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Darwin's frogs Rhinoderma darwinii and Rhinoderma rufum are the only known species of amphibians in which males brood their offspring in their vocal sacs. We propose these frogs as flagship species for the conservation of the Austral temperate forests of Chile and Argentina. This recommendation forms part of the vision of the Binational Conservation Strategy for Darwin's Frogs, which was launched in 2018. The strategy is a conservation initiative led by the IUCN SSC Amphibian Specialist Group, which in 2017 convened 30 governmental, non-profit and private organizations from Chile, Argentina and elsewhere. Darwin's frogs are iconic examples of the global amphibian conservation crisis: R. rufum is categorized as Critically Endangered (Possibly Extinct) on the IUCN Red List, and R. darwinii as Endangered. Here we articulate the conservation planning process that led to the development of the conservation strategy for these species and present its main findings and recommendations. Using an evidence-based approach, the Binational Conservation Strategy for Darwin's Frogs contains a comprehensive status review of Rhinoderma spp., including critical threat analyses, and proposes 39 prioritized conservation actions. Its goal is that by 2028, key information gaps on Rhinoderma spp. will be filled, the main threats to these species will be reduced, and financial, legal and societal support will have been achieved. The strategy is a multi-disciplinary, transnational endeavour aimed at ensuring the long-term viability of these unique frogs and their particular habitat.

Darwin's frogs (Rhinoderma darwinii and R. rufum) are two species of mouth brooding frogs from Chile and Argentina that have experienced marked population declines. Rhinoderma rufum has not been found in the wild since 1980. We investigated historical and current evidence of Batrachochytrium dendrobatidis (Bd) infection in Rhinoderma spp. to determine whether chytridiomycosis is implicated in the population declines of these species. Archived and live specimens of Rhinoderma spp., sympatric amphibians and amphibians at sites where Rhinoderma sp. had recently gone extinct were examined for Bd infection using quantitative real-time PCR. Six (0.9%) of 662 archived anurans tested positive for Bd (4/289 R. darwinii; 1/266 R. rufum and 1/107 other anurans), all of which had been collected between 1970 and 1978. An overall Bd-infection prevalence of 12.5% was obtained from 797 swabs taken from 369 extant individuals of R. darwinii and 428 individuals representing 18 other species of anurans found at sites with current and recent presence of the two Rhinoderma species. In extant R. darwinii, Bd-infection prevalence (1.9%) was significantly lower than that found in other anurans (7.3%). The prevalence of infection (30%) in other amphibian species was significantly higher in sites where either Rhinoderma spp. had become extinct or was experiencing severe population declines than in sites where there had been no apparent decline (3.0%; x(2) = 106.407, P<0.001). This is the first report of widespread Bd presence in Chile and our results are consistent with Rhinoderma spp. declines being due to Bd infection, although additional field and laboratory investigations are required to investigate this further.

Darwin's frogs (Rhinoderma darwinii and R. rufum) are two species of mouth-brooding frogs from Chile and Argentina. Here, we present evidence on the extent of declines, current distribution and conservation status of Rhinoderma spp.; including information on abundance, habitat and threats to extant Darwin's frog populations. All known archived Rhinoderma specimens were examined in museums in North America, Europe and South America. Extensive surveys were carried out throughout the historical ranges of R. rufum and R. darwinii from 2008 to 2012. Literature review and location data of 2,244 archived specimens were used to develop historical distribution maps for Rhinoderma spp. Based on records of sightings, optimal linear estimation was used to estimate whether R. rufum can be considered extinct. No extant R. rufum was found and our modelling inferred that this species became extinct in 1982 (95% CI, 1980-2000). Rhinoderma darwinii was found in 36 sites. All populations were within native forest and abundance was highest in Chiloé Island, when compared with Coast, Andes and South populations. Estimated population size and density (five populations) averaged 33.2 frogs/population (range, 10.2-56.3) and 14.9 frogs/100 m(2) (range, 5.3-74.1), respectively. Our results provide further evidence that R. rufum is extinct and indicate that R. darwinii has declined to a much greater degree than previously recognised. Although this species can still be found across a large part of its historical range, remaining populations are small and severely fragmented. Conservation efforts for R. darwinii should be stepped up and the species re-classified as Endangered.

Amphibian chytridiomycosis, caused by Batrachochytrium dendrobatidis (Bd), has been recognized as the infectious disease causing the most catastrophic loss of biodiversity known to science, with South America being the most impacted region. We tested whether Bd prevalence is distributed among host taxonomy, ecoregion, conservation status and habitat preference in South America. Here we provide a synthesis on the extent of Bd infection across South America based on 21 648 molecular diagnostic assays, roles of certain species in the epidemiology of Bd and explore its association with the reported amphibian catastrophic declines in the region. We show that Bd is widespread, with a continental prevalence of 23.2%. Its occurrence in the region shows a phylogenetic signal and the probability of infection is determined by ecoregion, preferred habitat and extinction risk hosts' traits. The taxa exhibiting highest Bd occurrence were mostly aquatic amphibians, including Ranidae, Telmatobiidae, Hylodidae, Calyptocephalellidae and Pipidae. Surprisingly, families exhibiting unusually low Bd prevalence included species in which lethal chytridiomycosis and population declines have been described (genera Atelopus, Rhinoderma and Eleutherodactylus). Higher than expected prevalence of Bd occurred mainly in amphibians living in association with mountain environments in the Andes and Atlantic forests, reflecting highly favourable Bd habitats in these areas. Invasive amphibian species (e.g. Lithobates catesbeianus and Xenopus laevis) exhibited high Bd prevalence; thus we suggest using these as sentinels to understand their potential role as reservoirs, vectors or spreaders of Bd that can be subjected to management. Our results guide on the prioritization of conservation actions to prevent further biodiversity loss due to chytridiomycosis in the world's most amphibian diverse region.

Sexual signals in different animals are expected to be dimorphic when both sexes signal, but cases of monomorphism are known to occur, and we lack a clear understanding about the factors that modulate the level of sexual dimorphism in signals. In this study, we evaluated the hypothesis that the lack of dimorphism in sexual signals might evolve in systems experiencing temporal changing conditions of intra-sexual competition. We used the Darwin’s frog (Rhinoderma darwinii), a species with paternal care, as a model. We compared advertisement calls and examined call distinctiveness among females, pregnant and non-pregnant males in a wild population from Chiloé island, Chile. We also recorded the vocal activity of both sexes along the reproductive season. Additionally, we compared the acoustic properties of their advertisement calls in terms of sexual distinctiveness and individual repeatability. We found that the proportion of females and pregnant males vocalizing changed over time following distinct patterns. Females produced calls with lower dominant frequency and longer note and call durations than males, and these acoustic differences were related to body size differences between sexes, but only dominant frequency contributed significantly to the distinctiveness of calls between sexes. Also, individual repeatability was high, indicating that calling can be relevant for social recognition. Overall, our results suggest that mutual selective pressures could be involved in the limited dimorphism of the advertisement calls in Darwin’s frogs, as the sex ratio of individuals vocalizing (i.e. females vs. reproductive males) is reversed along the breeding period.

Background Body size variation has played a central role in biogeographical research, however, most studies have aimed to describe trends rather than search for underlying mechanisms. In order to provide a more comprehensive understanding of the causes of intra-specific body size variation in ectotherms, we evaluated eight hypotheses proposed in the literature to account for geographical body size variation using the Darwin’s frog ( Rhinoderma darwinii ), an anuran species widely distributed in the temperate forests of South America. Each of the evaluated hypotheses predicted a specific relationship between body size and environmental variables. The level of support for each of these hypotheses was assessed using an information-theoretic approach and based on data from 1015 adult frogs obtained from 14 sites across the entire distributional range of the species. Results There was strong evidence favouring a single model comprising temperature seasonality as the predictor variable. Larger body sizes were found in areas of greater seasonality, giving support to the “starvation resistance” hypothesis. Considering the known role of temperature on ectothermic metabolism, however, we formulated a new, non-exclusive hypothesis, termed “hibernation hypothesis”: greater seasonality is expected to drive larger body size, since metabolic rate is reduced further and longer during colder, longer winters, leading to decreased energy depletion during hibernation, improved survival and increased longevity (and hence growth). Supporting this, a higher post-hibernation body condition in animals from areas of greater seasonality was found. Conclusions Despite largely recognized effects of temperature on metabolic rate in ectotherms, its importance in determining body size in a gradient of seasonality has been largely overlooked so far. Based on our results, we present and discuss an alternative mechanism, the “hibernation hypothesis”, underlying geographical body size variation, which can be helpful to improve our understanding of biogeographical patterns in ectotherms.

Darwin's frogs (Rhinoderma darwinii and R. rufum) are two species of mouth brooding frogs from Chile and Argentina that have experienced marked population declines. Rhinoderma rufum has not been found in the wild since 1980. We investigated historical and current evidence of Batrachochytrium dendrobatidis (Bd) infection in Rhinoderma spp. to determine whether chytridiomycosis is implicated in the population declines of these species. Archived and live specimens of Rhinoderma spp., sympatric amphibians and amphibians at sites where Rhinoderma sp. had recently gone extinct were examined for Bd infection using quantitative real-time PCR. Six (0.9%) of 662 archived anurans tested positive for Bd (4/289 R. darwinii; 1/266 R. rufum and 1/107 other anurans), all of which had been collected between 1970 and 1978. An overall Bd-infection prevalence of 12.5% was obtained from 797 swabs taken from 369 extant individuals of R. darwinii and 428 individuals representing 18 other species of anurans found at sites with current and recent presence of the two Rhinoderma species. In extant R. darwinii, Bd-infection prevalence (1.9%) was significantly lower than that found in other anurans (7.3%). The prevalence of infection (30%) in other amphibian species was significantly higher in sites where either Rhinoderma spp. had become extinct or was experiencing severe population declines than in sites where there had been no apparent decline (3.0%; x2 = 106.407, P<0.001). This is the first report of widespread Bd presence in Chile and our results are consistent with Rhinoderma spp. declines being due to Bd infection, although additional field and laboratory investigations are required to investigate this further.

The Southern Darwin’s frog Rhinoderma darwinii is a charismatic, mouth-brooding amphibian endemic to temperate forests of South America with a Vulnerable conservation status according to the IUCN Red List. We developed microsatellite markers from next generation sequence data that will aid genetic monitoring during and after re-introduction efforts. Using bioinformatics we characterized 3,521 perfect microsatellite repeats and designed primers for 35 of them. From these, 23 were polymorphic and amplified reliably. Number of alleles varied between 2 and 15, allele sizes varied between 84 and 299 bp, and observed heterozygosities varied between 0.105 and 0.904. These microsatellites represent a valuable resource to aid recovery of threatened Southern Darwin’s frog populations.

Larvae of Rhinoderma darwinii have a nonaquatic development inside the vocal sac of the male. Different authors have suggested the existence of a trophic relationship between the male and the larvae. In order to test this possibility, horseradish peroxidase, 3H valine and 3H leucine were injected into the paternal circulation and traced in the larvae at different postinjection intervals. The different tracers were demonstrated in several organs of the larvae thus indicating a possible paternal nourishment. Up to stage 10, the tracers were incorporated into the larvae through their skin and the cephalic and caudal parts of their digestive tract. At later stages, the intestines were the main route for the entry of tracers into the larvae.