009). Combining all sampling dates, there were 377 ponds and 1954 sample‐unit observations. Vegetation biomass was used as a proxy for productivity, and biodiversity was represented by species richness, evenness, and their interaction. Environmental variables comprised pond area, depth and internal heterogeneity. Productivity and species richness were not directly related at the metacommunity level, and were positively related at the community level. Taking environmental variables into account revealed positive species richness‐productivity relationships at the metacommunity level and positive quadratic relationships at the community level. Productivity showed both positive and negative linear and nonlinear relationships with the size and heterogeneity of ponds. We found a weak relationship between productivity and evenness. The identity of variables associated with productivity changed between spatial scales and through time. The pattern of relationships between productivity and diversity depends on spatial scale and environmental context, and changes idiosyncratically through time within the same ecosystem. Thus, the diversity‐productivity relationship is not only a property of the study system, but also a consequence of environmental variations and the temporal and spatial scale of analysis.     

Unravelling the correlates of species richness and ecological uniqueness in a metacommunity of urban pond insects

City ponds have the potential to harbour a rich biodiversity of aquatic insects despite being located in an urban landscape. However, our current knowledge on the correlates of pond biodiversity is limited and even less is known about the factors that influence the ecological uniqueness of urban ponds. The multiple environmental gradients, at different spatial scales, that may affect biodiversity and ecological uniqueness of urban ponds can thus be seen both as an opportunity and as a challenge for a study. In this study, we aimed to fill this gap by focusing on aquatic insect assemblages in 51 ponds in the Swedish city of Stockholm, using a metacommunity perspective. We found that species richness was primarily determined by the density of aquatic insects, water depth and proportion of buildings around the pond. The uniqueness of ponds was estimated as local contributions to beta diversity (LCBD), and it was primarily related to the proportion of arable land and industry around the ponds. With regard to the metacommunity we found two interesting patterns. First, there was a negative relationship between richness and LCBD. Second, biodiversity was spatially independent, suggesting that spatially-patterned dispersal did not structure species richness or LCBD. These last two patterns are important when considering conservation efforts of biodiversity in city ponds. We hence suggest that the conservation of insect biodiversity in urban pond should consider the surroundings of the ponds, and that high-richness ponds are not necessarily those that require most attention because they are not ecologically the most unique.     

Body size affects the spatial scale of habitat–beetle interactions

We used abundance data and the program Focus to determine the spatial scale at which 31 species of longhorned beetles (Coleoptera: Cerambycidae) respond to forest habitat amount. We predicted that the spatial scale at which species respond would increase with body size, and that species using ephemeral larval habitat would respond at larger spatial scales than species using more stable larval habitat. We found that forest cover was a better measure of the amount of habitat for polyphagous species than for oligophagous species. Larger species of longhorned beetles responded to forest cover at larger scales. We did not find evidence that species using more ephemeral larval habitat conditions responded at larger scales than species developing in more stable habitat conditions. Our results highlight the importance of accurately describing habitat in studies of species–environment relationships. While scales of response may be species-specific, some generalizations across species are possible.     

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.     

Sampling related individuals within ponds biases estimates of population structure in a pond‐breeding amphibian

Effective conservation and management of pond‐breeding amphibians depends on the accurate estimation of population structure, demographic parameters, and the influence of landscape features on breeding‐site connectivity. Population‐level studies of pond‐breeding amphibians typically sample larval life stages because they are easily captured and can be sampled nondestructively. These studies often identify high levels of relatedness between individuals from the same pond, which can be exacerbated by sampling the larval stage. Yet, the effect of these related individuals on population genetic studies using genomic data is not yet fully understood. Here, we assess the effect of within‐pond relatedness on population and landscape genetic analyses by focusing on the barred tiger salamanders (Ambystoma mavortium) from the Nebraska Sandhills. Utilizing genome‐wide SNPs generated using a double‐digest RADseq approach, we conducted standard population and landscape genetic analyses using datasets with and without siblings. We found that reduced sample sizes influenced parameter estimates more than the inclusion of siblings, but that within‐pond relatedness led to the inference of spurious population structure when analyses depended on allele frequencies. Our landscape genetic analyses also supported different models across datasets depending on the spatial resolution analyzed. We recommend that future studies not only test for relatedness among larval samples but also remove siblings before conducting population or landscape genetic analyses. We also recommend alternative sampling strategies to reduce sampling siblings before sequencing takes place. Biases introduced by unknowingly including siblings can have significant implications for population and landscape genetic analyses, and in turn, for species conservation strategies and outcomes.     

Fish assemblages in temporary ponds adjacent to 'terra-firme' streams in Central Amazonia

1. The effect of habitat structural features and physicochemical characteristics of the water on the composition and richness of fish assemblages in temporary ponds near streams were examined at three spatial scales: among ponds, among streams and between drainage basins, in a ‘terra‐firme’ (not subject to long‐term flooding) forest reserve in Central Amazonia. 2. The fish assemblage in temporary ponds was composed of subsets of 18 small‐bodied species widely distributed in the reserve. The assemblages had a nested subset structure, where smaller ponds contained subgroups of the species found in larger ponds. 3. Species composition and richness in temporary ponds were similar between drainage basins, although the fish assemblages in streams differed between basins. 4. Fish assemblage structure was influenced by local factors related to habitat structure, such as pond area and depth, canopy cover and hydroperiod. Physicochemical characteristics of the water in the ponds were similar between drainage basins and had little detectable effect on the structure of pond fish assemblages. 5. No correspondence was found between the composition, richness or abundance of fishes in the ponds and in stretches of the streams adjacent to the ponds. Therefore, it is not possible to predict the composition of these temporary pond fish assemblages from the fish assemblages found in adjacent streams.     

Abundance and composition of forest-dwelling anurans in cashew plantations in a tropical semi-evergreen forest landscape

Plantation crops in tropical human-modified landscapes provide alternative habitats to biodiversity outside protected areas. The Western Ghats of India are home to a mosaic of closely spaced habitats, including forests and agroecosystems. Cashew is a widely grown plantation crop in the northern Western Ghats and is known to provide economic and societal benefits. However, its role as a supplementary habitat for anurans is not well understood. We assessed the factors that influence understorey anuran composition and abundance in cashew plantations, forest edges, and forest interiors in Tillari Conservation Reserve, Maharashtra. Species composition of cashew plantations differed significantly from forests and was positively influenced by understorey and canopy cover. Cashew plantations had a near equal abundance of anurans as that of forest edges and interiors, which could be due to the preponderance of habitat generalists. Understorey positively influenced anuran abundance while ambient temperature had a negative influence. Reduced understorey and low canopy cover represent habitat modifications that occur in cashew plantations. Such structural changes could lead to reduced environmental refuges for anurans, thereby exposing them to large variations in temperature and moisture. Cashew plantations in Tillari Conservation Reserve serve as supplementary habitats for anurans. That said, cashew cultivation practices and markets must be understood before biodiversity-friendly plantation practices are proposed.     

Role of epibiotic diatoms isolated from the barnacle shell in the cyprid metamorphosis of <Emphasis Type="Italic">Balanus amphitrite</Emphasis>

The mechanisms structuring aquatic communities across environmental gradients are often difficult to distinguish from one another and can produce similar patterns of species distributions. In freshwater systems, the amount of canopy cover from surrounding trees is often associated with transitions in local community structure. These community changes could be driven by habitat selection prior to colonization of the aquatic habitat and/or species-sorting post-colonization. To assess the contributions of pre- versus post-colonization processes in structuring larval dragonfly assemblages, we tested the impact of artificial and natural canopy cover on the selection of experimental aquatic mesocosms by adult dragonflies, and monitored the performance (i.e. growth and survival) of larval dragonflies that were placed in mesocosms under a gradient of natural canopy cover. We found that greater levels of canopy cover resulted in fewer adult visits to mesocosms, and more natural canopy cover decreased the species richness of visitors. There were no effects of canopy cover on the growth and survival of larvae added to the mesocosms. Our results suggest that adult habitat selection plays a dominant role in structuring larval dragonfly assemblages across a canopy cover gradient, and that canopy cover can be an important environmental filter on species distributions.     

Multi-scale species density model for conserving an endangered songbird

Habitat modeling across a landscape that has gradients of habitat conditions requires potential predictor data that can be quantified at biologically relevant scales. We used remotely sensed data to develop a multi-scale density model in 2018 for the golden-cheeked warbler (Setophaga chrysoparia; warbler), a species that breeds in Ashe juniper (Juniperus ashei)-oak (Quercus spp.) woodlands in central Texas, USA. We first classified Ashe juniper and broadleaf tree cover at a 1-m resolution and used this to map potential habitat across the warbler's >67,000-km² breeding range. We then designed a survey for estimating warbler density based on hierarchical distance sampling. We used stratified random sampling to survey for male warblers at 1,804 points across the continuum of tree canopy cover and composition and detected 810 warblers during our surveys. We developed a suite of potential predictor variables for modeling warbler density that reflected vegetation, topography, climate, and anthropogenic land use conditions across the breeding range and developed these at 3 scales representing the territory, site, and landscape. We modeled warbler density and used the best fit model to produce a spatially explicit estimate. Predicted warbler density was influenced by tree canopy cover and canopy height at the territory scale (100-m radius); tree canopy cover, percent of the canopy comprised of juniper, and an interaction between canopy cover and compound topographic index at the site scale (1-km radius); and annual temperature range at the landscape scale (5-km radius). We estimated a population size of 217,444 male warblers (95% CI = 153,917–311,965) and >3,000 males in each recovery unit. After controlling for the duration of point count surveys, our estimate of population size was similar to that reported from the only previous breeding range survey conducted in 2008–2009. Our model results indicated that management activities to increase warbler density should promote woodlands with high tree canopy cover, approximately 60–80% Ashe juniper composition, and tree heights >3 m. In contrast to a patch-based approach, our treatment of habitat variables as continuous helped to credibly map the warbler distribution across areas with broad transitions from woodlands to shrublands. By measuring these predictor variables at biologically relevant scales, we allowed the warbler survey data to define habitat relationships instead of using anthropogenically defined habitat patches. Outcomes from our study show the benefits of developing spatial products tailored to individual species of interest for conservation and management decisions.     

The influence of niche and neutral processes on a neotropical anuran metacommunity

One of the most important questions in ecology is the relative importance of local conditions (niche processes) and dispersal ability (neutral processes) in driving metacommunity structure. Although many studies have been conducted in recent years, there is still much debate. We evaluated the processes (niche and neutral) responsible for variation in anuran composition in 28 lentic water bodies in southeastern Brazil. Because anurans depend heavily on environmental conditions, we hypothesized that environmental variables (niche processes) are the most important drivers of community composition. Additionally, as anurans have limited dispersal abilities, and the study region presents harsh conditions (high forest fragmentation, low rainfall and long dry season), we expected a lower, but significant, spatial signature in metacommunity structure, due to neutral dynamics. We used a partial redundancy analysis with variation partitioning to evaluate the relative influence of environmental and spatial variables as drivers of metacommunity structure. Additionally, we used a recently developed spatial autocorrelation analysis to test if neutral dynamics can be attributed to the pure spatial component. This analysis is based on predictions that species abundances are independent but similarly spatially structured, with correlograms similar in shape. Therefore, under neutral dynamics there is no expectation of a correlation between the pairwise distance of spatial correlograms and the pairwise correlation of species abundances predicted by the pure spatial component. We found that the environmental component explained 21.5%, the spatial component 10.2%, and the shared component 6.4% of the metacommunity structure. We found no correlation between correlograms and correlation of abundances predicted by the pure spatial component (Mantel test = ?0.109, P = 0.961). In our study, niche‐based processes are the dominant process that explained community composition. However, neutral processes are important because spatial variation can be attributed to pure neutral dynamics rather than to missing spatially structured environmental factors.     

Canopy Closure,Competition, and the Endangered Dusky Gopher Frog

Abstract: A major challenge facing wildlife biologists is understanding why some species go extinct while others persist in the same habitat. To address this question, we investigated whether tree canopy closure over ponds affects growth and survival of rare and common tadpoles within ponds and mediates competitive interactions among species. We conducted 2 experiments to test whether canopy closure and competition may have contributed to the decline of the endangered dusky gopher frog (Rana sevosa), but allowed the persistence of the southern leopard frog (R. sphenocephala). We explored the response of both species to canopy closure in single-species and mixed- (1:1) species treatments of identical total tadpole density. An experiment using aquatic enclosures in temporary ponds showed that canopy closure reduced tadpole growth approximately 20% for both species. Survival of dusky gopher frog tadpoles was higher in mixed-species enclosures than in single-species enclosures. In a complementary experiment using artificial ponds, dusky gopher frogs had lower survival to metamorphosis, reduced size at metamorphosis, and produced a lower total biomass of metamorphosed juveniles in shaded ponds. Southern leopard frogs exhibited reduced body size at metamorphosis only when shaded. These studies suggest that pond canopy closure, not larval competition, may be contributing to the decline of the dusky gopher frog. The different responses to canopy closure suggest a potential mechanism for the loss of dusky gopher frogs and the persistence of southern leopard frogs. Removal of trees from historically open-canopy ponds may help facilitate the recovery of dusky gopher frogs and benefit similar species.