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Hidden colors are a widespread phenomenon in the animal kingdom, particularly in anurans. In some cases, hidden colors are suddenly exposed during defensive displays to startle predators, others seemingly remain hidden—particularly from researchers. Amazonian species of Neotropical harlequin toads (genus Atelopus) show striking and consistent ventral sexual dichromatism where females show and males generally lack melanization. Inspired by these observations we undertook a deeper inquiry across this species-rich genus. We collected data on ventral sexual dichromatism in Atelopus species and scored expression of sex-specific ventral melanization (i.e. black, brown and/or grey coloration). Ventral sexual dichromatism was present throughout the entire range of the genus and in almost all phylogenetic groups. However, there was a clear geographic signal with this trait being most common and widespread in Amazonian Atelopus species. Ventral melanization was correlated with temperature and elevation. Focusing on the Amazonian species, we present hypotheses on potential functions of sexually dimorphic ventral patterns and sex-specific ventral melanization as a baseline to further investigate the dynamics of sexual and natural selection as potential drivers of these traits. Selective pressures on less exposed body parts, such as ventral sides, likely differ considerably from those on dorsal appearance. Given the amount of research on amphibian coloration, it is remarkable how little we know about the evolution, function and underlying mechanisms of ventral appearance. We hope our work will spark more interest in the flip side of amphibians, thereby broadening our understanding of animal coloration. [ABSTRACT FROM AUTHOR]

Species distribution models (SDM) are an important tool to predict the invasion risk of alien species and emerging infectious diseases. However, building reliable models in early stages of invasions is a challenging task. Katz and Zellmer (Biol Invasions 20:2107–2119, 2018) addressed this problem and presented a framework for model selection for data-poor newly invasive species. Based on data of the recently discovered and invasive amphibian chytrid fungus Batrachochytrium salamandrivorans (Bsal), they built SDM and gave concluding implications for model selection in general and for Bsal in its invasive range in particular. In our opinion the authors' SDM and their final processing show severe flaws and lead to a biologically implausible (although statistically best) model, which they used to promote their framework. We here intend to remind readers of the importance of biological relevance and plausibility of SDM, especially its input data, by highlighting some deficiencies and their implications for the biological relevance of the predictions. We further emphasize considerations and recommendations to improve the biological relevance and to make model evaluation more comprehensible. Though biological relevance is a basic SDM rationale, it is of particular importance when limited data impede a 'proper' model evaluation. If a prediction is evaluated as good or best by some statistical measure, but is obviously implausible in a biological sense, we should not ignore the biology of our model species. Especially in the face of an imminent threat such as the spreading Bsal, when immediate conservation or management actions are needed, wrong implications based on biologically implausible models may have severe counterproductive consequences. [ABSTRACT FROM AUTHOR]

Considerable scientific, politic and economic attention has been directed to biological invasions. Multiple pathways serve to introduce species to new environments and the release or escape of pets are among the most important sources for species invasions. Risk assessments help to identify species that are likely to become invasive and to set up preventive measures. Weighing the relative importance of ecological and human factors driving the establishment success of abandoned pets, we here present a new methodological guideline to help prioritising management activities for frequently traded pet reptiles. Climate match scores between the different distribution ranges as well as traits and niche axes shared by native and non-native species were assessed. Moreover, we tested for discrepancies in niche breadth between native and non-native ranges and estimated the ability of species to coexist with humans. Potentially moderate to high establishment success in most species was linked to appropriate climate match scores, broader niches with restrained human impacts and high similarities in reproductive traits with the native herpetofauna. Providing baseline information on the invasion potential of pet reptiles, this assessment calls for trade regulations and, to an even greater degree, for large scale education campaigns to prevent the establishment of non-native populations. [ABSTRACT FROM AUTHOR]

For regulatory and scientific purposes, there is a need to understand the sensitivity of a wider variety of wild species of amphibians and the sensitivities within their life stages to chemicals of widespread use such as herbicides. We investigated the acute toxic effects of the herbicide formulation Focus Ultra [with the active ingredient (a.i.) cycloxydim plus solvent naphtha and sodium dioctylsulphosuccinate as added substances] on embryos and early stage larvae of the Moroccan painted frog ( Discoglossus scovazzi). Different clinical signs (twitching, convulsion, and narcosis) occurred at 40 and 80 mg/L in embryos (4 and 8 mg a.i./L) and narcotic effects (total immobilization or irregular escape responses) at 10, 15, and 20 mg/L in larvae (1, 1.5, and 2 mg a.i./L). Growth inhibition (total length), starting at 20 mg/L in embryos and 2.5 mg/L in larvae (2 and 0.25 mg a.i./L, respectively) was understood as sign of toxicity (retardation) and not as sign of teratogenicity. However, the connection to teratogenesis remained unclear though total length reduction occurred at concentrations <20 % of the 96-h LC value and at a minimum concentration that inhibits growth of only 17 % of the 96-h LC value. Starting at 20 mg/L, mortality in embryos significantly increased and at 15 mg/L in early larvae (2 and 1.5 mg a.i./L, respectively). Mortality of larvae was enhanced during the first 24 h of exposure to 15 and 20 mg/L (1.5 and 2 mg a.i./L). Morphology of the embryos remained unobtrusive. In contrary, axial malformations significantly increased in the early larvae starting at 10 mg/L (1 mg a.i./L), a concentration free of lethal effects. In all considered end points, larvae were significantly more sensitive than embryos, probably because of developmental and physiological properties or different exposure and bioavailability of the compound. Focus Ultra induced comparable lethal and immobilization effects in D. scovazzi as it does to standard test organisms in pesticide approval. However, to validate the apparent safety in the field, which is based on calculated surface water concentrations of the a.i., more data on real contamination levels is necessary (e.g., peak concentrations, concentrations of added substances). Furthermore, sufficient buffer strips between the farmland and amphibian ponds must be considered, and the effects of the substance on terrestrial life stages have not been assessed yet. [ABSTRACT FROM AUTHOR]

Most genetically engineered herbicide-tolerant crops are still awaiting approval in Europe. There is, however, a recent trend for the cultivation of cycloxydim-tolerant maize hybrids for use in maize production. We studied the acute toxic effects of the complementary herbicide Focus Ultra and its active ingredient cycloxydim on embryos and early-stage larvae of the African clawed frog ( Xenopus laevis). The results indicate that the herbicide formulation is significantly more toxic than the active ingredient alone. Therefore, it is suggested that the added substances either solely or in a synergistic action with the active ingredient are responsible for adverse effects. The formulation was found to be moderately toxic to embryos but highly toxic to early larvae. Based on calculated teratogenic indices, both cycloxydim and Focus Ultra seem to be non-teratogenic and also the minimum Focus Ultra concentration to inhibit growth in embryos and larvae was close to the LC50 values. The data suggest that tests with the rainbow trout are not in all cases appropriate to assess the risk in aquatically developing anurans. This is demonstrated by 96-h LC50 values, which are for rainbow trout more than 50- to 20-fold higher than for early X. laevis larvae. However, based on worst-case predicted environmental concentrations for surface waters, there is apparently a large safety margin in field use of Focus Ultra if buffer strips between the farm land and the amphibian habitats are regarded. [ABSTRACT FROM AUTHOR]

The protection of offspring against predators and competitors is especially important in organisms using spatially separated breeding resources, impeding the offspring's chances to escape. One example of such isolated reproductive resources are phytotelmata (small water bodies in plant axils), exploited by the Neotropical poison frog Ranitomeya variabilis (Dendrobatidae) for both clutch and tadpole deposition. Because poison frog tadpoles are often cannibalistic, parents tend to avoid deposition with conspecifics. Previous studies have shown that this avoidance is based on chemical cues produced by conspecific tadpoles. Further, cues produced by phylogenetically less-related tadpoles (Bufonidae) were avoided for clutch but not tadpole depositions. We analyzed how the different responses to tadpole cues are triggered. We tested the reactions of parental R. variabilis to tadpole cues of species differing in two aspects: whether or not they are dendrobatids, and whether or not they reproduce in phytotelmata. We found that for clutch deposition, tadpole cues were always avoided, i.e., all tadpoles were treated by the frogs as if they pose a danger to the eggs. However, responses varied for tadpole depositions: while dendrobatid larvae living in phytotelmata were avoided, those breeding in streams were not. Non-poison frog tadpoles were ignored when associated with habitat other than phytotelmata, but they were preferred when living in phytotelmata. This suggests that both phylogeny and tadpole habitat are important triggers for the decisions made by R. variabilis. Only tadpoles using the same breeding resources are considered as relevant for the frog's own larvae (i.e., as a potential danger or food resource), while further decisions are related to evolutionary relationship. [ABSTRACT FROM AUTHOR]

Pesticide residues in breeding ponds can cause avoidance by at least some amphibian species. So far, outdoor experiments have been performed only with artificial pools in areas where the focus species usually occur and new colonization has been observed. Results of this kind of study are potentially influenced by natural disturbances and therefore are of limited comparability. We used an easily manufactured and standardizable arena approach, in which animals in reproductive condition for some hours had a choice among pools with different concentrations of a contaminant. Because there has been much debate on the potential environmental impacts of glyphosate-based herbicides, we investigated the impact of glyphosate isopropylamine salt (GLY-IS), Roundup LB PLUS (RU-LB-PLUS), and glyphosate's main metabolite aminomethylphosphonic acid (AMPA) on individual residence time in water. The following European amphibian species were tested: Common frog ( Rana temporaria), Palmate newt ( Lissotriton helveticus), and Alpine newt ( Ichthyosaura alpestris). The residence time in water was not significantly affected by concentrations below or slightly above the European Environmental Quality Standards for AMPA or the German 'worst-case' expected environmental concentrations for GLY-IS and RU-LB-PLUS. Occasionally, microclimatic cofactors (nightly minimum ground temperature, water temperature) apparently influenced the residence time. The major drawback of such quick behavior studies is that results can only be transferred to perception and avoidance of contaminated water but not easily to site selection by amphibians. For example, testing oviposition site selection requires more natural water bodies and more time. Hence, to develop a standard procedure in risk assessment, an intermediate design between an arena approach, as presented here, and previously performed field studies should be tested. [ABSTRACT FROM AUTHOR]

The quality of breeding sites is of great importance for the reproductive success and accordingly the fitness of many animal species. Hence, individuals should decide carefully where to rear their offspring. Often parents have to account for multiple characteristics of habitat quality at once, which in turn might change over time. Specimens confronted with such variability may evolve the ability to display context-dependant decision plasticity. Anuran amphibians breeding in ephemeral pools largely face two risks for their offspring: desiccation and predation. The Neotropical poison frog Ranitomeya variabilis deposits both eggs and tadpoles in phytotelmata. These small tadpole nurseries lower the risk of offspring predation. However, because most poison frog tadpoles are cannibalistic, even these pools need to be surveyed for predators, and parents tend to avoid deposition with conspecifics. We tested if this avoidance behaviour does change in parental R. variabilis depending on seasonal circumstances. Over several months we provided the frogs the option to deposit their eggs or tadpoles in pools that did and did not contain chemical cues of cannibalistic conspecifics, respectively. During the rainy season, frogs strongly avoided conspecific cues for both eggs and tadpoles. Anyway, with the change to the dry season, parental preferences changed such that parent frogs were more likely to deposit tadpoles (but not eggs) in pools containing cues of conspecific tadpoles. We suggest that R. variabilis, a species that typically isolates its cannibalistic offspring, has evolved a plastic feeding behaviour with regard to the risk of phytotelmata desiccation. We interpret that parents provide older tadpoles with younger 'trophic' tadpoles in order to accelerate their development and save them from impending desiccation. [ABSTRACT FROM AUTHOR]

Within the framework of the present study we test whether climatic niche similarity can be identified in a monophyletic group of species inhabiting remarkably restricted ranges by pooling presence data of all species into a single concatenated data set and subsequently jackknifing single species. We expect that, when the jackknifed species differs markedly in its climatic niche from all other species, this approach will result in increased niche homogeneity, allowing assessments of niche divergence patterns. To test our novel jackknife approach, we developed species distribution models for all members of Lycian salamanders (genus Lyciasalamandra), native to Turkey and the adjacent Aegean islands using Maxent. Degrees of niche similarity among species were assessed using Schoener's index. Significance of results was tested using null-models. The degree of niche similarity was generally high among all seven species, with only L. helverseni differing significantly from the others. Carstic lime stones providing specific microhabitat features may explain the high degree of niche similarity detected, since the variables with the highest explanative power in our models (i.e. mean temperature, and precipitation of the coldest quarter) corresponded well with salamander natural history observations, supporting the biologically plausibility of the results. We conclude that the jackknife approach presented here for the first time allows testing for niche similarity in species inhabiting restricted ranges and with few species records available. Our results strongly support the view that detailed natural history information and knowledge of microhabitats is crucial when assessing possible climate change impacts on species. [ABSTRACT FROM AUTHOR]

The use of species distribution models (SDMs) to predict potential distributions of species is steadily increasing. A necessary assumption when projecting models throughout space or time is that climatic niches are conservative, but recent findings of niche shifts during biological invasion of particular plant and animal species have indicated that this assumption is not categorically valid. One reason for observed shifts may relate to variable selection for modelling. In this study, we assess differences in climatic niches in the native and invasive ranges of the Greenhouse frog ( Eleutherodactylus planirostris). We analyze which variables are more ‘conserved’ in comparison to more ‘relaxed’ variables (i.e. subject to niche shift) and how they influence transferability of SDMs developed with Maxent on the basis of ten bioclimatic layers best describing the climatic requirements of the target species. We focus on degrees of niche similarity and conservatism using Schoener's index and Hellinger distance. Significance of results are tested with null models. Results indicate that the degrees of niche similarity and conservatism vary greatly among the predictive variables. Some shifts can be attributed to active habitat selection, whereas others apparently reflect variation in the availability of climate conditions or biotic interactions between the frogs' native and invasive ranges. Patterns suggesting active habitat selection also vary among variables. Our findings evoke considerable implications on the transferability of SDMs over space and time, which is strongly affected by the choice and number of predictors. The incorporation of ‘relaxed’ predictors not or only indirectly correlated with biologically meaningful predictors may lead to erroneous predictions when projecting SDMs. We recommend thorough assessments of invasive species' ecology for the identification biologically meaningful predictors facilitating transferability. [ABSTRACT FROM AUTHOR]

The disturbance vicariance hypothesis (DV) has been proposed to explain speciation in Amazonia, especially its edge regions, e.g. in eastern Guiana Shield harlequin frogs ( Atelopus) which are suggested to have derived from a cool-adapted Andean ancestor. In concordance with DV predictions we studied that (i) these amphibians display a natural distribution gap in central Amazonia; (ii) east of this gap they constitute a monophyletic lineage which is nested within a pre-Andean/western clade; (iii) climate envelopes of Atelopus west and east of the distribution gap show some macroclimatic divergence due to a regional climate envelope shift; (iv) geographic distributions of climate envelopes of western and eastern Atelopus range into central Amazonia but with limited spatial overlap. We tested if presence and apparent absence data points of Atelopus were homogenously distributed with Ripley’s K function. A molecular phylogeny (mitochondrial 16S rRNA gene) was reconstructed using Maximum Likelihood and Bayesian Inference to study if Guianan Atelopus constitute a clade nested within a larger genus phylogeny. We focused on climate envelope divergence and geographic distribution by computing climatic envelope models with MaxEnt based on macroscale bioclimatic parameters and testing them by using Schoener’s index and modified Hellinger distance. We corroborated existing DV predictions and, for the first time, formulated new DV predictions aiming on species’ climate envelope change. Our results suggest that cool-adapted Andean Atelopus ancestors had dispersed into the Amazon basin and further onto the eastern Guiana Shield where, under warm conditions, they were forced to change climate envelopes. [ABSTRACT FROM AUTHOR]

Amphibians are globally declining and approximately one-third of all species are threatened with extinction. Some of the most severe declines have occurred suddenly and for unknown reasons in apparently pristine habitats. It has been hypothesized that these “rapid enigmatic declines” are the result of a panzootic of the disease chytridiomycosis caused by globally emerging amphibian chytrid fungus. In a Species Distribution Model, we identified the potential distribution of this pathogen. Areas and species from which rapid enigmatic decline are known significantly overlap with those of highest environmental suitability to the chytrid fungus. We confirm the plausibility of a link between rapid enigmatic decline in worldwide amphibian species and epizootic chytridiomycosis. [ABSTRACT FROM AUTHOR]

Abstract: Poison frogs of the family Dendrobatidae contain cryptic as well as brightly colored, presumably aposematic species. The prevailing phylogenetic hypothesis assumes that the aposematic taxa form a monophyletic group while the cryptic species (Colostethus sensu lato) are basal and paraphyletic. Analysis of 86 dendrobatid sequences of a fragment of the 16S rRNA gene resulted in a much more complex scenario, with several clades that contained aposematic as well as cryptic taxa. Monophyly of the aposematic taxa was significantly rejected by SH-tests in an analysis with additional 12S and 16S rDNA fragments and reduced taxon sampling. The brightly colored Allobates femoralis and A. zaparo (Silverstone) comb. nov. (previously Epipedobates) belong in a clade with cryptic species of Colostethus. Additionally, Colostethus pratti was grouped with Epipedobates, and Colostethus bocagei with Cryptophyllobates. In several cases, the aposematic species have general distributions similar to those of their non-aposematic sister groups, indicating multiple instances of regional radiations in which some taxa independently acquired bright color. From a classificatory point of view, it is relevant that the type species of Minyobates, M. steyermarki, resulted as the sister group of the genus Dendrobates, and that species of Mannophryne and Nephelobates formed monophyletic clades, corroborating the validity of these genera. Leptodactylids of the genera Hylodes and Crossodactylus were not unambiguously identified as the sister group of the Dendrobatidae; these were monophyletic in all analyses and probably originated early in the radiation of Neotropical hyloid frogs. [Copyright &y& Elsevier]

An amendment to this paper has been published and can be accessed via a link at the top of the paper. [ABSTRACT FROM AUTHOR]

Many animals have evolved remarkable strategies to avoid predation. In diurnal, toxic harlequin toads (Atelopus) from the Amazon basin, we find a unique colour signal. Some Atelopus populations have striking red soles of the hands and feet, visible only when walking. When stationary, the toads are hard to detect despite their yellow-black dorsal coloration. Consequently, they switch between high and low conspicuousness. Interestingly, some populations lack the extra colour display of the soles. We found comprehensive support that the red coloration can act as an aposematic signal directed towards potential predators: red soles are significantly more conspicuous than soles lacking red coloration to avian predators and the presence of the red signal significantly increases detection. Further, toads with red soles show bolder behaviour by using higher sites in the vegetation than those lacking this signal. Field experiments hint at a lower attack risk for clay models with red soles than for those lacking the signal, in a population where the red soles naturally occur. We suggest that the absence of the signal may be explained by a higher overall attack risk or potential differences of predator community structure between populations. [ABSTRACT FROM AUTHOR]