The impacts of pathogen emergence in naïve hosts can be catastrophic, and pathogen spread now ranks as a major threat to biodiversity. However, pathogen impacts can persist for decades after epidemics and produce variable host outcomes. Chytridiomycosis in amphibians (caused by the fungal pathogen Batrachochytrium dendrobatidis, Bd) is an exemplar, with impacts ranging from rapid population crashes and extinctions, to population declines and subsequent recoveries. Here, we investigate long-term impacts associated with chytridiomycosis in Australia. We conducted a continent-wide assessment of the disease, reviewing data collected since the arrival of Bd in about 1978, to assess and characterize mechanisms driving past, present and future impacts. We found chytridiomycosis to be implicated in the extinction or decline of 43 of Australia's 238 amphibian species. Population trajectories of declined species are highly variable; six species are experiencing ongoing declines, eight species are apparently stable and 11 species are recovering. Our results highlight that while some species are expanding, Bd continues to threaten species long after its emergence. Australian case-studies and synthesis of the global chytridiomycosis literature suggests that amphibian reservoir hosts are associated with continued declines in endemically infected populations, while population stability is promoted by environmental conditions that restrict Bd impact, and maintenance of high recruitment capacity that can offset mortality. Host genetic adaptation or decreased pathogen virulence may facilitate species recovery, but neither has been empirically demonstrated. Understanding processes that influence Bd-host dynamics and population persistence is crucial for assessing species extinction risk and identifying strategies to conserve disease-threatened species.

for providing expert advice during the post-workshop review stage. We thank the experts involved in the New Guinea assessment workshop, for their information on species with distributions spanning Torres

Metapopulation persistence in fragmented landscapes depends on habitat patches that can support resilient local populations and sufficient connectivity between patches. Yet epidemiological theory for metapopulations has largely overlooked the capacity of particular patches to act as refuges from disease, and has suggested that connectivity can undermine persistence. Here, we show that relatively warm and saline wetlands are environmental refuges from chytridiomycosis for an endangered Australian frog, and act jointly with connectivity to sustain frog metapopulations. We coupled models of microclimate and infection probability to map chytrid prevalence, and demonstrate a strong negative relationship between chytrid prevalence and the persistence of frog populations. Simulations confirm that frog metapopulations are likely to go extinct when they lack environmental refuges from disease and lose connectivity between patches. This study demonstrates that environmental heterogeneity can mediate host-pathogen interactions in fragmented landscapes, and provides evidence that connectivity principally supports host metapopulations afflicted by facultative pathogens.

Identifying determinants of the probability and intensity of infections is important for understanding the epidemiology of wildlife diseases, and for managing their impact on threatened species. Chytridiomycosis, caused by the fungal pathogen Batrachochytrium dendrobatidis, has decimated populations of some amphibians. However, recent studies have identified important environmental constraints on the disease, related to the pathogen's physiological tolerances. In this study, we identified several intrinsic and extrinsic determinants of the probability and intensity of chytrid infections for the threatened growling grass frog (Litoria raniformis) in southeastern Australia, and used mark-recapture to estimate the effect of chytrid infections on the probability of survival of these frogs. Water temperature and salinity had negative effects on both the probability and intensity of chytrid infections. We coupled models of the infection process with a model of the effect of chytrid infections on the probability of survival to assess variation in the impact of chytridiomycosis between wetlands with differing temperature and salinity profiles. Our results suggest that warm, saline wetlands may be refuges from chytridiomycosis for L. raniformis, and should be priorities for protection. Our results also suggest that management actions that increase water temperature (e.g., reducing canopy shading) and salinity (e.g., complementing inflows with groundwater) could be trialed to reduce the impacts of chytridiomycosis on this species. This and other recent studies highlight the value of research on environmental risk factors for chytridiomycosis.

Amphibian chytridiomycosis, caused by infection with Batrachochytrium dendrobatidis (Bd)