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    Delayed metamorphosis of amphibian larvae facilitates Batrachochytrium dendrobatidis transmission and persistence

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    Highly virulent pathogens that cause host population declines confront the risk of fade-out, but if pathogen transmission dynamics are age-structured, pathogens can persist. Among other features of amphibian biology, variable larval developmental rates generate age-structured larval populations, which in theory can facilitate pathogen persistence. We investigated this possibility empirically in a population of Salamandra salamandra in Spain affected by Batrachochytrium dendrobatidis (Bd) at breeding sites that lacked alternative amphibian hosts. None of the adults presented infection by Bd. However, for the larvae, while environmental heterogeneity was the most important predictor of infection, the effect on infection dynamics was mediated by transmission from overwintered larvae to new larval recruits, which occurred only in permanent larval habitats. We suggest that interannual Bd maintenance in a host population that experiences mass mortality associated with infection can occur without an environmental reservoir or direct involvement of an alternative host in our study system. However the 2 aquatic habitat types that support intraspecific reservoirs, permanent streams and ponds, are not ideal habitats for long-term Bd maintenance, either due to poor transmission probability or low host survival, respectively. While intraspecific pathogen maintenance due to larval plasticity might be possible at our study sites, this transmission pattern is not without significant risk to the pathogen. The availability of alternative hosts nearby does indicate that permanent Bd fade-out is unlikely.Peer reviewe
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