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The fossorial life-history emerged as a response to hostile environmental conditions, such as extreme temperatures and dryness. It evolved independently in different lineages of amphibians, which are particularly vulnerable to dehydration and require a continuous level of humidity because of their permeable skin. For the first time, we test the hypothesis that amphibians are able to adapt their fossorial behaviour to substrate humidity, using the Pacific horned frogs (Ceratophrys stolzmanni) from the Ecuadorian dry forest as a model. When exposed to dry substrate, the animals burrowed much deeper than the ones in wet soil. The capacity to excavate was particularly high in this species: frogs from the dry treatment could be found at more than 1-m depth after just a few hours. Our results reveal a conditional strategy that contributes to explain the persistence of amphibians in dry environments.

The spatial distribution of organisms in patchy environments can be determined by the relationship between habitat quality and animal behaviour. In species with complex life cycles, such as pond-breeding amphibians, the selection of a suitable wetland is crucial. While the traditional view of amphibian ecology suggests strong site fidelity and low vagility, more recent research highlights mechanisms involving active site choice through avoidance behaviour and bet-hedging strategies in heterogeneous environments. The introduction of fish to the aquatic environment is one of the factors that may alter site selection and lead to local dispersal. In this context, we investigated the habitat choice of Alpine newts, Ichthyosaura alpestris, in networks of fish (Carassius auratus) and fishless aquatic patches. Using a laboratory design consisting of aquaria divided into two water tanks connected by a terrestrial platform, we assessed newt distribution and movement between patches. During the breeding period, we compared the reproductive success of individuals in two types of networks. We found that newts escaped fish by rapidly changing aquatic patches and then aggregating in safe aquatic patches that were free of fish. In the fish network, newts maintained reproduction, but the high local abundance resulted in decreased sexual activity and egg production and increased use of the terrestrial habitat. However, in the fishless network, newts moved between aquatic patches several times, exhibited more courtship behaviour and laid more eggs than they did in the fish networks. Our results showed both adaptive habitat switching due to environmental risks in the fish network and habitat supplementation (i.e. use of alternative resources) in the fishless network. Such studies on movement behaviour and habitat selection have conservation implications in showing that the persistence of native species in invaded networks depends on the rescue effect, with immigration to fish-free habitats potentially preventing local extinction.

Antipredator responses are a key determinant of the successful persistence of prey, and behavioural modifications are a frequent antipredator strategy. However, conspecific populations often inhabit heterogeneous environments. This can determine local adaptations, and might also induce variation in antipredator responses. Nevertheless, there is limited information on whether heterogeneity of predation risk among populations determines variation in antipredator response. Here we studied the fire salamander, Salamandra salamandra, a species that can breed in both surface streams and caves, habitats that are predator-rich and predator-free, respectively, and measured differences in antipredator responses across populations with different predation risk. We combined field surveys and laboratory experiments to understand the role of predation risk on the activity patterns of larvae, while measuring behavioural differences between populations. We reared larvae from different habitats in safe and risky conditions and tested their response to predator cues before and after rearing. In the field, predation risk was much higher in surface streams than in caves; larvae moved more in the absence of predators and when the light intensity was low. During laboratory experiments, larvae were less active if reared in risky conditions, but cave larvae showed a stronger response to risk than stream larvae. Therefore, larvae from sites without predators showed higher antipredator responses than those from risky habitats. This response fits the predictions of the risk allocation model, in which prey from habitats with a high background level of risk need to be active even when predators are present, to satisfy their energetic demands. Our findings show that antipredator behaviour may differ strongly between populations and stress the importance of integrating this variability in studies on predatory responses.

Resilience of organisms after threat removal is an essential feature to justify conservation efforts. Amphibians are particularly threatened with a worldwide decline, showing a low resistance to invaders such as fish. Previous research has shown that they could recover after fish extirpation due to metamorphosed colonisers. However, not all amphibian phenotypes are able to persist fish introduction and disperse. In many species of newts and salamanders, paedomorphs retain gills in the adult stage, which makes them fully aquatic. A proposed way to conserve this phenotype would be to remove introduced fish from their habitats. However, because paedomorphosis is usually not expressed in the presence of fish, it is unknown whether fish removal could allow the resilience of paedomorphs. This would be possible only if progenies of metamorphosed individuals could become paedomorphic in restored habitats. Through a quantitative survey in three types of ponds, including control ponds without fish, ponds in which fish were extirpated, and fish ponds, we determined abundances of paedomorphic and metamorphic palmate newts (Lissotriton helveticus). The results show that paedomorphosis resilience is possible and even highly frequent, as paedomorphs were found in 80% of ponds where fish disappeared. Abundances were similar between these ponds and control ponds whereas fish ponds had almost no newts, indicating a very low resistance to invaders. This shows that conserving common phenotypes can help to preserve endangered phenotypes, as paedomorphs were produced through the reproduction of metamorphs. There is thus hope of maintaining intraspecific biodiversity though conservation action involving threat removal.

Species with complex life cycles are good models to understand trade-offs between life in hostile and favourable habitats. Newts remain in breeding wetlands for a long period and are strongly affected by fish introduction; however, mechanisms of the exclusion observed in the field are still not well known. In particular, whether newts skip breeding and leave water for land in response to fish introduction and how aquatic shelter may influence their choice remain open questions. To investigate these questions, we experimentally studied the use of aquatic and terrestrial habitats during the breeding season of palmate newts, Lissotriton helveticus, in the presence and absence of goldfish, Carassius auratus. We determined the consequences of habitat choice on newt fitness by assessing sexual activity and number of eggs. There was a strong, significant avoidance of the aquatic environment in the presence of fish, particularly when no aquatic shelter was available. This escape from the water had an impact on reproduction: newts decreased their sexual activity and laid fewer eggs. The availability of shelters favoured coexistence but did not prevent a large proportion of the newts from leaving water and skipping reproduction. This study shows how the presence of fish and the absence of aquatic shelters can lead to newts forgoing aquatic life, thus improving our understanding of the mechanisms behind the coexistence and exclusion patterns found in the wild. More broadly, these data contribute to explaining aquatic versus terrestrial life in favourable and unfavourable environments.

Despite growing evidence of the detrimental effect of chemical substances on organisms, limited research has focused on changes in behavioral patterns, in part due to the difficulties to obtain detailed quantitative data. Recent developments in efficient computer-based video analyses have allowed testing pesticide effects on model species such as the zebrafish. However, these new techniques have not yet been applied to amphibians and directly to conservation issues, i.e., to assess toxicological risks on threatened species. We used video-tracking analyses to test a quantitative effect of an environmental contaminant on the locomotion of amphibian tadpoles (Rana temporaria) by taking into account cumulative effects. Because recent research has demonstrated effects of de-icing salts on survival and community structure, we used sodium chloride in our experimental design (25 replicates, 4 concentrations, 4 times) to test for an effect at the scale of behavior at environmentally relevant concentrations. Analysis of 372 1-h video-tracks (5 samples/s) showed a complex action of salts on behavioral patterns with a dose and cumulative response over time. Although no effects were found on mortality or growth, the highest salt concentrations reduced the speed and movement of tadpoles in comparison with control treatments. The reduced locomotor performance could have detrimental consequences in terms of tadpoles' responses to competition and predation and may be an indicator of the low concentration effect of the contaminant. On one hand, this study demonstrates the usefulness of examining behavior to address conservation issues and understand the complex action of environmental factors and, more particularly, pollutants on organisms. On the other hand, our results highlight the need of new computerized techniques to quantitatively analyze these patterns.

Increa ses of g loba l temperat u res have resu lted i n mea su rable sh i f ts i n t he d ist r ibut ion, phenolog y a nd survival of some plant and animal species. However, the mechanisms showing links between global warming and biodiversity declines remain unclear. The aim of this study was to examine whether a key parameter of fitness, i.e. offspring number, could be affected by a temperature increase. To this end, we compared egg-laying traits at naturally occurring temperatures (14°C, 18°C and 22°C) in palmate newts, Lissotriton helveticus . Our study suggests that water temperature increase has a negative effect on the fecu nd it y of fema le new ts. Fema les lay ha l f a s ma ny eg gs at h ig h temperat u res a s t hey do at low temperatures, which results in a lower number of hatchlings. This study shows that global warming would affect amphibian populations. It complements other studies in pointing out that changes in phenology may not be driven only by warmer earlier temperatures but also by counter-selec tion during late-breed i ng , pa r t icu la rly i n long-ter m breeders such a s new ts. More ex per i menta l st ud ies shou ld be carried out to understand the complex consequences of global warming and the proximate mechanisms of amphibian decline. 2009 Elsevier Masson SAS. All rights reserved.

Facultative paedomorphosis, a process in which newt larvae can opt for reproduction before or after metamorphosis, is geographically heterogeneous. Despite numerous ecological studies and recent evidence of declines in paedomorphic populations, however, no attempt to model environmental variables that explain the presence of paedomorphs has been made at a multi-scale level. Our aim was to fill this gap in studying three newt species (Lissotriton vulgaris, Mesotriton alpestris, and Triturus macedonicus) of the Montenegrin karst as model species. To this end, we used multivariate analysis on three scales of habitat: the breeding pond, the land use and the climatologic features. Results show that the study area is both an important hotspot for paedomorphosis and where intraspecific diversity is quickly disappearing (20-47% extirpation) because of fish introductions. Other habitat variables (water permanency, pH or the habitat origin) were shown to act on paedomorphosis but not consistently across species, confirming complexity of the evolutionary and ecological processes. This study appeals for more long-term and detailed landscape studies of polyphenisms, a neglected but promising topic, to better understand and protect alternative modes of development. Particularly, measures should be taken to identify hotspots of intraspecific diversity at a global scale and stop fish introductions before we reach a point of no-return.

Oophagy is a behavioural pattern that has been found in a large variety of predator species in the animal kingdom. In contrast to other modes of feeding, it is peculiar in that it involves the detection, capture and ingestion of immobile prey. Several hypotheses have been proposed to explain the evolutionary origin and persistence of this pattern, but they have rarely been tested. The aim of this study was to compare the benefits of a heterospecific oophagous tactic over a non-oophageous diet in terms of biomass intake. To this end, stomach contents were gathered by flushing the stomachs of male and female Alpine newts (Mesotriton alpestris) found in forestry ruts (i.e. pools caused by traffic) during their reproductive period. Prey items were identified, classified into functional categories and their dry mass determined. Frog (Rana temporaria) eggs are valuable prey items that give a higher biomass intake to individuals foraging on them than on those relying on invertebrates. Both sexes of newts practice oophagy but frog eggs are a transient resource that is only available during a part of their aquatic phase. Consequently, the newts adjust their diet to invertebrate predation later in the season after the peak of the frogs' breeding season. Oophagy is thus facultative and not obligate in the study species. The correlated occurrence of prey and predator, similarities between frog eggs and mobile potential prey (tadpoles), and high resource intake are all in favour of the occurrence and persistence of an oophagous feeding tactic.

Recent studies in population dynamics suggest that landscape processes and habitat quality act at different scales on population abundances, but very few have modelled their simultaneous effects. However, at a time of large declines in natural populations, it is essential to understand such multivariate components. We tested the hypothesis that natural populations of palmate newts (Triturus helveticus) are affected on three scales: breeding patch (pond), habitat complementation (terrestrial cover), and metapopulation structure (density of ponds, surrounding populations). We conducted our survey in 130 ponds from southern France (Larzac) and analysed data with generalized additive models (GAM). Two main novel results emerge from these models: (1) the three landscape scales have significant effects on newt abundance, with more newts in deep, vegetated ponds, devoid of fish and surrounded by wooded areas and inhabited ponds; (2) the quality of the surrounding breeding patches is of primary importance in determining the abundance at core sites in a complex way: high abundances are associated positively with high densities of inhabited ponds, but negatively with the number of surrounding ponds. Deforestation, invasive species and abandonment of ponds all have negative impacts on the persistence of palmate newt populations. Future studies should encompass landscapes at different scales and incorporate the habitat quality in surrounding sites to better understand population dynamics and provide adequate conservation measures.

Facultative paedomorphosis is a polyphenism in which individuals may express one of two alternative ontogenetic pathways (metamorphosis vs. paedomorphosis) depending on environmental cues. Previous laboratory experiments showed that drying can cause morph ratio change, suggesting that the maintenance of facultative paedomorphosis is highly dependent on environmental determinants. The aim of this study was to examine seasonal variation in morph ratios in eight ponds from Larzac (southern France) naturally inhabited by palmate newts and to relate it to pond drying. In some ponds, the relative proportion of paedomorphs (i.e. individuals retaining gills at the adult stage) increased after the breeding period, but it remained stable or decreased in other ponds. This seasonal variation in the abundance of the two morphs most probably reflects (1) the emigration of metamorphs leaving the pond to occupy terrestrial habitats and (2) metamorphosis of paedomorphic adults in response to drying of the ponds. This study shows that facultative paedomorphosis in palmate newts is a dynamic process that allows rapid change (i.e. within a single year) in morph ratio to fit environmental variation (i.e. risk of drying) within the aquatic habitats. Long-term studies are needed to model the evolution of the dimorphism according to environmental change.

Current ecological models suggest that the maintenance of trophic polymorphisms is favoured by a different resource use in alternative morphs. Facultative paedomorphosis in newts is an example of phenotypic variation as paedomorphs retain morphological larval traits, such as gills and gill slits. The aim of this study was to find out whether heterochronic morphs occupy particular micro-habitats and focus on specific prey items. Resource partitioning was found between morphs. It concerns mainly food selection with paedomorphs preying more on plankton and less on terrestrial invertebrates than metamorphs. Some habitat specializations were also found with metamorphs being more abundant at the water surface than paedomorphs. Diel variation in habitat use of the two different morphs was minimal. Polymorphism allows Alpine newts to exploit the different resources in the lakes in order to minimize intraspecific competition, but the extent of resource partitioning depends on habitat characteristics.

Paedomorphosis, in which individuals retain ancestral characteristics in the adult stage, is widespread in newts and salamanders and is suspected to play an important role in evolution. In some species, paedomorphosis is facultative with some individuals forgoing metamorphosis. Optimality models have been proposed to explain the maintenance of this polymorphism, but require the integration of reproductive patterns into the models. We investigated the frequencies of inbreeding and outbreeding in two syntopic heterochronic morphs of the Alpine newt. The two morphs are sexually compatible: encounters between and within morphs were equally successful in terms of spermatophore transfer. Behavioural observations were in agreement with the sexual compatibility observed. Nevertheless, paedomorphic males displayed to females less frequently than metamorphic males. The two morphs differ largely on the basis of sexual secondary characteristics, but the majority of these traits did not affect mating success. Because of the large flow of genes between the two heterochronic morphs and because of the absence of spatial and temporal isolation, these results do not support sympatric speciation models, but are in favour of the maintenance of polymorphism in natural populations.