Diet diversity and environment determine the intestinal microbiome and bacterial pathogen load of fire salamanders.

Diverse communities of symbiotic microbes inhabit the digestive systems of vertebrates and play a crucial role in animal health, and host diet plays a major role in shaping the composition and diversity of these communities. Here, we characterized diet and gut microbiome of fire salamander populations from three Belgian forests. We carried out DNA metabarcoding on fecal samples, targeting eukaryotic 18S rRNA of potential dietary prey items, and bacterial 16S rRNA of the concomitant gut microbiome. Our results demonstrated an abundance of soft-bodied prey in the diet of fire salamanders, and a significant difference in the diet composition between males and females. This sex-dependent effect on diet was also reflected in the gut microbiome diversity, which is higher in males than female animals. Proximity to human activities was associated with increased intestinal pathogen loads. Collectively, the data supports a relationship between diet, environment and intestinal microbiome in fire salamanders, with potential health implications.

Toxin variation among salamander populations: discussing potential causes and future directions.

Amphibians produce defensive chemicals which provide protection against both predators and infections. Within species, populations can differ considerably in the composition and amount of these chemical defenses. Studying intraspecific variation in toxins and linking it to environmental variables may help us to identify the selective drivers of toxin evolution, such as predation pressure and infection risk. Recently, there has been a renewed interest in the unique toxins produced by salamanders from the genus Salamandra: the samandarines. Despite this attention, intraspecific variation has largely been ignored within Salamandra-species. The aim of this study was to investigate whether geographic variation in profiles of samandarines exists, by sampling 4 populations of Salamandra atra over its range in the Dinaric Alps. In addition, we preliminary explored whether potential variation could be explained by predation (counting the number of snake species) and infection risk (cultivation and genomic analyses of collected soil samples). Salamanders from the 4 populations differed in toxin composition and in the size of their poison glands, although not in overall toxin quantity. Nor predation nor infection risk could explain this variation, as populations barely differed in these variables. Sampling over a much broader geographic range, using better estimators for predation and infection risk, will contribute to an improved understanding of how environment may shape variation in chemical defenses. Nevertheless, as the 4 populations of S. atra did differ in their toxin profiles, we propose that this species provides an interesting opportunity for further ecological and evolutionary studies on amphibian toxins.

Weighing density and kinship: Aggressive behavior and time allocation in fire salamander (Salamandra infraimmaculata).

Kin-biased behavior (that is responding differentially to kin and non-kin) is thought to be adaptive in many social interactions. One example of this kin bias is behaving less aggressively toward a relative than a non-relative, a behavior which yields inclusive fitness benefits. However, data are lacking about the ability of animals to weigh their preference for kinship and the density of conspecifics simultaneously and to respond accordingly. Fire salamanders (Salamandra infraimmaculata) larviposit in high densities in ponds. Thus, larvae of different females confront competition and predation by other larvae. We studied whether larvae prefer their kin over particular density or vice versa. We experimentally used a transparent glass aquarium with inner chambers to test the responses of a focal larva toward its siblings and non-siblings. Specifically, we quantified the time a focal larva spent near its siblings or non-siblings, presented in varying densities, and the aggression level it demonstrated. We found that focal larvae spent more time near non-siblings if non-sibling and sibling groups were of equal density. The focal larvae were also more aggressive toward non-siblings. The results may be explained by the cannibalistic nature of these larvae: high density may provide more opportunities for food, especially when non-siblings are present. Further explanations for these findings may include other advantages of staying in a larger group and/or the stronger olfactory and visual stimulation offered by groups compared to a single individual. These findings suggest that larvae make differential responses toward conspecifics depending simultaneously on the level of relatedness and the density of the group. Such responses have important implications for social-aggregation decisions and may especially affect the fitness of cannibalistic species.

The conspicuous postmetamorphic coloration of fire salamanders, but not their toxicity, is affected by larval background albedo.

Experimental work in the early 20th century showed that background albedo experienced by larvae of the fire salamander (Salamandra salamandra) induce a durable morphological modification of the postmetamorphic color pattern, which needed confirmation due to the controversies regarding Paul Kammerer's experiments. Such a carry-over effect would be relevant as the black and yellow pattern of the alkaloid-containing adult fire salamanders has been suggested to serve as an aposematic signal. Hence, we hypothesized that (a) adult coloration is conspicuous to potential predators under light conditions at night, given the nocturnal activity of this species, and (b) a condition affecting the salamander's coloration pattern would also affect its toxicity to maintain a quantitatively honest aposematic signal. To test the first hypothesis, we used spectrometry to model the vision of potential avian and snake predators and confirmed that fire salamander's black-and-yellow pattern is contrasting enough against the forest leaf litter to be considered conspicuous at night. To test the second hypothesis, we first confirmed the background carry-over effect on black and yellow proportions in the dorsal skin of experimentally reared fire salamanders, using a rigorous experimental design. Then, we calculated the conspicuousness and determined the alkaloid profiles of these individuals. We did not find a correlation between conspicuousness and toxicity at the intrapopulation level. Moreover, there was no background carry-over effect on the alkaloid profile. We discuss our results in a physiological, ecological, evolutionary, and historical context.

What shapes the trophic niche of European plethodontid salamanders?

The trophic niche is a life trait that identifies the consumer's position in a local food web. Several factors, such as ontogeny, competitive ability and resource availability contribute in shaping species trophic niches. To date, information on the diet of European Hydromantes salamanders are only available for a limited number of species, no dietary studies have involved more than one species of the genus at a time, and there are limited evidences on how multiple factors interact in determining diet variation. In this study we examined the diet of multiple populations of six out of the eight European cave salamanders, providing the first data on the diet for five of them. In addition, we assessed whether these closely related generalist species show similar diet and, for each species, we tested whether season, age class or sex influence the number and the type of prey consumed. Stomach condition (empty/full) and the number of prey consumed were strongly related to seasonality and to the activity level of individuals. Empty stomachs were more frequent in autumn, in individuals far from cave entrance and in juveniles. Diet composition was significantly different among species. Hydromantes imperialis and H. supramontis were the most generalist species; H. flavus and H. sarrabusensis fed mostly on Hymenoptera and Coleoptera Staphylinidae, while H. genei and H. ambrosii mostly consumed Arachnida and Endopterygota larvae. Furthermore, we detected seasonal shifts of diet in the majority of the species examined. Conversely, within each species, we did not find diet differences between females, males and juveniles. Although being assumed to have very similar dietary habits, here Hydromantes species were shown to be characterized by a high divergence in diet composition and in the stomach condition of individuals.

The effect of brumation on memory retention.

Long-term torpor is an adaptive strategy that allows animals to survive harsh winter conditions. However, the impact that prolonged torpor has on cognitive function is poorly understood. Hibernation causes reduced synaptic activity and experiments with mammals reveal that this can have adverse effects on memories formed prior to hibernation. The impact of brumation, the winter dormancy that is observed in ectotherms, on memory remains unknown. The aim of this study was to examine whether an amphibian, the fire salamander (Salamandra salamandra), was able to retain learned spatial information after a period of brumation. Twelve fire salamanders were trained to make a simple spatial discrimination using a T-maze. All subjects learned the initial task. Upon reaching criterion, half of the subjects were placed into brumation for 100 days while the other half served as controls and were maintained under normal conditions. A post-brumation memory retention test revealed that animals from both conditions retained the learned response. Control tests showed that they solved the task using learned information and not olfactory cues. This finding contrasts with much of the mammalian research and suggests that the processes involved in prolonged torpor may have a fundamentally different impact on memory in mammals and amphibians.

Salamandra.

Beukema et al. Introduce the genus salamandra comprising the well-known fire salamanders.

Thermoregulatory strategies in an aquatic ectotherm from thermally-constrained habitats: An evaluation of current approaches.

Many ectotherms employ diverse behavioral adjustments to effectively buffer the spatio-temporal variation in environmental temperatures, whereas others remain passive to thermal heterogeneity. Thermoregulatory studies are frequently performed on species living in thermally benign habitats, which complicate understanding of the thermoregulation-thermoconformity continuum. The need for new empirical data from ectotherms exposed to thermally challenging conditions requires the evaluation of available methods for quantifying thermoregulatory strategies. We evaluated the applicability of various thermoregulatory indices using fire salamander larvae, Salamandra salamandra, in two aquatic habitats, a forest pool and well, as examples of disparate thermally-constrained environments. Water temperatures in the well were lower and less variable than in the pool. Thermal conditions prevented larvae from reaching their preferred body temperature range in both water bodies. In contrast to their thermoregulatory abilities examined in a laboratory thermal gradient, field body temperatures only matched the mean and range of operative temperatures, showing thermal passivity of larvae in both habitats. Despite apparent thermoconformity, thermoregulatory indices indicated various strategies from active thermoregulation, to thermoconformity, and even thermal evasion, which revealed their limited applicability under thermally-constrained conditions. Salamander larvae abandoned behavioral thermoregulation despite varying opportunities to increase their body temperature above average water temperatures. Thermoconformity represents a favored strategy in these ectotherms living in more thermally-constrained environments than those examined in previous thermoregulatory studies. To understand thermal ecology and its impact on population dynamics, the quantification of thermoregulatory strategies of ectotherms in thermally-constrained habitats requires the careful choice of an appropriate method to avoid misleading results.

Exotic Fish in Exotic Plantations: A Multi-Scale Approach to Understand Amphibian Occurrence in the Mediterranean Region.

Globally, amphibian populations are threatened by a diverse range of factors including habitat destruction and alteration. Forestry practices have been linked with low diversity and abundance of amphibians. The effect of exotic Eucalyptus spp. plantations on amphibian communities has been studied in a number of biodiversity hotspots, but little is known of its impact in the Mediterranean region. Here, we identify the environmental factors influencing the presence of six species of amphibians (the Caudata Pleurodeles waltl, Salamandra salamandra, Lissotriton boscai, Triturus marmoratus and the anurans Pelobates cultripes and Hyla arborea/meridionalis) occupying 88 ponds. The study was conducted in a Mediterranean landscape dominated by eucalypt plantations alternated with traditional use (agricultural, montados and native forest) at three different scales: local (pond), intermediate (400 metres radius buffer) and broad (1000 metres radius buffer). Using the Akaike Information Criterion for small samples (AICc), we selected the top-ranked models for estimating the probability of occurrence of each species at each spatial scale separately and across all three spatial scales, using a combination of covariates from the different magnitudes. Models with a combination of covariates at the different spatial scales had a stronger support than those at individual scales. The presence of predatory fish in a pond had a strong effect on Caudata presence. Permanent ponds were selected by Hyla arborea/meridionalis over temporary ponds. Species occurrence was not increased by a higher density of streams, but the density of ponds impacted negatively on Lissotriton boscai. The proximity of ponds occupied by their conspecifics had a positive effect on the occurrence of Lissotriton boscai and Pleurodeles waltl. Eucalypt plantations had a negative effect on the occurrence of the newt Lissotriton boscai and anurans Hyla arborea/meridionalis, but had a positive effect on the presence of Salamandra salamandra, while no effect on any of the other species was detected. In conclusion, eucalypts had limited effects on the amphibian community at the intermediate and broad scales, but predatory fish had a major impact when considering all the scales combined. The over-riding importance of introduced fish as a negative impact suggests that forest managers should prevent new fish introductions and eradicate fish from already-occupied ponds whenever possible.

Landscape influences on dispersal behaviour: a theoretical model and empirical test using the fire salamander, Salamandra infraimmaculata.

When populations reside within a heterogeneous landscape, isolation by distance may not be a good predictor of genetic divergence if dispersal behaviour and therefore gene flow depend on landscape features. Commonly used approaches linking landscape features to gene flow include the least cost path (LCP), random walk (RW), and isolation by resistance (IBR) models. However, none of these models is likely to be the most appropriate for all species and in all environments. We compared the performance of LCP, RW and IBR models of dispersal with the aid of simulations conducted on artificially generated landscapes. We also applied each model to empirical data on the landscape genetics of the endangered fire salamander, Salamandra infraimmaculata, in northern Israel, where conservation planning requires an understanding of the dispersal corridors. Our simulations demonstrate that wide dispersal corridors of the low-cost environment facilitate dispersal in the IBR model, but inhibit dispersal in the RW model. In our empirical study, IBR explained the genetic divergence better than the LCP and RW models (partial Mantel correlation 0.413 for IBR, compared to 0.212 for LCP, and 0.340 for RW). Overall dispersal cost in salamanders was also well predicted by landscape feature slope steepness (76%), and elevation (24%). We conclude that fire salamander dispersal is well characterised by IBR predictions. Together with our simulation findings, these results indicate that wide dispersal corridors facilitate, rather than hinder, salamander dispersal. Comparison of genetic data to dispersal model outputs can be a useful technique in inferring dispersal behaviour from population genetic data.

The more the better - polyandry and genetic similarity are positively linked to reproductive success in a natural population of terrestrial salamanders (Salamandra salamandra).

Although classically thought to be rare, female polyandry is widespread and may entail significant fitness benefits. If females store sperm over extended periods of time, the consequences of polyandry will depend on the pattern of sperm storage, and some of the potential benefits of polyandry can only be realized if sperm from different males is mixed. Our study aimed to determine patterns and consequences of polyandry in an amphibian species, the fire salamander, under fully natural conditions. Fire salamanders are ideal study objects, because mating, fertilization and larval deposition are temporally decoupled, females store sperm for several months, and larvae are deposited in the order of fertilization. Based on 18 microsatellite loci, we conducted paternity analysis of 24 female-offspring arrays with, in total, over 600 larvae fertilized under complete natural conditions. More than one-third of females were polyandrous and up to four males were found as sires. Our data clearly show that sperm from multiple males is mixed in the female's spermatheca. Nevertheless, paternity is biased, and the most successful male sires on average 70% of the larvae, suggesting a 'topping off' mechanism with first-male precedence. Female reproductive success increased with the number of sires, most probably because multiple mating ensured high fertilization success. In contrast, offspring number was unaffected by female condition and genetic characteristics, but surprisingly, it increased with the degree of genetic relatedness between females and their sires. Sires of polyandrous females tended to be genetically similar to each other, indicating a role for active female choice.

Review of the systematics, distribution, biogeography and natural history of Moroccan amphibians.

The amphibian fauna of the Kingdom of Morocco was traditionally regarded as poor and closely related to its European counterpart. However, an increase in research during the last decades revealed a considerable degree of endemism amongst Moroccan amphibians, as well as phenotypic and genotypic inter- and intraspecific divergence. Despite this increase in knowledge, a comprehensible overview is lacking while several systematic issues have remained unresolved. We herein present a contemporary overview of the distribution, taxonomy and biogeography of Moroccan amphibians. Fourteen fieldtrips were made by the authors and colleagues between 2000 and 2012, which produced a total of 292 new distribution records. Furthermore, based on the results of the present work, we (i) review the systematics of the genus Salamandra in Morocco, including the description of a new subspecies from the Rif- and Middle Atlas Mountains, Salamandra algira splendens ssp. nov.; (ii) present data on intraspecific morphological variability of Pelobates varaldiiand Pleurodeles waltl in Morocco; (iii) attempt to resolve the phylogenetic position of Bufo brongersmai and erect a new genus for this species, Barbarophryne gen. nov.; (iv) summarize and assess the availability of tadpole-specific characteristics and bioacoustical data, and (v) summarize natural history data.

Brainstem reticulospinal neurons are targets for corticotropin-releasing factor-Induced locomotion in roughskin newts.

Stress-induced release or central administration of corticotropin-releasing factor (CRF) enhances locomotion in a wide range of vertebrates, including the roughskin newt, Taricha granulosa. Although CRF's stimulatory actions on locomotor behavior are well established, the target neurons through which CRF exerts this effect remain unknown. To identify these target neurons, we utilized a fluorescent conjugate of CRF (CRF-TAMRA 1) to track this peptide's internalization into reticulospinal and other neurons in the medullary reticular formation (MRF), a region critically involved in regulating locomotion. Epifluorescent and confocal microscopy revealed that CRF-TAMRA 1 was internalized by diverse MRF neurons, including reticulospinal neurons retrogradely labeled with Cascade Blue dextran. In addition, we immunohistochemically identified a distinct subset of serotonin-containing neurons, located throughout the medullary raphé, that also internalized the fluorescent CRF-TAMRA 1 conjugate. Chronic single-unit recordings obtained from microwire electrodes in behaving newts revealed that intracerebroventricular (icv) administration of CRF-TAMRA 1 increased medullary neuronal firing and that appearance of this firing was associated with, and strongly predictive of, episodes of CRF-induced locomotion. Furthermore, icv administered CRF-TAMRA 1 produced behavioral and neurophysiological effects identical to equimolar doses of unlabeled CRF. Collectively, these findings provide the first evidence that CRF directly targets reticulospinal and serotonergic neurons in the MRF and indicate that CRF may enhance locomotion via direct effects on the hindbrain, including the reticulospinal system.

Comparative skull osteology of Karsenia koreana (Amphibia, Caudata, Plethodontidae).

The recent discovery of a plethodontid salamander, Karsenia koreana, in Korea challenged our understanding of the biogeographic history of the family Plethodontidae, by far the largest family of salamanders, which otherwise is distributed in the New World with a few European species. Molecular studies suggest that Karsenia forms a clade with Hydromantes (sensu lato), which includes among its species the only other Old World plethodontids. We studied the skull of K. koreana and compared it with that of other plethodontid genera, especially members of the subfamily Plethodontinae, which it resembles most closely in general anatomy. The anatomy of its skull corresponds to the most generalized and apparently ancestral condition for plethodontids. No clearly autapomorphic states were detected, and no synapomorphies can be found that would link it to other genera. The Karsenia skull is cylindrical and well ossified, giving an impression of strength. In contrast, the skull of Hydromantes is highly derived; the skull is flattened and the bones are weakly ossified and articulated. Hydromantes and Karsenia share no unique anatomical features; differences between them are especially evident in the hyobranchial skeleton, which is generalized in Karsenia but highly modified in Hydromantes, which is well known for its highly projectile tongue. Plethodon and Plethodon-like species, including Karsenia and to a lesser degree Ensatina, represent the more generalized and apparently ancestral plethodontid morphology. Specialized morphologies have evolved along only a few morphological axes within the Plethodontidae, resulting in a pattern of rampant homoplasy. Our analysis of the anatomy of the new Asiatic lineage illuminates some potential mechanisms underlying adaptive morphological evolution within the Plethodontidae.

Context-dependent reproductive habitat selection: the interactive roles of structural complexity and cannibalistic conspecifics.

Structural complexity generally reduces predation and cannibalism rates. Although the benefits from this effect vary among environmental contexts and through time, it has been the common explanation for high species abundance in complex habitats. We hypothesized that oviposition habitat selection for structural complexity depends on the expected trophic function of the progeny. In Salamandra infraimmaculata larvae, expected trophic function is dictated by their sequence of deposition. First cohorts cannibalize later-arriving cohorts, while all compete for shared prey resources. In a mesocosm experiment, we show that gravid salamanders facing conspecific-free pools preferred structurally simple habitats (no rocks), while females facing only pools with older conspecific larvae preferred complex habitats (with rocks). Context-dependent preference of habitat complexity for managing food/safety trade-offs may be extended from classic foraging patch decisions to breeding habitat selection. These trade-offs vary with dynamic larval processes such as priority effects and ontogenetic diet shifts, potentially leading to complex maternal parturition behaviours.

Influence of landscape elements in riparian buffers on the conservation of semiaquatic amphibians.

Studies on riparian buffers have usually focused on the amount of land needed as habitat for the terrestrial life stages of semiaquatic species. Nevertheless, the landscape surrounding wetlands is also important for other key processes, such as dispersal and the dynamics of metapopulations. Multiple elements that influence these processes should therefore be considered in the delineation of buffers. We analyzed landscape elements (forest cover, density of roads, and hydrographic network) in concentric buffers to evaluate the scale at which they influence stream amphibians in 77 distinct landscapes. To evaluate whether our results could be generalized to other contexts, we determined whether they were consistent across the study areas. Amphibians required buffers of 100-400 m of suitable terrestrial habitat, but interspecific differences in the amount of habitat were large. The presence of amphibians was related to roads and the hydrographic network at larger spatial scales (300-1500 m), which suggests that wider buffers are needed with these elements. This pattern probably arose because these elements influence dispersal and metapopulation persistence, processes that occur at large spatial scales. Furthermore, in some cases, analyses performed on different sets of landscapes provided different results, which suggests caution should be used when conservation recommendations are applied to disparate areas. Establishment of riparian buffers should not be focused only on riparian habitat, but should take a landscape perspective because semiaquatic species use multiple elements for different functions. This approach can be complex because different landscape elements require different spatial extents. Nevertheless, a shift of attention toward the management of different elements at multiple spatial scales is necessary for the long-term persistence of populations.

Tracing the first step to speciation: ecological and genetic differentiation of a salamander population in a small forest.

Mechanisms and processes of ecologically driven adaptive speciation are best studied in natural situations where the splitting process is still occurring, i.e. before complete reproductive isolation is achieved. Here, we present a case of an early stage of adaptive differentiation under sympatric conditions in the fire salamander, Salamandra salamandra, that allows inferring the underlying processes for the split. Larvae of S. salamandra normally mature in small streams until metamorphosis, but in an old, continuous forest area near Bonn (the Kottenforst), we found salamander larvae not only in small streams but also in shallow ponds, which are ecologically very different from small streams. Common-environment experiments with larvae from both habitat types reveal specific adaptations to these different ecological conditions. Mitochondrial and microsatellite analyses show that the two ecologically differentiated groups also show signs of genetic differentiation. A parallel analysis of animals from a neighbouring much larger forest area (the Eifel), in which larvae mature only in streams, shows no signs of genetic differentiation, indicating that gene flow between ecologically similar types can occur over large distances. Hence, geographical factors cannot explain the differential larval habitat adaptations in the Kottenforst, in particular since adult life and mating of S. salamandra is strictly terrestrial and not associated with larval habitats. We propose therefore that the evolution of these adaptations was coupled with the evolution of cues for assortative mating which would be in line with models of sympatric speciation that suggest a co-evolution of habitat adaptations and associated mating signals.

Waterborne amitrole affects the predator-prey relationship between common frog tadpoles (Rana temporaria) and larval spotted salamander (Salamandra salamandra).

Within their aquatic habitats, larval amphibians are often subjected to multiple natural and anthropic stressors. Among these, predation and waterborne pollution represent two types of stressing factor that frequently co-occur. In this connection, the present laboratory study was designed to investigate the effects of amitrole, a commonly used triazole herbicide, on the predator-prey relationship between common frog tadpoles (Rana temporaria) and larval spotted salamander (Salamandra salamandra). Tadpoles were exposed for 3 days to 0, 0.01, 0.1, 1, and 10 mg/L amitrole, either in the absence or in the presence of larval salamanders. Tadpole behavior (refuge use, movements) was monitored every day, and the predation efficiency was assessed at the end of the experiment by counting the number of surviving tadpoles. In the absence of the predator, amitrole-exposed tadpoles (at 0.01, 0.1, and 1 mg/L) increased their refuge use and decreased their rate of movements. In the presence of the predator, amitrole contamination did not affect tadpole behavior, except on the first day, where tadpoles exposed to 10 mg/L were found to be significantly more active than unexposed control tadpoles. Throughout the experiment, control tadpoles were the only group to show significant reductions of activity and visibility in response to the predator's presence. In contrast, tadpoles exposed to 0.01 and 0.1 mg/L amitrole increased their refuge use in response to the predator, whereas their rate of movements remained unaffected. Furthermore, exposures of tadpoles to the two highest amitrole concentrations (1 and 10 mg/L) resulted in the loss of both behavioral responses to the predator's presence. Interestingly, the lack of antipredator behavior in amitrole-exposed tadpoles did not enhance their vulnerability to predation by the larval salamander. Moreover, tadpoles exposed to the two highest herbicide concentrations showed a better survival than unexposed controls, indicating that amitrole contamination also had detrimental effects on the predatory behavior of the larval salamander. These findings emphasize the need to consider the effects of contaminants on both predator and prey before drawing conclusions about the possible consequences of prey behavioral modifications on the predation risk.

Evolutionary ecology of facultative paedomorphosis in newts and salamanders.

Facultative paedomorphosis is an environmentally induced polymorphism that results in the coexistence of mature, gilled, and fully aquatic paedomorphic adults and transformed, terrestrial, metamorphic adults in the same population. This polymorphism has been of interest to scientists for decades because it occurs in a large number of caudate amphibian taxa as well as in a large diversity of habitats. Numerous experimental and observational studies have been conducted to explain the proximate and ultimate factors affecting these heterochronic variants in natural populations. The production of each alternative phenotype is based on a genotypexenvironment interaction and research suggests that differences in the environment can produce paedomorphs through several ontogenetic pathways. No single advantage accounts for the maintenance of this polymorphism. Rather, the interplay of different costs and benefits explains the success of the polyphenism across variable environments. Facultative paedomorphosis allows individuals to cope with habitat variation, to take advantage of environmental heterogeneity in the presence of open niches, and to increase their fitness. This process is expected to constitute a first step towards speciation events, and is also an example of biodiversity at the intraspecific level. The facultative paedomorphosis system is thus ripe for future studies encompassing ecology, evolution, behaviour, endocrinology, physiology, and conservation biology. Few other systems have been broad enough to provide varied research opportunities on topics as diverse as phenotypic plasticity, speciation, mating behaviour, and hormonal regulation of morphology. Further research on facultative paedomorphosis will provide needed insight into these and other important questions facing biologists.