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Influence of demography and environment on persistence in toad populations

Effective conservation of rare species requires an understanding of how potential threats affect population dynamics. Unfortunately, information about population demographics prior to threats (i.e., baseline data) is lacking for many species. Perturbations, caused by climate change, disease, or other stressors can lead to population declines and heightened conservation concerns. Boreal toads ( Anaxyrus boreas boreas ) have undergone rangewide declines due mostly to the amphibian chytrid fungus Batrachochytrium dendrobatidis (Bd), with only a few sizable populations remaining in the southern Rocky Mountains, USA, that are disease-free. Despite the apparent region-wide occurrence of Bd, our focal populations in central Colorado were disease free over a 14-year capture-mark-recapture study until the recent discovery of Bd at one of the sites. We used recapture data and the Pradel reverse-time model to assess the influence of environmental and site-specific conditions on survival and recruitment. We then forecast changes in the toad populations with 2 growth models; one using an average lambda value to initiate the projection, and one using the most recent value to capture potential effects of the incursion of disease into the system. Adult survival was consistently high at the 3 sites, whereas recruitment was more variable and markedly low at 1 site. We found that active season moisture, active season length, and breeding shallows were important factors in estimating recruitment. Population growth models indicated a slight increase at 1 site but decreasing trends at the 2 other sites, possibly influenced by low recruitment. Insight into declining species management can be gained from information on survival and recruitment and how site-specific environmental factors influence these demographic parameters. Our data are particularly useful because they provide baseline data on demographics in populations before a disease outbreak and enhance our ability to detect changes in population parameters potentially caused by the disease.

Journal of Wildlife Management

Enigmatic near‐extirpation in a boreal toad metapopulation in northwestern Montana

North America's protected lands harbor biodiversity and provide habitats where species threatened by a variety of stressors in other environments can thrive. Yet disease, climate change, and other threats are not limited by land management boundaries and can interact with conditions within protected landscapes to affect sensitive populations. We examined the population dynamics of a boreal toad ( Anaxyrus boreas boreas ) metapopulation at a wildlife refuge in northwestern Montana, USA, over a 16‐year period (2003–2018). We used robust design capture‐recapture models to estimate male population size, recruitment, and apparent survival over time and in relation to the amphibian chytrid fungus ( Batrachochytrium dendrobatidis ). We estimated female population size in years with sufficient captures. Finally, we examined trends in male and female toad body size and condition. We found no evidence of an effect of disease or time on male toad survival but detected a strong negative trend in recruitment of new males to the population. Estimates of male and female abundance decreased over time. Body size of males and females was inversely related to estimated population size, consistent with reduced recruitment to replace adults, but body condition of adult males was only weakly associated with abundance. Together, these results describe the demography of a near‐extirpation event, and point to dramatic decreases in the recruitment of new individuals to the breeding population as the cause of this decline. We surmise that processes related to the restoration of historical hydrology within the refuge adversely affected amphibian breeding habitat, and that these changes interacted with disease, life history, and other factors to restrict the recruitment of new individuals to the breeding population over time. Our results point to challenges in understanding and predicting factors that influence population change and highlight that current metrics for assessing population status can have limited predictive ability. Published 2021. This article is a U.S. Government work and is in the public domain in the USA.

Montana

Distribution and pathogenicity of Batrachochytrium dendrobatidis in boreal toads from the grand teton area of western wyoming

The pathogen Batrachochytrium dendrobatidis (Bd), which causes the skin disease chytridiomycosis, has been linked to amphibian population declines and extinctions worldwide. Bd has been implicated in recent declines of boreal toads, Bufo boreas boreas, in Colorado but populations of boreal toads in western Wyoming have high prevalence of Bd without suffering catastrophic mortality. In a field and laboratory study, we investigated the prevalence of Bd in boreal toads from the Grand Teton ecosystem (GRTE) in Wyoming and tested the pathogenicity of Bd to these toads in several environments. The pathogen was present in breeding adults at all 10 sites sampled, with a mean prevalence of 67%. In an experiment with juvenile toadlets housed individually in wet environments, 10 6 zoospores of Bd isolated from GRTE caused lethal disease in all Wyoming and Colorado animals within 35 days. Survival time was longer in toadlets from Wyoming than Colorado and in toadlets spending more time in dry sites. In a second trial involving Colorado toadlets exposed to 35% fewer Bd zoospores, infection peaked and subsided over 68 days with no lethal chytridiomycosis in any treatment. However, compared with drier aquaria with dry refuges, Bd infection intensity was 41% higher in more humid aquaria and 81% higher without dry refuges available. Our findings suggest that although widely infected in nature, Wyoming toads may escape chytridiomycosis due to a slight advantage in innate resistance or because their native habitat hinders Bd growth or provides more opportunities to reduce pathogen loads behaviorally than in Colorado. ?? 2009 International Association for Ecology and Health.

EcoHealth

Disease in a dynamic landscape: host behavior and wildfire reduce amphibian chytrid infection

Disturbances are often expected to magnify effects of disease, but these effects may depend on the ecology, behavior, and life history of both hosts and pathogens. In many ecosystems, wildfire is the dominant natural disturbance and thus could directly or indirectly affect dynamics of many diseases. To determine how probability of infection by the aquatic fungus Batrachochytrium dendrobatidis (Bd) varies relative to habitat use by individuals, wildfire, and host characteristics, we sampled 404 boreal toads ( Anaxyrus boreas boreas ) across Glacier National Park, Montana (USA). Bd causes chytridiomycosis, an emerging infectious disease linked with widespread amphibian declines, including the boreal toad. Probability of infection was similar for females and the combined group of males and juveniles. However, only 9% of terrestrial toads were infected compared to >30% of aquatic toads, and toads captured in recently burned areas were half as likely to be infected as toads in unburned areas. We suspect these large differences in infection reflect habitat choices by individuals that affect pathogen exposure and persistence, especially in burned forests where warm, arid conditions could limit Bd growth. Our results show that natural disturbances such as wildfire and the resulting diverse habitats can influence infection across large landscapes, potentially maintaining local refuges and host behaviors that facilitate evolution of disease resistance.

Montana

Livestock grazing, climatic variation, and breeding phenology jointly shape disease dynamics and survival in a wild amphibian

Wildlife responses to infectious disease can be influenced by environmental stressors that alter host-pathogen dynamics. We investigated how livestock grazing, climatic variation, and breeding phenology influence disease prevalence and annual survival in boreal toad ( Anaxyrus boreas boreas ) populations challenged with Batrachochytrium dendrobatidis ( Bd ), a fungal pathogen implicated in global amphibian declines. We conducted a five-year (2015–2019) capture-recapture study of boreal toads ( n = 1301) inhabiting pastures grazed by cattle in western Wyoming, USA. We employed structural equation models to determine whether the effects of climatic variation on Bd prevalence were direct or mediated through effects on breeding phenology and multi-state models to explore the interplay of grazing, weather, and Bd infection on adult survival. Higher winter snowpack was linked with shorter spring breeding seasons, which were associated with lower Bd prevalence. Boreal toads infected with Bd suffered increased mortality, but only at relatively cool temperatures. Although cattle grazing created warmer microclimates, likely by reducing vegetation cover, grazing-induced habitat changes did not scale up to influence adult survival. Our results suggest that boreal toads in cooler environments face increased risk of disease-induced mortality, possibly because infected individuals are not able to elevate body temperature to reduce or clear infection. More generally, we demonstrate that host-pathogen dynamics can be shaped jointly by independent and interactive effects of livestock grazing, breeding season length, and climatic variation. Future investigations of wildlife responses to disease therefore may benefit from considering anthropogenic land use and climatic regimes, including the effect of weather on host phenology.

Wyoming

Trade-offs in initial and long-term handling efficiency of PIT-tag and photographic identification methods

Individual identification is required for long-term investigations that examine population-level changes in survival or abundance, and mechanisms associated with these changes in wild populations. Such identification generally requires the application of a unique mark, or the documentation of characteristics distinctive to each individual animal. To minimize impacts to often declining populations, scientific and ethical concerns encourage marking strategies that minimize handling time (i.e., stress) for captured individuals. We examined the relative efficacy of passive integrated transponder (PIT)-tagging and photo-identification to identify individual Boreal toads ( Anaxyrus boreas boreas ) in field and indoor settings. We evaluated whether initial handling time was influenced by identification method (PIT-tag or photo-identification) or environment (field or indoor) and assessed the applicability of each method in long-term monitoring programs. Initial handling time was higher for PIT-tagging than photo-identification and higher in the field than in an indoor environment; however, handling time for previously PIT-tagged individuals was greatly reduced such that photo-identification led to > 5.5 times more handling time than PIT-tagging over the course of a toad's lifetime. Investigators must determine the trade-off between initial and subsequent handling times to minimize the expected (mean) cumulative handling time for an individual over the course of a study. Cumulative handling time is a function of the study design and the species’ survival and detection probabilities. We developed a Shiny Application to allow investigators to determine the identification method that minimizes handling time for their own study system.

Ecological Indicators

Integrating biology, field logistics, and simulations to optimize parameter estimation for imperiled species

Conservation of imperiled species often requires knowledge of vital rates and population dynamics. However, these can be difficult to estimate for rare species and small populations. This problem is further exacerbated when individuals are not available for detection during some surveys due to limited access, delaying surveys and creating mismatches between the breeding behavior and survey timing. Here we use simulations to explore the impacts of this issue using four hypothetical boreal toad ( Anaxyrus boreas boreas ) populations, representing combinations of logistical access (accessible, inaccessible) and breeding behavior (synchronous, asynchronous). We examine the bias and precision of survival and breeding probability estimates generated by survey designs that differ in effort and timing for these populations. Our findings indicate that the logistical access of a site and mismatch between the breeding behavior and survey design can greatly limit the ability to yield accurate and precise estimates of survival and breeding probabilities. Simulations similar to what we have performed can help researchers determine an optimal survey design(s) for their system before initiating sampling efforts.

Ecological Modelling

Thermal characteristics of amphibian microhabitats in a fire-disturbed landscape

Disturbance has long been a central issue in amphibian conservation, often regarding negative effects of logging or other forest management activities, but some amphibians seem to prefer disturbed habitats. After documenting increased use of recently burned forests by boreal toads (Bufo boreas), we hypothesized that burned habitats provided improved thermal opportunities in terrestrial habitats. We tested this hypothesis by conducting a radio telemetry study of habitat use (reported previously) and by using physical models that simulated the temperature of adult toads. We deployed 108 physical models in and adjacent to a 1-year old burn using a fully-replicated design with three burn severities (unburned, partial, high severity) and four microhabitats (open surface, under vegetation, under log, in burrow). Model temperatures were compared to a range of preferred temperatures in published studies. We found 70% more observations within the preferred temperature range of B. boreas in forests burned with high severity than in unburned areas. Burned forest was warmer than unburned forest across all microhabitats, but the largest relative difference was in burrows, which averaged 3 ??C warmer in high-severity burn areas and remained warmer though the night. More than twice as many observations were within the preferred temperature range in high-severity burrows than in unburned burrows. Areas burned with high severity were still warmer than unburned forest 3 years after the fire. Habitat use of toads during the concurrent radio telemetry study matched that predicted by the physical models. These results suggest there are fitness-linked benefits to toads using burned habitats, such as increased growth, fertility, and possibly disease resistance. However, increased soil temperatures that result from wildfire may be detrimental to other amphibian species that prefer cooler temperatures and stable environments. More broadly, our data illustrate the use of physical models to measure and interpret changes that amphibians may experience from disturbance, and highlight the need for research linking vital rates such as growth and survival to disturbance.

Forest Ecology and Management

Infection status as the basis for habitat choices in a wild amphibian

Animals challenged with disease may select specific habitat conditions that help prevent or reduce infection. Whereas preinfection avoidance of habitats with a high risk of disease exposure has been documented in both captive and free-ranging animals, evidence of switching habitats after infection to support the clearing of the infection is limited to laboratory experiments. The extent to which wild animals proximately modify habitat choices in response to infection status thus remains unclear. We investigated preinfection behavioral avoidance and postinfection habitat switching using wild, radio-tracked boreal toads ( Anaxyrus boreas boreas ) in a population challenged with Batrachochytrium dendrobatidis ( Bd ), a pathogenic fungus responsible for a catastrophic panzootic affecting hundreds of amphibian species worldwide. Boreal toads did not preemptively avoid microhabitats with conditions conducive to Bd growth. Infected individuals, however, selected warmer, more open habitats, which were associated with elevated body temperature and the subsequent clearing of infection. Our results suggest that disease can comprise an important selective pressure on animal habitat and space use. Habitat selection models, therefore, may be greatly improved by including variables that quantify infection risk and/or the infection status of individuals through time.

Wyoming

Informed breeding dispersal following stochastic changes to patch quality in a pond-breeding amphibian

The unidirectional movement of animals between breeding patches (i.e. breeding dispersal) has profound implications for the ecological and evolutionary dynamics of spatially structured populations. In spatiotemporally variable environments, individuals are expected to adjust their dispersal decisions according to information gathered on the environmental and/or social cues that reflect the fitness prospects in a given breeding patch (i.e. informed dispersal). A paucity of empirical work limited our understanding of the ability of animals to depart from low-quality breeding patches and settle in high-quality breeding patches. We examined the capacity of individuals to respond to stochastic changes in habitat quality via informed breeding dispersal in a pond-breeding amphibian. We conducted a 5-year (2015–2019) capture–recapture study of boreal toads Anaxyrus boreas boreas ( n = 1,100) that breed in beaver ponds in western Wyoming, USA. During early spring of 2017, an extreme flooding event destroyed several beaver dams and resulted in the loss of breeding habitat. We used multi-state models to investigate how temporal changes in pond characteristics influenced breeding dispersal, and determine whether movement decisions were in accordance with prospects for reproductive fitness. Boreal toads more often departed from low-quality breeding ponds (without successful metamorphosis) and settled in high-quality breeding ponds (with successful metamorphosis). Movement decisions were context-dependent and associated with pond characteristics altered by beaver dam destruction. Individuals were more likely to depart from shallow ponds with high vegetation cover and settle in deep ponds with low vegetation cover. The probability of metamorphosis was related to the same environmental cues, suggesting that boreal toads assess the fitness prospects of a breeding patch and adjust movement decisions accordingly (i.e. informed breeding dispersal). We demonstrated that stochastic variability in environmental conditions and habitat quality can underpin dispersal behaviour in amphibians. Our study highlighted the mechanistic linkages between habitat change, movement behaviour and prospects for reproductive performance, which is critical for understanding how wild animals respond to rapid environmental change.

Wyoming

Quantifying intraspecific variation in host resistance ad tolerance to a lethal pathogen

Testing for intraspecific variation for host tolerance or resistance in wild populations is important for informing conservation decisions about captive breeding, translocation, and disease treatment. Here, we test the importance of tolerance and resistance in multiple populations of boreal toads ( Anaxyrus boreas boreas ) against Batrachochytrium dendrobatidis (Bd), the amphibian fungal pathogen responsible for the greatest host biodiversity loss due to disease. Boreal toads have severely declined in Colorado (CO) due to Bd, but toad populations challenged with Bd in western Wyoming (WY) appear to be less affected. We used a common garden infection experiment to expose post-metamorphic toads sourced from four populations (2 in CO and 2 in WY) to Bd and monitored changes in mass, pathogen burden and survival for 8 weeks. We used a multi-state modelling approach to estimate weekly survival and transition probabilities between infected and cleared states, reflecting a dynamic infection process that traditional approaches fail to capture. We found that WY boreal toads are more tolerant to Bd infection with higher survival probabilities than those in CO when infected with identical pathogen burdens. WY toads also appeared more resistant to Bd with a higher probability of infection clearance and an average of 5 days longer to reach peak infection burdens. Our results demonstrate strong intraspecific differences in tolerance and resistance that likely contribute to why population declines vary regionally across this species. Our multi-state framework allowed us to gain inference on typically hidden disease processes when testing for host tolerance or resistance. Our findings demonstrate that describing an entire host species as ‘tolerant’ or ‘resistant’ (or lack thereof) is unwise without testing for intraspecific variation.

Colorado, Wyoming

Compensatory recruitment unlikely in high-elevation amphibian populations challenged with disease

Understanding the causes of population variation in host response to disease, and the mechanisms of persistence, can serve as vital information for species conservation. One such mechanism of population persistence that has gained support is the demographic process of compensatory recruitment. Host populations may persist by increasing recruitment to compensate for reduced survival due to infection, thus limiting the negative effects of the disease on population trajectories. However, high-elevation populations are inherently vulnerable to stochastic processes and may be limited in their ability to exhibit compensatory recruitment relative to lower elevation populations. We use long-term mark–recapture data from five populations of boreal toads Anaxyrus boreas boreas , across an elevational gradient in Colorado, before and after pathogen arrival to assess whether populations can persist with Batrachochytrium dendrobatidis ( Bd ) via compensatory recruitment. Prior to pathogen arrival, we found a life-history trade-off between survival and recruitment across elevations, where high-elevation toads have high survival but lower recruitment and vice versa at lower elevations. Pathogen arrival had a strong negative effect on apparent annual survival and recruitment leading to negative population growth rates and dramatically reduced host abundances. The data did not support the occurrence of compensatory recruitment. Synthesis and applications . Our unique dataset indicates that demographic responses to pathogens may be environmentally (i.e. elevationally) context dependent and highlights the value of long-term monitoring. We recommend that practitioners verify that potential persistence mechanisms occur across multiple populations and relevant environmental gradients to counter any assumptions of the mechanism existing species-wide. Quantifying variation in population responses to disease will aid in understanding the bounds of such persistence mechanisms and identify particularly vulnerable populations where mechanisms are nonexistent.

Journal of Applied Ecology

Optimal management decisions are robust to unknown dynamics in an amphibian metapopulation plagued by disease

Identifying conservation actions to recover threatened species can be challenging due to many ecological uncertainties. For example, major threats to a species' conservation are commonly known or suspected, but the specific impacts on population or metapopulation dynamics can be uncertain. This is frequently the case with emerging infectious diseases, including chytridiomycosis, a global driver of amphibian population declines caused by the fungal pathogens Batrachochytrium dendrobatidis (Bd) and Batrachochytrium salamandrivorans . While these diseases are known to cause amphibian declines and extirpations, the mechanisms of their landscape-scale spread are still largely unknown. Such uncertainty can lead to inaction which may jeopardize timely recovery of a species. Decision analysis is a pragmatic approach to making transparent and defensible decisions while dealing with uncertainties. We investigated whether optimal actions aimed at recovering boreal toad ( Anaxyrus boreas boreas ) metapopulations in the southern Rocky Mountains are robust to the unknown dynamics of Bd spread using value of information and regret analyses. Value of information is a decision-analytic tool for calculating the value of new information in terms of performance on management objectives, while regret measures the cost of acting under incorrect information. We further conducted a stochastic sensitivity analysis to identify the relative effects of metapopulation parameters on system dynamics. We found optimal actions were robust to the unknown dynamics of Bd spread. While boreal toad breeding occurrence is highly sensitive to Bd distribution, the optimal decision is not. Resolving the unknown dynamics of Bd spread would lead to a minimal gain of less than one breeding toad subpopulation at the end of 50 years, given the currently available management actions. Applying a decision-analytic framework coupled with value of information and regret analyses can help frame how uncertainties affect decisions in a way that empowers decision makers.

Animal Conservation

Identifying species conservation strategies to reduce disease-associated declines

Emerging infectious diseases (EIDs) are a salient threat to many animal taxa, causing local and global extinctions, altering communities and ecosystem function. The EID chytridiomycosis is a prominent driver of amphibian declines, which is caused by the fungal pathogen Batrachochytrium dendrobatidis (Bd). To guide conservation policy, we developed a predictive decision-analytic model that combines empirical knowledge of host-pathogen metapopulation dynamics with expert judgment regarding effects of management actions, to select from potential conservation strategies. We apply our approach to a boreal toad ( Anaxyrus boreas boreas ) and Bd system, identifying optimal strategies that balance tradeoffs in maximizing toad population persistence and landscape-level distribution, while considering costs. The most robust strategy is expected to reduce the decline of toad breeding sites from 53% to 21% over 50 years. Our findings are incorporated into management policy to guide conservation planning. Our online modeling application provides a template for managers of other systems challenged by EIDs.

Conservation Letters

Hydrology and geomorphology of the Taiya River near the West Creek Tributary, southeast Alaska

The Taiya River flows through the Chilkoot Trail Unit of Klondike Gold Rush National Historical Park in southeast Alaska, which was founded to preserve cultural and historical resources and further understanding of natural processes active in the surrounding coastal-to-subarctic basin. Riverine processes exert an important influence on ecologically important boreal toad ( Anaxryus boreas boreas ), salmon [chum salmon ( Oncorhynchus keta ), pink salmon ( O. gorbushca ), and coho salmon ( O. kisutch )], and eulachon ( Thaleichthys pacificus ) habitats, erosion of the historic ghost town of Dyea and other cultural and historical artifacts, and recreational opportunities in the lower 7.5 kilometers (km) of the Taiya River valley bottom. Recurrent consideration of hydroelectric development in West Creek upstream of the park since the 1980s has included proposals for damming and diverting West Creek, which could alter the delivery of water and sediment to this section of the Taiya River. To improve understanding of the hydrologic dependence of park resources for the purposes of guiding effective monitoring and conservation, this study, conducted by the U.S. Geological Survey in cooperation with the National Park Service, used a review of hydrologic data, collection of discrete suspended sediment data, geomorphic mapping, and analysis of historical aerial and ground photographs in a reconnaissance of formative geomorphic processes and hydrologic conditions in the lower 7.5 km of the Taiya River valley bottom. Streamflow and suspended sediment data collected at the U.S. Geological Survey streamgages on the Taiya River and West Creek, combined with historical data, document conditions consistent with streams draining strongly glacierized basins in Alaska. Suspended sediment concentrations from samples collected concurrently over six varying flow levels during 2017–18 ranged from 6 to 284 milligrams per liter (mg/L) for the Taiya River and 13 to 162 mg/L for West Creek, which are similar to or slightly higher than historical values. For the common period of record (1970–77), correlation of daily mean discharge between the two streams was strongest (Pearson’s r = 0.97) during the prolonged May–October high-flow season and weakest ( r = 0.90) during the November–April low-flow season, when West Creek daily mean discharge was proportionally higher. For the Taiya River, streamflow data compared between the available periods of record (1970–77 and 2004–17) showed no decadal-scale patterns in mean annual discharge but did show a shift toward an earlier spring snowmelt pulse. Notable flooding in the Taiya River Basin includes glacial lake outburst floods from the Nourse River valley prior to and during the 1897–98 Gold Rush, a 2002 glacial lake outburst flood from the West Creek valley, and a 1967 rainfall-generated flood. Geomorphic mapping identified four categories of surfaces in the valley bottom—active main stem, abandoned main stem, alluvial fans, and emergent tidal surfaces. Using the maps, main-stem surfaces were subdivided into age categories to identify channel migration patterns from prior to 1940s to 2018. The valley bottom is dominated by active or abandoned channels of the Taiya River except at the extensive low-angle West Creek fan. The active main stem presently supports a mostly single-thread channel with bars and a few sloughs, but the channel actively moved and sometimes was braided within multiple, wider unvegetated corridors in 1894 and earlier. An inventory of 29 off-main-stem channels identified for the study indicates that abandoned main stem channels provide local topographic lows that can intercept groundwater or sustain tributary flow, facilitating the formation of most nonestuarine wetlands in the valley and sustaining important boreal toad breeding habitat. Within the active main stem corridor, the channel has episodically built and reworked meanders and bars, eroding more than one-half of the historic Dyea townsite, in response to glacially controlled delivery of water and sediment, flooding, inputs from West Creek, local features including large woody debris and beaver dams, and rapid uplift from isostatic rebound. West Creek has constructed a large, persistent fan, provoked kilometer-scale Taiya River channel change near the confluence, constructively added to high-season streamflow that affects Taiya River channel migration capacity, disproportionately contributed early-season streamflow, and possibly contributed to groundwater levels in the valley bottom. The progressive narrowing and stability of the main stem corridor, possibly a result of reduction in the magnitude or frequency of glacial lake outburst floods or glacial sediment delivery to streams, indicates less active future reworking of abandoned main-stem surfaces or regeneration of wetland features. The fluvial history of the Taiya River valley bottom collectively indicates continued channel change within a limited corridor, relative stability in wetland locations but uncertainty in stability of groundwater supply to them, and channel incision and extension in response to uplift.

Alaska

Temperature, hydric environment, and prior pathogen exposure alter the experimental severity of chytridiomycosis in boreal toads

Prevalence of the pathogen Batrachochytrium dendrobatidis (Bd), implicated in amphibian population declines worldwide, is associated with habitat moisture and temperature, but few studies have varied these factors and measured the response to infection in amphibian hosts. We evaluated how varying humidity, contact with water, and temperature affected the manifestation of chytridiomycosis in boreal toads Anaxyrus (Bufo) boreas boreas and how prior exposure to Bd affects the likelihood of survival after re-exposure, such as may occur seasonally in long-lived species. Humidity did not affect survival or the degree of Bd infection, but a longer time in contact with water increased the likelihood of mortality. After exposure to ~10 6 Bd zoospores, all toads in continuous contact with water died within 30 d. Moreover, Bd-exposed toads that were disease-free after 64 d under dry conditions, developed lethal chytridiomycosis within 70 d of transfer to wet conditions. Toads in unheated aquaria (mean = 15°C) survived less than 48 d, while those in moderately heated aquaria (mean = 18°C) survived 115 d post-exposure and exhibited behavioral fever, selecting warmer sites across a temperature gradient. We also found benefits of prior Bd infection: previously exposed toads survived 3 times longer than Bd-naïve toads after re-exposure to 10 6 zoospores (89 vs. 30 d), but only when dry microenvironments were available. This study illustrates how the outcome of Bd infection in boreal toads is environmentally dependent: when continuously wet, high reinfection rates may overwhelm defenses, but periodic drying, moderate warming, and previous infection may allow infected toads to extend their survival.

Diseases of Aquatic Organisms

Health evaluation of amphibians in and near Rocky Mountain National Park (Colorado, USA)

We conducted a health survey of amphibians in and adjacent to Rocky Mountain National Park (RMNP) to document current disease presence inside RMNP and identify disease outside RMNP with the potential to spread to the Park's amphibians. Amphibians from five sites within RMNP and seven sites within 60 km of Park boundaries were collected and examined. Necropsies (n - 238), virus isolation, bacterial and fungal cultures, and histological examinations were carried out on amphibian egg masses (outside RMNP/within RMNP: 26/22), larvae (30/42), imagos (recently metamorphosed individuals) (0/3) and adults (61/67) of five species. Marked infections by a pathogenic chytrid fungus (chyridiomycosis), Batrachochytrium dendrobatidis, were detected in three species (Bufo boreas, Pseudacris maculata and Rana sylvatica) from three of five sites within RMNP and in one of three species (P. maculata) from three sites outside RMNP. Of the fully metamorphosed individuals tested (B. boreas, P. maculata and R. sylvatica), chytridiomycosis was found in 60 % (n = 3), 46 % (n = 37) and 54 % (n = 7), respectively. Chytridiomycosis was the principal lethal pathogenic infectious disease detected in three amphibian species within or adjacent to RMNP. Higher fungi were isolated from the cloaca and skin of all five amphibian species. Watermolds (Oomycetes) were isolated from amphibian eggs or skin of all five species. No evidence of Ranavirus was found in cultures and histological examinations of 176 and 142 amphibians, respectively. Fifteen genera of bacteria were identified in larval and just metamorphosed amphibians, and a potentially pathogenic lungworm, Rhabdias sp, was identified in 61.1 % (n = 11) of B. woodhousii outside RMNP, but in only 2 (15.4 %) R. sylvatica within the Park.

Colorado

A framework for guiding management decisions for amphibians in an uncertain future

Managing species in a rapidly changing climate requires knowledge of how species will respond to climate change and other threats while simultaneously developing management actions to reduce threats. Amphibians are one of the most threatened taxa on earth and often serve as the ‘canary in the coalmine’ for the health of ecosystems that countless other species and humans rely on. To understand the status of and guide management for the boreal toad (Anaxyrus boreas boreas), an imperiled amphibian species in the North Central region, we coproduced several products with the Boreal Toad Conservation Team. These products included 1) reconstructed seasonal hydrology patterns for historical boreal toad breeding wetlands and high elevation watersheds in the Southern Rocky Mountain Region (SRMR) from remotely sensed data, 2) current and future predictions of drying rates for historical breeding wetlands, 3) current and future predictions on the status of the boreal toad in the SRMR, and 4) a web tool to guide management actions. While the boreal toad is considered a ‘data rich’ species given data collection efforts that span multiple decades, many amphibian species are considered ‘data poor’, meaning managers lack data on the biology, ecology, or status of the species needed to make sound decisions. To address this knowledge gap, we also quantified drying patterns across watersheds for two ‘data poor’ species in the North Central region at risk from climate change: the Great Basin spadefoot toad ( Spea intermontana ) and the wood frog ( Lithobates sylvaticus ). These new data can guide management decisions for these species by allowing managers to understand habitat changes with respect to water availability, a crucial element for amphibian survival and persistence. Together, these products demonstrate how cutting-edge technology and analytical methods can produce a range of useful information to support amphibian conservation.

Colorado, New Mexico, Wyoming