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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  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.

In an era of shrinking budgets yet increasing demands for conservation, the value of existing (i.e., historical) data is elevated. Lengthy time-series on common, or previously common, species are particularly valuable and may be available only through the use of historical information. We provide first estimates of the probability of survival and longevity (https://0.67-0.79; 5-7 yr) for a subalpine population of a small-bodied, ostensibly common amphibian, the boreal chorus frog, using historical data and contemporary, hypothesis-driven information-theoretic analyses. We also test a priori hypotheses about the effects of color morph (as suggested by early reports) and of drought (as suggested by recent climate predictions on survival). Using robust mark-recapture models, we find some support for early hypotheses regarding the effect of color on survival, but we find no effect of drought. The congruence between early findings and our analyses highlights the usefulness of historical information by providing raw data for contemporary analyses and context for conservation and management decisions.

Despite prevalent awareness of global amphibian declines, there is still little information on trends for many widespread species. To inform land managers of trends on protected landscapes and identify potential conservation strategies, we collected occurrence data for five wetland-breeding amphibian species in four national parks in the U.S. Rocky Mountains during 2002–2011. We used explicit dynamics models to estimate variation in annual occupancy, extinction, and colonization of wetlands according to summer drought and several biophysical characteristics (e.g., wetland size, elevation), including the influence of North American beaver (Castor canadensis). We found more declines in occupancy than increases, especially in Yellowstone and Grand Teton national parks (NP), where three of four species declined since 2002. However, most species in Rocky Mountain NP were too rare to include in our analysis, which likely reflects significant historical declines. Although beaver were uncommon, their creation or modification of wetlands was associated with higher colonization rates for 4 of 5 amphibian species, producing a 34% increase in occupancy in beaver-influenced wetlands compared to wetlands without beaver influence. Also, colonization rates and occupancy of boreal toads (Anxyrus boreas) and Columbia spotted frogs (Rana luteiventris) were ≥2 times higher in beaver-influenced wetlands. These strong relationships suggest management for beaver that fosters amphibian recovery could counter declines in some areas. Our data reinforce reports of widespread declines of formerly and currently common species, even in areas assumed to be protected from most forms of human disturbance, and demonstrate the close ecological association between beaver and wetland-dependent species.

Environmental stochasticity can have profound effects on the dynamics and viability of wild populations, and habitat heterogeneity provides one mechanism by which populations may be buffered against the negative effects of environmental fluctuations. Heterogeneity in breeding pond hydroperiod across the landscape may allow amphibian populations to persist despite variable interannual precipitation. We examined recruitment dynamics over 10 yr in a high elevation Columbia spotted frog (Rana luteiventris) population that breeds in ponds with a variety of hydroperiods. We combined these data with matrix population models to quantify the consequences of heterogeneity in pond hydroperiod on net recruitment (i.e. number of metamorphs produced) and population growth rates. We compared our heterogeneous system to hypothetical homogeneous environments with (1) only ephemeral ponds, (2) only semi-permanent ponds, and (3) only permanent ponds. We also examined the effect of breeding pond habitat loss on population growth rates. Most eggs were laid in permanent ponds each year, but survival to metamorphosis was highest in the semi-permanent ponds. Recruitment success varied by both year and pond type. Net recruitment and stochastic population growth rate were highest under a scenario with homogeneous semi-permanent ponds, but variability in recruitment was lowest in the scenario with the observed heterogeneity in hydroperiods. Loss of pond habitat decreased population growth rate, with greater decreases associated with loss of permanent and semi-permanent habitat. The presence of a diversity of pond hydroperiods on the landscape will influence population dynamics, including reducing variability in recruitment in an uncertain climatic future.

1. Negative effects of global warming are predicted to be most severe for species that occupy a narrow range of temperatures, have limited dispersal abilities, or have long generation times. These are characteristics typical of many species that occupy small, cold streams.
2. Habitat use, vulnerabilities and mechanisms for coping with local conditions can differ among populations and ontogentically within populations, potentially affecting species-level responses to climate change. However, we still have little knowledge of mean thermal performance for many vertebrates, let alone variation in performance among populations. Assessment of these sources of variation in thermal performance is critical for projecting the effects of climate change on species and for identifying management strategies to ameliorate its effects.
3. To gauge how populations of the Rocky Mountain tailed frog (Ascaphus montanus) might respond to long-term effects of climate change, we measured the ability of tadpoles from six populations in Glacier National Park (Montana, USA) to acclimate to a range of temperatures. We compared survival among populations according to tadpole age (age 1-yr or 2-yr) and according to the mean and variance of late-summer temperatures in natal streams.
4. The ability of tadpoles to acclimate to warm temperatures increased with age and with variance in late-summer temperature of natal streams. Moreover, performance differed among populations from the same catchment.
5. Our experiments with a cold-water species show that population-level performance varies across small geographic scales and is linked to local environmental heterogeneity. This variation could influence the rate and mode of species-level responses to climate change, both by facilitating local persistence in the face of changes in thermal conditions, and by providing thermally-tolerant colonists to neighbouring populations.

Though a third of amphibian species worldwide are thought to be imperiled, existing assessments simply categorize extinction risk, providing little information on the rate of population losses. We conducted the first analysis of the rate of change in the probability that amphibians occupy ponds and other comparable habitat features across the United States. We found that overall occupancy by amphibians declined 3.7% annually from 2002 to 2011. Species that are Red-listed by the International Union for Conservation of Nature (IUCN) declined an average of 11.6% annually. All subsets of data examined had a declining trend including species in the IUCN Least Concern category. This analysis suggests that amphibian declines may be more widespread and severe than previously realized.

Despite the high profile of amphibian declines and the increasing threat of drought and frag- mentation to aquatic ecosystems, few studies have examined long-term rates of change for a single species across a large geographic area. We analyzed growth in annual egg-mass counts of the Columbia spotted frog (<i>Rana luteiventris</i>) across the northwestern United States, an area encompassing 3 genetic clades. On the basis of data collected by multiple partners from 98 water bodies between 1991 and 2011, we used state-space and linear-regression models to measure effects of patch characteristics, frequency of summer drought, and wetland restoration on population growth. Abundance increased in the 2 clades with greatest decline history, but declined where populations are consideredmost secure. Population growthwas negatively associated with temporary hydroperiods and landscape modification (measured by the human footprint index), but was similar in modified and natural water bodies. The effect of drought was mediated by the size of the water body: populations in large water bodies maintained positive growth despite drought, whereas drought magnified declines in small water bodies. Rapid growth in restored wetlands in areas of historical population declines provided strong evidence of successful management. Our results highlight the importance of maintaining large areas of habitat and underscore the greater vulnerability of small areas of habitat to environmental stochasticity. Similar long-term growth rates inmodified and natural water bodies and rapid, positive responses to restoration suggest pond construction and other forms of management can effectively increase population growth. These tools are likely to become increasingly important to mitigate effects of increased drought expected from global climate change.
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More than 25 years ago, scientists began to identify unexplained declines in amphibian populations around the world. Much has been learned since then, but amphibian declines have not abated and the interactions among the various threats to amphibians are not clear. Amphibian decline is a problem of local, national, and international scope that can affect ecosystem function, biodiversity, and commerce. This fact sheet provides a snapshot of the state of the amphibians and introduces examples to illustrate the range of issues in the United States.

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.

Projected increases in wildfire and other climate-driven disturbances will affect
populations and communities worldwide, including host–parasite relationships. Research in
temperate forests has shown that wildfire can negatively affect amphibians, but this research
has occurred primarily outside of managed landscapes where interactions with human
disturbances could result in additive or synergistic effects. Furthermore, parasites represent a
large component of biodiversity and can affect host fitness and population dynamics, yet they
are rarely included in studies of how vertebrate hosts respond to disturbance. To determine
how wildfire affects amphibians and their parasites, and whether effects differ between
protected and managed landscapes, we compared abundance of two amphibians and two
nematodes relative to wildfire extent and severity around wetlands in neighboring protected
and managed forests (Montana, USA). Population sizes of adult, male long-toed salamanders
(Ambystoma macrodactylum) decreased with increased burn severity, with stronger negative
effects on isolated populations and in managed forests. In contrast, breeding population sizes
of Columbia spotted frogs (Rana luteiventris) increased with burn extent in both protected and
managed protected forests. Path analysis showed that the effects of wildfire on the two species
of nematodes were consistent with differences in their life history and transmission strategies
and the responses of their hosts. Burn severity indirectly reduced abundance of soil-transmitted
Cosmocercoides variabilis through reductions in salamander abundance. Burn severity also
directly reduced C. variabilis abundance, possibly though changes in soil conditions. For the
aquatically transmitted nematode Gyrinicola batrachiensis, the positive effect of burn extent on
density of Columbia spotted frog larvae indirectly increased parasite abundance. Our results
show that effects of wildfire on amphibians depend upon burn extent and severity, isolation,
and prior land use. Through subsequent effects on the parasites, our results also reveal how
changes in disturbance regimes can affect communities across trophic levels.

Climate change is expected to increase the frequency and severity of drought and wildfire. Aquatic and moisture-sensitive species, such as amphibians, may be particularly vulnerable to these modified disturbance regimes because large wildfires often occur during extended droughts and thus may compound environmental threats. However, understanding of the effects of wildfires on amphibians in forests with long fire-return intervals is limited. Numerous stand-replacing wildfires have occurred since 1988 in Glacier National Park (Montana, U.S.A.), where we have conducted long-term monitoring of amphibians. We measured responses of 3 amphibian species to fires of different sizes, severity, and age in a small geographic area with uniform management. We used data from wetlands associated with 6 wildfires that burned between 1988 and 2003 to evaluate whether burn extent and severity and interactions between wildfire and wetland isolation affected the distribution of breeding populations. We measured responses with models that accounted for imperfect detection to estimate occupancy during prefire (0–4 years) and different postfire recovery periods. For the long-toed salamander (Ambystoma macrodactylum) and Columbia spotted frog (Rana luteiventris), occupancy was not affected for 6 years after wildfire. But 7–21 years after wildfire, occupancy for both species decreased &#8805;25% in areas where >50% of the forest within 500 m of wetlands burned. In contrast, occupancy of the boreal toad (Anaxyrus boreas) tripled in the 3 years after low-elevation forests burned. This increase in occupancy was followed by a gradual decline. Our results show that accounting for magnitude of change and time lags is critical to understanding population dynamics of amphibians after large disturbances. Our results also inform understanding of the potential threat of increases in wildfire frequency or severity to amphibians in the region.

Monitoring of natural resources is crucial to ecosystem conservation, and yet it can pose many challenges. We developed an amphibian monitoring program as a means for examining ecological conditions in Yellowstone and Grand Teton National Parks. Annual surveys for amphibian breeding occupancy were conducted over a 4-yr period (2006 – 2009) at two scales: catchments (portions of watersheds) and individual wetland sites. Catchments were selected in a stratified random sample with habitat quality and ease of access serving as strata. All known wetland sites with suitable habitat were surveyed within selected catchments. Changes in breeding occurrence of tiger salamanders, boreal chorus frogs and Columbia-spotted frogs were assessed using multi-season occupancy estimation. Numerous a priori models were considered within an information theoretic framework including those with catchment and site-level covariates. Habitat quality was the most important predictor of occupancy. Boreal chorus frogs demonstrated the greatest change (increase) in breeding occupancy at the catchment level. Tiger salamander breeding occurrence increased slightly and Columbia-spotted frogs decreased slightly over the 4-yr period. Larger changes for all 3 species were detected at the finer site-level scale. Use of covariates (e.g., connectivity of sites) offered improvements over simpler models, and may improve understanding of the dynamic processes occurring among wetlands within this ecosystem. Our results suggest monitoring occupancy at two spatial scales within large study areas such as ours is feasible and informative.

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 106 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.

1. Altered global climates in the 21st century pose serious threats for biological systems and practical actions are needed to mount a response for species at risk.<br />
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2. We identify management actions from across the world and from diverse disciplines that are applicable to minimizing loss of amphibian biodiversity under climate change. Actions were grouped under three thematic areas of intervention: (i) installation of microclimate and microhabitat refuges; (ii) enhancement and restoration of breeding sites; and (iii) manipulation of hydroperiod or water levels at breeding sites.<br />
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3. Synthesis and applications. There are currently few meaningful management actions that will tangibly impact the pervasive threat of climate change on amphibians. A host of potentially useful but poorly tested actions could be incorporated into local or regional management plans, programmes and activities for amphibians. Examples include: installation of irrigation sprayers to manipulate water potentials at breeding sites; retention or supplementation of natural and artificial shelters (e.g. logs, cover boards) to reduce desiccation and thermal stress; manipulation of canopy cover over ponds to reduce water temperature; and, creation of hydrologoically diverse wetland habitats capable of supporting larval development under variable rainfall regimes. We encourage researchers and managers to design, test and scale up new initiatives to respond to this emerging crisis.

A dedicated session highlighting studies conducted by the USGS Amphibian Research and Monitoring Initiative (ARMI) in the West was held at the joint annual meeting of the Society for Northwestern Vertebrate Biology and Washington Chapter of The Wildlife Society at Gig Harbor, Washington on March 25th. The session, organized by Steve Corn, Research Zoologist at the Northern Rocky Mountain Science Center in Missoula, Montana, included presentations by ARMI scientists from five different science centers and addressed the main themes of ARMI since its inception in 2000: monitoring of status and trends of amphibian populations, research into causes of amphibian declines, and development of new sampling methods.

Presenters, followed by the title of their talks, included: Gary Fellers, Western Ecological Research Center, Point Reyes National Seashore, CA (Population size, survival, longevity, and movements of Rana draytonii, and R. sierrae at two closely monitored sites: tracking population trends); Mike Adams, Forest and Rangeland Ecosystem Science Center, Corvallis, OR (Does the probability of local extinction for northern red-legged frogs relate to introduced fish or bullfrogs?); Nate Chelgren, Forest and Rangeland Ecosystem Science Center, Corvallis, OR (Spatial and temporal variation in the demography of coastal tailed frogs (Ascaphus truei): isolating aquatic from terrestrial stage dynamics); Erin Muths, Fort Collins Science Center, Fort Collins, CO (Compensatory effects of recruitment and survival on population persistence); Tara Chestnut, Oregon Water Science Center, Portland, OR, (The ecology of the amphibian chytrid fungus, Batrachochytrium dendrobatidis in the aquatic environment); Blake Hossack, Northern Rocky Mountain Science Center, Missoula, MT (Wildfire and fragmentation: effects on amphibian populations and associated nematodes); David Pilliod, Forest and Rangeland Ecosystem Science Center, Boise, ID (Introducing an automated pattern recognition program for leopard frogs); and Steve Corn, Northern Rocky Mountain Science Center, Missoula, MT (How well do call indices represent abundance of breeding anurans?).

Call surveys used to monitor breeding choruses of anuran amphibians generate index values that are frequently used to represent the number of male frogs present, but few studies have quantified this relationship. We compared abundance of male Boreal Chorus Frogs (Pseudacris maculata), estimated using capture-recapture methods in two populations in Colorado, to call index values derived from automated recordings. Single index values, such as might result from large monitoring efforts, were unrelated to population size. A synthetic call saturation index (CSI), the daily proportion of the maximum possible sum of index values derived from multiple recordings, was greater in larger populations, but the relationship was not highly predictive.

The US Geological Survey’s Amphibian Research and Monitoring Initiative (ARMI) is a national project with goals to monitor the status and trends of amphibians, conduct research on causes of declines, and provide information and support to management agencies for conservation of amphibian populations. ARMI activities are organized around extensive inventories and place-based monitoring (such as collaboration with the Greater Yellowstone I&M Network), and intensive population studies and research at selected locations (apex sites). One such is an oxbow pond on the Buffalo Fork near the Black Rock Ranger Station east of Grand Teton National Park. We have been conducting capture-mark-recapture of boreal toads (Anaxyrus boreas) at Black Rock since 2002. In concert with studies of other toad populations in the Rocky Mountains, we have documented a high rate of incidence of the chytrid fungus Batrachochytrium dendrobatidis (Bd) and a negative rate of growth of the toad population, but not the population crash or extinction observed in other populations with high occurrence of Bd.
Long-term observations at other ARMI apex sites have proven invaluable for studying effects of climate change on amphibian behavior, and the Black Rock site has been upgraded with on-site recording of temperature and precipitation, and auditory monitoring of other amphibian species. Continued research at Black Rock will be critical for understanding the interrelated effects of climate and disease on amphibians in the Greater Yellowstone Ecosystem.

Two populations of boreal toads (Bufo boreas) experienced drastic declines in abundance in the late-1990s. Evidence supported the hypothesis of disease (the chytrid fungus, Batrachochytrium dendrobatidis) as the cause of these declines, but other hypotheses had not been evaluated. We used an 11- year capture-recapture data set to evaluate weather and disease as causes of these declines. We developed sets of mathematical models that reflected hypothesized relationships between several weather variables and annual survival rates of adult males in these populations. In addition, models that reflected the possibility that the declines were caused by an introduced fungus were developed. All models were fit to the data and evaluated using a model selection criterion (QAICc). Our analysis provided strong support for the hypothesis of an introduced fungus and little support for the hypothesis that weather conditions caused the declines. Our results also suggest a strong, negative ‘marking effect’ on survival rates of boreal toads. Model-averaged estimates of survival rate are presented.

Chytridiomycosis is an amphibian disease linked with population declines worldwide, yet there is little information about its effects on the demography of infected populations. We used capture-recapture data from two infected populations and one uninfected population of boreal toads (Bufo boreas) in the Rocky Mountains, U.S.A. to examine a priori hypotheses about the effect of the amphibian chytrid fungus (Batrachochytrium dendrobatidis, Bd) on survival probability (&amp;#934;) and population growth rate (&amp;#61548;). Bd-positive toads had lower average annual &amp;#934; than Bd-negative individuals at sites where Bd was detected suggesting that chytridiomycosis may reduce survival by 31-42% in wild boreal toads. Bd-negative toads at infected sites had comparable survival probabilities as Bd-negative toads at the uninfected site, further supporting observed differences. Model results indicated weak evidence that environmental covariates (particularly cold temperatures during the breeding season) influenced toad survival. The diseased populations were declining by 5 - 7% per year over the 6 years of this study whereas the Bd-free population had comparatively stable (&amp;#61548; &amp;#8776; 1) population growth. Our data suggest that Bd is a low-level chronic mortality factor in these toad populations rather than an acute factor causing rapid population declines. These results show how some amphibian populations may be coexisting with the amphibian chytrid fungus and highlight the importance of quantitative assessments of survival in wildlife populations challenged with disease.