Genomics-informed delineation of conservation units in a desert amphibian.

Delineating conservation units (CUs, e.g., evolutionarily significant units, ESUs, and management units, MUs) is critical to the recovery of declining species because CUs inform both listing status and management actions. Genomic data have strengths and limitations in informing CU delineation and related management questions in natural systems. We illustrate the value of using genomic data in combination with landscape, dispersal and occupancy data to inform CU delineation in Nevada populations of the Great Basin Distinct Population Segment of the Columbia spotted frog (Rana luteiventris). R. luteiventris occupies naturally fragmented aquatic habitats in this xeric region, but beaver removal, climate change and other factors have put many of these populations at high risk of extirpation without management intervention. We addressed three objectives: (i) assessing support for ESUs within Nevada; (ii) evaluating and revising, if warranted, the current delineation of MUs; and (iii) evaluating genetic diversity, effective population size, adaptive differentiation and functional connectivity to inform ongoing management actions. We found little support for ESUs within Nevada but did identify potential revisions to MUs based on unique landscape drivers of connectivity that distinguish these desert populations from those in the northern portion of the species range. Effective sizes were uniformly small, with low genetic diversity and weak signatures of adaptive differentiation. Our findings suggest that management actions, including translocations and genetic rescue, might be warranted. Our study illustrates how a carefully planned genetic study, designed to address priority management goals that include CU delineation, can provide multiple insights to inform conservation action.

Diverse aging rates in ectothermic tetrapods provide insights for the evolution of aging and longevity.

Comparative studies of mortality in the wild are necessary to understand the evolution of aging; yet, ectothermic tetrapods are underrepresented in this comparative landscape, despite their suitability for testing evolutionary hypotheses. We present a study of aging rates and longevity across wild tetrapod ectotherms, using data from 107 populations (77 species) of nonavian reptiles and amphibians. We test hypotheses of how thermoregulatory mode, environmental temperature, protective phenotypes, and pace of life history contribute to demographic aging. Controlling for phylogeny and body size, ectotherms display a higher diversity of aging rates compared with endotherms and include phylogenetically widespread evidence of negligible aging. Protective phenotypes and life-history strategies further explain macroevolutionary patterns of aging. Analyzing ectothermic tetrapods in a comparative context enhances our understanding of the evolution of aging.

Genomic signatures of thermal adaptation are associated with clinal shifts of life history in a broadly distributed frog.

Temperature is a critical driver of ectotherm life-history strategies, whereby a warmer environment is associated with increased growth, reduced longevity and accelerated senescence. Increasing evidence indicates that thermal adaptation may underlie such life-history shifts in wild populations. Single nucleotide polymorphisms (SNPs) and copy number variants (CNVs) can help uncover the molecular mechanisms of temperature-driven variation in growth, longevity and senescence. However, our understanding of these mechanisms is still limited, which reduces our ability to predict the response of non-model ectotherms to global temperature change. In this study, we examined the potential role of thermal adaptation in clinal shifts of life-history traits (i.e. life span, senescence rate and recruitment) in the Columbia spotted frog Rana luteiventris along a broad temperature gradient in the western United States. We took advantage of extensive capture-recapture datasets of 20,033 marked individuals from eight populations surveyed annually for 14-18 years to examine how mean annual temperature and precipitation influenced demographic parameters (i.e. adult survival, life span, senescence rate, recruitment and population growth). After showing that temperature was the main climatic predictor influencing demography, we used RAD-seq data (50,829 SNPs and 6,599 putative CNVs) generated for 352 individuals from 31 breeding sites to identify the genomic signatures of thermal adaptation. Our results showed that temperature was negatively associated with annual adult survival and reproductive life span and positively associated with senescence rate. By contrast, recruitment increased with temperature, promoting the long-term viability of most populations. These temperature-dependent demographic changes were associated with strong genomic signatures of thermal adaptation. We identified 148 SNP candidates associated with temperature including three SNPs located within protein-coding genes regulating resistance to cold and hypoxia, immunity and reproduction in ranids. We also identified 39 CNV candidates (including within 38 transposable elements) for which normalized read depth was associated with temperature. Our study indicates that both SNPs and structural variants are associated with temperature and could eventually be found to play a functional role in clinal shifts in senescence rate and life-history strategies in R. luteiventris. These results highlight the potential role of different sources of molecular variation in the response of ectotherms to environmental temperature variation in the context of global warming.

Sex-related differences in aging rate are associated with sex chromosome system in amphibians.

Sex-related differences in mortality are widespread in the animal kingdom. Although studies have shown that sex determination systems might drive lifespan evolution, sex chromosome influence on aging rates have not been investigated so far, likely due to an apparent lack of demographic data from clades including both XY (with heterogametic males) and ZW (heterogametic females) systems. Taking advantage of a unique collection of capture-recapture datasets in amphibians, a vertebrate group where XY and ZW systems have repeatedly evolved over the past 200 million years, we examined whether sex heterogamy can predict sex differences in aging rates and lifespans. We showed that the strength and direction of sex differences in aging rates (and not lifespan) differ between XY and ZW systems. Sex-specific variation in aging rates was moderate within each system, but aging rates tended to be consistently higher in the heterogametic sex. This led to small but detectable effects of sex chromosome system on sex differences in aging rates in our models. Although preliminary, our results suggest that exposed recessive deleterious mutations on the X/Z chromosome (the "unguarded X/Z effect") or repeat-rich Y/W chromosome (the "toxic Y/W effect") could accelerate aging in the heterogametic sex in some vertebrate clades.

Thermal conditions predict intraspecific variation in senescence rate in frogs and toads.

Variation in temperature is known to influence mortality patterns in ectotherms. Even though a few experimental studies on model organisms have reported a positive relationship between temperature and actuarial senescence (i.e., the increase in mortality risk with age), how variation in climate influences the senescence rate across the range of a species is still poorly understood in free-ranging animals. We filled this knowledge gap by investigating the relationships linking senescence rate, adult lifespan, and climatic conditions using long-term capture-recapture data from multiple amphibian populations. We considered two pairs of related anuran species from the Ranidae (Rana luteiventris and Rana temporaria) and Bufonidae (Anaxyrus boreas and Bufo bufo) families, which diverged more than 100 Mya and are broadly distributed in North America and Europe. Senescence rates were positively associated with mean annual temperature in all species. In addition, lifespan was negatively correlated with mean annual temperature in all species except A. boreas In both R. luteiventris and A. boreas, mean annual precipitation and human environmental footprint both had negligible effects on senescence rates or lifespans. Overall, our findings demonstrate the critical influence of thermal conditions on mortality patterns across anuran species from temperate regions. In the current context of further global temperature increases predicted by Intergovernmental Panel on Climate Change scenarios, a widespread acceleration of aging in amphibians is expected to occur in the decades to come, which might threaten even more seriously the viability of populations and exacerbate global decline.

From satellites to frogs: Quantifying ecohydrological change, drought mitigation, and population demography in desert meadows.

Increasing frequency and severity of droughts have motivated natural resource managers to mitigate harmful ecological and hydrological effects of drought, but drought mitigation is an emerging science and evaluating its effectiveness is difficult. We examined ecohydrological responses of drought mitigation actions aimed at conserving populations of the Columbia spotted frog (Rana luteiventris) in a semi-arid valley in Nevada, USA. Abundance of this rare frog had declined precipitously after multiple droughts. Mitigation included excavating ponds to increase available surface water and installing earthen dams to raise water tables. We assessed responses of riparian vegetation to mitigation using a 30-year time series of satellite-derived Normalized Difference Vegetation Index (NDVI) and gridded weather data. We then analyzed a 23-year mark-recapture dataset to evaluate the effects of drought mitigation and NDVI on the probability of frog survival and rates of recruitment. After accounting for interannual precipitation variability, we found that NDVI increased significantly from before to after drought mitigation, suggesting that mitigation influenced the hydrology and vegetation of the meadows. Frog survival increased with NDVI, but mitigation had a stronger effect than NDVI suggesting that excavated mitigation ponds were particularly important for frog survival during drought. In contrast, frog recruitment was associated with NDVI more than mitigation, but only in meadows where NDVI was dependent on precipitation. At meadows with available groundwater, recruitment was associated with mitigation ponds. These findings suggest that mitigation ponds are critical for juvenile frogs to recruit into the adult population, but recruitment can also be increased by raising water tables in meadows lacking groundwater sources. Lagged recruitment (i.e., effects on larvae and juveniles) was negatively associated with NDVI. This study illustrates the ecohydrological complexity of drought mitigation and demonstrates novel ways to assess the effectiveness of drought mitigation using time series of readily available satellite imagery and organismal data.

A National-Scale Assessment of Mercury Bioaccumulation in United States National Parks Using Dragonfly Larvae As Biosentinels through a Citizen-Science Framework.

We conducted a national-scale assessment of mercury (Hg) bioaccumulation in aquatic ecosystems, using dragonfly larvae as biosentinels, by developing a citizen-science network to facilitate biological sampling. Implementing a carefully designed sampling methodology for citizen scientists, we developed an effective framework for a landscape-level inquiry that might otherwise be resource limited. We assessed the variation in dragonfly Hg concentrations across >450 sites spanning 100 United States National Park Service units and examined intrinsic and extrinsic factors associated with the variation in Hg concentrations. Mercury concentrations ranged between 10.4 and 1411 ng/g dry weight across sites and varied among habitat types. Dragonfly total Hg (THg) concentrations were up to 1.8-fold higher in lotic habitats than in lentic habitats and 37% higher in waterbodies with abundant wetlands along their margins than those without wetlands. Mercury concentrations in dragonflies differed among families but were correlated (r2 > 0.80) with each other, enabling adjustment to a consistent family to facilitate spatial comparisons among sampling units. Dragonfly THg concentrations were positively correlated with THg concentrations in both fish and amphibians from the same locations, indicating that dragonfly larvae are effective indicators of Hg bioavailability in aquatic food webs. We used these relationships to develop an integrated impairment index of Hg risk to aquatic ecosytems and found that 12% of site-years exceeded high or severe benchmarks of fish, wildlife, or human health risk. Collectively, this continental-scale study demonstrates the utility of dragonfly larvae for estimating the potential mercury risk to fish and wildlife in aquatic ecosystems and provides a framework for engaging citizen science as a component of landscape Hg monitoring programs.

Extreme Arsenic and Antimony Uptake and Tolerance in Toad Tadpoles during Development in Highly Contaminated Wetlands.

The effects of extreme concentrations of toxic metalloids, such as arsenic (As) and antimony (Sb), on larval amphibians are not well-understood. We sampled Western Toad tadpoles (Anaxyrus boreas) living in As- and Sb-contaminated wetlands throughout their development. Although the tadpoles completed metamorphosis, they accumulated among the highest concentrations of As and Sb ever reported for a living vertebrate (3866.9 mg/kg; 315.0 mg/kg (dry weight), respectively). Ingestion of contaminated sediment had a more important role in metalloid accumulation than aqueous exposure alone. Metalloids were initially concentrated in the gut; however, by metamorphosis, the majority were found in other tissues. These concentrations subsequently decreased with the onset of metamorphosis, yet remained quite elevated. Sublethal effects, including delayed development and reduced size at metamorphosis, were associated with elevated metalloid exposure. The presence of organic arsenicals in tadpole tissues suggests they have the ability to biomethylate inorganic As compounds. The arsenical trimethyl arsine oxide accounted for the majority of extractable organic As, with lesser amounts of monomethylarsonic acid and dimethylarsinic acid. Our findings demonstrate remarkable tolerance of toad tadpoles to extreme metalloid exposure and implicate physiological processes mediating that tolerance.

Harvester ant seed removal in an invaded sagebrush ecosystem: Implications for restoration.

A better understanding of seed movement in plant community dynamics is needed, especially in light of disturbance-driven changes and investments into restoring degraded plant communities. A primary agent of change within the sagebrush-steppe is wildfire and invasion by non-native forbs and grasses, primarily cheatgrass (Bromus tectorum). Our objectives were to quantify seed removal and evaluate ecological factors influencing seed removal within degraded sagebrush-steppe by granivorous Owyhee harvester ants (Pogonomyrmex salinus Olsen). In 2014, we sampled 76 harvester ant nests across 11 plots spanning a gradient of cheatgrass invasion (40%-91% cover) in southwestern Idaho, United States. We presented seeds from four plant species commonly used in postfire restoration at 1.5 and 3.0 m from each nest to quantify seed removal. We evaluated seed selection for presented species, monthly removal, and whether biotic and abiotic factors (e.g., distance to nearest nest, temperature) influenced seed removal. Our top model indicated seed removal was positively correlated with nest height, an indicator of colony size. Distance to seeds and cheatgrass canopy cover reduced seed removal, likely due to increased search and handling time. Harvester ants were selective, removing Indian ricegrass (Achnatherum hymenoides) more than any other species presented. We suspect this was due to ease of seed handling and low weight variability. Nest density influenced monthly seed removal, as we estimated monthly removal of 1,890 seeds for 0.25 ha plots with 1 nest and 29,850 seeds for plots with 15 nests. Applying monthly seed removal to historical restoration treatments across the western United States showed harvester ants can greatly reduce seed availability at degraded sagebrush sites; for instance, fourwing saltbush (Atriplex canescens) seeds could be removed in <2 months. Collectively, these results shed light on seed removal by harvester ants and emphasize their potential influence on postfire restoration within invaded sagebrush communities.

Transient population dynamics impede restoration and may promote ecosystem transformation after disturbance.

The apparent failure of ecosystems to recover from increasingly widespread disturbance is a global concern. Despite growing focus on factors inhibiting resilience and restoration, we still know very little about how demographic and population processes influence recovery. Using inverse and forward demographic modelling of 531 post-fire sagebrush populations across the western US, we show that demographic processes during recovery from seeds do not initially lead to population growth but rather to years of population decline, low density, and risk of extirpation after disturbance and restoration, even at sites with potential to support long-term, stable populations. Changes in population structure, and resulting transient population dynamics, lead to a > 50% decline in population growth rate after disturbance and significant reductions in population density. Our results indicate that demographic processes influence the recovery of ecosystems from disturbance and that demographic analyses can be used by resource managers to anticipate ecological transformation risk.

Quantifying climate sensitivity and climate-driven change in North American amphibian communities.

Changing climate will impact species' ranges only when environmental variability directly impacts the demography of local populations. However, measurement of demographic responses to climate change has largely been limited to single species and locations. Here we show that amphibian communities are responsive to climatic variability, using >500,000 time-series observations for 81 species across 86 North American study areas. The effect of climate on local colonization and persistence probabilities varies among eco-regions and depends on local climate, species life-histories, and taxonomic classification. We found that local species richness is most sensitive to changes in water availability during breeding and changes in winter conditions. Based on the relationships we measure, recent changes in climate cannot explain why local species richness of North American amphibians has rapidly declined. However, changing climate does explain why some populations are declining faster than others. Our results provide important insights into how amphibians respond to climate and a general framework for measuring climate impacts on species richness.

Regional variation in drivers of connectivity for two frog species (Rana pretiosa and R. luteiventris) from the U.S. Pacific Northwest.

Comparative landscape genetics has uncovered high levels of variability in which landscape factors affect connectivity among species and regions. However, the relative importance of species traits versus environmental variation for predicting landscape patterns of connectivity is unresolved. We provide evidence from a landscape genetics study of two sister taxa of frogs, the Oregon spotted frog (Rana pretiosa) and the Columbia spotted frog (Rana luteiventris) in Oregon and Idaho, USA. Rana pretiosa is relatively more dependent on moisture for dispersal than R. luteiventris, so if species traits influence connectivity, we predicted that connectivity among R. pretiosa populations would be more positively associated with moisture than R. luteiventris. However, if environmental differences are important drivers of gene flow, we predicted that connectivity would be more positively related to moisture in arid regions. We tested these predictions using eight microsatellite loci and gravity models in two R. pretiosa regions and four R. luteiventris regions (n = 1,168 frogs). In R. pretiosa, but not R. luteiventris, connectivity was positively related to mean annual precipitation, supporting our first prediction. In contrast, connectivity was not more positively related to moisture in more arid regions. Various temperature metrics were important predictors for both species and in all regions, but the directionality of their effects varied. Therefore, the pattern of variation in drivers of connectivity was consistent with predictions based on species traits rather than on environmental variation.

Adapting management to a changing world: Warm temperatures, dry soil, and interannual variability limit restoration success of a dominant woody shrub in temperate drylands.

Restoration and rehabilitation of native vegetation in dryland ecosystems, which encompass over 40% of terrestrial ecosystems, is a common challenge that continues to grow as wildfire and biological invasions transform dryland plant communities. The difficulty in part stems from low and variable precipitation, combined with limited understanding about how weather conditions influence restoration outcomes, and increasing recognition that one-time seeding approaches can fail if they do not occur during appropriate plant establishment conditions. The sagebrush biome, which once covered over 620,000 km2 of western North America, is a prime example of a pressing dryland restoration challenge for which restoration success has been variable. We analyzed field data on Artemisia tridentata (big sagebrush) restoration collected at 771 plots in 177 wildfire sites across its western range, and used process-based ecohydrological modeling to identify factors leading to its establishment. Our results indicate big sagebrush occurrence is most strongly associated with relatively cool temperatures and wet soils in the first spring after seeding. In particular, the amount of winter snowpack, but not total precipitation, helped explain the availability of spring soil moisture and restoration success. We also find considerable interannual variability in the probability of sagebrush establishment. Adaptive management strategies that target seeding during cool, wet years or mitigate effects of variability through repeated seeding may improve the likelihood of successful restoration in dryland ecosystems. Given consistent projections of increasing temperatures, declining snowpack, and increasing weather variability throughout midlatitude drylands, weather-centric adaptive management approaches to restoration will be increasingly important for dryland restoration success.

An analytical framework for estimating aquatic species density from environmental DNA.

Environmental DNA (eDNA) analysis of water samples is on the brink of becoming a standard monitoring method for aquatic species. This method has improved detection rates over conventional survey methods and thus has demonstrated effectiveness for estimation of site occupancy and species distribution. The frontier of eDNA applications, however, is to infer species density. Building upon previous studies, we present and assess a modeling approach that aims at inferring animal density from eDNA. The modeling combines eDNA and animal count data from a subset of sites to estimate species density (and associated uncertainties) at other sites where only eDNA data are available. As a proof of concept, we first perform a cross-validation study using experimental data on carp in mesocosms. In these data, fish densities are known without error, which allows us to test the performance of the method with known data. We then evaluate the model using field data from a study on a stream salamander species to assess the potential of this method to work in natural settings, where density can never be known with absolute certainty. Two alternative distributions (Normal and Negative Binomial) to model variability in eDNA concentration data are assessed. Assessment based on the proof of concept data (carp) revealed that the Negative Binomial model provided much more accurate estimates than the model based on a Normal distribution, likely because eDNA data tend to be overdispersed. Greater imprecision was found when we applied the method to the field data, but the Negative Binomial model still provided useful density estimates. We call for further model development in this direction, as well as further research targeted at sampling design optimization. It will be important to assess these approaches on a broad range of study systems.

Survey of Beaver-related Restoration Practices in Rangeland Streams of the Western USA.

Poor condition of many streams and concerns about future droughts in the arid and semi-arid western USA have motivated novel restoration strategies aimed at accelerating recovery and increasing water resources. Translocation of beavers into formerly occupied habitats, restoration activities encouraging beaver recolonization, and instream structures mimicking the effects of beaver dams are restoration alternatives that have recently gained popularity because of their potential socioeconomic and ecological benefits. However, beaver dams and dam-like structures also harbor a history of social conflict. Hence, we identified a need to assess the use of beaver-related restoration projects in western rangelands to increase awareness and accountability, and identify gaps in scientific knowledge. We inventoried 97 projects implemented by 32 organizations, most in the last 10 years. We found that beaver-related stream restoration projects undertaken mostly involved the relocation of nuisance beavers. The most common goal was to store water, either with beaver dams or artificial structures. Beavers were often moved without regard to genetics, disease, or potential conflicts with nearby landowners. Few projects included post-implementation monitoring or planned for longer term issues, such as what happens when beavers abandon a site or when beaver dams or structures breach. Human dimensions were rarely considered and water rights and other issues were mostly unresolved or addressed through ad-hoc agreements. We conclude that the practice and implementation of beaver-related restoration has outpaced research on its efficacy and best practices. Further scientific research is necessary, especially research that informs the establishment of clear guidelines for best practices.

Refining the cheatgrass-fire cycle in the Great Basin: Precipitation timing and fine fuel composition predict wildfire trends.

Larger, more frequent wildfires in arid and semi-arid ecosystems have been associated with invasion by non-native annual grasses, yet a complete understanding of fine fuel development and subsequent wildfire trends is lacking. We investigated the complex relationships among weather, fine fuels, and fire in the Great Basin, USA. We first modeled the annual and time-lagged effects of precipitation and temperature on herbaceous vegetation cover and litter accumulation over a 26-year period in the northern Great Basin. We then modeled how these fine fuels and weather patterns influence subsequent wildfires. We found that cheatgrass cover increased in years with higher precipitation and especially when one of the previous 3 years also was particularly wet. Cover of non-native forbs and native herbs also increased in wet years, but only after several dry years. The area burned by wildfire in a given year was mostly associated with native herb and non-native forb cover, whereas cheatgrass mainly influenced area burned in the form of litter derived from previous years' growth. Consequently, multiyear weather patterns, including precipitation in the previous 1-3 years, was a strong predictor of wildfire in a given year because of the time needed to develop these fine fuel loads. The strong relationship between precipitation and wildfire allowed us to expand our inference to 10,162 wildfires across the entire Great Basin over a 35-year period from 1980 to 2014. Our results suggest that the region's precipitation pattern of consecutive wet years followed by consecutive dry years results in a cycle of fuel accumulation followed by weather conditions that increase the probability of wildfire events in the year when the cycle transitions from wet to dry. These patterns varied regionally but were strong enough to allow us to model annual wildfire risk across the Great Basin based on precipitation alone.

Methodological considerations of terrestrial laser scanning for vegetation monitoring in the sagebrush steppe.

Terrestrial laser scanning (TLS) provides fast collection of high-definition structural information, making it a valuable field instrument to many monitoring applications. A weakness of TLS collections, especially in vegetation, is the occurrence of unsampled regions in point clouds where the sensor's line-of-sight is blocked by intervening material. This problem, referred to as occlusion, may be mitigated by scanning target areas from several positions, increasing the chance that any given area will fall within the scanner's line-of-sight from at least one position. Because TLS collections are often employed in remote regions where the scope of sampling is limited by logistical factors such as time and battery power, it is important to design field protocols which maximize efficiency and support increased quantity and quality of the data collected. This study informs researchers and practitioners seeking to optimize TLS sampling methods for vegetation monitoring in dryland ecosystems through three analyses. First, we quantify the 2D extent of occluded regions based on the range from single scan positions. Second, we measure the efficacy of additional scan positions on the reduction of 2D occluded regions (area) using progressive configurations of scan positions in 1 ha plots. Third, we test the reproducibility of 3D sampling yielded by a 5-scan/ha sampling methodology using redundant sets of scans. Analyses were performed using measurements at analysis scales of 5 to 50 cm across the 1-ha plots, and we considered plots in grass and shrub-dominated communities separately. In grass-dominated plots, a center-scan configuration and 5 cm pixel size sampled at least 90% of the area up to 18 m away from the scanner. In shrub-dominated plots, sampling at least 90% of the area was only achieved within a distance of 12 m. We found that 3 and 5 scans/ha are needed to sample at least ~ 70% of the total area (1 ha) in the grass and shrub-dominated plots, respectively, using 5 cm pixels to measure sampling presence-absence. The reproducibility of 3D sampling provided by a 5 position scan layout across 1-ha plots was 50% (shrub) and 70% (grass) using a 5-cm voxel size, whereas at the 50-cm voxel scale, reproducibility of sampling was nearly 100% for all plot types. Future studies applying TLS in similar dryland environments for vegetation monitoring may use our results as a guide to efficiently achieve sampling coverage and reproducibility in datasets.

Quantitative evidence for the effects of multiple drivers on continental-scale amphibian declines.

Since amphibian declines were first proposed as a global phenomenon over a quarter century ago, the conservation community has made little progress in halting or reversing these trends. The early search for a "smoking gun" was replaced with the expectation that declines are caused by multiple drivers. While field observations and experiments have identified factors leading to increased local extinction risk, evidence for effects of these drivers is lacking at large spatial scales. Here, we use observations of 389 time-series of 83 species and complexes from 61 study areas across North America to test the effects of 4 of the major hypothesized drivers of declines. While we find that local amphibian populations are being lost from metapopulations at an average rate of 3.79% per year, these declines are not related to any particular threat at the continental scale; likewise the effect of each stressor is variable at regional scales. This result - that exposure to threats varies spatially, and populations vary in their response - provides little generality in the development of conservation strategies. Greater emphasis on local solutions to this globally shared phenomenon is needed.

Effects of changing climate on aquatic habitat and connectivity for remnant populations of a wide-ranging frog species in an arid landscape.

Amphibian species persisting in isolated streams and wetlands in desert environments can be susceptible to low connectivity, genetic isolation, and climate changes. We evaluated the past (1900-1930), recent (1981-2010), and future (2071-2100) climate suitability of the arid Great Basin (USA) for the Columbia spotted frog (Rana luteiventris) and assessed whether changes in surface water may affect connectivity for remaining populations. We developed a predictive model of current climate suitability and used it to predict the historic and future distribution of suitable climates. We then modeled changes in surface water availability at each time period. Finally, we quantified connectivity among existing populations on the basis of hydrology and correlated it with interpopulation genetic distance. We found that the area of the Great Basin with suitable climate conditions has declined by approximately 49% over the last century and will likely continue to decline under future climate scenarios. Climate conditions at currently occupied locations have been relatively stable over the last century, which may explain persistence at these sites. However, future climates at these currently occupied locations are predicted to become warmer throughout the year and drier during the frog's activity period (May - September). Fall and winter precipitation may increase, but as rain instead of snow. Earlier runoff and lower summer base flows may reduce connectivity between neighboring populations, which is already limited. Many of these changes could have negative effects on remaining populations over the next 50-80 years, but milder winters, longer growing seasons, and wetter falls might positively affect survival and dispersal. Collectively, however, seasonal shifts in temperature, precipitation, and stream flow patterns could reduce habitat suitability and connectivity for frogs and possibly other aquatic species inhabiting streams in this arid region.

Persistence at distributional edges: Columbia spotted frog habitat in the arid Great Basin, USA.

A common challenge in the conservation of broadly distributed, yet imperiled species is understanding which factors facilitate persistence at distributional edges, locations where populations are often vulnerable to extirpation due to changes in climate, land use, or distributions of other species. For Columbia spotted frogs (Rana luteiventris) in the Great Basin (USA), a genetically distinct population segment of conservation concern, we approached this problem by examining (1) landscape-scale habitat availability and distribution, (2) water body-scale habitat associations, and (3) resource management-identified threats to persistence. We found that areas with perennial aquatic habitat and suitable climate are extremely limited in the southern portion of the species' range. Within these suitable areas, native and non-native predators (trout and American bullfrogs [Lithobates catesbeianus]) are widespread and may further limit habitat availability in upper- and lower-elevation areas, respectively. At the water body scale, spotted frog occupancy was associated with deeper sites containing abundant emergent vegetation and nontrout fish species. Streams with American beaver (Castor canadensis) frequently had these structural characteristics and were significantly more likely to be occupied than ponds, lakes, streams without beaver, or streams with inactive beaver ponds, highlighting the importance of active manipulation of stream environments by beaver. Native and non-native trout reduced the likelihood of spotted frog occupancy, especially where emergent vegetation cover was sparse. Intensive livestock grazing, low aquatic connectivity, and ephemeral hydroperiods were also negatively associated with spotted frog occupancy. We conclude that persistence of this species at the arid end of its range has been largely facilitated by habitat stability (i.e., permanent hydroperiod), connectivity, predator-free refugia, and a commensalistic interaction with an ecosystem engineer. Beaver-induced changes to habitat quality, stability, and connectivity may increase spotted frog population resistance and resilience to seasonal drought, grazing, non-native predators, and climate change, factors which threaten local or regional persistence.