Parental effects influence life history traits and covary with an environmental cline in common frog populations

Oecologia - Across latitudinal clines, the juvenile developmental rates of ectotherms often covary with the length of the growing season, due to life-history trade-offs imposed by the...     

Responses to competition effects of two anuran tadpoles according to life-history traits

Experimental manipulations of the densities of two larval anurans, Pelodytes punctatus and Bufo bufo , showed that these species compete asymmetrically in semi-natural conditions. Growth, mass at metamorphosis, date of metamorphosis, and survival were used as measures of response to interspecific competition. A mechanistic approach was used to collect information on the behaviour of the two species in different conditions. The competitive superiority of Pelodytes at individual level was correlated with a larger body, faster growth rate, increased per capita competitive impact on conspecifics, and greater reduction in the availability of trophic and spatial resources. In the presence of Pelodytes, Bufo showed slower growth, smaller size at metamorphosis and reduced survival. In the interspecific treatments Bufo individuals modified their behaviour by increasing activity and use of the water column while Pelodytes did not change their foraging activity or space use in the aquaria. However, the presence of Bufo resulted in a reduced larval period and smaller size at metamorphosis. We hypothesise that the presence of Bufo act as a signal of environmental degradation and shorten the larval period of Pelodytes, a typical temporal pond breeder . The smaller Bufo tadpoles are potentially stronger competitors at population level because they use relatively large amounts of energy (greater densities and higher metabolic rates). Consequently, they use larger proportions of the shared resources than their larger competitor. A possible evolutionary response for larger tadpoles is the development of interference mechanisms or "escaping" from ephemeral ponds where mortality by drying represent a high risk.     

Morphological Consequences of Developmental Plasticity in <Emphasis Type="Italic">Rana temporaria</Emphasis> are not Accommodated into Among-Population or Among-Species Variation

Environmental induced developmental plasticity occurs in many organisms and it has been suggested to facilitate biological diversification. Here we use ranid frogs to examine whether morphological changes derived from adaptive developmental acceleration in response to pool drying within a species are mirrored by differences among populations and across species. Accelerated development in larval anurans under pool drying conditions is adaptive and often results in allometric changes in limb length and head shape. We examine the association between developmental rate and morphology within population, among populations in divergent environments, and among species inside the Ranidae frog family, combining experimental approaches with phylogenetic comparative analyses. We found that frogs reared under decreasing water conditions that simulated fast pool drying had a faster development rate compared to tadpoles reared on constant water conditions. This faster developmental rate resulted in different juvenile morphologies between the two pool drying conditions. The association between developmental rate and morphology found as a result of plasticity was not mirrored by differences among populations that differed in development, neither was it mirrored among species that differed in development rate. We conclude that morphological differences among populations and species were not driven by variation in developmental time per se. Instead, selective factors, presumably operating on locomotion and prey choice, seem to have had a stronger evolutionary effect on frog morphology than evolutionary divergences in developmental rate in the ranid populations and species studied.     

Do anuran larvae respond behaviourally to chemical cues from an invasive crayfish predator? A community-wide study

Antipredator behaviour is an important fitness component in most animals. A co-evolutionary history between predator and prey is important for prey to respond adaptively to predation threats. When non-native predator species invade new areas, native prey may not recognise them or may lack effective antipredator defences. However, responses to novel predators can be facilitated by chemical cues from the predators’ diet. The red swamp crayfish Procambarus clarkii is a widespread invasive predator in the Southwest of the Iberian Peninsula, where it preys upon native anuran tadpoles. In a laboratory experiment we studied behavioural antipredator defences (alterations in activity level and spatial avoidance of predator) of nine anurans in response to P. clarkii chemical cues, and compared them with the defences towards a native predator, the larval dragonfly Aeshna sp. To investigate how chemical cues from consumed conspecifics shape the responses, we raised tadpoles with either a tadpole-fed or starved crayfish, or dragonfly larva, or in the absence of a predator. Five species significantly altered their behaviour in the presence of crayfish, and this was largely mediated by chemical cues from consumed conspecifics. In the presence of dragonflies, most species exhibited behavioural defences and often these did not require the presence of cues from predation events. Responding to cues from consumed conspecifics seems to be a critical factor in facilitating certain behavioural responses to novel exotic predators. This finding can be useful for predicting antipredator responses to invasive predators and help directing conservation efforts to the species at highest risk.     

Evolution and plasticity of anuran larval development in response to desiccation. A comparative analysis

Anurans breed in a variety of aquatic habitats with contrasting levels of desiccation risk, which may result in selection for faster development during larval stages. Previous studies suggest that species in ephemeral ponds reduce their developmental times to minimize desiccation risks, although it is not clear how variation in desiccation risk affects developmental strategies in different species. Employing a comparative phylogenetic approach including data from published and unpublished studies encompassing 62 observations across 30 species, we tested if species breeding in ephemeral ponds (High risk) develop faster than those from permanent ponds (Low risk) and/or show increased developmental plasticity in response to drying conditions. Our analyses support shorter developmental times in High risk, primarily by decreasing body mass at metamorphosis. Plasticity in developmental times was small and did not differ between groups. However, accelerated development in High risk species generally resulted in reduced sizes at metamorphosis, while some Low risk species were able compensate this effect by increasing mean growth rates. Taken together, our results suggest that plastic responses in species breeding in ephemeral ponds are constrained by a general trade-off between development and growth rates.     

Are protected areas truly protected? The impact of road traffic on vertebrate fauna

The extension of road networks is considered one of the major factors affecting fauna survival. Roadkill has been documented widely and affects all taxonomic groups. Although roadkill is associated mainly with traffic density, some life-history traits of species and the area surrounding roads are expected to modify number of roadkills both taxonomically and geographically. Here we studied the number of roadkills of vertebrates in an extensive region in the northeastern Iberian Peninsula. We surveyed 820?km of 41 roads in two different seasons (spring and autumn), that differ in traffic intensity. In addition, we covered zones with distinct climatic characteristics and levels of protection of the surrounding habitats. Amphibians showed the highest number of roadkills whereas reptiles, birds and mammals had similar rates. General Linear Model tests showed no differences in roadkills by climatic region; however, differences in number of roadkills were linked to protection status, with the highest number of casualties in highly protected areas. Redundancy Analysis demonstrated that the number of amphibians and reptiles killed was associated with roads in highly protected areas whereas that of mammals and birds was linked to unprotected areas. Protected areas often receive many visitors, which in turn may increase wildlife casualties as a result of greater traffic density. We recommend that correction measures be taken to reduce the high number of vertebrate fauna killed along roads that cross protected areas.     

Genetic variability in geographic populations of the natterjack toad (Bufo calamita)

Across altitudinal and latitudinal gradients, the proportion of suitable habitats varies, influencing the individual dispersal that ultimately can produce differentiation among populations. The natterjack toad (Bufo calamita) is distributed across a wide geographic range that qualifies the species as interesting for a geographic analysis of its genetic variability. Five populations of B. calamita in the Sierra de Gredos (Spain) were studied in an altitudinal gradient ranging from 750 to 2270 m using microsatellite markers. In addition, we analyzed the latitudinal genetic variation in B. calamita within a global European distribution using genetic diversity parameters (mean number of alleles per locus [M(a)] and expected heterozygosity [H(E)]) obtained from our results and those published in the literature. The low level of genetic differentiation found between populations of B. calamita (F(st) ranging from 0.0115 to 0.1018) and the decreases in genetic diversity with altitude (M(a) from 13.6 to 8.3, H(E) from 0.82 to 0.74) can be interpreted by the combined effects of discontinuous habitat, produced mainly by the high slopes barriers and geographic distance. In the latitudinal gradient, genetic diversity decreases from south to north as a consequence of the colonization of the species from the Pleistocene refugium. We conclude that the genetic variability in B. calamita along its wide altitudinal and latitudinal geographic distribution mainly reflects the colonization history of the species after the last glacial period.     

Trophic Strategies of a Non-Native and a Native Amphibian Species in Shared Ponds

One of the critical factors for understanding the establishment, success and potential impact on native species of an introduced species is a thorough knowledge of how these species manage trophic resources. Two main trophic strategies for resource acquisition have been described: competition and opportunism. In the present study our objective was to identify the main trophic strategies of the non-native amphibian Discoglossus pictus and its potential trophic impact on the native amphibian Bufo calamita. We determine whether D. pictus exploits similar trophic resources to those exploited by the native B. calamita (competition hypothesis) or alternative resources (opportunistic hypothesis). To this end, we analyzed the stable isotope values of nitrogen and carbon in larvae of both species, in natural ponds and in controlled laboratory conditions. The similarity of the δ¹⁵N and δ¹³C values in the two species coupled with isotopic signal variation according to pond conditions and niche partitioning when they co-occurred indicated dietary competition. Additionally, the non-native species was located at higher levels of trophic niches than the native species and B. calamita suffered an increase in its standard ellipse area when it shared ponds with D. pictus. These results suggest niche displacement of B. calamita to non-preferred resources and greater competitive capacity of D. pictus in field conditions. Moreover, D. pictus showed a broader niche than the native species in all conditions, indicating increased capacity to exploit the diversity of resources; this may indirectly favor its invasiveness. Despite the limitations of this study (derived from potential variability in pond isotopic signals), the results support previous experimental studies. All the studies indicate that D. pictus competes with B. calamita for trophic resources with potential negative effects on the fitness of the latter.     

Genetic analysis of differentiation among breeding ponds reveals a candidate gene for local adaptation in Rana arvalis

One of the main questions in evolutionary and conservation biology is how geographical and environmental features of the landscape shape neutral and adaptive genetic variation in natural populations. The identification of genomic polymorphisms that account for adaptive variation can aid in finding candidate loci for local adaptation. Consequently, a comparison of spatial patterns in neutral markers and loci under selection may help disentangle the effects of gene flow, genetic drift and selection at the landscape scale. Many amphibians breed in wetlands, which differ in environmental conditions and in the degree of isolation, enhancing the potential for local adaptation. We used microsatellite markers to measure genetic differentiation among 17 local populations of Rana arvalis breeding in a network of wetlands. We found that locus RC08604 deviated from neutral expectations, suggesting that it is a good candidate for directional selection. We used a genetic network analysis to show that the allele distribution in this locus is correlated with habitat characteristics, whereas this was not the case at neutral markers that displayed a different allele distribution and population network in the study area. The graph approach illustrated the genomic heterogeneity (neutral loci vs. the candidate locus for directional selection) of gene exchange and genetic divergence among populations under directional selection. Limited gene flow between wetlands was only observed at the candidate genomic region under directional selection. RC08604 is partially located inside an up‐regulated thyroid‐hormone receptor (TRβ) gene coordinating the expression of other genes during metamorphosis and appears to be linked with variation in larval life‐history traits found among R. arvalis populations. We suggest that directional selection on genes coding larval life‐history traits is strong enough to maintain the divergence in these genomic regions, reducing the effective recombination of locally adapted alleles but not in other regions of the genome. Integrating this knowledge into conservation plans at the landscape scale will improve the design of management strategies to preserve adaptive genetic diversity in wetland networks.     

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.