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Current international legislation regarding agrochemicals requires thorough toxicological testing mainly of the active ingredients. In a 96-h acute toxicity test we exposed Rana dalmatina and Bufo bufo tadpoles to either one of three concentrations of glyphosate, three concentrations of the surfactant (POEA), three concentrations of the two components together, or to non-contaminated water (control), and subsequently assessed mortality and body mass. To investigate whether simultaneous exposure to another stress factor influences effects of the contaminants, we performed tests both in the presence or absence of predator chemical cues. We found that the surfactant had significant harmful effects on tadpoles; survival was lowered by the highest concentration of the surfactant in case of R. dalmatina, while in B. bufo tadpoles it reduced survival already at medium concentrations. Body mass was significantly influenced by medium and high surfactant concentrations in both species. The presence of glyphosate did not have a significant effect by itself, but it slightly increased mortality in tadpoles exposed to medium concentrations of the surfactant in both species. The presence of chemical cues did not have an effect on the examined variables. Our study confirms that the toxicity of glyphosate-based herbicides is mainly due to the examined surfactant. Nonetheless, we found that glyphosate can enhance the harmful effect of the surfactant. These results stress that during the authorization process of new pesticide formulations, not only the active ingredients would need to be examined but the excipients should also be taken into account in an obligatory and systematic manner.

1. Behavioural consistency within and across behaviours (animal personality and behavioural syndrome, respectively) has been vigorously studied in the last decade, leading to the emergence of \"animal personality\" research. It has been proposed recently that not only mean behaviour (behavioural type), but the environmentally induced behavioural change (behavioural plasticity) might also differ between individuals within populations. 2. While case studies presenting between-individual variation in behavioural plasticity have started to accumulate, the mechanisms behind its emergence are virtually unknown. We have recently demonstrated that ecologically relevant environmental stimuli during ontogeny are necessary for the development of animal personality and behavioural syndromes. However, it is unknown whether between-individual variation in behavioural plasticity is hard-wired or induced. 3. Here, we tested whether experience with predation during development affected predator-induced behavioural plasticity in Rana dalmatina tadpoles. We ran a common garden experiment with two ontogenetic predation treatments: tadpoles developed from hatching in either the presence or absence of olfactory predator stimuli. Then, we assayed all tadpoles repeatedly for activity and risk-taking both in the absence and presence of olfactory predator stimuli. 4. We found that (a) between-individual variation in predator-induced behavioural plasticity was present only in the group that developed in the presence of olfactory stimuli from predators and (b) previous experience with predatory stimuli resulted in lower plastic response at the group level. The latter pattern resulted from increased between-individual variation and not from universally lower individual responses. We also found that experience with predation during development increased the predictability (i.e. decreased the within-individual variation unrelated to environmental change) of activity, but not risk-taking. In line with this, tadpoles developing under perceived predatory risk expressed their activity with higher repeatability. 5. We suggest that ecologically relevant environmental stimuli are not only fundamental for the development of animal personality and behavioural syndromes, but also for individual variation in behavioural plasticity. Thus, experience is of central importance for the emergence of individual behavioural variation at many levels.

Predation is a major source of selection and prey are known to modify their behavior depending on their past experiences and the current perceived risk. Within a species, variation in experience and in the response to perceived risk combine to explain variation in personality and individual plasticity. Between species, variation in personality and plasticity might also be the evolutionary consequence of different selective regimes. In this study, we describe the anti-predator behavior of two closely related brown frogs, Rana dalmatina and Rana latastei, and compare their structures of personality variation. We raised tadpoles in a common garden experiment with either fish, dragonfly larvae, or no predators. Tadpoles were then repeatedly tested in the presence of the three acute stimuli and their behavioral variation was described in terms of quantity and quality of movements and of path sinuosity. In these tests, tadpoles of both species and ontogenetic treatments responded flexibly to predators by moving less, faster, and with more tortuous movements, and tadpoles raised with predators tended to move even faster. Independent of the acute treatment, R. dalmatina moved more and faster than R. latastei and the differences were larger without than with predators, demonstrating its higher plasticity. At the individual level, the two species showed qualitatively similar but quantitatively different structures of personality variation. R. dalmatina, more active, faster, and more plastic than R. latastei, showed also higher repeatability and a larger behavioral variation both among and within individuals.Significance statementPredators are a major source of selection and preys have evolved the ability to flexibly respond to them. These responses often vary among species, because of their different evolutionary histories, and among individuals, because of their different experiences. We analyzed both these sources of behavioral variation in two closely related brown frogs, Rana dalmatina and R. latastei. We raised tadpoles either with or without predators and tested them in open field trials both with and without predators. The effects of the raising environment were similar in the two species, whereas the effects of the testing arena differed. Both species decreased activity and increased speed and sinuosity with predators, but R. dalmatina moved always more and faster than R. latastei and it showed higher plasticity, larger variation among and within individuals, and relatively higher values in repeatability.

We studied how individuals modify their behavior in response to inter- and intraspecific competitors and how these changes affected the pattern of variation between populations and species. As study models, we used tadpoles of two brown frogs, Rana latastei and R. dalmatina. Since R. latastei is always sympatric to R. dalmatina, whereas R. dalmatina is sympatric to R. latastei only in the periphery of its range, we predicted a stronger response to heterospecifics in R. latastei than in R. dalmatina and, within each species, in syntopic than in allotopic populations. To test these predictions, we raised tadpoles, from either syntopic or allotopic populations, in either syntopy or allotopy and repeatedly tested them in open field trials in the presence of a caged conspecific, a caged heterospecific, or an empty cage. As predicted, we found that, on average, R. latastei tadpoles modified their behavior across treatments more than R. dalmatina tadpoles and individuals from the syntopic population changed more than their conspecifics from the allotopic population. In both species, the pattern of variation at the individual level mirrored that at the population and species levels providing no evidence for an individual-by-environment interaction (I×E). Besides these differences, however, individuals of the two species also showed unpredicted and context-independent behavioral differences, suggesting that there might be more to interspecific behavioral variation than the effect of selection by heterospecific competitors.Significance statementDoes the distribution range of a species influence the evolution of plastic behaviors to heterospecific competitors? And how do differences in plasticity affect animal personality? To answer these questions, we raised tadpoles of two brown frog species, Rana dalmatina and R. latastei, and studied how the amount and the type of their swimming varied with the presence of the other species. R. latastei, whose small distribution range fully overlaps with that of R. dalmatina, plastically responds to it, whereas R. dalmatina, which is sympatric to R. latastei only in the periphery of its broader range, does not. These interspecific differences mirrored those among individuals: tadpoles of both species show repeatable behaviors, but only those of R. latastei plastically changed their behavior with the presence of the other species; however, neither R. latastei nor R. dalmatina show among-individual variation in plasticity.

Aim Understanding which factors determine the variation in population size across space and time is crucial to plan sound conservation interventions. Amphibians are often characterized by large demographic changes, therefore a better understanding of factors driving these changes could help mitigate their global crisis. We investigated drivers of abundance dynamics of two similar frog species to understand why they met different fates in the same study area. Location Northern Italy. Methods In seven different years between 2004 and 2020, we performed repeated counts of egg masses of two similar frog species, the agile frog (Rana dalmatina) and the Italian agile frog (Rana latastei), in 31 wetlands, and used Bayesian models to estimate the relationships between frog abundance and candidate drivers acting at the scale of (i) site: wetland surface, shading percentage and the presence of the invasive red swamp crayfish (Procambarus clarkii); (ii) landscape: forest cover around wetlands; (iii) climate: yearly precipitation and (iv) spatially structured population (SSP): clutch incidence of the focal species and crayfish incidence in the surrounding landscape. Results The two species showed sharp differences in population size and trends: R. dalmatina was abundant and showed a stable trend throughout the entire study period; R. latastei showed low abundance in the first years and then almost disappeared. The abundance of R. dalmatina was positively related to forest cover, shading, wetland area and precipitation, while negatively related to the occurrence of invasive crayfish at both local and SSP level. R. latastei abundance increased with wetland area and precipitation, while models were not able to detect relationships with other factors. Main conclusions The high sensitivity to drought and demographic stochasticity could have contributed to the quasi‐extinction of R. latastei in the study area, highlighting that similar species can meet different fates even under the same environmental conditions.

Predation and competition are two major factors that drive natural selection. When they vary unpredictably, selection has promoted the evolution of plastic responses in behavioral, morphological and life-history traits. These responses are not independent of each other and often represent a trade-off between conflicting interests. We conducted a common-garden experiment to study the effects of predation and competition on the morphology and life history of R. dalmatina and R. latastei tadpoles. The experiment used a randomized-block design, where tadpoles were raised either with or without siblings and either with or without predators. Regardless of the treatments, R. dalmatina grew faster, developed proportionally larger tails, proportionally smaller bodies, and completed metamorphosis earlier than R. latastei. Both species developed relatively larger bodies with competitors, and relatively larger tails with predators. While the relative increase in body size with competitors was similar in the two species, the increase in tail size with predators was greater in R. dalmatina, suggesting that this species invested more in defense than R latastei. Competitors delayed metamorphosis in both species and in R. latastei they negatively affected froglet body size and shape. Conversely, predators delayed the metamorphosis only in R. dalmatina. This delay was the long-term cost paid by R. dalmatina for the short-term benefits of developing more effective behavioral and morphological defensive traits.Significance statementTadpoles develop plastic morphological and life-history traits in responses to predators and competitors. This plasticity is costly in terms of both performance and function. Our study shows that two closely related brown frogs have evolved different trade-offs in their plastic responses to predators and competitors. Rana dalmatina, which grew and developed faster than R. latastei, was more sensitive to predators and developed more effective morphological defenses. Both species delayed metamorphosis at high density, but R. dalmatina did it more than R. latastei. Predators did not affect the duration of larval development in R. latastei, whereas in R. dalmatina, they further delayed metamorphosis. This delay was the long-term cost of the R. dalmatina’s larger investment in short-term defensive traits.

The focus of evolutionary behavioural ecologists has recently turned towards understanding the causes and consequences of behavioural consistency, manifesting either as animal personality (consistency in a single behaviour) or behavioural syndrome (consistency across more behaviours). Behavioural type (mean individual behaviour) has been linked to life-history strategies, leading to the emergence of the integrated pace-of-life syndrome (POLS) theory. Using Rana dalmatina tadpoles as models, we tested if behavioural consistency and POLS could be detected during the early ontogenesis of this amphibian. We targeted two ontogenetic stages and measured activity, exploration and risk-taking in a common garden experiment, assessing both individual behavioural type and intraindividual behavioural variation. We observed that activity was consistent in all tadpoles, exploration only became consistent with advancing age and risk-taking only became consistent in tadpoles that had been tested, and thus disturbed, earlier. Only previously tested tadpoles showed trends indicative of behavioural syndromes. We found an activity—age at metamorphosis POLS in the previously untested tadpoles irrespective of age. Relative growth rate correlated positively with the intra-individual variation of activity of the previously untested older tadpoles. In previously tested older tadpoles, intra-individual variation of exploration correlated negatively and intra-individual variation of risk-taking correlated positively with relative growth rate. We provide evidence for behavioural consistency and POLS in predator- and conspecific-naive tadpoles. Intraindividual behavioural variation was also correlated to life history, suggesting its relevance for the POLS theory. The strong effect of moderate disturbance related to standard behavioural testing on later behaviour draws attention to the pitfalls embedded in repeated testing.

Invasive alien predators (IAP) are spreading on a global scale—often with devastating ecological effects. One reason for their success may be that prey species fail to recognize them due to a lack of co-evolutionary history. We performed a comprehensive test of this \"prey naiveté\" hypothesis using a novel approach: we tested whether predator-naive tadpoles of the agile frog (Rana dalmatina) display antipredator behavior upon encountering chemical cues produced by native, invasive (established or recent) or allopatric fishes (four perciforms, four siluriforms, and two cypriniforms). We studied the influence of population origin on predator-detection ability by presenting chemical cues to predator-naive tadpoles that originated from fishless hill-ponds or fish-infested floodplain populations. Before trials, we fed fishes with tadpoles or an alternative food to test whether direct chemical cues from the predator's diet influences the tadpoles' recognition of potential predators. Tadpoles reduced their activity upon exposure to cues from native and long-established invasive perciforms, but not in response to recent invaders, allopatric predators, or to any siluriforms. Also, predators that were previously fed with tadpoles did not universally induce behavioral defensedefenses upon first encounter. Finally, tadpoles originating from isolated hill-ponds exhibited higher baseline activity and responded in weaker fashion than their conspecifics from floodplain populations, which co-exist with predatory fishes. Our results indicate that tadpoles may be vulnerable to invading predatory fishes due to their inability to recognize them as dangerous, though their ability to recognize invasive IAP may evolve rapidly, in fewer than 30 generations.

For 29 consecutive years, the populations of three species of brown frogs, Rana arvalis , R. dalmatina and R. temporaria , were monitored in an open area in South Zealand, Denmark, with no direct influence of agriculture. Population sizes were recorded by counting egg clumps and showed large variations from year to year. The total population of R. arvalis differed by a factor of 100 between the years with the lowest and the highest numbers. A total of 19 initially unoccupied suitable waterbodies could potentially be colonised by the frogs. Rana dalmatina colonised all of the ponds, mostly in the very first year of existence. Rana arvalis colonised 17 ponds after an average of 10.5 years. Rana temporaria colonised eight ponds after an average of 13.4 years. Colonisation by R. dalmatina was independent of changes in total population size, whereas colonisation by R. arvalis predominantly occurred in years with considerable population increases. The results are discussed in relation to the movement patterns and philopatry of juvenile frogs of the three species. Juvenile R. dalmatina disperse far from the breeding site, but most individuals return to their natal site. This allows the species to be an efficient coloniser of new waterbodies and, at the same time, to have stable occurrence at the original site. The two other species show a more erratic type of dispersal and especially R. temporaria often shifts breeding site from year to year.

Understanding population dynamics is critical for the management of animal populations. Comparatively little is known about the relative importance of endogenous (i.e. density-dependent) and exogenous (i.e. density-independent) factors on the population dynamics of amphibians with complex life cycles. We examined the potential effects of density-dependent and -independent (i.e. climatic) factors on population dynamics by analyzing a 15-yr time series data of the agile frog Rana dalmatina population from Târnava Mare Valley, Romania. We used two statistical models: 1) the partial rate correlation function to identify the feedback structure and the potential time lags in the time series data and 2) a Gompertz state-space model to simultaneously investigate direct and delayed density dependence as well as climatic effects on population growth rate. We found evidence for direct negative density dependence, whereas delayed density dependence and climate did not show a strong influence on population growth rate. Here we demonstrated that direct density dependence rather than delayed density dependence or climate determined the dynamics of our study population. Our results confirm the findings of many experimental studies and suggest that density dependence may buffer amphibian populations against environmental stress. Consequently, it may not be easy to scale up from individual-level effects to population-level effects.