Relationship between environmental, ecological and genetic drivers of emergence in amphibian chytridiomycosis.

Of the animal groups for which comprehensive assessments have been made, amphibians rank as the most threatened major taxon, with nearly three times as many amphibian species threatened with extinction as bird species. Disease has been identified as one of the major contributors to amphibian declines and extinctions, with one pathogen singled out as the most dangerous amphibian disease identified to date. Batrachochytrium dendrobatidis, a chytridiomycete fungus, has been implicated in mass mortality events, population declines and species extinctions around the globe. The Amphibian Conservation Summit recently organized by the IUCN recognized that the current state of knowledge regarding this fungus is manifestly inadequate. In response to this research gap, the Summit produced a Declaration calling for immediate research to determine the distribution of the disease, how this distribution was achieved and the consequences of invasion of the pathogen for local amphibian communities. Funded by NERC, we have completed three years of monitoring B. dendrobatidis and chytridiomycosis across Europe and have shown for the first time that infection is widespread, is present in the wild in the UK, and is unfortunately present in the Mallorcan midwife toad, one of the most critically endangered species of amphibian. We now need to add to our current descriptions of the prevalence of infection and test hypotheses generated by these patterns in order to ascertain the processes by which B. dendrobatidis causes populations to collapse. We will do this by using a combination of field surveys and laboratory experiments in order to understand how the dynamics of infection in natural populations lead to extinction. Firstly, we will intensively survey five focal study sites where the disease is present, but is causing different effects in the amphibian populations. We will sample these communities over three years in order to track how infection moves through the various species, and how infection differentially affects larval (tadpole), metamorph and adult stages within a community of amphibians. Using our newly developed environmental molecular assay, we are now able to test the density of infectious stages in the environment. This gives us the ability to directly measure the exposure-levels of amphibians and to assess whether the fungus can persist in the environment in the absence of its host. The idea that there are multiple reservoirs of infection is very important, as extinction is more likely in susceptible species when pathogen 'spillover' occurs from disease reservoirs. We will therefore test these ideas in laboratory and mesocosm systems where we are able to manipulate the density and type of potential reservoirs of B. dendrobatidis. Results from our first NERC grant have shown that there is strong evidence that the international trade in amphibians (specifically Xenopus and North American bullfrogs) is causing multiple introductions of B. dendrobatidis into the UK and mainland Europe. We need to know whether there is variation between these different strains of the pathogen in their ability to cause disease, and to test this idea we will perform challenge experiments in our model species, the common toad Bufo bufo. Once these comparisons have been completed, we will develop mathematical epidemiological models with the explicit goal of predicting the dynamics of disease emergence across several scales. We have strong evidence that temperature is a key determinant of chytrid-driven mortality, and we will develop statistical models to determine the potential effects of global-warming on the distribution of the disease within Europe. Our aim is that these short and long term research goals will eventually enable us to control fungal spread or manage amphibian populations in order to prevent the population declines that are associated with the emergence of this pathogen.