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Many aquatic species in the arid USA-Mexico borderlands region are imperiled, but limited information on distributions and threats often hinders management. To provide information on the distribution of the Western Tiger Salamander (Ambystoma mavortium), including the USA-federally endangered Sonoran Tiger Salamander (Ambystoma mavortium stebbinsi), we used traditional (seines, dip-nets) and modern (environmental DNA [eDNA]) methods to sample 91 waterbodies in northern Sonora, Mexico, during 2015-2018. The endemic Sonoran Tiger Salamander is threatened by introgressive hybridization and potential replacement by another sub-species of the Western Tiger Salamander, the non-native Barred Tiger Salamander (A. m. mavortium). Based on occupancy models that accounted for imperfect detection, eDNA sampling provided a similar detection probability (0.82 [95% CI: 0.56-0.94]) as seining (0.83 [0.46-0.96]) and much higher detection than dip-netting (0.09 [0.02-0.23]). Volume of water filtered had little effect on detection, possibly because turbid sites had greater densities of salamanders. Salamanders were estimated to occur at 51 sites in 3 river drainages in Sonora. These results indicate tiger salamanders are much more widespread in northern Sonora than previously documented, perhaps aided by changes in land and water management practices. However, because the two subspecies of salamanders cannot be reliably distinguished based on morphology or eDNA methods that are based on mitochondrial DNA, we are uncertain if we detected only native genotypes or if we documented recent invasion of the area by the non-native sub-species. Thus, there is an urgent need for methods to reliably distinguish the subspecies so managers can identify appropriate interventions.

Small waterbodies are numerically dominant in many landscapes and provide several important ecosystem services, but automated measurement of waterbodies smaller than a standard Landsat pixel (0.09 ha) remains challenging. To further evaluate sub‐Landsat pixel techniques for estimating inundation extent of small waterbodies (basin area: 0.06–1.79 ha), we used a partial spectral unmixing method with matched filtering applied to September 1985–2018 Landsat 5 and eight imagery from southern Arizona, USA. We estimated trends in modeled surface water area each September and evaluated the ability of several common drought indices to explain variation in mean water area. Our methods accurately classified waterbodies as dry or inundated (Landsat 5: 91.3%; Landsat 8: 98.9%) and modeled and digitized surface water areas were strongly correlated (R2 = 0.70–0.92; bias = −0.024 to −0.015 ha). Estimated surface water area was best explained by the 3‐month seasonal standardized precipitation index (SPI03; July‒September). We found a wide range of estimated relationships between drought indices (e.g. SPI vs. Palmer Drought Severity Index) and estimated water area, even for different durations of the same drought index (e.g. SPI01 vs. SPI12). Mean waterbody surface area decreased by ~14% from September 1985 to September 2018, which matches declines in local annual precipitation and regional trends of reduced inundation extent of larger waterbodies. These results emphasize the importance of understanding local systems when relying on drought indices to infer variation in past or future surface water dynamics. Several challenges remain before widespread application of sub‐pixel methods is feasible, but our results provide further evidence that partial spectral unmixing with matched filtering provides reliable measures of inundation extent of small waterbodies.

For species with variable phenology, it is often challenging to produce reliable estimates of population dynamics or changes in occupancy. The Arizona Toad (Anaxyrus microscaphus) is a southwestern USA endemic that has been petitioned for legal protection, but status assessments are limited by a lack of information on population trends. Also, timing and consistency of Arizona Toad breeding varies greatly, making it difficult to predict optimal survey times or effort required for detection. To help fill these information gaps, we conducted breeding season call surveys during 2013–2016 and 2019 at 86 historically occupied sites and 59 control sites across the species' range in New Mexico. We estimated variation in mean dates of arrival and departure from breeding sites, changes in occupancy, and site-level extinction since 1959 with recently developed multi-season staggered-entry models, which relax the within-season closure assumption common to most occupancy models. Optimal timing of surveys in our study areas was approximately 5–30 March. Averaged across years, estimated probability of occupancy was 0.58 (SE = 0.09) for historical sites and 0.19 (SE = 0.08) for control sites. Occupancy increased from 2013 through 2019. Notably, even though observer error was trivial, annual detection probabilities varied from 0.23 to 0.75 and declined during the study; this means naive occupancy values would have been misleading, indicating apparent declines in toad occupancy. Occupancy was lowest during the first year of the study, possibly due to changes in stream flows and conditions in many waterbodies following extended drought and recent wildfires. Although within-season closure was violated by variable calling phenology, simple multi-season models provided nearly identical estimates as staggered-entry models. Surprisingly, extinction probability was unrelated to the number of years since the first or last record at historically occupied sites. Collectively, our results suggest a lack of large, recent declines in occupancy by Arizona Toads in New Mexico, but we still lack population information from most of the species' range.

Many natural wetlands around the world have disappeared or been replaced, resulting in the dependence of many wildlife species on small, artificial earthen stock ponds. These ponds provide critical wildlife habitat, such that the accurate detection of water and assessment of inundation extent is required. We applied a full (linear spectral mixture analysis; LSMA) and partial (matched filtering; MF) spectral unmixing algorithm to a 2007 Landsat 5 and a 2014 Landsat 8 satellite image to determine the ability of a time-intensive (i.e., more spectral input; LSMA) vs. a more efficient (less spectral input; MF) spectral unmixing approach to detect and estimate surface water area of stock ponds in southern Arizona, USA and northern Sonora, Mexico. Spearman rank correlations (rs) between modeled and actual inundation areas less than a single Landsat pixel ( 900 m2 (rs range = 0.34 to 0.70). Our results demonstrate that the MF approach can model ranked inundation extent of known pond locations with results comparable to or better than LSMA, but further refinement is required for estimating absolute inundation areas and mapping wetlands

First-order dynamic occupancy models (FODOMs) are a class of state-space model in which the true state (occurrence) is observed imperfectly. An important assumption of FODOMs is that site dynamics only depend on the current state and that variations in dynamic processes are adequately captured with covariates or random effects. However, it is often difficult to understand and/or measure the covariates that generate ecological data, which are typically spatio-temporally correlated. Consequently, the non-independent error structure of correlated data causes underestimation of parameter uncertainty and poor ecological inference. Here, we extend the FODOM framework with a second-order Markov process to accommodate site memory when covariates are not available. Our modeling framework can be used to make reliable inference about site occupancy, colonization, extinction, turnover, and detection probabilities. We present a series of simulations to illustrate the data requirements and model performance. We then applied our modeling framework to 13 years of data from an amphibian community in southern Arizona, USA. In this analysis, we found residual temporal autocorrelation of population processes for most species, even after accounting for long-term drought dynamics. Our approach represents a valuable advance in obtaining inference on population dynamics, especially as they relate to metapopulations.

American bullfrogs (Lithobates catesbeianus) are significant invaders in many places and can negatively impact native species. Despite their impact and wide distribution, little is known about their demography. We used five years of capture mark-recapture data to estimate annual apparent survival of post-metamorphic bullfrogs in a population on the Buenos Aires National Wildlife Refuge in their invaded range in Arizona, U.S.A. This population is a potential source of colonists into breeding ponds used by the federally threatened Chiricahua leopard frog (L. chiricahuensis). Results from robust-design Cormack-Jolly-Seber models suggested that survival of bullfrogs was influenced by sex and precipitation but not body condition. Survival was higher for females (mean = 0.37; 95% , 0.72) than males (mean = 0.17; 95% , 0.49), and declined with reduced annual precipitation (mean = −0.36, 95% = −2.09, 0.84). These survival estimates can be incorporated into models of population dynamics and to help predict spread of bullfrogs.

Understanding the additive or interactive threats of habitat transformation and invasive species is critical for conservation, especially where climate change is expected to increase the severity or frequency of drought. In the arid southwestern USA, this combination of stressors has caused widespread declines of native aquatic and semi-aquatic species. Achieving resilience to drought and other effects of climate change may depend upon continued management, so understanding the combined effects of stressors is important. We used Bayesian hierarchical models fitted with 10-years of pond-based monitoring surveys for the federally-endangered Sonoran Tiger Salamander (Ambystoma mavortium stebbinsi) and invasive predators (fishes and American Bullfrogs, Lithobates catesbeianus) that threaten native species. We estimated trends in occupancy of salamanders and invasive predators while accounting for hydrological dynamics of ponds, then used a two-species interaction model to directly estimate how invasive predators affected salamander occupancy. We also tested a conceptual model that predicted that drought, by limiting the distribution of invasive predators, could ultimately benefit native species. Even though occupancy of invasive predators was stationary and their presence in a pond reduced the probability of salamander presence by 23%, occupancy of Sonoran Tiger Salamanders increased, annually, by 2.2%. Occupancy of salamanders and invasive predators both declined dramatically following the 5th consecutive year of drought. Salamander occupancy recovered quickly after return to non-drought conditions, while occupancy of invasive predators remained suppressed. Models that incorporated three time-lagged periods (1 to 4 years) of local moisture conditions confirmed that salamanders and invasive predators responded differently to drought, reflecting how life-history strategies shape responses to disturbances. The positive 10-year trend in salamander occupancy and their rapid recovery after drought provided partial support for the hypothesis of drought-mediated coexistence with invasive predators. These results also suggest management opportunities for conservation of the Sonoran Tiger Salamander and other imperiled organisms in human-transformed landscapes.

Global amphibian declines have been attributed to a number of factors including disease, invasive species, habitat degradation, and climate change. Reintroduction is one management action that is commonly used with the goal of recovering imperiled species. The success of reintroductions varies widely, and evaluating their efficacy requires estimates of population viability metrics, such as underlying vital rates and trends in abundance. Although rarely quantified, assessing vital rates for recovering populations provides a more mechanistic understanding of population growth than numerical trends in population occupancy or abundance. We used three years of capture-mark-recapture data from three breeding ponds and a Cormack-Jolly-Seber model to estimate annual apparent survival for reintroduced populations of the federally threatened Chiricahua Leopard Frog (Lithobates chiricahuensis) at the Buenos Aires National Wildlife Refuge (BANWR), in the Altar Valley, Arizona, USA. To place our results in context, we...

Conservation of at-risk species is aided by reliable forecasts of the consequences of environmental change and management actions on population viability. Forecasts from conventional population viability analysis (PVA) are made using a two-step procedure in which parameters are estimated, or elicited from expert opinion, and then plugged into a stochastic population model without accounting for parameter uncertainty. Recently developed statistical PVAs differ because forecasts are made conditional on models fitted to empirical data. The statistical forecasting approach allows for uncertainty about parameters, but it has rarely been applied in metapopulation contexts where spatially explicit inference is needed about colonization and extinction dynamics and other forms of stochasticity that influence metapopulation viability. We conducted a statistical metapopulation viability analysis (MPVA) using 11 yr of data on the federally threatened Chiricahua leopard frog (Lithobates chiricahuensis) to forecast responses to landscape heterogeneity, drought, environmental stochasticity, and management. We evaluated several future environmental scenarios and pond restoration options designed to reduce extinction risk. Forecasts over a 50-yr time horizon indicated that metapopulation extinction risk was

Managing species with intensive tools such as reintroduction may focus on single sites or entire landscapes. For vagile species, long-term persistence will require colonization and establishment in neighboring habitats. Therefore, both suitable colonization sites and suitable dispersal corridors between sites are required. Assessment of landscapes for both requirements can contribute to ranking and selection of reintroduction areas, thereby improving management success. Following eradication of invasive American Bullfrogs (Lithobates catesbeianus) from most of Buenos Aires National Wildlife Refuge (BANWR; Arizona, United States), larval Chiricahua Leopard Frogs (Lithobates chiricahuensis) from a private pond were reintroduced into three stock ponds. Populations became established at all three reintroduction sites followed by colonization of neighboring ponds in subsequent years. Our aim was to better understand colonization patterns by the federally threatened L. chiricahuensis which could help i...

Invasive species are a major threat to the persistence of native species, particularly in systems where ephemeral aquatic habitats have been replaced by permanent water and predators, such as fish, have been introduced. Within the Altar Valley, Arizona, the invasive American bullfrog, Lithobates catesbeianus (formerly Rana catesbeianus), has been successfully eradicated to help recover Chiricahua leopard frogs (Lithobates chiricahuensis). However, other nonnative predators including sunfish (Lepomis) are present in some permanent water bodies. During four consecutive years (2014–2017), we detected both the federally threatened Chiricahua leopard frog and sunfish at one permanent water body in the Altar Valley. This suggests that despite the potential negative effect of predatory fish on amphibians, there may be conditions where the Chiricahua leopard frog can co-occur with this nonnative predator. A better understanding of rare situations of co-occurrence with nonnative predators may contribute to our understanding of why co-occurrence happens in some, but not all, systems and whether conservation strategies can be developed in situations where eradication of nonnative predators is infeasible.

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. John Wesley Powell Center for Analysis and Synthesis - US Geological Survey U.S. Geological Survey-Amphibian Research and Monitoring Initiative (ARMI) U.S. Fish and Wildlife Service National Park Service U.S. Forest Service National Science Foundation [DEB-0841758, DEB-1149308] National Institutes of Health [R01GM109499] National Geographic Society Morris Animal Foundation David and Lucille Packard Foundation This work was conducted as part of the Amphibian Decline Working Group supported by the John Wesley Powell Center for Analysis and Synthesis, funded by the US Geological Survey. Funding and logistical support for field data collection came from a range of sources including the U.S. Geological Survey-Amphibian Research and Monitoring Initiative (ARMI), U.S. Fish and Wildlife Service, National Park Service, U.S. Forest Service, National Science Foundation (DEB-0841758, DEB-1149308), National Institutes of Health (R01GM109499), National Geographic Society, Morris Animal Foundation, and David and Lucille Packard Foundation. Data are deposited at the U.S. Geological Survey's John Wesley Powell Center for Analysis and Synthesis. This manuscript is contribution 654 of USGS ARMI. Any use of trade, firm, or product names is for descriptive purposes only and does not imply endorsement by the U.S. Government.

Summary 1. The reintroduction of a species into its historic range is a critical component of conservation programmes designed to restore extirpated metapopulations. However, many reintroduction efforts fail, and the lack of rigorous monitoring programmes and statistical models have prevented a general understanding of the factors affecting metapopulation viability following reintroduction. 2. Spatially explicit metapopulation theory provides the basis for understanding the dynamics of fragmented populations linked by dispersal, but the theory has rarely been used to guide reintroduction programmes because most spatial metapopulation models require presence– absence data from every site in the network, and they do not allow for observation error such as imperfect detection. 3. We develop a spatial occupancy model that relaxes these restrictive assumptions and allows for inference about metapopulation extinction risk and connectivity. We demonstrate the utility of the model using six years of data on the Chiricahua leopard frog Lithobates chiricahuensis, a threatened desert-breeding amphibian that was reintroduced to a network of sites in Arizona USA in 2003. 4. Our results indicate that the model can generate precise predictions of extinction risk and produce connectivity maps that can guide conservation efforts following reintroduction. In the case of L. chiricahuensis, many sites were functionally isolated, and 82% of sites were characterized by intermittent water availability and high local extinction probabilities (0� 84, 95% CI: 0� 64–0� 99). However, under the current hydrological conditions and spatial arrangement of sites, the risk of metapopulation extinction is estimated to be

Metapopulation ecology and landscape ecology aim to understand how spatial structure influences ecological processes, yet these disciplines address the problem using fundamentally different modeling approaches. Metapopulation models describe how the spatial distribution of patches affects colonization and extinction, but often do not account for the heterogeneity in the landscape between patches. Models in landscape ecology use detailed descriptions of landscape structure, but often without considering colonization and extinction dynamics. We present a novel spatially explicit modeling framework for narrowing the divide between these disciplines to advance understanding of the effects of landscape structure on metapopulation dynamics. Unlike previous efforts, this framework allows for statistical inference on landscape resistance to colonization using empirical data. We demonstrate the approach using 11 yr of data on a threatened amphibian in a desert ecosystem. Occupancy data for Lithobates chiricahuensis (Chiricahua leopard frog) were collected on the Buenos Aires National Wildlife Refuge (BANWR), Arizona, USA from 2007 to 2017 following a reintroduction in 2003. Results indicated that colonization dynamics were influenced by both patch characteristics and landscape structure. Landscape resistance increased with increasing elevation and distance to the nearest streambed. Colonization rate was also influenced by patch quality, with semi-permanent and permanent ponds contributing substantially more to the colonization of neighboring ponds relative to intermittent ponds. Ponds that only hold water intermittently also had the highest extinction rate. Our modeling framework can be widely applied to understand metapopulation dynamics in complex landscapes, particularly in systems in which the environment between habitat patches influences the colonization process.

The salamander chytrid fungus (Batrachochytrium salamandrivorans [Bsal]) is causing massive mortality of salamanders in Europe. The potential for spread via international trade into North America and the high diversity of salamanders has catalyzed concern about Bsal in the U.S. Surveillance programs for invading pathogens must initially meet challenges that include low rates of occurrence on the landscape, low prevalence at a site, and imperfect detection of the diagnostic tests. We implemented a large-scale survey to determine if Bsal was present in North America designed to target taxa and localities where Bsal was determined highest risk to be present based on species susceptibility and geography. Our analysis included a Bayesian model to estimate the probability of occurrence of Bsal given our prior knowledge of the occurrence and prevalence of the pathogen. We failed to detect Bsal in any of 11,189 samples from 594 sites in 223 counties within 35 U.S. states and one site in Mexico. Our modeling indicates that Bsal is highly unlikely to occur within wild amphibians in the U.S. and suggests that the best proactive response is to continue mitigation efforts against the introduction and establishment of the disease and to develop plans to reduce impacts should Bsal establish.