In the toad Bufo calamita, among-population variation of size follows roughly a converse Bergmann cline, but populations exist that do not fit this pattern. We propose that latitudinal body size variation is a byproduct of adaptive covariation among the life-history traits juvenile growth rate, longevity and lifetime fecundity. We choose five populations (two in Andalusia, two in Catalonia and one in Rhineland-Palatinate) representing a variation of adult size from 39 mm to 95 mm snout-vent length, a latitudinal gradient from 37 to 501 and an altitudinal gradient from sea level to 420 m. Skeletochronology was used to estimate the age-related life-history traits of 313 toads and their lifetime pattern of growth. At southern latitudes, toads matured and reproduced earlier than those at northern latitudes, but had a reduced potential reproductive lifespan due to lower longevity. Ageadjusted adult size depended mainly on the size achieved between metamorphosis and first hibernation or aestivation, which in turn was influenced by local factors. We propose that first-year size corresponds to the duration of the aboveground activity period, temperature during the activity period and the type of shelter sites and hibernacula available in the habitat. After attaining sexual maturity, the growth rates did not differ among populations. Interactions of multiple environmental factors during the first year of life determine age at maturity, adult size and size variation among populations. Local body size and potential reproductive lifespan covary to optimize lifetime fecundity throughout the geographical range. The presence of a small-sized population in southern Spain does not fit the pattern predicted by a converse Bergmann cline, but is compatible with the hypothesis that body size variation among B. calamita populations may be the evolutionary byproduct of optimized lifetime fecundity.

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The delimitation of the spatial equivalent of isolated populations or interacting sets of local populations is crucial for conservation management. In Amphibia, spatial organization in metapopulations requires knowledge on the individual variation of annual migratory capacity within local populations to delimit core habitats around the breeding site and to estimate the connectivity among neighbouring populations. The migratory behaviour of 143 adult Bufo calamita was monitored using radio‐telemetry at eight localities covering a latitudinal range from 41° to 54°N. Aims of the study were to assess the geographic variation of migratory distance moved by males and females and to model their capacity for dispersal assuming exclusively unidirectional movements. Migratory range was not sex‐biased, but was three times lower in population inhabiting sandy areas than in those on clay soils, probably because of the scarcity of moist shelters causing more frequent and more distant movements. For conservation management of local natterjack populations, we propose to use the migratory capacity of the 50% most sedentary individuals to delimit the core area around a given breeding site. To estimate the potential genetic connectivity between neighbouring local population, we propose to use the minimum migratory capacity of the 5% of individuals that moved most. Estimates obtained for populations in central Europe and the UK indicate a core area of 600 m around the breeding site and a maximum distance of 2250 m between the breeding ponds to maintain connectivity. Thus, the principal conservation problem in the UK is that most populations are isolated by distance and prone to local extinction. In contrast, core areas of populations in Spain extend to distances of about 5 km and connectivity is maintained up to a distance 12 km between neighbouring breeding ponds.

The Pyrenees represent a natural laboratory for biogeographic, evolutionary and ecological research of mountain fauna as a result of the high variety of habitats and the profound effect of the glacial and interglacial periods. There is a paucity of studies providing a detailed insight into genetic processes and better knowledge on the patterns of genetic diversity and how they are maintained under high altitude conditions. This is of particular interest when considering the course of past climate conditions and glaciations in a species which is considered site tenacious, with long generation times. Here we analyzed the genetic patterns of diversity and structure of the endemic Pyrenean brook newt (Calotriton asper) along its distribution range, with special emphasis on the distinct habitat types (caves, streams, and lakes), and the altitudinal and geographical ranges, using a total set of 900 individuals from 44 different localities across the Pyrenean mountain range genotyped for 19 microsatellite loci. We found evidence for a negative longitudinal and positive altitudinal gradient of genetic diversity in C. asper populations. The fact that genetic diversity was markedly higher westwards is in accordance with other Pyrenean species. However, the impact of altitudinal gradient on the genetic diversity seems to differ from other species, and mostly from other amphibians. We found that lower altitudes can act as a barrier probably because the lowlands do not provide a suitable habitat for C. asper. Regarding the distinct habitat types, caves had significantly lower values of genetic diversity compared to streams or lakes. The mean FST value was relatively high (0.304) with maximum values as high as 0.771, suggesting a highly structured total population. Indeed, populations were grouped into five subclusters, the eastern populations (cluster 1) remained grouped into two subclusters and the central-western Pyrenees (cluster 2) into three subclusters. The increase of isolation with geographical distance is consistent with the population structure detected. In conclusion, C. asper seems to be adapted to high altitude mountain habitats, and its genetic diversity is higher in the western Pyrenees. In terms of conservation priority, we consider more relevant the populations that represent a reservoir of genetic diversity.

The Pyrenean brook newt (Calotriton asper) is a salamandrid that mostly lives in fast running and cold mountain-streams, although some populations are also found in lakes. In the present work, we report in detail on the occurrence of facultative paedomorphosis traits in a population from a Pyrenean high altitude lake. We compare its morphology, life history traits and mitochondrial DNA variation with a nearby lotic metamorphic population. Our results indicate that the lacustrine newts are smaller and present a less developed sexual dimorphism, smooth skin, and that 53% of the adults retain gills at different degrees of development, but not gill slits. Although both populations and sexes have the same age at sexual maturity (four years), the lacustrine population presents higher longevity (12 and 9 years for males and females, respectively) than the one living in the stream (8 and 9 years). The variation on the climatic conditions at altitudinal scale is probably the main cause of the differences in life history traits found between the two populations. The food availability, which could to be limiting in the lacustrine population, is another factor that can potentially affect body size. These results are congruent with the significant mitochondrial DNA genetic isolation between populations, probably a consequence of the lack of juvenile dispersal. We found low cytochrome b variability and significant genetic structuring in the lake population that is very remarkably considering the short distance to the nearby stream population and the whole species’ pattern. We suggest that a bottleneck effect and/or phenotypic plasticity may have resulted in the appearance of a paedomorphic morph in the lake.

Paedomorphosis and metamorphosis are two major developmental processes that characterize the evolution of complex life cycles in many lineages. Whereas these processes were fixed in some taxa, they remained facultative in others, with alternative phenotypes expressed in the same populations. From a genetic perspective, it is still unknown whether such phenotypes form a single population or whether they show some patterns of isolation in syntopy. This has deep implications for understanding the evolution of the phenotypes, i.e. towards their persistence or their fixation and speciation. Newts and salamanders are excellent models to test this hypothesis because they exhibit both developmental processes in their populations: the aquatic paedomorphs retain gills, whereas the metamorphs are able to colonize land. Using microsatellite data of coexisting paedomorphic and metamorphic palmate newts (Lissotriton helveticus), we found that they formed a panmictic population, which evidences sexual compatibility between the two phenotypes. The high gene flow could be understood as an adaptation to unstable habitats in which phenotypic plasticity is favored over the fixation of developmental alternatives. This makes then possible the persistence of a polyphenism: only metamorphosis could be maintained in case of occasional drying whereas paedomorphosis could offer specific advantages in organisms remaining in water.

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 [Ma] and expected heterozygosity [HE]) obtained from our results and those published in the literature. The low level of genetic differentiation found between populations of B. calamita (Fst ranging from 0.0115 to 0.1018) and the decreases in genetic diversity with altitude (Ma from 13.6 to 8.3, HE 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.

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