Larval life history and anti-predator strategies are affected by breeding phenology in an amphibian

Seasonal time constraints can pose strong selection on life histories. Time-constrained animals should prioritise fast development over predation risk to avoid unfavourable growing conditions. However, changes in phenology could alter the balance between anti-predator and developmental needs. We studied variation of anti-predator strategies in common frog (Rana temporaria) tadpoles in four populations from the two extremes of a latitudinal gradient across Sweden. We examined, under common conditions in the laboratory, the anti-predator responses and life histories of tadpoles raised with predatory Aeshna dragonfly larvae in two consecutive years with a difference of 20 days in breeding time in the north, but no difference in breeding time in the nouth. In a year with late breeding, northern tadpoles did not modify their behaviour and morphology in the presence of predators, and metamorphosed faster and smaller than tadpoles born in a year with early breeding. In the year with early breeding, northern tadpoles showed a completely different anti-predator strategy by reducing activity and developing morphological defences in the presence of predators. We discuss the possible mechanisms that could activate these responses (likely a form of environmentally-mediated parental effect). To our knowledge, this is the first study to show that a vertebrate modifies the anti-predator strategy of its offspring in response to natural variation in reproductive phenology, which highlights the need to consider phenology in studies of life-history evolution.     

Compensating for delayed hatching reduces offspring immune response and increases life‐history costs

Organisms are exposed to multiple sources of stress in nature. When confronted with a stressful period affecting growth and development, compensatory responses allow the restoration of individual fitness, providing an important buffering mechanism against climatic and other environmental variability. However, tradeoffs between increased growth/development and other physiological traits are predicted to prevent these high growth and development rates from becoming constitutive. Here, we investigated how compensatory responses in growth and development affect immune responses. By using low temperature to stop embryonic development, we exposed moor frog Rana arvalis tadpoles to two levels of time‐constraints: non‐delayed hatching and 12‐day delayed hatching. In a common garden experiment, we recorded larval growth and development, as well as their immune response, measured as the inflammatory reaction after the injection of phytohaemagglutinin (PHA). Tadpoles originating from delayed hatching treatments had a lower immune response to PHA challenge than those from the non‐delayed hatching treatment. In general, tadpoles from the delayed hatching treatment reached metamorphosis faster and at a smaller size than control tadpoles. However, immune‐challenged tadpoles were not able to accelerate their development in response to delayed hatching. Our results indicate that 1) the innate immune response can be reduced in organisms undergoing compensatory developmental responses in growth and development and 2) compensatory capacity can be reduced when organisms are immunologically challenged. These dual findings reveal the complexity of handling multiple stressors and highlight the importance of examining the costs and limits of mounting an immune response in the context of increasing phenological instability ascribed to climate change.     

Differential reproductive responses to stress reveal the role of life-history strategies within a species

Life-history strategies describe that ‘slow’- in contrast to ‘fast’-living species allocate resources cautiously towards reproduction to enhance survival. Recent evidence suggests that variation in strategies exists not only among species but also among populations of the same species. Here, we examined the effect of experimentally induced stress on resource allocation of breeding seabirds in two populations with contrasting life-history strategies: slow-living Pacific and fast-living Atlantic black-legged kittiwakes. We tested the hypothesis that reproductive responses in kittiwakes under stress reflect their life-history strategies. We predicted that in response to stress, Pacific kittiwakes reduce investment in reproduction compared with Atlantic kittiwakes. We exposed chick-rearing kittiwakes to a short-term (3-day) period of increased exogenous corticosterone (CORT), a hormone that is released during food shortages. We examined changes in baseline CORT levels, parental care and effects on offspring. We found that kittiwakes from the two populations invested differently in offspring when facing stress. In response to elevated CORT, Pacific kittiwakes reduced nest attendance and deserted offspring more readily than Atlantic kittiwakes. We observed lower chick growth, a higher stress response in offspring and lower reproductive success in response to CORT implantation in Pacific kittiwakes, whereas the opposite occurred in the Atlantic. Our findings support the hypothesis that life-history strategies predict short-term responses of individuals to stress within a species. We conclude that behaviour and physiology under stress are consistent with trade-off priorities as predicted by life-history theory. We encourage future studies to consider the pivotal role of life-history strategies when interpreting inter-population differences of animal responses to stressful environmental events.     

Influence of seasonal time constraints on growth and development of common frog tadpoles: a photoperiod experiment

Organisms living in seasonal environments are often limited by the time available to complete their development. Especially individuals in northern populations may face severe time constraints in their need of completing development before the end of the growth season. Larval amphibians have been widely used in studies of phenotypic plasticity. However, their responses to changes in photoperiod, the main seasonal cue in many organisms, are unknown. In a laboratory experiment, we studied whether common frog (Rana temporaria) tadpoles originating from two populations (separated latitudinally by 1600 km) adjust their growth and development according to the progress of the season by using photoperiodic cues, and whether these responses are temperature dependent. We hypothesised that if frogs use photoperiod as a cue, they should increase growth and development rates as a response to photoperiodic treatments mimicking progressing season. Although our predictions were not verified in either of the populations, photoperiod manipulations had effects on larval life history in both populations. When exposed to progressing season treatments and high temperature, tadpoles from the southern population ceased feeding, which led to delayed metamorphosis and increased mortality. In the northern population, age at metamorphosis was unaffected by the photoperiod treatments, but growth rate until metamorphosis and metamorphic size were slightly larger in the treatments with shorter (increasing or decreasing) day length. Irrespective of photoperiod treatments, growth and development rates, size at metamorphosis and food consumption were higher in the northern as compared to the southern population. These results indicate that in contrast to several insect species, the critical life history decisions in amphibian larvae may not be strongly influenced by photoperiodic cues, but different populations seem to differ in this respect. However, the strong temperature×photoperiod interactions in several traits in the southern population suggest that the role of photoperiodic cues may be affected by other environmental factors, although the ecological significance of these differences remains unclear.     

Proximate causes of adaptive growth rates: growth efficiency variation among latitudinal populations of Rana temporaria

In ectothermic organisms, declining season length and lower temperature towards higher latitudes often select for latitudinal variation in growth and development. However, the energetic mechanisms underlying this adaptive variation are largely unknown. We investigated growth, food intake and growth efficiency of Rana temporaria tadpoles from eight populations along a 1500 km latitudinal gradient across Sweden. To gain an insight into the mechanisms of adaptation at organ level, we also examined variation in tadpole gut length. The tadpoles were raised at two temperatures (16 and 20 degrees C) in a laboratory common garden experiment. We found increased growth rate towards higher latitudes, regardless of temperature treatment. This increase in growth was not because of a higher food intake rate, but populations from higher latitudes had higher growth efficiency, i.e. they were more efficient at converting ingested food into body mass. Low temperature reduced growth efficiency most strongly in southern populations. Relative gut length increased with latitude, and tadpoles at low temperature tended to have longer guts. However, variation in gut length was not the sole adaptive explanation for increased growth efficiency as latitude and body length still explained significant amounts of variation in growth efficiency. Hence, additional energetic adaptations are probably involved in growth efficiency variation along the latitudinal gradient.     

Are high-latitude individuals superior competitors? A test with <Emphasis Type="Italic">Rana temporaria</Emphasis> tadpoles

Species with a wide distribution over latitudinal gradients often exhibit increasing growth and development rates towards higher latitudes. Ecological theory predicts that these fast-growing genotypes are, in the absence of trade-offs with fast growth, better competitors than low-latitude conspecifics. While knowledge on key ecological traits along latitudinal clines is important for understanding how these clines are maintained, the relative competitive ability of high latitude individuals against low latitude conspecifics has not been tested. Growth and development rates of the common frog Rana temporaria increase along the latitudinal gradient across Scandinavia. Here we investigated larval competition over food resources within and between two R. temporaria populations originating from southern and northern Sweden in an outdoor common garden experiment. We used a factorial design, where southern and northern tadpoles were reared either as single populations or as mixes of the two populations at two densities and predator treatments (absence and non-lethal presence of Aeshna dragonfly larvae). Tadpoles from the high latitude population grew and developed faster and in the beginning of the experiment they hid less and were more active than tadpoles from the low latitude population. When raised together with high latitude tadpoles the southern tadpoles had a longer larval period, however, the response of high latitude tadpoles to the competition by low latitude tadpoles did not differ from their response to intra-population competition. This result was not significantly affected by density or predator treatments. Our results support the hypothesis that high latitude populations are better competitors than their low latitude conspecifics, and suggest that in R. temporaria fast growth and development trade off with other fitness components along the latitudinal gradient across Scandinavia.     

Plasticity in age and size at metamorphosis in Rana temporaria - comparison of high and low latitude populations

Effects of different combinations of stressors (viz. temperature, food level) on growth, developmental and survival rates of Rana temporaria tadpoles from two geographically widely (∼ 1500 km) separated populations were studied in a common garden experiment. In both populations, low temperature and low food level lead to towered growth rates and delayed metamorphosis, whereas high temperature and high food level had the opposite effect. Tadpoles from north metamorphosed earlier and exhibited higher growth rates than tadpoles from south, suggesting local adaptation to shorter growth period and cooler ambient temperature in north. Size at metamorphosis did not differ between the two populations, but when the differences in metamorphic age were accounted for, then the tadpoles from north were larger than those from south. These results suggest considerable adaptive genetic differentiation in growth rates, size and timing of metamorphosis between northern and southern R. temporaria populations. In both populations, high food levels tended to reduce tadpole survival rates and there was a negative correlation between growth and survival rates across different treatments in both populations. In general, tadpoles from north experienced high mortality rates in high food level - low temperature treatments, whereas southern tadpoles experienced high mortality in high food level-high temperature treatments. This suggest that there may be genetic differences among different populations as how they would be influenced by high nutrient loads, such as brought along for example by fertilization of forest or agricultural soils.     

Limits to compensatory responses to altered phenology in amphibian larvae

Changes in phenology are among the most pervasive effects of current climate change. Modifications in the timing of life-cycle events can affect the behavior, physiology and life-history of wildlife. However, organisms can develop compensatory strategies in order to reduce the costs of phenological alterations. Here, we examine the extent and limits of compensatory developmental responses in amphibian larvae exposed to variation in hatching timing. Using a common-garden experiment, we analyze how changes in temperature and food conditions alter compensatory responses to hatching delay, paying particular attention at how adverse environmental conditions can constrain these responses. We found that under benign conditions (warm temperature, unrestricted food) larvae fully compensate for the hatching delay, without cost in mass. However, under detrimental conditions (cold temperature and restricted food) these responses were prevented, and the combination of adverse conditions with long hatching delay completely disrupt compensatory responses. This study highlights the need of examining ecological responses to climate variation across a broad spectrum of environmental conditions in order to accurately predict the putative effect that climatic alterations can have on the life-histories and survival of wildlife.     

Behaviour and physiology shape the growth accelerations associated with predation risk, high temperatures and southern latitudes in Ischnura damselfly larvae

1.?To better predict effects of climate change and predation risk on prey animals and ecosystems, we need studies documenting not only latitudinal patterns in growth rate but also growth plasticity to temperature and predation risk and the underlying proximate mechanisms: behaviour (food intake) and digestive physiology (growth efficiency). The mechanistic underpinnings of predator-induced growth increases remain especially poorly understood. 2.?We reared larvae from replicated northern and southern populations of the damselfly Ischnura elegans in a common garden experiment manipulating temperature and predation risk and quantified growth rate, food intake and growth efficiency. 3.?The predator-induced and temperature-induced growth accelerations were the same at both latitudes, despite considerably faster growth rates in the southern populations. While the higher growth rates in the southern populations and the high rearing temperature were driven by both an increased food intake and a higher growth efficiency, the higher growth rates under predation risk were completely driven by a higher growth efficiency, despite a lowered food intake. 4.?The emerging pattern that higher growth rates associated with latitude, temperature and predation risk were all (partly or completely) mediated by a higher growth efficiency has two major implications. First, it indicates that energy allocation trade-offs and the associated physiological costs play a major role both in shaping large-scale geographic variation in growth rates and in shaping the extent and direction of growth rate plasticity. Secondly, it suggests that the efficiency of energy transfer in aquatic food chains, where damselfly larvae are important intermediate predators, will be higher in southern populations, at higher temperatures and under predation risk. This may eventually contribute to the lengthening of food chains under these conditions and highlights that the prey identity may determine the influence of predation risk on food chain length.     

Non-equivalence of growth arrest induced by predation risk or food limitation: context-dependent compensatory growth in anuran tadpoles

1. To gain insight into the evolution of compensatory growth, we studied the growth patterns of anuran (Rana temporaria) larvae following either a period of exogenous growth depression (food restriction) or a period of endogenous depression (exposure to predators). We also investigated the potential deferred costs that larval compensatory growth could impose on post-metamorphic individuals. 2. Food-deprived larvae exhibited full compensatory growth in response to reduced growth rates caused by food limitation, and the growth trajectories of low- and high-rations tadpoles converged before the onset of metamorphosis. 3. According to our predictions, individuals exposed to larval predators did not show growth compensation following predator removal despite undergoing a significant reduction in growth rate associated with low activity levels. 4. Jumping ability of individuals exposed to predators during only 20 days from the commencement of the larval phase was equivalent to that of non-exposed animals, and greater than the jumping capacity of those maintained with predators until the time of metamorphosis. This pattern was consistent with the pattern observed for variation in relative leg length. 5. These results support the suggestion that submaximum and compensatory growth could have evolved to minimize the overall growth/mortality costs in environments with high spatiotemporal variation in predation intensity.     

Plasticity in food assimilation,retention time and coprophagy allow herbivorous cavies (Microcavia australis) to cope with low food quality in the Monte desert

Energy balance depends on the efficiency with which organisms make use of their trophic resources, and has direct impact on their fitness. There are environmental variations that affect the availability as well as the quality of such resources; energy extraction also depends on the design of the digestive tract. It is expected that features associated with food utilization will be subjected to selective pressures and show some adjustment to the variability of the environment. Since energetic constraints challenge animals to display digestive compensatory mechanisms, the objective of this study is to determine the physiological and behavioral responses to spatial and seasonal heterogeneity in food quality. We investigated digestive strategies (digestive efficiency and coprophagy) in cavies inhabiting two different populations, and hence naturally experiencing different levels of diet quality. Cavies under experimentally different quality diets showed changes in dry matter digestibility and intake, digesta retention time and coprophagy. Our results partially support the expectations from theory and also reveal interpopulation differences in the ability to cope with changes in food quality, and may explain the capability of Microcavia australis to colonize extreme habitats.     

A profile of carbohydrate metabolites in the fasting northern elephant seal

Northern elephant seals endure prolonged periods of food deprivation at multiple life-history stages and simultaneous with energetically costly activities—including reproduction and development. Most mammals decrease their energy expenditure while fasting, with simultaneous reductions in gluconeogenesis and circulating glucose concentration. Paradoxically, elephant seals maintain high rates of both energy expenditure and gluconeogenesis, and high blood glucose concentrations throughout fasting. We therefore characterized the suite of changes that occur in carbohydrate metabolites during fasting in northern elephant seals. Using a broad-based metabolomics platform we investigated fasting during two states—lactation in adult females and the post-weaning developmental period in pups. A total of 227 metabolites were detected in seal plasma; 31 associated with carbohydrate metabolism were analyzed in the present study. Several compounds showed similar responses during lactation and the post-weaning fast (e.g. glycerol and mesaconate) whereas other compounds displayed quite different abundances between groups (e.g. citrate and pyruvate). This work found that, while the changes that occur with fasting were frequently similar in lactating females and developing pups, the relative abundance of compounds often varied markedly. These differences suggest that the metabolic strategies used to endure prolonged fasts are influenced by life-history or nutrient constraints.     

Compensatory growth for free? A field experiment on brown trout,Salmo trutta

Laboratory studies suggest that animals may be capable of compensatory growth after periods of food shortage. There is, however, a lack of field experiments investigating the incidence and consequences of compensatory growth in the wild, and the relevance of compensatory responses in natural populations has recently been questioned. Here we addressed the hypotheses that (1) food restriction during critical growth periods can induce compensatory growth, and (2) that compensatory growth is associated with delayed costs in natural populations. These hypotheses were addressed by (1) manipulating the food intake of brown trout in spring, (2) measuring growth rate responses over the first month following release, and (3) measuring growth and mortality (i.e. recapture rate) over the subsequent fall and winter. We found that brown trout restored lost body weight and condition within a month, providing the first experimental demonstration of compensatory growth in the wild. However, no delayed costs of the compensatory response could be detected within the timespan of the experiment. We suggest that wild brown trout have an adapted "buffer capacity" to withstand fluctuations in food supply, allowing restoration of lost lipid reserves when feeding conditions improve. However, when prolonged food deprivation affect structural components, compensation may not be possible without compromising long-term performance.     

Life history traits associated with body size covary along a latitudinal gradient in a generalist grasshopper

Animal body size often varies systematically along latitudinal gradients, where individuals are either larger or smaller with varying season length. This study examines ecotypic responses by the generalist grasshopper Melanoplus femurrubrum (Orthoptera: Acrididae) in body size and covarying, physiologically based life history traits along a latitudinal gradient with respect to seasonality and energetics. The latitudinal compensation hypothesis predicts that smaller body size occurs in colder sites when populations must compensate for time constraints due to short seasons. Shorter season length requires faster developmental and growth rates to complete life cycles in one season. Using a common garden experimental design under laboratory conditions, we examined how grasshopper body size, consumption, developmental time, growth rate and metabolism varied among populations collected along an extended latitudinal gradient. When reared at the same temperature in the lab, individuals from northern populations were smaller, developed more rapidly, and showed higher growth rates, as expected for adaptations to shorter and generally cooler growing seasons. Temperature-dependent, whole organism metabolic rate scaled positively with body size and was lower at northern sites, but mass-specific standard metabolic rate did not differ among sites. Total food consumption varied positively with body size, but northern populations exhibited a higher mass-specific consumption rate. Overall, compensatory life history responses corresponded with key predictions of the latitudinal compensation hypothesis in response to season length.     

Bridging Developmental Boundaries: Lifelong Dietary Patterns Modulate Life Histories in a Parthenogenetic Insect

Determining the effects of lifelong intake patterns on performance is challenging for many species, primarily because of methodological constraints. Here, we used a parthenogenetic insect (Carausius morosus) to determine the effects of limited and unlimited food availability across multiple life-history stages. Using a parthenogen allowed us to quantify intake by juvenile and adult females and to evaluate the morphological, physiological, and life-history responses to intake, all without the confounding influences of pair-housing, mating, and male behavior. In our study, growth rate prior to reproductive maturity was positively correlated with both adult and reproductive lifespans but negatively correlated with total lifespan. Food limitation had opposing effects on lifespan depending on when it was imposed, as it protracted development in juveniles but hastened death in adults. Food limitation also constrained reproduction regardless of when food was limited, although decreased fecundity was especially pronounced in individuals that were food-limited as late juveniles and adults. Additional carry-over effects of juvenile food limitation included smaller adult size and decreased body condition at the adult molt, but these effects were largely mitigated in insects that were switched to ad libitum feeding as late juveniles. Our data provide little support for the existence of a trade-off between longevity and fecundity, perhaps because these functions were fueled by different nutrient pools. However, insects that experienced a switch to the limited diet at reproductive maturity seem to have fueled egg production by drawing down body stores, thus providing some evidence for a life-history trade-off. Our results provide important insights into the effects of food limitation and indicate that performance is modulated by intake both within and across life-history stages.     

Microgeographic variation in temperature-induced plasticity in an isolated amphibian metapopulation

Variation in local environments may lead to variation in the selection pressures and differentiation among local populations even at microgeographic scale. We investigated variation in temperature-induced plasticity in larval life-history traits among populations of an isolated pool frog (Rana lessonae) metapopulation in Central Sweden. Successful breeding of this northern fringe metapopulation is highly dependent on early summer temperature, however, the metapopulation shows very little variation in molecular genetic markers suggesting limited potential for local differentiation. We exposed larvae from three closely-located populations to two temperatures (20 and 25°C) in laboratory to investigate their growth and development responses to temperature variation. In general, larvae exposed to warmer temperature experienced higher survival and metamorphosed faster, but at a smaller size than those at low temperature. We found differences among the populations in both trait mean values and in the plastic responses. Among-family variation within populations was found in growth rate and time to metamorphosis, as well as in plasticity suggesting that these traits have a capacity to evolve. Our results indicate ample phenotypic variation within and among these closely-located populations despite the low molecular genetic variation. The differences in pond temperature characteristics detected in the study in the three localities may suggest that differential selection is acting in the populations. The strong differentiation found in the larval traits implies that understanding the factors that influence the potential of the populations to adapt to environmental changes may be essential for successful conservation strategies.     

Life-history variation in relation to time constraints in a damselfly

Although variation within populations in plasticity to time constraints is expected with regard to hatching date, empirical studies are largely lacking. We studied life-history responses to time constraints manipulated by photoperiod and associated with hatching date in larvae of the damselfly Lestes viridis for two populations with a different hydroperiod. In a common garden experiment, early- and late-hatched larvae from both populations were reared at two photoperiods mimicking the start and the end of the egg-hatching season. In a reciprocal transplant experiment, early- and late-hatched larvae from both populations were reared in both ponds. In all these experiments, larvae were reared from egg hatching until adult emergence. Within both populations, larvae reared at the photoperiod indicating a late time point in the growing season, reduced development time to compensate for their perceived shorter development period. Growth rate, however, did not respond to photoperiod, resulting in a lower mass at emergence. As expected, both in the laboratory and in the field, larvae from eggs that hatched later in the season generally had a shorter development time and a faster growth rate, resulting in a higher mass at emergence compared to early-hatched larvae. This may explain the intriguing seasonal increase in mass at emergence in this species, and affect the predictions of optimality models. None of these life-history responses differed between the two populations, despite clear differences in time constraints linked to hydroperiod, suggesting the robustness of the observed patterns. Given the ubiquity of asynchronous hatching in nature, and the adaptive value of the observed differences between early- and late-hatched larvae, we expect the effects of hatching date on life-history plasticity to be widespread.     

Phenotypic clines, plasticity, and morphological trade-offs in an intertidal snail

Understanding the genetic and environmental bases of phenotypic variation and how they covary on local and broad geographic scales is an important goal of evolutionary ecology. Such information can shed light on how organisms adapt to different and changing environments and how life-history trade-offs arise. Surveys of phenotypic variation in 25 Littorina obtusata populations across an approximately 400-km latitudinal gradient in the Gulf of Maine revealed pronounced clines. The shells of snails from northern habitats weighed less and were thinner and weaker in compression than those of conspecifics from southern habitats. In contrast, body size (as measured by soft tissue mass) followed an opposite pattern; northern snails weighed more than southern snails. A reciprocal transplant between a northern and southern habitat revealed substantial plasticity in shell form and body mass and their respective measures of growth. Southern snails transplanted to the northern habitat produced lighter, thinner shells and more body mass than controls raised in their native habitat. In contrast, northern snails transplanted to the southern site produced heavier, thicker shells and less body mass than controls raised in their native habitat. Patterns of final phenotypic variation for all traits were consistent with cogradient variation (i.e., a positive covariance between genetic and environmental influences). However, growth in shell traits followed a countergradient pattern (i.e., a negative covariance between genetic and environmental influences). Interestingly, body growth followed a cogradient pattern, which may reflect constraints imposed by cogradient variation in final shell size and thickness. This result suggests the existence of potential life-history trade-offs associated with increased shell production. Differences in L. obtusata shell form, body mass, and their respective measures of growth are likely induced by geographic differences in both water temperature and the abundance of an invading crab predator (Carcinus maenas). Water temperatures averaged 6.8 degrees C warmer during the transplant experiment and C. maenas abundance is greater in the southern Gulf of Maine. Because both increased water temperature and crab effluent affect shell form in the same way, future experiments are needed to determine the relative importance of each. Nevertheless, it is clear that phenotypic plasticity has an important role in producing geographic variation in L. obtusata shell form. Moreover, the evolution of phenotypic plasticity in L. obtusata and other marine gastropods may be driven by architectural constraints imposed by shell form on body mass and growth.     

Embryonic and larval defensive responses of agile frog (Rana dalmatina) to alien crayfish

Red swamp crayfish Procambarus clarkii, a widespread invasive alien crayfish, represents a serious threat for several freshwater species, including amphibians, which are declining at a global scale. As a shared coevolutionary history is the main factor determining the emergence of antipredator responses, Anuran tadpoles may not be able to cope effectively with this introduced predator. We performed two experiments to assess agile frog's (Rana dalmatina) defensive responses to both P. clarkii and native dragonfly larvae (Anax imperator). First, we conditioned embryos (collected from two ponds 30 km away from each other) with predators’ chemical cues to explore possible variation in hatching time caused by predation risk. In the second experiment, to evaluate how predators’ diet affects tadpole behavior, we conditioned tadpoles for a 5‐week period with cues from tadpole‐fed and gammarid‐fed predators and recorded behavioral and morphological responses. Embryos did not alter hatching time in the presence of any predator cue, while tadpoles from both populations strongly reduced activity and visibility when raised in the presence of tadpole‐fed dragonfly larvae. Morphological changes were less straightforward and were induced only in one population, for which broader tails and a slight increase in body size of tadpoles exposed to tadpole‐fed predators were observed. The lack of defensive responses in crayfish‐exposed tadpoles suggests that the spreading of this invasive species in agricultural lowlands of northern Italy may represent a further threat to their conservation.     

Seasonality determines patterns of growth and age structure over a geographic gradient in an ectothermic vertebrate

Environmental variation connected with seasonality is likely to affect the evolution of life-history strategies in ectotherms, but there is no consensus as to how important life-history traits like body size are influenced by environmental variation along seasonal gradients. We compared adult body size, skeletal growth, mean age, age at first reproduction and longevity among 11 common frog (Rana temporaria) populations sampled along a 1,600-km-long latitudinal gradient across Scandinavia. Mean age, age at first reproduction and longevity increased linearly with decreasing growth season length. Lifetime activity (i.e. the estimated number of active days during life-time) was highest at mid-latitudes and females had on average more active days throughout their lives than males. Variation in body size was due to differences in lifetime activity among populations??individuals (especially females) were largest where they had the longest cumulative activity period??as well as to differences between populations in skeletal growth rate as determined by skeletochronological analyses. Especially, males grew faster at intermediate latitudes. While life-history trait variation was strongly associated with latitude, the direction and shape of these relationships were sex- and trait-specific. These context-dependent relationships may be the result of life-history trade-offs enforced by differences in future reproductive opportunities and time constraints among the populations. Thus, seasonality appears to be an important environmental factor shaping life-history trait variation in common frogs.