A global reptile assessment highlights shared conservation needs of tetrapods.

Comprehensive assessments of species' extinction risks have documented the extinction crisis1 and underpinned strategies for reducing those risks2. Global assessments reveal that, among tetrapods, 40.7% of amphibians, 25.4% of mammals and 13.6% of birds are threatened with extinction3. Because global assessments have been lacking, reptiles have been omitted from conservation-prioritization analyses that encompass other tetrapods4-7. Reptiles are unusually diverse in arid regions, suggesting that they may have different conservation needs6. Here we provide a comprehensive extinction-risk assessment of reptiles and show that at least 1,829 out of 10,196 species (21.1%) are threatened-confirming a previous extrapolation8 and representing 15.6 billion years of phylogenetic diversity. Reptiles are threatened by the same major factors that threaten other tetrapods-agriculture, logging, urban development and invasive species-although the threat posed by climate change remains uncertain. Reptiles inhabiting forests, where these threats are strongest, are more threatened than those in arid habitats, contrary to our prediction. Birds, mammals and amphibians are unexpectedly good surrogates for the conservation of reptiles, although threatened reptiles with the smallest ranges tend to be isolated from other threatened tetrapods. Although some reptiles-including most species of crocodiles and turtles-require urgent, targeted action to prevent extinctions, efforts to protect other tetrapods, such as habitat preservation and control of trade and invasive species, will probably also benefit many reptiles.

Intra and interspecific variation in thermal performance and critical limits in anurans from southern Chile.

The relationship between temperature and performance can be illustrated through a thermal performance curve (TPC), which has proven useful in describing various aspects of ectotherms' thermal ecology and evolution. The parameters of the TPC can vary geographically due to large-scale variations in environmental conditions. However, only some studies have attempted to quantify how thermal performance varies over relatively small spatial scales, even in the same location or consistently among individuals within a species. Here, we quantified individual and species variation in thermal sensitivity of locomotor performance in five amphibia Eupsophus species found in the temperate rainforests of southern Chile and compared their estimates against co-occurring species that exhibit a substantially more extensive distributional range. We measured critical thermal limits and jumping performance under five different temperatures. Our results suggest that thermal responses are relatively conserved along the phylogeny, as the locomotor performance and thermal windows for activity remained narrow in Eupsophus species when compared against results observed for Batrachyla taeniata and Rhinella spinulosa. Additionally, we found significant individual differences in locomotor performance within most species, with individual consistency in performance observed across varied temperatures. Further analyses explored the influence of body size on locomotor performance and critical thermal limits within and between species. Our results suggest a trade-off scenario between thermal tolerance breadth and locomotor performance, where species exhibiting broader thermal ranges might have compromised performance. Interestingly, these traits seem partly mediated by body size variations, raising questions about potential ecological implications.

Morphological and molecular identification of the invasive freshwater snail Physa acuta (Gastropoda: Physidae) into Llanquihue Lake, Chilean Patagonia.

The sewage snail Physa acuta, native to North America, is an effective invasive species around the world. In Chile, it was first reported in 2014 in the north central area of the country. So far, the species has not been recorded in southern Chile. Sampling performed in 2015 in three localities from Llanquihue Lake, Chilean Patagonia, only provided native freshwater snails. However, new collections performed in February 2018 in the same three sites were successful for physid specimens suggesting a biological invasion entailing a large southward range expansion of these snails. Here we performed morphological, microstructural and phylogenetic analyses to investigate whether the new samples belong to Physa acuta. The shell morphology, male copulatory complex and radula microstructure of the new material agree with those of the sewage snail. The molecular phylogenetic analyses using the cytochrome c oxidase subunit 1 (COI) gene confirmed morphological identification. We suggest to take prompt measures to prevent the expansion of Physa acuta in Llanquihue Lake or nearby aquatic ecosystems.

Modeling warming predicts a physiological threshold for the extinction of the living fossil frog Calyptocephalella gayi.

Global climate change will have a greater impact on ectotherms in tropical and subtropical communities than at higher latitudes, because ambient temperatures are closer to the upper thermal limits of species. Amphibian species are highly dependent on external weather conditions, and the effect of global warming on these has been evaluated recently. The Great Chilean frog (Calyptocephalella gayi) is an endemic, monotypic species and genus whose conservation status is considered Vulnerable because of high extraction pressure for human consumption, lack of regulatory measures and comprehension by its consumers. Their populations have also declined due to the loss and destruction of their habitats. C. gayi has not been considered as an object of physiological study, so this large species is not known as one that can adapt to current environmental changes. In this study we analyze the thermoregulatory capacity and thermal efficiency of C. gayi to determine its potential for climatic adaptation. The results indicate that this species is strictly a thermal-conformer; its thermal efficiency and its ability to withstand high temperatures allow it to sustain itself under a climate change scenario, however, it has thermal constraints that do not allow it to withstand temperatures greater than 30°C. By modeling its ontogenetic conditions mathematically, we project that the larvae are not in danger, although there is a group of around 4% which is very close to 30°C, which is the highest temperature recorded for the species. However, about 40% of subadults and approximately 47% of adult frogs will not survive the change of ~7°C projected for the following 85 years, which will affect future generations.

Identifying Strategies for Effective Biodiversity Preservation and Species Status of Chilean Amphibians.

Resources are limited in global biodiversity conservation efforts, which emphasizes the significance of setting conservation priorities. Using standardized criteria, we evaluated 58 amphibian species in Chile to determine their conservation priority (CP). Species with insufficient historical data had their values marked as missing. With a median value of p = 1.67, the results demonstrated CP values ranging from p = 0.48 to p = 3.0, classifying species into priority and non-priority groups. Four levels were established for the priority categories: no priority, low priority, medium priority, and high priority. Additionally, the Telmatobiidae and Alsodidae families were identified as two more priority families. Notably, the species with the highest priority were found to be T. halli, T. fronteriensis, T. philippii, T. chusmisensis, A. pehuenche, and Alsodes tumultuosus, where T. philippii and T. fronteriensis have equal priority for conservation at the national level according to the conservation priority analysis. Eight priority families-the Alsodidae, Batrachylidae, Bufonidae, Ceratophryidae, Leptodactylidae, Rhinodermatidae, and Telmatobiidae-were determined, and 14 species-or 24% of the species examined-need further study. Based on the conservation priority analysis, the species T. fronteriensis and T. philippii share the highest priority for conservation at the national level (p = 2.50). With 70% of the amphibians under study being threatened mainly by habitat loss, pollution, and emerging diseases, the creation of conservation categories made the threat assessment process easier. Due to a lack of information on geographic distribution and abundance, quantitatively classifying amphibians in Chile remains difficult. The analysis of conservation priorities and potential extinction threats informs appropriate management strategies.

Genetic Diversity, Morphometric Characterization, and Conservation Reassessment of the Critically Endangered Freshwater Snail, Heleobia atacamensis, in the Atacama Saltpan, Northern Chile.

Evaporitic ecosystems of the Atacama Desert contain a rich endemic fauna, including mollusk species. A recent study performed in the freshwater snail Heleobia atacamensis, endemic to the Atacama Saltpan, revealed a strong interdependence of genetic patterns with climatic fluctuations and landscape physiography. The species is currently listed as Critically Endangered at regional scale and as Data Deficient on the International Union for Conservation of Nature (IUCN) Red List. Here, we studied genetic diversity and demographic history of several populations of the species occurring on a connectivity gradient, including snails from new peripherical localities (Peine and Tilomonte), which were compared with topotype specimens. In addition, we reassessed the conservation status using the IUCN Red List categories and criteria considering species-specific idiosyncrasy. Phylogenetic and phylogeographical analyses indicated that snails from Peine and Tilomonte belong to H. atacamensis. We discovered significant differentiation in shell morphology, which was generally greater in geographically isolated populations. We also inferred six genetic clusters and a demographic expansion congruent with the wet periods that occurred at the end of the Pleistocene. Considering the highest risk category obtained, H. atacamensis was reassessed as Endangered at regional scale. Future conservation plans should consider the genetic assemblages as conservation units.

Amphibian responses in experimental thermal gradients: Concepts and limits for inference.

The interpretation of thermal-gradient data depends on the behavioral drives reported or assumed, and on the underlying behavioral models explaining how such drives operate. The best-known example is positive thermotaxis, a thermoregulatory behavioral drive frequently linked to a dual set-point model of thermoregulation around a target range. This behavioral drive is often assumed as dominant among 'ectotherms', including amphibians. However, we argue that, because amphibians are extremely diverse, they may exhibit alternative behavioral drives in thermal gradients, and tackle this idea from two perspectives. First, we provide a historical review of original definitions and proposed limits for inference. Second, although caveats apply, we propose that a cross-study analysis of data of temperature settings of gradients and the temperatures selected by amphibians would corroborate alternative behavioral drives, including negative thermotaxis. Therefore, we analyzed published data focusing on such relationships and show that gradient temperature settings influence the temperatures selected by amphibians, with further effects of phylogeny and ontogeny. We conclude that thermal gradient experiments are outstanding tools to investigate behavioral drives, but no given drive can be assumed a priori unless additional information about thermoregulation is available. Based on the historical debate, we propose using selected temperatures and preferred temperatures as different concepts, the former merely operational and the second explicitly linked to positive thermotaxis (and thus compatible with dual set-point thermoregulation). Under this view, thermal preferences would stand for a hypothesis of a behavioral drive (positive thermotaxis) requiring formal testing. These considerations impact the scope for inference based on thermal gradient experiments, particularly ecological modeling and emerging disease.

Geography, Temperature, and Water: Interaction Effects in a Small Native Amphibian.

Amphibian locomotor capacity is strongly linked to temperature and hydration. However, organisms in nature experience covariation of multiple environmental factors, and thus to better understand the effects of thermal and hydric conditions on physiological performance, it is critical not only to experimentally disentangle them but also to incorporate potential interactive effects due to geographic variation. To this end, we selected two populations of the small amphibian Pleurodema thaul inhabiting highly contrasting temperatures and precipitation regimens. With these two populations, we evaluated the thermal and hydric sensitivities of locomotor performance. For both factors, performance increased with temperature as well as with hydration level, although performance reached a plateau between 25° and 30°C. In addition, the influence of dehydration on performance was independent of the temperature at which it was tested. Our results also showed that the population from the warmer environment has lower sensitivity of locomotor performance to dehydration, probably as a consequence of thermal adaptation, although further studies might be required to fully understand this.

Morphological and molecular analysis of cryptic native and invasive freshwater snails in Chile.

Species delimitation in minute freshwater snails is often difficult to perform using solely shell morphology. The problem intensifies when invasive species spread within the distribution range of morphologically similar native species. In Chile, the Truncatelloidean snails are represented by the native genera Heleobia and Potamolithus plus the invasive mudsnail Potamopyrgus antipodarum, which can easily be confused. Using an integrative approach, we performed molecular phylogenetic analysis and studied reproductive and morphological features to identify superficially similar forms inhabiting the central area of the country. Truncatelloidean snails were identified in 40 of 51 localities sampled, 10 containing Potamopyrgus antipodarum, 23 Heleobia and 7 Potamolithus. Based on these results and previously published data, the known distribution of the mudsnail in Chile encompasses 6 hydrological basins, including 18 freshwater ecosystems. The finding of the mudsnails in several type localities of native species/subspecies of "Heleobia" that were not find in situ suggests species replacement or significant extinction of native fauna, a hypothesis supported by the restudy of type material that shows that endemic forms belong to the genus Potamolithus. This study shows the usefulness of integrative taxonomy not only resolving complex taxa with cryptic morphology but also measuring the extent of an ongoing invasion.

Geometric morphometric analysis of the head of Microlophus atacamensis (Tropiduridae) in a latitudinal gradient.

BACKGROUND: Clinal variation is defined as gradual variation in a character associated with geographic distance among sites. Microlophus atacamensis is a medium large lizard species which inhabits the intertidal zone of northern Chile, distributed from Antofagasta (23° 39' S) to Arrayán, La Serena (29° 41' S) in a gradient which could show clinal variation. Geometric morphometrics analyzes differences in shape independent of size; information about shape allows a more complete biological interpretation than information on morphological variation. This studyevaluated clinal variation in the head shape of M.atacamensis fromfive localities (Antofagasta, Paposo, Pan de Azúcar, Caldera, and Tres Playitas) using dorsal and lateral views, comparing form variation with latitudinal distribution. The heads of adults collected were photographed in lateral and dorsal views. RESULTS: The analysis did not find significant differences in form among the five localities, in contrast to the proposal of earlier studies, and no differences were recorded between the sexes. Possible reasons why these populations are not differentiated in the latitudinal gradient are discussed. CONCLUSIONS: Our results show that there are no differences between the studied, among the explanations are that populationsof this species are subjected to similar environments that promote convergence of the structures analyzed.

Stable isotopes document mainland-island divergence in resource use without concomitant physiological changes in the lizard Liolaemus pictus.

Shifts in feeding ecology are believed to promote island-mainland divergence. The lizard Liolaemus pictus has several different subspecies on Chilean islands and mainland. These subspecies inhabit contrastingly different habitats both in different islands and mainland, which suggests the potential for habitat related dietary variation. We investigated the dietary habits of L. pictus by both stomach content analyses and by nitrogen stable isotope analyses (delta(15)N), which we used as a proxy variable for trophic level. We also compared the morphology of the digestive tract and the activity of intestinal digestive enzymes of mainland and island lizards. We hypothesized differences in diet and trophic level among populations and that these differences would predict the expression of the morphological and biochemical features of the digestive tract. More specifically, we predicted shorter intestines and higher levels of peptidases in more insectivorous than in more frugivorous/herbivorous lizards. The diet of L. pictus was characterized by the consumption of a wide diversity of food types, including fruit and insects, in all populations. Stable isotopes revealed higher trophic level, and hence probably higher protein intake, in mainland than in island populations, but contrary to our prediction, they had shorter intestines and higher relative activity of intestinal peptidases than mainland lizards. Furthermore, the proportion of fruit items in the stomach content was higher in the population that exhibited the lowest tropic level. These results suggest that morphological and physiological differences among populations of L. pictus are not correlated with feeding ecology, suggesting that the lizard's first responses to the selective pressure represented by a diet shift are behavioral.