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In metapopulations, dispersal and population growth rates are influenced by patch quality, spatial structure, and local population density. Recently developed spatial metapopulation models allow inferences about distance effects on dispersal, but these models typically focus on patch occupancy rather than abundance of animals. Spatial occupancy models are useful for studying colonization-extinction dynamics, but richer insights can be gained from estimating abundance and density-dependent demographic rates. We used presence-absence and count data from an 11-year study of a reintroduced metapopulation of federally-threatened Chiricahua leopard frogs (Lithobates chiricahuensis) to develop an integrated abundance-based metapopulation model to draw inferences about the processes contributing to spatiotemporal variation in density. Pond-specific population growth was influenced by pond hydroperiod and frog density, such that permanent and semi-permanent ponds with low densities of adult frogs experienced the highest annual population growth. Immigration rate declined as the distance among ponds increased. Metapopulation-level abundance increased from 2004 until 2015, when it stabilized around 1323 adult frogs (95% CI, 1166–1539). Further, changes in metapopulation abundance were driven mostly by changes in abundance at a few ponds. These high-density populations, which would not have been identifiable with traditional metapopulation models, are likely especially important for species recovery in the area. Our study extends existing statistical models of metapopulation dynamics by focusing on abundance and making it possible to test hypotheses regarding the influence of pond quality and density on local dynamics and colonization.

Habitat loss and degradation are leading causes of biodiversity declines, therefore assessing the capacity of created mitigation wetlands to replace habitat for wildlife has become a management priority. We used single season occupancy models to compare occurrence of larvae of four species of pond-breeding amphibians in wetlands created for mitigation, wetlands impacted by road construction, and unimpacted reference wetlands along a highway corridor in the Greater Yellowstone Ecosystem, U.S.A. Created wetlands were shallow and had less aquatic vegetation and surface area than impacted and reference wetlands. Occupancy of barred tiger salamander (Ambystoma mavortium) and boreal chorus frog (Pseudacris maculata) larvae was similar across wetland types, whereas boreal toads (Anaxyrus boreas) occurred more often in created wetlands than reference and impacted wetlands. However, the majority of created wetlands (> 80%) dried partially or completely before amphibian metamorphosis occurred in both years of our study, resulting in heavy mortality of larvae and, we suspect, little to no recruitment. Columbia spotted frogs (Rana luteiventris), which require emergent vegetation that is not common in newly-created wetlands, occurred commonly in impacted and reference wetlands but were found in only one created wetland. Our results show that shallow created wetlands with little aquatic vegetation may be attractive breeding areas for some amphibians, but may result in high mortality and little recruitment if they fail to hold water for the entire larval period.

Success in reintroducing amphibians may be more context- than detail-dependent such that a slavish adherence to protocol may not foster success better than a more intuitive approach. We provide two reintroduction case studies for boreal toads where the approach was different, but where both resulted in gains in understanding, including first estimates of survival for boreal toads from a reintroduced population. Given the effects of disease on amphibian populations and the potential for disease to remain in a system after extirpation, there is a need to restructure reintroduction guidelines. Maintaining populations on the landscape through reintroductions provides an opportunity for the development of resistance and may facilitate species persistence into the future. But to be effective, care in understanding the context of the reintroduction and a re-envisioning of guidelines is necessary.

Numerous papers have highlighted the need to integrate amphibian research and conservation across multiple scales. Despite this, most amphibian movement studies focus on a single level of organization (e.g., local population) and a single life stage (e.g., adults) and many suggest potential conservation actions or imply that the information is useful to conservation, yet these presumptions are rarely clarified or tested. Movement studies to date provide little information to guide conservation decisions directly because they fail to integrate movement across scales with individual or population parameters (i.e., fitness metrics); this is exacerbated by a general failure to set movement studies in a probabilistic context. An integrative approach allows prediction of population or metapopulation responses to environmental changes and different management actions, thus directly informing conservation decisions and ‘moving the needle’ towards an informed application of conservation actions. To support this perspective we: 1) revisit reviews of amphibian movement to illustrate the focus on single scales and to underscore the importance of movement – at all scales – to conservation; 2) make the case that movement, breeding, and other demographic probabilities are intertwined and studies executed at different temporal and spatial scales can aid in understanding species’ responses to varying environmental and/or management conditions; 3) identify limitations of existing movement-related research to predict conservation action outcomes and inform decision-making; and 4) highlight under-utilized quantitative approaches that facilitate research that either connects movement to fitness metrics (individual-level studies) or estimates population and metapopulation vital rates in addition to, or associated with, movement probabilities.

Successfully addressing complex conservation problems requires attention to pattern and process at multiple spatial scales. This is challenging from a logistical and organizational perspective. In response to indications of worldwide declines in amphibian populations, the Amphibian Research and Monitoring Initiative (ARMI) of the U.S. Geological Survey was established in 2000. This national program is unique in its structure, organization, and success in integrating information at multiple scales. ARMI works under the principle that a good study design is tailored to specific questions, but stipulates the use of methods that result in unbiased parameter estimates (e.g., occupancy). This allows studies to be designed to address local questions but also to produce data that can easily be scaled up to accomplish the objectives of a broad-scale monitoring program. Here we describe how the implementation of the Amphibian Research and Monitoring Initiative results in research that is applicable across scales – global, in contributing to the understanding of amphibian decline phenomena; continental, in synthesizing local data to understand large-scale drivers; regional, by characterizing threats and assessing status of species at the range scale; and local, by working with National Park, Wildlife Refuge, and other Federal and State land managers to identify research needs and serve conservation-relevant research results to inform management decisions.

Despite increasing interest in determining the population-level effects of emerging infectious diseases on wildlife, estimating effects of disease on survival rates remains difficult. Even for a well-studied disease such as amphibian chytridiomycosis (caused by the fungus Batrachochytrium dendrobatidis [Bd]), there are few estimates of how survival of wild hosts is affected. We applied hierarchical models to long-term capture-mark-recapture data (mean = 10.6 yrs, range = 6?15 yrs) from >5500 uniquely-marked individuals to estimate the effect of Bd on apparent survival of four threatened or endangered ranid frog species (Rana draytonii, R. muscosa, R. pretiosa, R. sierrae) at 14 study sites in California and Oregon (USA) and one bufonid toad (Anaxyrus boreas) at two study sites in Wyoming and Montana. Our models indicated that the presence of Bd on an individual reduced apparent survival of ranid frogs by ~6?15% depending on species and sex. The estimated difference between toads with and without Bd was 19% for the Montana population and 55% for the Wyoming population; however, the 95% Credible Interval of these estimates included zero. These results provide evidence for negative effects of Bd on survival in wild populations even in the absence of obvious die-offs. Determining what factors influence the magnitude of the effects of Bd on wildlife populations is an important next step toward identifying management actions. These estimates of Bd effects are important for understanding the extent and severity of disease, whether disease effects have changed over time, and for informing management actions.

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 theWestern 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 (https://0.82 [95% CI: 0.56-0.94]) as seining (https://0.83 [0.46-0.96]) and much higher
detection than dip-netting (https://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.

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) were low for both techniques (rs range = https://0.22 to 0.62), but improved for inundation areas > 900 m2 (rs range = https://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 < 1 Landsat pixel.

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 that are 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 years of data on the federally-threatened Chiricahua leopard frog 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 <8% for all scenarios, but uncertainty was high. Without pond restoration, extinction risk is forecasted to be 5.6% (95% CI: 0?60%) by year 2060. Restoring six ponds by increasing hydroperiod reduced extinction risk to 1.0% (0 ? 11%) in year 2060. We found little evidence that drought influences metapopulation viability when managers have the ability to maintain ponds that hold water throughout the year and are free of invasive species. Our study illustrates the utility of the spatially explicit statistical forecasting approach to MPVA in conservation planning efforts.

Relocations are increasingly popular among wildlife managers despite often low rates of relocation success in vertebrates. In this context, understanding the influence of extrinsic (e.g., relocation design, habitat characteristics) and intrinsic factors (e.g., age and sex) on demographic parameters, such as survival, that regulate the dynamics of relocated populations is critical to improve relocation protocols and better predict relocation success. We investigated survival rates in naturally established and relocated populations of yellow-bellied toads (Bombina variegata), an amphibian that was nearly extinct in Belgium by the late 1980s. We quantified survival at three ontogenetic stages (juvenile, subadult, and adult) in the relocated population, the source population, and a control population. In the relocated population, we quantified survival in captive bred individuals and their locally born descendants. Then, using simulations, we examined how survival cost to relocation affects the self-sustainability of the relocated population. We showed that survival at juvenile and subadult stages was relatively similar in all populations. In contrast, relocated adult survival was lower than adult survival in the source and control populations. Despite this, offspring of relocated animals (the next generation, regardless of life stage) survived at similar rates to individuals in the source and control populations. Simulations revealed that the relocated population was self-sustaining under different scenarios and that the fate (e.g., stability or increase) of the simulated populations was highly dependent on the fecundity of relocated adults and their offspring. our results indicate that survival in relocated individuals is lower than in non-relocated individuals but that this cost (i.e., reduced survival) disappears in the second generation. A finer understanding of how relocation affects demographic processes is an important step in improving relocation success of amphibians and other animals.

Research is necessary to identify patterns in nature, to understand how a system functions, and to make predictions about the future state of an ecosystem. Applied research in conservation biology can identify effective strategies to maintain biodiversity, though many papers end with the conclusion that more research is needed. However, more research does not necessarily lead to solutions. We use the ongoing global decline of amphibians as a salient example to highlight limitations in current conservation research, and to focus on finding solutions which are directly relevant for conservation. While research has been conducted since declines were first detected in the 1990s, outside a few specific examples, little progress in conservation has been achieved. We suggest that the case of amphibian declines is relevant to conservation science in general, as the current paradigm for conservation is that management is planned after research is completed; research and management are not effectively (and not directly) connected. This disconnect illustrates the knowledge-action divide which has been noted recently as a serious deficiency in conservation. Accordingly, we use this introductory paper to the Special Issue (Amphibian conservation in the Anthropocene: Progress and challenges) to describe amphibians as a conservation dilemma, and to make the case for a different, more pragmatic, and more solutions-focused, view of conservation research.

Understanding how natural and anthropogenic processes affect population dynamics of species with patchy distributions is critical to predicting their responses to environmental changes. Despite considerable evidence that demographic rates and dispersal patterns vary temporally in response to an array of biotic and abiotic processes, few applications of metapopulation theory have sought to explore factors that explain spatio-temporal variation in extinction or colonization rates. To facilitate exploring these factors, we extended a spatially explicit model of metapopulation dynamics to create a framework that requires only binary presence-absence data, makes few assumptions about the dispersal process, and accounts for imperfect detection. We apply this framework to 22 years of biannual survey data for lowland leopard frogs, Lithobates yavapaiensis, an amphibian that inhabits arid stream systems in the southwestern U.S. and northern Mexico. Our results highlight the importance of accounting for factors that govern temporal variation in transition probabilities, as both extinction and colonization rates varied with hydrologic conditions. Specifically, local extinctions were more frequent during drought periods, particularly at sites without reliable surface water. Colonization rates increased when larval and dispersal periods were wetter than normal, which increased the probability that potential emigrants metamorphosed and reached neighboring sites. Extirpation of frogs from one watershed during a period of severe drought demonstrated the influence of site-level features, as frogs persisted only in areas where most sites held water consistently and where the amount of sediment deposited from high-elevation wildfires was low. Application of our model provided novel insights into how climate-related processes affected the distribution and population dynamics of an arid-land amphibian. The approach we describe has application to a wide array of species that inhabit patchy environments, can improve our understanding of factors that govern metapopulation dynamics, and can inform strategies for conservation of imperiled species.

A research priority can be defined as a knowledge gap that, if resolved, identifies the optimal course of conservation action. We (a group of geographically distributed and multidisciplinary research scientists) used tools from nominal group theory and decision analysis to collaboratively identify and prioritize information needs within the context of disease-associated amphibian decline, in order to develop a strategy that would support US management agency needs. We developed iterated influence diagrams to create and assess a unified research strategy. We illustrated a transparent process for identifying specific knowledge gaps in amphibian disease ecology relevant to environmental management, and then constructed a research plan to address these uncertainties. The resulting priorities include a need to: (1) understand the drivers of the community-disease relationship, (2) determine the mechanisms by which exposure to contaminants influence disease outcomes, (3) identify elements of terrestrial and aquatic habitats that stabilize host-pathogen dynamics, (4) discuss how metapopulations may be managed to reduce the speed and intensity of disease outbreaks, and (5) define the relationship between habitat management and the environmental and host microbiomes. Along with identifying research priorities for disease management, we present the details of the process used to develop a consensus plan for addressing disease-related declines in amphibians on federally managed lands of the United States.

USGS and Iowa State University scientists quantified Northern Leopard frog exposure to pesticides in aquatic and terrestrial habitats using a novel combination of radio telemetry and passive sampling techniques to better understand factors affecting frog health and survival in agricultural landscapes. The results of this newly published research can inform conservation strategies by providing information about when and where the frogs are most likely exposed to pesticides. This unique approach can be utilized in other land-use settings and with other amphibian species to better understand if contaminant exposure affects growth, development, fitness, and survival.

Full article is available here: https://www.nature.com/articles/s41598-018-28132-3

Movement of individuals has been described as one of the best studied, but least understood concepts in ecology. The magnitude of movements, routes, and probability of movement, has significant application to conservation. Information about movement can inform efforts to model species persistence and is particularly applicable in situations where specific threats (e.g., disease) may depend on the movement of hosts and potential vectors. We estimated the probability of movement (breeding dispersal and permanent emigration) in a metapopulation of 16 breeding sites for boreal toads (Anaxyrus boreas boreas). We used a multi-state mark-recapture approach unique in its complexity (16 sites over 18 years) to address questions related to these movements and variation in resident survival. We found that individuals had a 1-2% probability of dispersing in a particular year and that approximately 10-20% of marked individuals were transient and observed in the metapopulation only once. Resident survival probabilities differed by season, with 71-90% survival from emergence from hibernation through early post-breeding and > 97% survival from mid/late active season through hibernation. Movement-related probabilities are needed to predict species range expansions and contractions, estimate population and metapopulation dynamics, understand host-pathogen and native-invasive species interactions, and to evaluate the relative effects of proposed management actions.

Invasive species are a major threat to the persistence of native species, particularly in systems where ephemeral aquatic habitats have been converted to or replaced by permanent water and predators such as fish have been introduced. Within the Altar Valley, Arizona, USA, the invasive American bullfrog (Lithobates [=Rana] catesbeianus) has been successfully eradicated to help recover Chiricahua leopard frogs (Lithobates chiricahuensis). However, other non-native predators including sunfish (Lepomis spp) 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 may be able to co-occur with this non-native predator. A better understanding of rare situations of co-occurrence with non-native 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 complete eradication of non-native predators is infeasible.

Identifying population declines before they reach crisis proportions is imperative given the current global decline in vertebrate fauna and the associated challenges and expense of recovery. Understanding life-histories and how the environment influences demography are critical aspects of this challenge, as is determining the biological relevance of covariates that are best supported by data. We used 29 years of data on chorus frogs at two sites to estimate demographic parameters, examine life-history, assess weather-related covariates, and determine the magnitude of process variation in target parameters. Average estimates of survival probabilities were https://0.51 (SE=0.04) and https://0.43 (SE=0.04), and average estimates of recruitment probabilities were https://0.64 (SE=0.07) and https://0.44 (SE=0.04). Process variation accounted for &#61619; 76% of the total temporal variation in both parameters at one pond and in survival probability alone at the other, suggesting that the covariates in our top models were explaining predominantly process rather than sampling variation. Estimates of population growth rates indicated a declining population at one pond (i.e., negative population growth rates in 15 of 18 years) and comparisons with historical estimates suggested declines in survival probability at the other. The amount of deviance explained was low, providing little support for the influence of covariates on target parameters, despite model selection support. Synthesis and applications: This analysis illustrates the value of disentangling components of variance when assessing demographic drivers and highlights the need for adequate demographic information in assigning conservation labels.

We quantified the response of amphibian communities to climatic variability across the United States and Canada using more than 500,000 observations for 81 species across 86 study areas. We estimated the relationships between annual variation in climate variables and local colonization and persistence probabilities across more than 5000 surveyed sites. This allowed us to estimate sensitivity to change in five climate variables. Climate sensitivity differs greatly among eco-regions and depends on local climate, species life-history, and phylogeny. Local species richness was especially sensitive to changes in water availability during breeding and changes in winter temperature. These results allowed us to ask whether changing climate explains strong overall rates of decline in species richness observed in our data set. We found that recent change in the climate variables we measured does not explain why North American amphibian richness is rapidly declining, but does explain why some populations decline faster than others.&#8195;